Journal of Fish and Wildife Management Vol 1 #2

Volume 1, Number 2, November 2010 ISSN 1944-687X
JOURNAL OF FISH
AND WILDLIFE
MANAGEMENT
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snakes Nerodia sipedon insularum, western pearlshell data archiving policies).
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woodpeckers, and turtles. This issue also contains two Reviewers and Editors Wanted
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Some of these services should be active before this References
editorial is published, but the registration process is
extremely time consuming and rigorous, requiring up to Pullin AS, Salafsky N. 2010. Save the whales? Save the
three published issues and many months to complete for rainforest? Save the data! Conservation Biology 24:
some service providers. Rest assured that all of the past 915–917.
and future content from both journals will be incorpo- Whitlock MC, McPeek MA, Rausher MD, Rieseberg L,
rated into these services as soon as possible and new Moore AJ. 2010. Data archiving. The American
content will be uploaded almost immediately upon Naturalist 175:145–146.
Journal of Fish and Wildlife Management | www.fwspubs.org November 2010 | Volume 1 | Issue 2 | 62
J.K. Wenburg
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Articles
Initial Effects of Prescribed Burning and Understory
Fertilization on Browse Production in Closed-Canopy
Hardwood Stands
Christopher E. Shaw,* Craig A. Harper, Michael W. Black, Allan E. Houston
C.E. Shaw, C.A. Harper
Department of Forestry, Wildlife and Fisheries, University of Tennessee, Knoxville, Tennessee 37996
M.W. Black
Aerospace Testing Alliance Conservation, Arnold Air Force Base, Tennessee 37389
Present address: Northern Bobwhite Conservation Initiative, University of Tennessee Institute of Agriculture, Knoxville,
Tennessee 37996
A.E. Houston
Ames Plantation, Grand Junction, Tennessee 38039
Abstract
Forage production for white-tailed deer Odocoileus virginianus is often limited in closed-canopy forests. We measured
browse production and nutritional carrying capacity after prescribed burning and understory fertilization in closed-
canopy hardwood stands one growing season after treatment in two physiographic regions of Tennessee. Nutritional
carrying capacity estimates for prescribed burning, fertilization, and prescribed burning with fertilization were greater
than in controls on the Cumberland Plateau. However, the cost per pound of forage produced after fertilization
exceeded US$26. In the Coastal Plain, fertilization did not affect nutritional carrying capacity, and prescribed burning
and prescribed burning with fertilization lowered nutritional carrying capacity from controls. At both sites, prescribed
fire had minimal effect on soil pH or soil phosphate and potash levels. Our results suggest prescribed fire and
fertilization are of limited utility for increasing browse production in closed-canopy hardwood forests.
Keywords: fertilizer; fire; nutrient; Odocoileus virginianus; white-tailed deer

Received: October 18, 2009; Accepted: July 12, 2010; Published Online Early: August 2010; Published: November 2010
Citation: Shaw CE, Harper CA, Black MW, Houston AE. 2010. Initial effects of prescribed burning and understory
fertilization on browse production in closed-canopy hardwood stands. Journal of Fish and Wildlife Management 1(2):
64–72; e1944-687X. doi: 10.3996/102009-JFWM-016
Copyright: All material appearing in the Journal of Fish and Wildlife Management is in the public domain and may be
reproduced or copied without permission. Citation of the source, as given above, is requested.
The findings and conclusions in this article are those of the author(s) and do not necessarily represent the views of the
U.S. Fish and Wildlife Service.
* Corresponding author: cshaw2@utk.edu
Introduction Regeneration methods, such as clearcutting, and

timber stand improvement practices can improve forage
An increasing number of nonindustrial private land- availability for deer (Blymyer and Mosby 1977; Miller et al.
owners in the eastern United States actively manage 2009). However, many landowners are not interested in
their property for wildlife (Measells et al. 2005, 2006). The harvesting their timber or removing any trees. Prescribed
majority of these landowners manage for white-tailed fire also has been used to enhance forage availability for
deer Odocoileus virginianus (hereafter deer), and the deer in forested areas (Dills 1970); however, most work
most popular land management practice is planting food concerning use of prescribed fire for increased deer
plots (Schweiss and Dwyer 2008). Acreage dedicated to browse has followed some level of canopy removal to
food plots, however, is a small fraction of the property, increase available sunlight (Masters et al. 1993; Jackson
and practices to improve forested areas could increase et al. 2007). Fertilization has been shown to affect
nutritional carrying capacity (NCC). production (Segelquist and Rogers 1975; Dyess et al.
Burning and Fertilization in Closed-Canopy Hardwood Stands C.E. Shaw et al.
Figure 1. Location of two study areas in Tennessee: Rocky River Hunting Club is located within the Cumberland Plateau
physiographic province and Ames Plantation is in the Coastal Plain physiographic province.
1994) and nutritional quality (Wood 1986; Harlow et al. Ames Plantation
1993) of deer forages, but data evaluating effects of We selected an oak–hickory stand (12.80 acre) within
fertilization in closed-canopy hardwood stands are the Coastal Plain physiographic province at Ames
lacking. Plantation in Fayette County (Figure 1). White oak,
We are not aware of any published evaluations of the yellow-poplar Liriodendron tulipifera, southern red oak
efficacy of prescribed fire and fertilization in closed- Quercus falcata, blackgum, and sweetgum Liquidambar
canopy hardwood forests. Evaluation of practices that do styraciflua were common in the overstory. Midstory
not alter the forest overstory is warranted because many species included winged elm Ulmus alata, black cherry
landowners are interested in improving forage availabil- Prunus serotina, and flowering dogwood. Poison ivy
ity for deer without altering the forest canopy. We Toxicodendron radicans, Japanese honeysuckle Lonicera
conducted this field study to evaluate the effects of japonica, Virginia creeper Parthenocissus quinquefolia,
prescribed fire, understory fertilization, and prescribed and supplejack Berchemia scandens were common in the
fire with understory fertilization on nutrient availability understory. Soils were primarily Ruston sandy loam that
and browse production in closed-canopy hardwood are well drained and acidic, with 12–30% slopes (Flowers
stands in two distinct physiographic regions of Tennes- 1964). Site index for shortleaf pine was 50–70 (Flowers
see. Our objectives were to determine deer use of 1964). Deer density estimates obtained with infrared-
browse species and production and nutritional quality of triggered cameras (Jacobson et al. 1997) indicated a
browse after treatments. minimum density of 21 deer/mi2.
Study Areas Methods

We selected two closed-canopy hardwood stands with Sampling methodology and treatment application
no recent fire histories. Stands were chosen to ensure We systematically located sixteen 100-yd (91.44-m)
uniformity (e.g., similar species composition, soils) within transects 100 ft (30.48 m) apart within each 12.80-acre
a site. (5.18-ha) stand during summer 2004. We measured
woody leaf biomass (pounds per acre) and herbaceous
Rocky River forage within sixty-four 60-ft2 (5.57-m2) sampling plots
Within the Cumberland Plateau physiographic prov- systematically placed every 25 yd along each transect
ince, we selected a shortleaf pine–oak Pinus echinata– (Figure 2). We tallied woody browse plants within
Quercus spp. stand (12.80 acre [5.18 ha]) known as Rocky sample plots to species (stem count tally), and stems
River, in Sequatchie County (Figure 1). Overstory species were noted as browsed or unbrowsed. We also noted
included scarlet oak Quercus coccinea, white oak Quercus browsing on herbaceous plants along the line transect.
alba, shortleaf pine, black oak Quercus velutina, and For woody vines, we used a measure of inches covered
mockernut hickory Carya tomentosa. Midstory species along the line transect in a regression equation to
included mockernut hickory, sassafras Sassafras albidum, estimate total stem counts of these species from their
sourwood Oxydendrum arboreum, blackgum Nyssa sylva- coverage. We collected and sorted leaves of woody
tica, red maple Acer rubrum, pignut hickory Carya glabra, vegetation and all above-ground growth of herbaceous
and flowering dogwood Cornus florida. Soils were plants #4 ft. We placed samples in a forced-air oven at
primarily Lonewood silt loam and Lily loam that are well 50uC until cessation of weight loss and then weighed
drained and acidic, with 2–12% slopes (Prater 2003). Site samples to determine dry-matter weights (grams).
index for shortleaf pine was 70 (Prater 2003). Deer After pretreatment data collection, we divided stands
density estimates obtained with infrared-triggered cam- into four 3.2-acre (1.3-ha) sections, each containing four
eras (Jacobson et al. 1997) indicated a minimum of 28 of the established transects (Figure 2). We collected soil
deer/mi2 (deer/2.59 km2). samples along the four transects within each section;
phate/acre and 131 lb potash/acre. We collected soil

samples in June and August 2005 to track responses in
pH, phosphate, and potash levels posttreatment.
During July and August 2005, we located plots
between areas sampled in 2004 to avoid previously
sampled areas. Sample plots in summer 2005 were 4 ft in
width 6 10 ft in length. We recorded evidence of
browsing on woody plants in sample plots by using a
stem tally. We collected all woody leaves and herbaceous
plants #4 ft and sorted them by species or species
groups (i.e., hickory, red oak, or white oak group). We
placed samples in a forced-air oven at 50uC until
cessation of weight loss and weighed to determine
dry-matter weights. We combined samples of species or
species groups within the same treatment into a
composite sample and ground with a Wiley mill until
particles passed through a 1-mm screen. We analyzed
composite samples for nitrogen (N) with a LECO FP-2000
nitrogen analyzer (LECO Corp., St. Joseph, MI) by using
the Dumas combustion method (method 990.03; AOAC
1998) to obtain estimates of crude protein (CP) for
species or species groups. We conducted fiber analyses
(neutral and acid detergent; Jung 1997) with an ANKOM
200 fiber analyzer (ANKOM Technology, Macedon, NY).
Figure 2. Schematic illustrating sampling and treatment
application (understory fertilization, prescribed burning, and Statistical analyses
prescribed burning with understory fertilization) in each of two We collected browse and herbaceous forage in both
closed-canopy hardwood stands (Rocky River Hunting Club and
Ames Plantation) in Tennessee.
years to compare production in control and treatment
areas within the closed-canopy hardwood stands. There-
fore, we used a completely randomized split-plot design
combined them to form a composite sample; and for a mixed model analysis of variance. Fixed effects were
submitted them to the University of Tennessee Soil, treatment, year, and the treatment 6 year interaction.
Plant and Pest Laboratory for analysis of pH, phosphorus Random effects were transect (treatment) and sample plot
(P), and potassium (K) levels. (transect 6 treatment). Log or log + 0.5 transformations
We burned two sections in each stand during the were used when necessary to address normality and
dormant season (Rocky River, March 30, 2005; Ames homogeneity of variance. When the interaction term was
Plantation, April 5, 2005) by using low-intensity fire under significant (P , 0.05), we used the least significant
the following conditions: 6–20uC, 20–40% relative humid- difference method for mean separation. We chose 10
ity, wind speed of 3–6 mi/h (4.83–9.66 km/h), and a mixing browse species or species groups for biomass compari-
height of .1,640 ft. For all controlled burns, backing fires sons after treatments based on deer selectivity (see
were set initially and the remainder of the units were description below) and contribution of each species or
burned using relatively low-intensity strip-heading fires species group to total biomass at each site. We compared
generating 6–18-in. (15.24–45.72 cm) flame heights. individual browse species or species group biomass
We applied fertilizer in late spring 2005 (Rocky River, among treatments by using a completely randomized
May 16, 2005; Ames Plantation, May 12, 2005). To avoid design for the mixed model analysis of variance. Burn and
issues with pseudoreplication, fertilizer was applied to fertilizer treatment were the treatment factors. Before
each individual transect (replicate) instead of across the using the log transformation for the 10 individual species
entire burned section. Before application, we calibrated a or species groups, 0.5 was added to all biomass values to
hand spreader to ensure proper distribution for each retain observations with 0 values. For testing treatment
nutrient according to pretreatment soil test results. effects, we used a Bonferroni-corrected a level of 0.01
We fertilized four transects within one burned and one (0.10/10 species tested). When significant (P , 0.10)
unburned section with ammonium nitrate (34–0–0 [N–P– differences were found, we used the least significant
K]) at 45 lb N/acre. Triple superphosphate (0–46–0) and difference method to detect differences among means.
muriate of potash (0–0–60) were applied to raise Using pretreatment (2004) data, we calculated a
phosphate (P2O5) and potash (K2O) to levels where a selection index (Chesson index; Chesson 1978, 1983) by
plant response would be expected based upon soil test dividing the ratio of use and availability for a given
results. At Rocky River, we applied 72 lb (32.66 kg) species or species group by the sum of ratios for all
phosphate/acre and 205 lb potash/acre. In the burned species or species groups for woody plants and browse
section at Ames Plantation, we applied 52 lb phosphate/ species having stem counts $25 (Supplemental Material,
acre and 101 lb potash/acre. For the fertilized-only Table S1, http://dx.doi.org/10.3996/102009-JFWM-016.S1).
transects at Ames Plantation, we applied 72 lb phos- We combined species or species groups with ,25 stems
Table 1. Soil test results (2-in. [5.1 cm] depth) before and after implementation of understory fertilization (fert), prescribed
burning (burn), and prescribed burning with understory fertilization (burn/fert) in two closed-canopy hardwood stands (Rocky River
Hunting Club [RR] and Ames Plantation [AP]) in Tennessee, September 2004–August 2005.
Treatment
Site Sample date Comparison Control Fert Burn Burn/fert
RR September 23, 2004 pH 4.4 4.6 4.3 4.5
RR June 23, 2005 pH 4.3 4.4 4.4 4.6
RR July 28, 2005 pH 4.4 4.3 4.3 4.4
AP February 12, 2005 pH 6.3 6.4 6.1 6.2
AP June 16, 2005 pH 6.1 6.1 6.0 6.6
AP August 17, 2005 pH 6.2 6.0 6.1 6.1
RR September 23, 2004 Phosphate 4 4 8 4
RR June 23, 2005 Phosphate 4 24 4 24
RR July 28, 2005 Phosphate 4 20 12 24
AP February 12, 2005 Phosphate 4 4 4 12
AP June 16, 2005 Phosphate 4 16 4 24
AP August 17, 2005 Phosphate 8 28 8 28
RR September 23, 2004 Potash 48 71 49 46
RR June 23, 2005 Potash 98 220 78 280
RR July 28, 2005 Potash 51 160 53 170
AP February 12, 2005 Potash 180 140 140 160
AP June 16, 2005 Potash 120 200 100 240
AP August 17, 2005 Potash 200 300 170 300
into an ‘‘other’’ category. We could not calculate a Effects on forage production

selection index for herbaceous forage species. Cutoff Effects of treatments on forage production varied
values indicating no selection depended on the number among study sites. Herbaceous forage increased in all
of species or species group compared at each site (Ames treatments as well as controls at Rocky River from 2004
Plantation, 1/25 = 0.04; Rocky River, 1/11 = 0.09). Values to 2005 (Table 2). At Ames Plantation, herbaceous
above and below these values indicate greater and lesser forage was increased after prescribed fire and prescribed
use, respectively, than expected at a given site. fire with fertilization (Table 2). Browse production at
We calculated estimates of NCC in 2005 with the Rocky River did not increase after fertilization but did
explicit nutritional constraints model (Hobbs and Swift increase after prescribed fire and prescribed fire with
1985). Following criteria used by Edwards et al. (2004), fertilization (Table 2). No treatment increased browse
we estimated NCC for deer by using constraints of 12% production at Ames Plantation (Table 2), and there was
CP and a dry matter intake of 3 lb/d. We determined no meaningful effect on biomass of individual browse
nutritional values for individual browse species for each species or species groups after treatments at either site
species collected within each treatment or control area. (Table 3). Crude protein and fiber content were variable
Because samples of each species within each treatment among species or species groups and treatments
or control area were combined for nutritional analyses, (Table 4).
we report absolute values for CP, neutral detergent fiber,
and acid detergent fiber. We based browse species Deer selection
included in the NCC estimate upon selection indices Greenbrier Smilax spp., blackgum, and blackberry
calculated at each site. We used a completely random- Rubus spp. were used more than expected based on
ized design for a mixed model analysis of variance to availability at Rocky River. Hickory, blueberry Vaccinium
compare NCC estimates among treatments, with an a spp., red maple, sourwood, sassafras, white oak group,
level of 0.05. We log-transformed data when necessary to and red oak group were used less than expected. No use
address normality and variance problems. was recorded for species in the red oak group. At Ames
Plantation, greenbrier, supplejack, blackgum, rose Rosa
Results spp., and winged elm were browsed more than expected
based on availability. Species used less than expected
Effects on soil based on availability included slippery elm Ulmus rubra,
Soil pH remained similar across all treatments and sugar maple Acer saccharum, blackberry, red oak group,
sampling periods at both study sites (Table 1). As black cherry, white oak group, eastern redbud Cercis
expected, soil phosphate and potash levels increased canadensis, red maple, hickory, grape Vitis spp., Japanese
after fertilization treatments, but they were not influ- honeysuckle, ash Fraxinus spp., Virginia creeper, and
enced by prescribed fire. poison ivy. No browsing was recorded for yellow-poplar,
Table 2. Woody leaf biomass and herbaceous forage production (dry matter pounds/acre) before and after implementation of
understory fertilization (fert), prescribed burning (burn), and prescribed burning with understory fertilization (burn/fert) in two
closed-canopy hardwood stands (Rocky River Hunting Club [RR] and Ames Plantation [AP]) in Tennessee, July–August 2004 and
2005. Within each comparison at a site, means with the same letter are not different (P . 0.05).
Treatment
Control Fert Burn Burn/fert
Site Year Comparison x̄ (SE) x̄ (SE) x̄ (SE) x̄ (SE)
a
RR 2004 (pre-) Woody leaf 59.4 (7.1) E 135.8 (18.0) BC 108.8 (11.6) CD 90.7 (16.9) D
RR 2005 (post-) Woody leafa 71.4 (11.7) E 106.4 (11.1) CD 195.7 (23.7) AB 232.4 (21.8) A
RR 2004 (pre-) Herbaceousb 9.5 (2.0) 13.0 (4.6) 8.2 (3.3) 10.4 (2.6)
RR 2005 (post-) Herbaceousb 13.2 (3.0) 23.4 (7.6) 18.0 (6.8) 51.0 (13.3)
AP 2004 (pre-) Woody leafc 162.8 (18.8) 139.4 (12.9) 140.3 (26.4) 169.1 (19.5)
AP 2005 (post-) Woody leafc 188.3 (20.5) 163.6 (22.1) 104.3 (16.1) 204.4 (30.3)
AP 2004 (pre-) Herbaceousd 9.6 (4.3) BC 28.8 (8.7) A 7.3 (3.0) BC 1.3 (0.7) C
d
AP 2005 (post-) Herbaceous 13.4 (7.8) B 21.3 (4.5) A 15.6 (3.0) A 49.1 (11.6) A
a
Treatment effect significant (P = 0.02) for woody leaf production at RR.
b
Treatment effect not significant (P = 0.61) for herbaceous forage production at RR; year effect significant (P , 0.01).
c
Treatment effect not significant (P = 0.21) for woody leaf production at AP.
d
Treatment effect significant (P = 0.01) for herbaceous forage production at AP.
Table 3. Woody leaf biomass production (dry matter pounds/acre) of individual species or species groups after implementation
of understory fertilization (fert), prescribed burning (burn), and prescribed burning with understory fertilization (burn/fert) in two
closed-canopy hardwood stands (Rocky River Hunting Club [RR] and Ames Plantation [AP]) in Tennessee, July–August 2005.
Asterisks indicate significant burning 6 fertilization interaction at P , 0.01. Within the row, means are not different (P . 0.05) if
followed by the same letter. Raw means are reported for red oak group, but analysis was conducted on transformed data.
Treatment
Control Fert Burn Burn/fert Effect
Speciesa x̄ (SE) x̄ (SE) x̄ (SE) x̄ (SE) Burning Fertilization
RR
Blueberry 22.1 (5.4) 35.0 (5.5) 61.9 (13.0) 42.7 (9.9) F1,12 = 0.26, P = 0.62 F1,12 = 0.17, P = 0.69
Sassafras 7.1 (1.6) 19.0 (5.2) 55.1 (11.0) 94.7 (19.5) F1,60 = 27.12, P , 0.01 F1,60 = 1.19, P = 0.28
Sourwood 5.2 (2.7) 4.1 (2.1) 39.1 (20.5) 21.8 (8.5) F1,60 = 2.48, P = 0.12 F1,60 = 0.18, P = 0.67
Greenbrier 6.8 (1.7) 7.3 (1.4) 10.3 (1.5) 7.4 (1.4) F1,60 = 1.60, P = 0.21 F1,60 = 1.21, P = 0.28
Blackberry 1.6 (0.5) 5.5 (1.7) 0.5 (0.0) 3.4 (1.4) F1,12 = 2.98, P = 0.11 F1,12 = 7.42, P = 0.02
Red maple 7.7 (2.1) 10.0 (3.9) 4.4 (2.0) 21.4 (11.3) F1,12 = 4.57, P = 0.05 F1,12 = 0.16, P = 0.70
Blackgum 1.6 (0.4) 2.6 (1.1) 4.4 (1.3) 9.7 (2.7) F1,12 = 8.14, P = 0.02 F1,12 = 2.23, P = 0.16
Red oak group* 4.3 (1.3) AB 11.2 (3.6) A 12.1 (6.0) A 8.8 (7.9) B F1,60 = 2.88, P = 0.10 F1,60 = 0.04, P = 0.85
White oak group 12.8 (7.4) 7.4 (3.0) 7.7 (5.0) 8.0 (2.1) F1,12 = 0.05, P = 0.82 F1,12 = 1.29, P = 0.28
Hickory 3.7 (1.3) 7.7 (2.3) 2.0 (1.2) 9.7 (4.4) F1,60 = 1.55, P = 0.22 F1,60 = 5.88, P = 0.02
AP
Poison ivy* 84.9 (19.0) A 29.7 (7.3) B 23.6 (6.1) B 96.5 (19.0) A F1,12 = 0.03, P = 0.86 F1,12 = 0.37, P = 0.56
Grape 10.2 (4.8) 11.7 (3.8) 16.7 (8.9) 18.3 (7.0) F1,12 = 0.22, P = 0.64 F1,12 = 0.84, P = 0.38
Virginia creeper 9.9 (2.3) 6.9 (2.5) 11.6 (3.6) 16.7 (5.2) F1,60 = 2.53, P = 0.12 F1,60 = 0.08, P = 0.78
Ash 7.2 (2.4) 22.2 (6.4) 11.7 (5.9) 3.2 (1.4) F1,12 = 3.26, P = 0.10 F1,12 = 0.09, P = 0.77
Honeysuckle 6.6 (1.9) 16.8 (4.6) 3.3 (0.9) 5.9 (1.5) F1,12 = 2.95, P = 0.11 F1,12 = 2.86, P = 0.12
Slippery elm 11.3 (7.2) 7.1 (2.4) 7.4 (3.0) 6.2 (2.6) F1,12 = 0.11, P = 0.75 F1,12 = 0.08, P = 0.78
Blackgum 8.1 (3.4) 6.6 (5.1) 2.1 (0.9) 6.2 (3.9) F1,60 = 0.08, P = 0.78 F1,60 = 0.27, P = 0.61
Winged elm 12.0 (4.0) 7.2 (1.9) 3.0 (0.8) 0.6 (0.5) F1,12 = 9.18, P = 0.01 F1,12 = 2.89, P = 0.12
Greenbrier 6.3 (3.7) 6.9 (1.9) 1.2 (0.5) 1.5 (0.7) F1,60 = 8.89, P , 0.01 F1,60 = 1.25, P = 0.27
Supplejack 0.3 (0.3) 4.8 (1.7) 0.3 (0.3) 4.4 (1.2) F1,60 = 0.12, P = 0.73 F1,60 = 19.60, P , 0.01
a
Blueberry Vaccinium spp., sassafras Sassafras albidum, sourwood Oxydendrum arboreum, greenbrier Smilax spp., blackberry Rubus spp., red maple
Acer rubrum, blackgum Nyssa sylvatica, red oak Quercus falcata, white oak Quercus alba, hickory Carya spp., poison ivy Toxicodendron radicans,
grape Vitis spp., Virginia creeper Parthenocissus quinquefolia, ash Fraxinus spp., honeysuckle Lonicera spp., slippery elm Ulmus rubra, winged elm
Ulmus alata, supplejack Berchemia scandens.
Table 4. Nutritional content of species after implementation of understory fertilization (fert), prescribed burning (burn), and
prescribed burning with understory fertilization (burn/fert) in two closed-canopy hardwood stands (Rocky River Hunting Club [RR]
and Ames Plantation [AP]) in Tennessee, July–August 2005. Dashes indicate composite sample of species collected in a given
treatment was not large enough for nutritional analysis. Rose also not included at AP because sample was not large enough for
nutritional analysis.
Crude protein (%) Neutral detergent fiber (%) Acid detergent fiber (%)
a
Species Control Fert Burn Burn/fert Control Fert Burn Burn/fert Control Fert Burn Burn/fert
RR
Blackgum 11.9 13.7 14.3 13.4 40.5 38.3 36.7 38.8 20.4 26.2 16.3 21.9
Red maple 11.3 13.3 15.2 14.7 44.0 41.2 39.2 43.0 28.3 25.4 25.3 31.7
Blackberry 13.5 14.8 — 11.9 43.9 43.0 — 41.6 21.9 24.3 — 23.8
Sassafras 15.2 15.1 14.9 14.0 57.3 53.4 53.5 55.7 42.1 46.1 36.0 40.2
Greenbrier 12.6 14.9 13.3 14.3 47.1 46.8 44.1 46.6 30.7 28.4 31.6 44.0
Sourwood 11.5 12.2 11.7 14.4 39.3 38.3 37.3 39.7 23.8 22.2 21.8 25.1
Blueberry 9.4 10.6 9.7 11.0 48.3 51.0 45.6 50.1 33.2 37.6 30.0 34.4
AP
Ash 12.4 12.8 14.5 15.2 56.6 57.3 58.7 54.5 38.7 38.4 40.3 39.8
Blackgum 10.8 11.6 13.5 13.1 37.9 38.5 37.7 37.0 21.9 21.2 23.4 22.2
Japanese honeysuckle 12.6 11.7 13.3 12.1 38.8 37.0 35.9 35.8 26.7 24.5 26.1 25.9
Poison ivy 12.2 11.8 13.4 12.4 45.1 45.0 48.5 46.4 29.7 30.8 32.8 32.9
Greenbrier 12.6 13.2 12.7 14.4 50.2 48.8 50.4 45.2 30.4 29.9 35.0 34.0
Supplejack — 13.0 — 15.0 — 50.8 — 42.6 — 20.7 — 25.0
Virginia creeper 10.4 10.4 12.6 11.8 47.7 42.9 40.9 40.5 32.9 33.7 30.4 32.7
Grape 12.4 11.4 13.6 13.2 45.4 41.1 41.5 40.3 33.1 33.2 32.5 29.2
Winged elm 12.8 12.7 15.8 17.7 68.2 64.0 66.9 62.0 28.2 31.0 29.8 23.8
a
Blackgum Nyssa sylvatica, red maple Acer rubrum, blackberry Rubus spp., sassafras Sassafras albidum, greenbrier Smilax spp., sourwood
Oxydendrum arboreum, blueberry Vaccinium spp., ash Fraxinus spp., Japanese honeysuckle Lonicera japonica, poison ivy Toxicodendron radicans,
supplejack Berchemia scandens, Virginia creeper Parthenocissus quinquefolia, grape Vitis spp., winged elm Ulmus alata.
sassafras, Carolina buckthorn Rhamnus caroliniana, com- Discussion

mon persimmon Diospyros virginiana, or devil’s walking-
stick Aralia spinosa. Although others have noted changes in pH after
prescribed fire (Binkley 1986; Blankenship and Arthur
Effect on NCC 1999), our results did not reveal an effect of fire on pH,
Although no treatment effects on individual species which was consistent with Franklin et al. (2003).
(Table 3) or nutritional quality (Table 4) were detected, Although using ammonium nitrate fertilizers may lower
prescribed fire and fertilization increased NCC estimates pH if used annually, pH changes after infrequent
at Rocky River (Table 5). Conversely, although no fertilization are usually negligible (Fisher and Binkley
treatment significantly affected forage production at 2000).
Ames Plantation, estimates of deer days/acre were Differences in soil potash responses at Ames Planta-
decreased after prescribed fire. tion and Rocky River may be a result of differences in soil
Table 5. Nutritional carrying capacity (deer days/acre, assuming 3 lb/d consumption) of selected white-tailed deer Odocoileus
virginianus forages combined to average 12% crude protein after implementation of understory fertilization (fert), prescribed
burning (burn), and prescribed burning with understory fertilization (burn/fert) in closed-canopy hardwood stands at Rocky River
Hunting Club (RR) in Sequatchie County, Tennessee, and Ames Plantation (AP) in Fayette County, Tennessee, July–August 2005.
Interaction between burning and fertilization not significant (P . 0.05) at either site.
Treatment
Control Fert Burn Burn/fert Effect
Site Deer days x̄ (SE) x̄ (SE) x̄ (SE) x̄ (SE) Burning Fertilization
RR per acre 2.8 (0.6) 4.6 (0.8) 4.6 (0.6) 6.3 (1.2) F1,60 = 4.44, P = 0.04 F1,60 = 4.70, P = 0.03
AP per acre 6.9 (1.7) 8.5 (2.6) 2.1 (0.5) 4.2 (1.2) F1,60 = 6.11, P = 0.02 F1,60 = 1.96, P = 0.17
texture. The sandy loam at Ames Plantation contained treatments were slightly higher than those in control
less clay than the silt loam at Rocky River. As summer areas, only control areas at Ames Plantation had
progressed, the greater clay content and cation ex- reductions in NCC estimates attributable to the mini-
change capacity of the silt loam probably contributed to mum criteria for CP (12%). Managers should only use
the observed decline in potash ratings during the late results from diet studies as general guidelines for deer
summer sampling period because clay particles attract use of various species and evaluate treatment effects on
more of the free K ions. Plant response to fertilization can browse species in relation to actual deer use on specific
be expected to vary among different soil types. areas.
Past research has documented increases in browse The response of herbaceous species to treatments
production after fertilization, but any increase is certainly suggested their inclusion in NCC estimates would not
buffered by available sunlight. Segelquist and Rogers have affected our results. At both study areas, sites that
(1975) and Dyess et al. (1994) reported increased were burned and fertilized produced greater amounts of
production of Japanese honeysuckle after applications herbaceous forages than other treatments. However,
of lime and N fertilization, but their plots were located in nonpreferred species, such as American burnweed
cleared openings. Production of Japanese honeysuckle Erechtites hieraciifolia and grasses, contributed almost
did not increase after fertilization at Ames Plantation. all of this production. On other sites with a different
Increases in NCC estimates at Rocky River after fertiliza- seedbank, a response by desirable forage species may
tion were significant but arguably not worth the cost. increase NCC.
Fertilizers in our study were US$0.22/lb (34–0–0),
US$0.31/lb (0–46–0), and US$0.28/lb (0–0–60). Average Management Recommendations
fertilization costs for rates of N (US$28.46/acre), P
(US$41.78/acre), and K (US$72.17/acre) applied totaled Prescribed burning and understory fertilization pro-
US$142.40/acre. The only increase in browse selected by duced mixed effects in two physiographic regions with
deer after fertilization was blackberry, which increased different soil types in Tennessee one growing season
4.0 lb/acre at Rocky River, costing US$35.60/lb in after treatment. Therefore, we caution against the use of
fertilized sections. With the greatest increase of 3.5 deer low-intensity prescribed fire in closed-canopy stands
days/acre (6.3 deer days/acre after prescribed burning with the objective of increasing browse for deer.
with understory fertilization compared to 2.8 deer days/ Although browse production may increase during
acre in control) at Rocky River and associated fertilization subsequent growing seasons or after additional fire
rates and costs (US$173.67/acre) used in our study, it prescription, we recommend some canopy reduction
would cost US$49.62 for each additional deer day. treatment (e.g., retention cutting and thinnings) to allow
Substantive changes in the structure and composition additional sunlight into the stand before burning (Healy
of understory vegetation usually necessitate several 1997; Jackson et al. 2007), especially if a relatively quick
successive fires and are also influenced by season of fire and positive treatment effect is desired. We do not
and fire intensity (Brockway and Lewis 1997; Sparks et al. recommend understory fertilization in closed-canopy
1998; Hutchinson and Sutherland 2000; Peterson and hardwood stands because plant response was minimal,
Reich 2001; Glasgow and Matlack 2007; Jackson et al. and the relatively small increase makes it difficult to
2007). Substantive changes after repeated burning are justify the cost of fertilization. Liming before fertilization
strongly correlated with increased sunlight entering the could improve pH and nutrient availability, but applica-
forest canopy. Our data represent the initial effect of tion of lime in forested areas is generally not practical
prescribed fire in closed-canopy stands. With repeated because of difficulty spreading lime in the woods,
low-intensity burning, mortality of the midstory may amount of lime needed to correct soil acidity, and
allow increased sunlight, which could lead to increased associated costs. We believe money spent on liming and
browse production. However, it is likely that landowners fertilization would be much more efficiently and
managing their property for wildlife would like to see a effectively spent on food plot plantings.
more timely response to their management efforts.
It is important to understand the approach we used to Supplemental Material
estimate NCC is not an absolute measure of carrying Please note: The Journal of Fish and Wildlife Management
capacity. However, it does allow relative comparisons is not responsible for the content or functionality of any
among treatments by using a biologically defensible supplemental material. Queries should be directed to the
diet-quality target using species or species groups corresponding author.
selected by deer during the growing season. This
approach is important because forage quality has a Table S1. Selection index value data.
tremendous influence on available nutrition. By combin- Found at DOI: 10.3996/102009-JFWM-016.S1 (182 KB
ing selected deer forages to average a minimum of 12% XLS).
CP, our data suggested NCC was actually negatively
influenced by burning at Ames Plantation, although the Acknowledgments
woody leaf biomass was not significantly decreased.
Although forage quality is important, browse species Funding and support for this study were provided by the
selected by deer influences NCC estimates more than Department of Forestry, Wildlife, and Fisheries at the
increases in CP values. Although CP values in most University of Tennessee; Hobart Ames Foundation;
Sequatchie Forest and Wildlife; Quality Deer Manage- and function. Forest Ecology and Management 184:
ment Association; and Tennessee Wildlife Resources 315–335.
Agency. Logistic support was provided by Benny Glasgow LS, Matlack GR. 2007. Prescribed burning and
Bowers, Carla Dilling, Jesus Gamboa, John Gruchy, Greg understory composition in a temperate deciduous
Julian, James McDonald, Larry Teague, and Shelton forest, Ohio, USA. Forest Ecology and Management
Whittington. We thank the Subject Editor and two 238:54–64.
anonymous reviewers, who helped improve the quality Harlow RF, Pinkerton BW, Guynn Jr DC, Williams Jr JG.
of this manuscript. 1993. Fertilizer effects on the quality of white-tailed
deer forages on utility rights-of-way. Southern Journal
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Articles
Match–Mismatch Regulation for Bluegill and Yellow
Perch Larvae and Their Prey in Sandhill Lakes
Jeffrey C. Jolley,* David W. Willis, Richard S. Holland
J.C. Jolley, D.W. Willis
Department of Wildlife and Fisheries Sciences, South Dakota State University, SNP 138, Box 2140B, Brookings,
South Dakota 57007
Present address of J.C. Jolley: U.S. Fish and Wildlife Service, Columbia River Fisheries Program Office, 1211 SE Cardinal
Court, Vancouver, Washington 98683
R.S. Holland
Nebraska Game and Parks Commission, P.O. Box 30370, Lincoln, Nebraska 68701
Abstract
Food availability may regulate fish recruitment, both directly and indirectly. The availability of zooplankton, especially to
newly hatched larvae, is thought to be crucial to their early growth and survival. We examined stomach contents of larval
bluegill Lepomis macrochirus and yellow perch Perca flavescens in Pelican Lake and Cameron Lake, Nebraska, in 2004 and
2005. We also determined zooplankton availability and calculated prey selection using Chesson’s a. In addition, we
investigated potential match–mismatch regulation of recruitment from 2004 to 2008. Bluegill positively selected
copepod nauplii and Bosmina spp., and yellow perch often selected copepods. Abundant zooplankton populations were
available for consumption. Matches of both larval bluegill and yellow perch abundance to zooplankton abundance were
detected in all years; exact matches were common. Mismatches in predator and prey production were not observed.
Predation by age-0 yellow perch on age-0 bluegill was not observed, even though yellow perch hatched 2 mo prior to
bluegill. Given that zooplankton were abundant and well-timed to larval fish relative abundance over the time span of
this study, the match–mismatch hypothesis alone may not fully account for observed recruitment variability in these
populations. Environmental conditions may also affect recruitment and warrant further investigation.
Keywords: Sandhills lakes; Valentine National Wildlife Refuge; predator–prey dynamics; recruitment; phenology;
survival; food habits
Received: June 30, 2010; Accepted: August 25, 2010; Published Online Early: August 2010; Published: November 2010
Citation: Jolley JC, Willis DW, Holland RS. 2010. Match–mismatch regulation for bluegill and yellow perch larvae and
their prey in Sandhill lakes. Journal of Fish and Wildlife Management 1(2):73–85; e1944-687X. doi: 10.3996/062010-
JFWM-018
* Corresponding author: jeffrey_ jolley@fws.gov
Introduction bottlenecks, of high mortality may exist for some species

(Hjort 1914; May 1974) and researchers commonly
Recruitment dynamics of populations will ultimately incorporate this factor when describing recruitment
structure fish communities (Diana 1995) and several processes (Marr 1956). This critical period is thought to
factors are believed to regulate recruitment, both directly occur early in age-0 yellow perch Perca flavescens and
and indirectly. These include abiotic factors such as bluegill Lepomis macrochirus cohort development (Toetz
physical habitat, temperature, and weather (Beard 1982; 1966; Forney 1971; Clady 1976; Anderson et al. 1998).
Jackson and Noble 2000), biotic factors such as food Large annual variation in yellow perch year-class
availability and competition (Prout et al. 1990; Welker et strength is common (Hamley et al. 1983; Henderson
al. 1994; Ludsin and DeVries 1997; Bunnell et al. 2003), 1985) although recruitment patterns may vary among
and predation (Forney 1971; Houde 1987; Rice et al. water bodies within a localized region (Lucchesi 1991;
1987; Santucci and Wahl 2003). Critical time periods, or Isermann et al. 2007). The early life-stages are commonly
Match–Mismatch of Bluegill and Yellow Perch and Their Prey J.C. Jolley et al.
reported as the time when year-class strength is formed 2009). The fish communities of both lakes were relatively
(Clady 1976; Mills and Forney 1981). Variable recruitment simple. Cameron Lake contained yellow perch, green
is often observed in bluegill populations. Previous sunfish Lepomis cyanellus, black bullhead Ameiurus melas,
research in South Dakota indicated that bluegill recruit- common carp Cyprinus carpio, fathead minnow Pime-
ment was asynchronous among four small impound- phales promelas, and golden shiner Notemigonus cryso-
ments (Edwards et al. 2007) and the authors suggested leucas. Pelican Lake contained bluegill, largemouth bass
that biotic factors likely affected bluegill recruitment. Micropterus salmoides, yellow perch, northern pike Esox
Although bluegill recruitment is generally consistent (i.e., lucius, black bullhead, common carp, and fathead
missing year-classes are rare) in Nebraska Sandhill lakes, minnow. The watersheds for both lakes were primarily
there appears to be a moderate level of variability in mixed- and tall-grass prairie and were used for limited
relative year-class strength (Paukert et al. 2002a; Jolley livestock grazing (Bleed and Flowerday 1989).
2009).
The availability of prey when larval fish begin Methods
exogenous feeding has been proposed as a potential
regulator of recruitment variability (match–mismatch We obtained larval yellow perch from Cameron and
hypothesis; Cushing 1975, 1990). Suitable growth and Pelican lakes and larval bluegill from Pelican Lake using
prey conditions for larval fish are largely determined by a single 1,000-mm-mesh conical ichthyoplankton net
the physical environment (e.g., water temperature). (mouth : net length ratio = 1:3) with a 0.76-m–diameter
Water temperature may indirectly affect larval fish via circular steel frame and 500-mL collection bucket (with
its influence on the food chain (e.g., zooplankton growth; 500-mm mesh). Isermann et al. (2002) found no
Sommer et al. 1986; Taylor et al. 1987) in addition to difference in yellow perch density estimates between
directly mediating spawning and hatching (Beard 1982). a 500- and 1,000-mm-mesh trawl, although the 1,000-mm
The match–mismatch hypothesis consists of two size was less likely to become fouled with algae and
assumptions: first, that fish at temperate latitudes spawn zooplankton. We sampled larvae approximately every
at approximately the same time each year and, second, 10 d from late April to early September in Pelican Lake,
that the larvae are released during the spring or autumn 2004–2008 and from late April to mid-June in Cameron
peaks in plankton production (Cushing 1990). A match Lake, 2004–2005. Randomly chosen, paired locations
occurs when production of fish larvae and their prey is (i.e., nearshore and offshore) were trawled in Pelican
synchronous or nearly so; conversely, a mismatch occurs Lake (n = 5) and Cameron Lake (n = 4) on each
when there is a large temporal difference in these two occasion. The density of recently hatched (i.e., , 8 mm)
variables (Cushing 1990). bluegill and yellow perch larvae in the lakes was
Although the food habits of age-0 yellow perch have indexed using a flowmeter (Ocean Test Equipment,
been extensively studied in Midwestern waters (e.g., Inc.) in the mouth of the trawl, which allowed
Weber and Les 1982; Wahl et al. 1993; Fisher and Willis determination of water volume towed. We collected
1997), lake-specific prey-selection patterns in the face of zooplankton at the time of each trawling sample during
variable prey densities is a topic of importance. Similarly, the daytime as two replicates at each site using a 2-m-
age-0 bluegill food habits are well-studied (Werner 1969; long tube sampler (Rabeni 1996). Samples were filtered
Mittelbach 1981; Werner and Hall 1988), although data through a 65-mm-mesh net. Replicate samples were
specific to Sandhill lakes have not been collected. collected and processed separately. All samples were
Furthermore, the panfish populations of Nebraska Sand- preserved in 90% ethanol and transported to the
hill lakes are high quality (i.e., relatively high abundance laboratory for identification and diet analysis.
of large fish; Paukert et al. 2002b) and relatively unique We tracked the same cohorts by sampling juvenile
to the Great Plains. Therefore, examinations of processes yellow perch from Cameron Lake in August and juvenile
that affect recruitment of these populations are of direct bluegill and yellow perch from Pelican Lake in August or
interest to managers. The objectives of this study were to September and the following April or May (age 1) using
1) describe prey selection for larval and juvenile yellow cloverleaf traps. Each three-lobed cloverleaf trap was
perch in Cameron and Pelican lakes and bluegill in constructed of galvanized 6.4-mm-bar mesh, with three
Pelican Lake, Nebraska, to better understand the prey 12.7-mm-wide openings between lobes to accommo-
utilization in these lakes, and 2) examine potential date entrance of small yellow perch (Brown and St.
match–mismatch regulation of yellow perch and bluegill Sauver 2002). Each lobe was 50 cm in diameter with a 41-
recruitment. cm height. Collected fishes were preserved in 90%
ethanol and returned to the laboratory. Autumn (age 0)
and spring (age 1) juvenile abundance was indexed as
Study Area
the mean number per cloverleaf trap-night.
Cameron Lake (39 ha) and Pelican Lake (322 ha) are Year-class strength was assessed in a concurrent study
shallow (mean depth = 1.2 m and 1.3 m, respectively), and information from the adult populations was used to
windswept natural lakes in the Sandhills region of north- examine recruitment; methods are described by Jolley
central Nebraska (McCarraher 1977). Submergent and (2009). Adult bluegill (i.e., age 2 and older) and yellow
emergent vegetation coverage was low (, 17% total perch (i.e., age 1 and older) were collected annually from
coverage) in Cameron Lake and was moderate (40–52% each lake using randomly located, overnight sets of
total coverage) in Pelican Lake in 2004 and 2005 (Jolley double-throated trap (i.e., modified fyke) nets with
Table 1. Number (No.) and mean total length (TL; mm) of yellow perch Perca flavescens and bluegill Lepomis macrochirus
examined for food items from Cameron and Pelican lakes, Nebraska, 2004–2005.
No. No.
Lake Date (with prey) examined x– TL SE Range % Empty
Cameron
Yellow perch
2004 13 May 30 30 11.5 0.2 9.9–13.1 0
18 May 30 30 12.5 0.2 9.1–14.9 0
27 May 30 30 15.7 0.5 10.0–19.6 0
7 June 30 30 25.0 0.5 19.4–30.0 0
2005 15 May 18 18 8.3 0.4 4.2–12.8 0
24 May 30 30 14.5 0.2 12.2–18.3 0
2 June 12 12 19.0 0.6 16.0–23.1 0
Pelican
Yellow perch
2004 8 May 30 35 8.8 0.2 6.8–13.8 14
17 May 30 30 12.7 0.2 10.4–14.5 0
26 May 30 30 17.1 0.4 12.7–20.6 0
6 June 23 24 23.1 0.5 18.1–26.9 4
16 June 30 31 32.2 0.7 19.5–37.3 3
26 June 17 17 28.0 1.7 18.9–39.5 0
7 July 7 7 31.8 2.5 22.0–36.8 0
17 July 3 3 41.8 3.7 34.6–46.5 0
27 July 9 9 49.3 1.0 46.0–53.0 0
6 August 14 14 57.6 2.3 40.2–69.5 0
16 August 13 13 59.8 0.9 56.2–67.1 0
26 August 9 9 59.9 2.6 44.6–74.1 0
2005 3 May 16 16 5.4 0.1 4.2–6.5 100
Pelican
Bluegill
2004 26 June 3 11 7.5 0.4 6.6–11.0 73
7 July 9 14 9.1 0.6 6.1–12.4 36
17 July 30 47 10.8 0.4 3.8–15.5 36
27 July 30 30 15.0 0.3 8.6–18.5 0
6 August 30 32 15.0 0.6 7.3–23.3 6
16 August 30 30 19.1 1.0 10.9–29.1 0
27 August 30 30 20.7 1.0 7.9–29.3 0
5 September 30 31 19.5 1.5 11.2–40.1 3
2005 30 June 30 32 8.3 0.2 7.2–12.3 6
11 July 30 31 10.7 0.4 7.8–14.7 3
21 July 30 31 10.4 0.4 7.4–14.1 3
1 August 30 35 11.4 0.4 8.0–15.6 14
11 August 30 30 11.2 0.4 7.3–15.3 0
22 August 30 30 12.9 0.4 9.8–18.4 0
16-mm-bar measure mesh, 1.1 6 1.5-m frames, and values in May or June for age-1 yellow perch and age-2
22-m leads. Age was estimated by two independent bluegill were selected as life-stages recruited to the adult
readers viewing the sagittal otolith in whole view for fish population.
of ages 4 and younger; older fish were aged after Larval fish samples were sorted and identified using
cracking and sanding the otoliths. Discrepancies in age identification keys (Auer 1982; Holland-Bartels et al.
estimates were reconciled by reading the otolith in 1990). Larvae were counted and up to 200 fish per
concert. If agreement could not be achieved, the otolith sample were measured (mm total length [TL]) from each
was omitted from the analysis. Mean catch per unit effort site. Digestive tracts were removed from up to 30
Table 2. Percent occurrence and percent by number for zooplankton prey items found in larval yellow perch Perca flavescens
stomachs in Cameron Lake, Nebraska, in 2004 and 2005.
2004 2005
Taxon 13 May 18 May 27 May 7 June 15 May 24 May 2 June
% Occurrence
Bosminidae 3.3 33.3 30.0 20.0 0 3.3 16.7
Chydoridae 6.7 20.0 26.7 66.7 0 0 16.7
Daphnidae 86.7 100.0 100.0 100.0 33.3 90.0 100.0
Sididae 0 23.3 30.0 0.0 0 0 0
Copepoda 100.0 96.7 66.7 60.0 94.4 100.0 91.7
Nauplii 0 0 0 0 0 0 0
Ostracoda 0 3.3 0 3.3 0 0 0
Rotifera 33.3 66.7 3.3 0 0 0 0
% by number
Bosminidae 0.2 2.3 2.5 0.3 0 0.2 0.3
Chydoridae 0.5 1.5 1.1 34.4 0 0 0.5
Daphnidae 26.8 37.5 81.7 60.2 24.4 28.8 38.6
Sididae 0 1.3 2.5 0 0 0 0
Copepoda 61.1 33.2 12.1 5.1 75.6 71.0 60.6
Nauplii 0 0 0 0 0 0 0
Ostracoda 0 0.2 0 0.1 0 0 0
Rotifera 11.3 24.1 0.1 0 0 0 0
randomly selected larvae of each species per sampling in 2004 and 2005 were examined and used to select prey
occasion during 2004 and 2005 at Cameron and Pelican items included in the match–mismatch analysis.
lakes. Diet items were identified using a microscope, Temporal predator and prey density curves for yellow
counted, and measured (mm TL). Zooplankton were perch and bluegill larvae and their prey were constructed
enumerated and identified to family for common for each year and the mean and standard deviation of
cladocerans (i.e., Bosminidae, Chydoridae, and Daphni- the peak density was calculated. Prey types used in the
dae), and as cyclopoid or calanoid copepods, copepod analyses were chosen as those showing positive
nauplii (copepodites), ostracods, and rotifers. Each selection by fish during first-feeding (earlier in the
sample was diluted with water to a volume of 30 mL. season). The width and overlap of the density curves
Three subsamples were then taken with a 5-mL Hensen– for predator and prey were calculated. Methods outlined
Stempel pipette and placed in a Ward counting wheel. by Mertz and Myers (1994) and Johnson (2000) were
Up to 20 individuals of each category were measured utilized for this analysis. The following parameters were
and all individuals were counted. The total number of first calculated:
zooplankton of each taxon in a sample was calculated by
dividing the number of organisms counted by the t0 ~timing between peaks of larval production and
proportion of the sample volume processed. Density food supplyðdaysÞ;
was then calculated by dividing the number of
zooplankton of each taxon by the volume of the water Dt0 ~annual differences inðt0 Þfrom its mean value;
filtered with the tube sampler.
To determine prey selectivity, mean Chesson’s (1983) d~one half width of the density curve for larvae; and
coefficient of selectivity (a) was calculated for individual
larval bluegill and yellow perch from each sampling s~one half width of the density curve for
occasion in 2004 and 2005: zooplankton:
ri =ni
a~ These parameters were used to calculate 1) variability
X
m
ri =ni in peak timing from the mean for individual species (s
i~1
and d), 2) variability in timing between larval abundance
and peak zooplankton production (t0), and 3) year-to-
where ri is the number of food type i in the predator diet, year variability in peak spawning and production (Dt0).
ni is the number of food type i in the environment, and When t0 = 0, the match is exact (Mertz and Myers 1994).
m is the number of prey types available. Mean a values A mismatch occurs when one-half the width of the larval
(6 95% CI) were compared with nonselective feeding density curve (i.e., d) does not overlap one-half the width
(1/m) to determine selectivity. Patterns in prey selection of the zooplankton density curve (i.e., s). Correlation
season and later shifted to cladocerans. In 2005, copepods

were exclusively positively selected (Figure 1). Empty
stomachs were not encountered in either year and
copepodites were not observed in stomachs in any year.
In Pelican Lake in 2004, the most abundant zooplank-
ters were generally daphnids, copepods, and copepod-
ites. In 2005, the most abundant zooplankters were
generally cladocerans (i.e., daphnids, chydorids, and
bosminids). In 2004, 222 yellow perch larvae were
examined from May through August for prey items
(Table 1). Seven empty stomachs were encountered
(3%). In 2005, 16 larval yellow perch were captured, all
with empty stomachs. In 2004, copepods, daphnids, and
chydorids were the most abundant prey items found in
yellow perch stomachs (Table 3). Yellow perch in Pelican
Lake selected copepods earliest, shifted to cladocerans,
and later exhibited neutral prey selection for most prey
items (Figure 2). Copepodites were not present and
rotifers were uncommon in yellow perch stomachs.
Temporal patterns of negative, neutral, and positive
prey selection (Chesson’s a . 0.143 indicated positive
prey selection) were also found for bluegill in Pelican
Lake in 2004 and 2005 (Figure 3). Bluegill larvae had 15%
and 5% empty stomachs in 2004 and 2005, respectively
(Table 1). Bluegill consumed cladocerans, copepods,
copepodites, ostracods, and rotifers. Copepodites and
Bosmina were common in first-feeding bluegill stomachs
while cladocerans and copepods became more common
later in the season (Table 4). Copepodites were initially
neutrally selected and bluegill eventually preferred
Figure 1. Mean (with 95% CI) prey selection (Chesson’s a) by cladocerans followed by copepods as prey items. Rotifers
yellow perch Perca flavescens in Cameron Lake, Nebraska, 2004– and ostracods were consistently neutrally or negatively
2005 by prey category of zooplankton. Confidence intervals selected (Figure 3) indicating opportunistic feeding or
above the random feeding (dashed) line indicate positive avoidance of these prey items.
selection, values below the line indicate negative selection, and Copepods were elected as an important (i.e., positively
values overlapping the line indicate neutral selection.
selected) prey resource for first-feeding yellow perch in
both lakes. The duration of larval yellow perch (TL ,
analysis was used to examine the relationship between 13 mm) abundance varied from 1 d in 2005 (Pelican Lake)
predator–prey overlap (t0) and indices of fish recruitment to 31 d in 2008 (Pelican Lake). The duration of peak
(i.e., larval abundance, juvenile abundance, and adult copepod abundance varied from 9 (Cameron Lake, 2005)
abundance) for yellow perch and bluegill in Pelican Lake to 38 d (Pelican Lake, 2004). The mean annual difference in
from 2004 to 2008. Yellow perch from Cameron Lake day of peak abundance for yellow perch larvae and their
were omitted from this analysis due to inadequate copepod prey (t0) was 7.1 d (Table 5). Copepods peaked
sample size (N = 2 y). after yellow perch larvae in three instances and exactly
matched in four instances (Figures 4 and 5; Table 5).
Results Copepodites and Bosmina were important (i.e., posi-
tively selected) prey item for first-feeding bluegill and the
In Cameron Lake, the most abundant zooplankters were timing of their combined abundance was examined. The
daphnids (Supplemental Material, Table S1, http:// duration of larval bluegill abundance in Pelican Lake
dx.doi.org/10.3996/062010-JFWM-018.S1). Chesson’s a . ranged from 21 (2004) to 63 d (2006; Figure 6). The
0.125 indicated positive prey selection by yellow perch. duration of peak copepodites–Bosmina abundance
Stomachs from 120 yellow perch larvae in May and June of ranged from 42 (2006) to 72 d (2004). The mean annual
2004 and 60 stomachs in 2005 were examined for prey difference in peak abundance date for bluegill predators
items (Table 1; Supplemental Material, Table S2, http:// and their prey (t0) was 14 d. Larval bluegill peaked in
dx.doi.org/10.3996/062010-JFWM-018.S2). Copepods and abundance before their prey in most years. In 2004, there
daphnids were consistently the most abundant prey item were two peaks in larval bluegill abundance and
found in stomachs (Table 2). Temporal patterns of zooplankton prey peaked between these two dates
negative, neutral, and positive prey selection (Chesson’s a (Figure 6; Table 5).
. 0.143 indicated positive prey selection) were found for Matches between larval abundance and zooplankton
yellow perch in Cameron Lake (Figure 1) in 2004 and 2005. prey occurred in all years (i.e., d and s overlapped;
In 2004, yellow perch selected copepods earliest in the Table 5) for both species. There were exact matches (i.e.,
Table 3. Percent occurrence and percent by number for zooplankton prey items found in larval yellow perch Perca flavescens
stomachs in Pelican Lake, Nebraska, in 2004.
8 17 26 6 16 26 7 17 27 6 16 26
Taxon May May May June June June July July July August August August
% Occurrence
Bosminidae 0 0 0 0 0 0 14.3 33.3 37.5 57.1 100.0 77.8
Chydoridae 0 3.3 0 0 50.0 64.7 71.4 66.7 75.0 50.0 100.0 77.8
Daphnidae 0 70.0 100.0 100.0 26.7 41.2 42.9 33.3 62.5 42.9 100.0 77.8
Sididae 0 0 0 0 0 0 0 0 12.5 21.4 76.9 0
Copepoda 100.0 100.0 13.3 17.4 70.0 58.8 71.4 66.7 75.0 57.1 76.9 22.2
Nauplii 0 6.7 0 0 0 0 0 0 0 0 0 0
Ostracoda 0 0 0 0 50.0 47.1 14.3 0 62.5 57.1 38.5 11.1
Rotifera 0 0 0 0 0 0 0 0 0 0 0 0
% by number
Bosminidae 0 0 0 0 0 0 0.6 19.2 39.5 74.0 84.8 73.0
Chydoridae 0 0.8 0 0 10.3 44.5 19.5 28.3 10.2 3.3 4.5 9.5
Daphnidae 0 15.5 97.0 98.8 2.3 33.2 18.5 12.1 31.3 14.1 9.3 16.9
Sididae 0 0 0 0 0 0 0 0 0.8 1.2 0.1 0
Copepoda 100.0 83.0 3.0 1.2 50.1 15.0 60.8 40.4 16.4 6.6 1.3 0.6
Nauplii 0 0.8 0 0 0 0 0 0 0 0 0 0
Ostracoda 0 0 0 0 37.3 7.4 0.6 0 1.8 0.8 0 0
Rotifera 0 0 0 0 0 0 0 0 0 0 0 0
t0 = 0) in 4 y for yellow perch. No correlations between Discussion

predator–prey overlap (t0) and recruitment indices were
significant for yellow perch or bluegill (P . 0.05; Table 6). Larval fish predators and their zooplankton prey were
For both species the highest observed abundance temporally well-matched in our study. Prey densities
occurred in a year where the predators temporally were adequate for larval yellow perch and bluegill.
matched their prey (Table 5). Although a low number of yellow perch were detected in
Figure 2. Mean (with 95% CI) prey selection (Chesson’s a) by yellow perch Perca flavescens in Pelican Lake, Nebraska, 2004 by prey
category of zooplankton. Confidence intervals above the random feeding (dashed) line indicate positive selection, values below the
line indicate negative selection, and values overlapping the line indicate neutral selection.
occur than in marine environments because freshwater

fish larvae are generally larger, thus conferring greater
energy reserves and resistance to starvation (Miller et al.
1988). Previous research examining the relation of
zooplankton biomass and abundance to larval growth
and survival of panfish has had mixed results (Pope and
Willis 1998; Garvey et al. 2002; Bunnell et al. 2003).
In 2005, yellow perch hatched over a very narrow time
frame in Pelican Lake (4 d) and larvae were only collected
in low numbers on one day, likely indicating a relatively
weak initial year-class. Concurrently, Pelican Lake expe-
rienced a drop in water temperature from nearly 17uC to
nearly 6uC over a period of 2 wk in late April (Jolley 2009).
This corresponded to the time period when yellow perch
eggs would have been incubating and hatching. It is
unclear whether the eggs or newly hatched larvae were
negatively affected by this cold front. Jansen et al. (2009)
simulated the effect of a cold front on yellow perch eggs
and found no decrease in egg survival, suggesting that
the newly hatched larval stage may be more susceptible
than eggs to these extreme weather events. Sandhill
lakes are shallow and windswept; they, thus, are
susceptible to erratic temperature changes and can
warm and cool quickly. Jolley (2009) examined the
relation of recruitment (i.e., year-class strength) of
bluegill and yellow perch to climatic variables in several
Sandhill lakes and found asynchronous recruitment
among the study lakes. In addition, limited support for
the concept of climatic influence on bluegill and yellow
perch was found over the years examined.
No mismatches in predator and prey abundance were
Figure 3. Mean (with 95% CI) prey selection (Chesson’s a) by detected over the years examined in our study; thus, we
bluegill Lepomis macrochirus in Pelican Lake, Nebraska, 2004 cannot determine if severe mismatches in the appear-
and 2005 by prey category of zooplankton. Confidence
intervals above the random feeding (dashed) line indicate
ance of fish larvae and their zooplankton prey would lead
positive selection, values below the line indicate negative to depressed survival of age-0 bluegill and yellow perch
selection, and values overlapping the line indicate neutral in Nebraska Sandhill lakes. Density of larval fish and
selection. zooplankton was variable among years but exact
matches occurred frequently for bluegill and yellow
perch. Although zooplankton density can vary spatially
both lakes in 2005, prey was seemingly available for (Young et al. 2009) leading to potential spatial mis-
consumption. The observation of all larval yellow perch matches (Chick and Van Den Avyle 1999), differences in
collected with empty stomachs in 2005 in Pelican Lake zooplankton densities between inshore and offshore
was notable. It is unknown if these fish were in a stressed strata were not apparent (Jolley 2009). Intrastation
condition upon their collection and were unable to variability of zooplankton density was generally less than
adequately seek prey items or if they may not have interstation variability, and density differences were
started first-feeding (yolk-sacs were not present). Sparse rarely detected among regions of the lake (unpublished
prey resources for first-feeding fish larvae have been data). Young et al. (2009) reported that small-scale
commonly implicated as a cause of high mortality (May patchiness (i.e., , 1 m) accounted for the majority of the
1974; Cushing 1975, 1990; Hart and Werner 1987) and variation in zooplankton abundance, which could
Toetz (1966) experimentally reported mass starvation contribute to spatial mismatches considering the small
of larval bluegill between 5 and 6 mm TL, which search volume reported for many larval fish species (,
corresponded to initiation of exogenous feeding. Iser- 2.5 L; Blaxter 1986; Pepin 2004). They suggested that
mann and Willis (2008) suggested that factors prior to, measures of average prey density made over larger
during, or immediately following hatching may play a scales may be independent of the feeding of individuals.
critical role in the recruitment of yellow perch under the Determination of the scale of zooplankton patchiness
constraint of a narrow spawning window. Our results are was beyond the scope of our study but may be a topic
contrary to multiple experimental studies in which worth revisiting in future studies.
survival of larval fishes is most influenced by zooplankton Although copepodites and rotifers have also been
biomass (Hart and Werner 1987; Welker et al. 1994). reported (Whiteside et al. 1985; Schael et al. 1991; Wahl
Conversely, Houde (1994) predicted that starvation of et al. 1993; Fisher and Willis 1997) as a preferred prey
larval fishes in freshwater environments was less likely to item of newly hatched yellow perch, we found rare
Table 4. Percent occurrence and percent by number for zooplankton prey items found in larval bluegill Lepomis macrochirus
stomachs in Pelican Lake, Nebraska, in 2004–2005.
2004 2005
26 7 17 27 6 16 27 5 30 11 21 1 11 22
Taxon June July July July August August August Sept June July July August August August
% Occurrence
Bosminidae 0 0 63.3 50.0 80.0 83.3 96.7 83.3 46.7 60.0 60.0 76.7 63.3 70.0
Chydoridae 0 22.2 10.0 3.3 6.7 40.0 53.3 60.0 6.7 56.7 63.3 33.3 40.0 50.0
Daphnidae 0 0 46.7 83.3 60.0 83.3 96.7 80.0 26.7 63.3 76.7 60.0 96.7 83.3
Sididae 0 0 0 3.3 3.3 13.3 10.0 43.3 0 0.0 3.3 6.7 3.3 43.3
Copepoda 33.3 11.1 30.0 96.7 76.7 83.3 86.7 90.0 60.0 33.3 36.7 20.0 23.3 63.3
Nauplii 66.7 44.4 10.0 3.3 3.3 10.0 6.7 6.7 40.0 6.7 16.7 0 3.3 0
Ostracoda 0 0 3.3 0 0 0 0 6.7 10.0 6.7 3.3 0 0 0
Rotifera 0 66.7 16.7 0 0 3.3 0 3.3 0 20.0 0 0 10.0 0
% by number
Bosminidae 0 0 46.7 15.3 42.6 46.2 47.9 34.6 21.7 28.3 32.9 51.2 44.5 32.8
Chydoridae 0 8.8 3.5 0.2 0.5 9.3 5.3 19.2 1.6 22.7 23.9 5.7 14.6 12.9
Daphnidae 0 0 38.1 21.9 22.7 12.9 18.4 23.4 10.1 34.0 31.2 39.2 33.0 28.9
Sididae 0 0 0 0.2 0.2 0.7 0.2 3.2 0 0 0.4 1.0 0.2 3.6
Copepoda 10.0 5.9 5.4 61.9 33.6 30.4 28.0 19.0 34.1 11.5 9.0 2.9 6.4 21.8
Nauplii 90.0 20.6 1.6 0.4 0.5 0.3 0.2 0.4 27.9 0.4 2.1 0 0.2 0
Ostracoda 0 0 0.4 0 0 0 0 0.2 4.7 0.7 0.4 0 0 0
Rotifera 0 64.7 4.3 0 0 0.1 0 0.1 0 2.4 0 0 1.1 0
consumption of these taxa despite their availability. although daphnids were first consumed when yellow
Increased predation on daphnids by larger (i.e., 30 mm) perch were between 15 and 17 mm TL and Wahl et al.
yellow perch has been previously reported (Whiteside et (1993) reported consumption of daphnids by larval
al. 1985; Prout et al. 1990) and positive selection for these yellow perch as small as 9 mm TL.
diet items has been shown (Mills et al. 1984; Schael et al. Yellow perch hatched 2 mo before bluegill and had
1991). Our observations corroborate such findings the potential to either prey upon or compete with
Table 5. Mean peak larval abundance (n/100 m3), parameters used in match–mismatch examination, and result for yellow perch
Perca flavescens and copepods in Cameron and Pelican Lake, Nebraska, and for bluegill Lepomis macrochirus and combined
copepodite–Bosmina in Pelican Lake, Nebraska, 2004–2008. Parameters are t0 (timing between peaks of larval production and food
supply in days), Dt0 (annual differences in [t0] from its mean value), d (one-half width of the production period for larvae), and s
(one-half width of the production period for zooplankton).
Parameter
Predator–prey Peak larval
Lake combination Year abundance t0 Dt0 d s Result
Cameron Yellow perch–copepods 2004 804 19 11.9 9.5 7 Match
2005 2 0 7.1 0 4.5 Exact match
Pelican Yellow perch–copepods 2004 23 0 7.1 0 19 Exact match
2005 6 0 7.1 0 10 Exact match
2006 46 21 13.9 4.5 16 Weak match
2007 89 0 7.1 4 9 Exact match
2008 238 10 2.9 15.5 26 Match
Bluegill–copepodite–Bosmina 2004a 66 20 8.0 20.5 36 Match
2004b 57 9 3.0 10.5 36 Match
2005 169 21 9.0 21 26.5 Match
2006 315 10 2.0 31.5 21 Match
2007 377 11 1.0 21 26 Match
2008 1,269 11 1.0 19 25 Match
a
First peak in larval bluegill abundance in 2004.
b
Second peak in larval bluegill abundance in 2004.
Figure 4. Larval yellow perch Perca flavescens density (solid

line) and copepod density (broken line) in Cameron Lake,
Nebraska, in 2004 and 2005 (n = number).
bluegill larvae. We did not observe any larval bluegill

consumed by juvenile yellow perch in this study (up to
74 mm TL). Although Graeb et al. (2006) demonstrated
experimentally that yellow perch begin a shift to fish
prey at 80 mm TL, most field studies do not report
consistent piscivory by yellow perch until they attain
130–150 mm TL (Clady 1974; Keast 1985; Fullhart et al.
2002), a larger size than examined in our study.
Total zooplankton densities in 2004 were low on the
first day that bluegill larvae were collected, which may
explain the lack of positive prey selection for any
particular group of zooplankton by newly hatched
bluegill larvae in 2004. Partridge and DeVries (1999)
noted a high proportion of rotifers in larval bluegill diets,
which may lead to suboptimal bluegill growth. Rotifers
were rarely consumed in our study, although they were
remarkably abundant in Pelican Lake in 2004. The
availability of energetically profitable prey (i.e., copepods
and cladocerans) likely precluded bluegill larvae from
consuming rotifers.
Although densities of yellow perch larvae varied
considerably (. 300%) among years, it appears that
Figure 5. Larval yellow perch Perca flavescens density (solid
recruitment of yellow perch was relatively consistent in line) and copepod density (broken line) in Pelican Lake,
Pelican and Cameron lakes over the years examined and Nebraska, 2004–2008 (n = number).
the observed densities of larval yellow perch were
generally higher than reported values of density in six
South Dakota glacial lakes monitored for over 8 y (Jansen and space is no doubt an important factor to the
2008). Many other Sandhill lakes concurrently studied survival and recruitment of age-0 fish and is likely a
exhibited relatively consistent recruitment (Jolley 2009). component of an integrated process acting throughout
The life history and reproductive and spawning early life-stages to explain components of recruitment
behavior of many species have evolved in ecosystems variability (Houde 2008). Many studies that provide
where environmental variability is unpredictable (Wine- support for the match–mismatch hypothesis involved
miller and Rose 1993). A combination of trophodynamic marine species and systems (Cushing 1990; Fortier and
and physical factors may interact in complex ways, over Gagné 1990; Gotceitas et al. 1996; Johnson 2000); it is
multiple temporal and spatial scales, to affect larval fish lesser studied in freshwater systems, with some support
survival, growth, and recruitment (Fitzgerald et al. 2001; for landlocked striped bass Morone saxatilis (Chick and
Houde 2008). Availability of appropriate prey in time Van Den Avyle 1999), threadfin shad Dorosoma pete-
Table 6. Bivariate correlations between predator–prey

overlap (t0) and fish abundance indices for yellow perch
Perca flavescens and bluegill Lepomis macrochirus. Abundance
indices are peak larval abundance (mean n/m3), autumn and
spring juvenile catch per unit effort (CPUE; mean n/cloverleaf
trap-night), and age-1 and age-2 CPUE (mean n/trap-net
night). Number of data pairs (N), correlation coefficient (r), and
P-value are given for each bivariate correlation.
Yellow perch Bluegill
Independent
variable N r P N r P
Peak larval abundance 7 20.19 0.68 6 20.34 0.51
Autumn juvenile CPUE 7 0.13 0.77 6 0.08 0.88
Spring juvenile CPUE 7 0.32 0.48 6 20.36 0.49
Age 1 CPUE 7 20.51 0.38
Age 2 CPUE 5 20.43 0.47
be able to account for observed recruitment variability

in the populations that we studied.
Given the importance of copepods, copepodites,
daphnids, and bosminids as prey for larval yellow perch
and bluegill, more in-depth examination of these
relationships is warranted. Abundance indices of these
zooplankton taxa may be used as surrogates for prey
availability. In addition, the dynamic nature of zooplank-
ton populations may suggest important consequences
via timing (e.g., match–mismatch regulation; Cushing
1975, 1990), which could be further explored. Our
sampling interval of 10 d may lack the required
resolution to fully understand the relation between
zooplankton and larval bluegill and yellow perch
recruitment. If catastrophic mortality events happen in
a short time (Hjort 1914; May 1974) then more frequent
sampling may be required to pinpoint the timing and
explanation for it. Multiple life-stage abundance indices
of yellow perch and bluegill were not correlated to
zooplankton abundance indices, although those results
were based on a low number of observations (Jolley
2009). Continued stage-specific investigations of the
relationship of zooplankton to larval fish growth and
recruitment may produce important insights into the
Figure 6. Larval bluegill Lepomis macrochirus density (solid dynamics of bluegill and yellow perch in temperate lakes.
line) and combined copepodite and Bosmina density (broken
In addition, examination of later life-stages (e.g., juve-
line) in Pelican Lake, Nebraska, 2004–2008 (n = number).
niles) of bluegill and yellow perch likely is also necessary.
Studies that incorporate multiple life-stages (Ludsin and
DeVries 1997; Jolley 2009) and include ecosystem
nense (Betsill and Van Den Avyle 1997) and yellow perch processes are especially valuable (Cury et al. 2008).
(Fitzgerald et al. 2001). Houde (1994) found that marine
larvae (compared to freshwater) may be more suscep- Supplemental Material
tible to starvation mortality due to high metabolic
demand and small size at hatch. These traits may be Please note: The Journal of Fish and Wildlife Management
related to the applicability of the match–mismatch is not responsible for the content or functionality of any
hypothesis for marine fishes. Houde (1994) found that supplemental material. Queries should be directed to the
freshwater fish larvae may be more susceptible to corresponding author.
episodic mortalities that affect recruitment, but that the
juvenile stage may be equally important in regulating Table S1. Zooplankton availability.
and controlling recruitment. In light of these complex- Found at DOI: 10.3996/062010-JFWM-018.S1 (292 KB XLS).
ities, and given that zooplankton were abundant and Table S2. Larval fish diets.
well-timed to larval fish abundance over the initial years Found at DOI: 10.3996/062010-JFWM-018.S2 (1757 KB
of this study, the match–mismatch hypothesis may not XLS).
Acknowledgments flavescens. Journal of the Fisheries Research Board of

Canada 33:1887–1893.
We thank A. Husman, E. Lorenzen, J. Rydell, D. Coulter,
Cury PM, Shin Y-J, Planque B, Durant JM, Fromentin J-M,
W. Bauer, and C. Longhenry for field and laboratory
assistance. M. Lindvall and Valentine National Wildlife Kramer-Schadt S, Stenseth NC, Travers M, Grimm V.
Refuge provided access to Pelican Lake and R. Lackaff 2008. Ecosystem oceanography for global change in
provided access to Cameron Lake. We thank D. Graham, fisheries. Trends in Ecology and Evolution 23:338–346.
D. Hartmann, D. Kruger, and the Valentine State Fish Cushing DH. 1975. Marine ecology and fisheries. United
Hatchery for assistance. We also thank the three Kingdom: Cambridge University Press.
anonymous reviewers and the Subject Editor for Cushing DH. 1990. Plankton production and year-class
valuable improvement to this manuscript. Funding for strength in fish populations: an update of the match/
this project was provided by the Nebraska Game and mismatch hypothesis. Advances in Marine Biology 26:
Parks Commission through Federal Aid in Sport Fish 249–293.
Restoration Project F-118-R. Diana JS. 1995. Biology and ecology of fishes. Carmel,
Indiana: Cooper.
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Articles
Acclimation Improves Short-Term Survival of Hatchery
Lahontan Cutthroat Trout in Water From Saline, Alkaline
Walker Lake, Nevada
John P. Bigelow,* Wendy M. Rauw, Luis Gomez-Raya
J.P. Bigelow
U.S. Fish and Wildlife Service, Lahontan National Fish Hatchery Complex, Reno, Nevada 89502
Present address: 615 Jill Drive, Gardnerville, Nevada 89460
J.P. Bigelow, W.M. Rauw, L. Gomez-Raya
Department of Animal Biotechnology, University of Nevada, Reno, 89557, Nevada USA
Abstract
We investigated the effectiveness of two acclimation protocols for 8-month-old Lahontan cutthroat trout Oncorhynchus
clarkii henshawi, reared at Lahontan National Fish Hatchery in terms of survival during a week-long challenge in water
from saline, alkaline Walker Lake, Nevada. Fish were acclimated for 0 (control), 3, and 8 d by increasing the ratio of lake
water to hatchery water. For the 3-d acclimation treatment, 50% of the tank water was replaced with lake water each day.
For the 8-d treatment, 33% of the water was replaced with lake water on the first through fourth day of acclimation.
Survival during acclimation (i.e., prior to the challenge) was lowest for fish acclimated 3 d. Median survival time during the
lake water challenge was 8 h for unacclimated fish, and 8 and 12 h for fish surviving the 3- and 8-d acclimation treatments,
respectively. No fish survived the entire week-long challenge. Compared with no acclimation, 3- and 8-d acclimation
decreased the hazard of mortality during the challenge. Increased fork length also reduced the hazard of death. Our
results indicate acceptable survival rates cannot be achieved for subyearling, hatchery-reared Lahontan cutthroat trout
stocked in Walker Lake without acclimation or with the acclimation methods employed in this study. Our results indicate
that the acclimation method might be improved by the use of longer fish, longer acclimation, and better control of water
temperature, ammonia concentration, and alkalinity.
Keywords: Lahontan cutthroat trout; alkalinity; Walker Lake; survival; acclimation; fork length; salinity
Received: January 19, 2010; Accepted: September 27, 2010; Published Online Early: October 2010; Published:
November 2010
Citation: Bigelow JP, Rauw WM, Gomez-Raya L. 2010. Acclimation improves short-term survival of hatchery Lahontan
cutthroat trout in water from saline, alkaline Walker Lake, Nevada. Journal of Fish and Wildlife Management 1(2):86–92;
e1944–687X. doi: 10.3996/012010-JFWM-001
* Corresponding author: jybglw@aol.com
Introduction dissolved solids [TDS]) for other trout species (Behnke

1992; Dickerson and Vinyard 1999). As a result of
Lahontan cutthroat trout Oncorhynchus clarkii hen- increased human use of ground and surface water in
shawi, a federally listed ‘‘threatened’’ species (USOFR the Walker River Basin, the volume of Walker Lake
1975), was extirpated from Walker Lake subsequent to decreased from 9 million acre-feet in 1882 to 4.5 million
the construction of Derby Dam in 1933, which blocked acre-feet in 2003 (USFWS 2003); TDS showed a corre-
access of the lake population to important spawning sponding increase from 2,500 mg/L to 14,600 mg/L
habitat in the Walker River (La Rivers 1962; Sigler and (USFWS 2003).
Sigler 1987; Behnke 1992). Prior to extirpation, Lahontan For the past several decades, state and federal
cutthroat trout grew to sizes exceeding 18 kg and hatcheries have maintained a fishery in Walker Lake by
thrived in waters too saline (i.e., about 2,000 mg/L total annually stocking Lahontan cutthroat trout derived from
Acclimation of Lahontan Cutthroat Trout to Walker Lake, Nevada J.P. Bigelow et al.
out-of-basin strains. After 1995, hatchery Lahontan and Kirchoff 2007). All fish were reared in 12uC artesian
cutthroat trout were acclimated in water of intermediate well water under natural photoperiod. Fish were raised
salinity at Pyramid Lake, Nevada, prior to release in Walker on Abernathy trout diet (Rangen Inc., Buhl, ID). Fish were
Lake. After 2005, reduced relative inflows to Walker Lake taken off feed 2 d prior to transport and remained off
compared to Pyramid Lake increased the disparity feed throughout acclimation and the lake water chal-
between their salinities to the point where acclimation lenge to reduce ammonia excretion rates in the static
at Pyramid Lake was considered ineffective. Bioassays in tanks. Fish were transported from the hatchery to a
Walker Lake indicated short-term survival rates of shoreline location (town of Walker Lake) in three 800-fish
Lahontan cutthroat trout acclimated in Pyramid lake batches corresponding to 8- and 3-d acclimation
decreased from .90% in 1995 to ,5% in 2005 (NDOW treatments and unacclimated fish (0-d acclimation) on
2005). The Lahontan National Fish Hatchery Complex December 10, 15, and 18, 2008, respectively. Each batch
produces Lahontan cutthroat trout for recovery efforts in of 800 fish was divided (with no known bias) into four
Walker Lake, a saline and alkaline terminal lake in 2,320-L tanks containing static hatchery water supplied
northwestern Nevada. The salinity and alkalinity of Walker with compressed oxygen. Tank walls and lids were
Lake (.17,400 mg/L TDS and .9.5 pH, respectively) are so insulated with 1.9-cm-thick R-MatteH Plus-3 (RMAX,
much higher than the well water utilized to produce trout Dallas, TX). Fish were allowed to adjust to the lakeside
at the Lahontan National Fish Hatchery in Gardnerville, tanks in hatchery water for 24 h before any exposure to
Nevada (291–296 mg/L TDS and 7.5–7.8 pH, respectively), lake water, during which time there was no mortality.
that new methods of acclimation are necessary. Dickerson
and Vinyard (1999) found that challenges in Walker Lake Acclimation
water concentrated to 15,467 mg/L TDS quickly killed Lake water was introduced into the 8- and 3-d
unacclimated subyearling Lahontan cutthroat trout de- acclimation treatment tanks beginning 24 h after
rived from the contemporary Pyramid Lake stock and transport on December 11 and 16, 2008, respectively.
reared at Lahontan National Fish Hatchery. Acclimation in The ratio of lake to hatchery water was increased by
static water and increased fish size or age improved pumping portions of water out of each tank and
survival rates in unaltered lake water (then at 12,400 mg/L replacing it with lake water. For the 8-d treatment, 33%
TDS and 9.4 pH; Dickerson and Vinyard 1999). of the water was replaced with lake water on the first
The U.S. Fish and Wildlife Service is evaluating the through fourth day of acclimation. For the 3-d acclima-
feasibility of constructing an acclimation facility on the tion treatment, 50% of the water was replaced with lake
shore of Walker Lake. The acclimation facility would water on each day during acclimation. As a result, salinity
provide a means of exposing hatchery Lahontan and alkalinity increased at rates associated with acclima-
cutthroat trout to increasing concentrations of water tion treatment (Figure 1). All lake water was collected
from Walker Lake prior to their release. The simplest from the lake surface near the shore. No lake water was
method of acclimation would be to transport fish and added to the 0-d acclimation tanks until the beginning of
hatchery water from the hatchery to static lakeside tanks the challenge 24 h after transport. Temperatures in all
and repeatedly replace a portion of the hatchery water tanks were monitored to the nearest 0.01uC at 30-min
with lake water to increase the ratio of lake water to intervals with Stowaway Tidbit Temperature Loggers
hatchery water. The most efficient rate, pattern, and (Onset, Bourne, MA). Total ammonia was measured with
duration of increasing the lake to hatchery water ratio a Nesslerization and colorimeter kit (LaMotte Co.,
need to be empirically determined. The objective of this Chestertown, MD) in each tank 24 h after the addition
study was to test the effect of fish size on survival, and of fish. Dissolved oxygen concentrations were monitored
the effectiveness of two acclimation protocols for 8- with a Hach HQ10 oxygen meter (Hach Co., Loveland,
month-old Lahontan cutthroat trout reared at Lahontan CO) and maintained between 8 and 14 mg/L in all tanks.
National Fish Hatchery, in terms of reducing the hazard Electrical conductivity (EC) and pH were monitored once
of mortality compared to that of unacclimated fish a day with a Hach sensION156 multiparameter meter.
during a week-long challenge in water from saline, Electrical conductivity was standardized for temperature
alkaline Walker Lake, Nevada. according to the linear relationship determined on site
(EC = 0.4177 6 temperature + 11.776; N = 79; r2 =
Methods 0.997). Electrical conductivity was converted to TDS as in
Tracy (2004; TDS [mg/L] = 0.8466 6 EC [microsiemens/
Two thousand four hundred 8-month-old Lahontan cm] at 25.0uC 2 1,470.4). Mortalities were enumerated
cutthroat trout with a mean fork length (FL) of 143 mm daily.
(617.6 mm SD) were randomly selected from the
2008 year class at Lahontan National Fish Hatchery, Challenge
Gardnerville, Nevada. The fish were of the same age and On December 19 at 1800 h, 76 fish were selected from
size typically stocked in Walker Lake during February and each acclimation tank (with no known bias) to participate
March. All fish were fourth-generation progeny of trout in a week-long lake water challenge. One extra tank was
captured from wild populations in the Pilot Mountains of placed next to tank 1 in the row of 12 acclimation tanks.
western Utah confirmed by phylogenetic analysis as The extra tank was filled with 1,935 L of Walker Lake
recent descendants from the extirpated Truckee River water and supplied with 76 fish netted from tank 1. Tank
subbasin populations in Nevada and California (Peacock 1 was then drained, filled with 1,935 L of lake water, and
Statistical analysis
The effects of the acclimation treatments and FL on
the hazard of death were compared using a Cox
proportional hazards model:

^ ^ ^
hi (t)~l0 (t)exp b3 xi3 z b8 xi8 z bFL xiFL ð1Þ
where hi(t) was the hazard of death for the ith individual
at time t; l0 was an unspecified baseline hazard function;
b̂3 and b̂8 were the estimated effect parameters for 3-
and 8-d acclimation treatments, respectively; b̂FL was the
effect parameter for fork length; xi3 was equal to 1 if
individual i was in the 3-d acclimation treatment and 0
otherwise; xi8 was equal to 1 if individual i was in the 8-d
acclimation treatment and 0 if otherwise; and xiFL was
the fork length of individual i. This model placed
unacclimated fish in the reference group with both xi3
and xi8 equal to 0.
Raw data were arranged in a matrix with rows
dedicated to individuals and a column designated for
each variable (xi3, xi8, xiFL, and time of death). Effect
parameters were estimated by maximizing an equation
based on the partial likelihood equation:
" Pp #
exp k~1 bk Z(i)k
D
L(b)~ P P Pp ð2Þ
j[R(ti ) exp k~1 bk Zjk
i~1
where t1 , t2 , … , tD denoted the ordered times of

death; bk was the kth of p effect parameters, Z(i)k was the
kth of p covariates associated with the individual whose
death was at time ti; R(ti) was the set of all individuals j
who were still alive at time i 2 1; and Zjk was the kth of p
covariates associated with individual j (Klein and
Moeschberger 1997) but modified to accommodate ties
in time of death according to the EXACT method as
described by Allsion (1992).
Figure 1. (A) Total dissolved solids (mg/L), (B) pH, and (C) The hazard ratio for each indicator variable was
temperature (uC) of 2,320-L tanks in 0-, 3-, and 8-d acclimation calculated as exp(b̂) and denoted the relative risk of
treatments of subyearling Lahontan cutthroat trout raised at death suffered by individuals for which that indicator
the Lahontan National Fish Hatchery to Walker Lake water
during December 2008 by days before lake water challenge.
variable was equal to 1 compared to individuals for
Values in A and B are means (6SE) of daily measurements for which the indicator variable was equal to 0 when all
four replicate tanks. Values in C are daily means (6SD) of 48 other variables were of the same value. For example, if
measurements taken from each of four replicate tanks. Daily b̂8 = 21.2, then the hazard ratio (e21.2) = 0.3, indicating
measurements of total dissolved solids, pH, and temperature of fish in the 8-d acclimation treatment experienced
Walker Lake surface water ranged from 17,344 to 17,854 mg/L, 0.3 times the hazard of death suffered by unacclimated
from 9.56 to 9.70, and from 7.2 to 9.1uC, respectively. fish with the same fork length. For quantitative
covariates (e.g., xiFL) subtracting 1 from the hazard
supplied with 76 fish from tank 2. This process was ratio yielded the estimated portion by which the hazard
repeated until fish were transferred from all of the of death changes per unit increased in the covariate.
acclimation tanks into lake water in adjacent tanks. The The direction of change is indicated by the sign of
number of fish per tank and tank volume were chosen to the parameter estimate (Allison 1995). For example, if
match a preliminary study performed in static hatchery b̂FL = 20.105, then the hazard ratio (e20.105) is 0.90
water, in which total ammonia (as nitrogen) concentra- indicating each millimeter increase in fork length
tions in three tanks remained less than 0.2 mg/L for 1 wk. decreases the hazard of death by a portion of 0.10 (i.e.,
Fish not selected for the challenge were removed from 0.90–1.00).
the tanks and discarded. Temperature, pH, and EC The Cox proportional hazards model was performed
measurements continued at the same frequency as using SAS 9.2 PROC PHREG (SAS Institute, Cary, NC).
during acclimation. Mortalities were removed and Significance was set at a , 0.05. Pair-wise comparisons
counted at 2-h intervals. Fork lengths of all fish were between tanks within treatments were performed using
measured upon removal from the tanks. the PROC PHREG TEST function in SAS 9.2 as in Allison
Figure 2. Survival, stratified by duration of acclimation, of subyearling Lahontan cutthroat trout raised at Lahontan National Fish
Hatchery by hours after commencement of a week-long challenge with saline water from Walker Lake, Nevada, during
December 2008.
(1995). Tanks with dissimilar survival functions within resulting increase in TDS (from ,300 to .10,000 mg/L)
treatments were omitted from the final model. One and pH (from ,7.6 to .9.5), including surface swimming
omitted tank (under the 0-d acclimation treatment) with their heads out of water, loss of equilibrium, air-
exhibited a higher mortality rate than the other control gulping, flared opercles, and sporadic bursts of rapid
tanks apparently due to a water leak. The remaining swimming resulting in collisions with the tank walls.
omitted tanks (one under the 3-d acclimation treatment These fish died during acclimation. Percent survival
and two under the 8-d acclimation treatment) exhibited during acclimation (i.e., before the lake-water challenge)
lower mortality rates than the other tanks in their was not independent of treatment (mean 6 SE: 0-d =
respective treatments, apparently because snow inunda- 98.9 6 1.13%, 3-d = 75.8 6 11.3%, and 8-d = 96.4 6
tion reduced their salinities after their covers were 0.45%; x2 = 266.288; P , 0.001).
shifted by the wind. All omitted tanks would have No fish survived the entire week-long challenge,
favored the alternative hypothesis; therefore, their during which mean salinity ranged from 17,459 to
omission was conservative and slightly increased the 17,819 mg/L TDS, and mean alkalinity ranged from 9.59
hazard ratios and P-values associated with the 3- and 8-d to 9.67 pH across treatments. Median challenge survival
acclimation treatments. time for the 0-, 3-, and 8-d acclimation treatments was 8,
8, and 12 h, respectively. Maximum challenge survival
Results time for the 0-, 3-, and 8-d acclimation treatments was
20, 36, and 110 h, respectively. Survival curves during the
We investigated the effectiveness of two simple challenge, stratified by acclimation treatment, reflect a
acclimation protocols for 8-month-old Lahontan cut- moderately beneficial effect of acclimation (Figure 2).
throat trout reared at Lahontan National Fish Hatchery in Fish acclimated for 3 and 8 d experienced significantly
terms of reducing the hazard of mortality, compared to reduced hazard of death compared to unacclimated fish
that of unacclimated fish, during a week-long challenge (P , 0.0001; Table 1). Longer FL was associated with
in water from saline, alkaline Walker Lake, Nevada, while lower hazard of death during the challenge at an
controlling for differences in fork length. Water temper- estimated rate of 20.027/mm increase in FL (P ,
ature decreased from 11.0uC to 2.5uC prior to the lake 0.0001; Table 1). A few of the mortalities appeared to
water challenge due to cold ambient temperatures. The be precocious males; however, their incidence did not
temperature decrease was more rapid in the 0- and 3-d appear to be associated with treatment. Our results
acclimation treatments than in the 8-d acclimation indicate the methods of acclimation used in this study
treatment (Figure 1). Total ammonia was less than will not produce acceptable survival rates for Lahontan
0.2 mg/L in all tanks 24 h after the addition of fish. cutthroat trout stocked in Walker Lake; however, the
Several fish in the 3-d acclimation treatment tanks apparent beneficial effect of the 8-d acclimation treat-
exhibited aberrant behavior soon after the replacement ment indicates the possibility of achieving higher survival
of 50% of the hatchery water with lake water and the rates pending improvements to the acclimation method.
Table 1. Cox regression proportional hazards model of results suggest the increase in TDS, pH, and/or the
survival of subyearling Lahontan cutthroat trout raised at the decrease in temperature was too rapid in the 3-d
Lahontan National Fish Hatchery; acclimated for 0, 3, or 8 d; acclimation treatment. Future investigations should
and challenged with saline water from Walker Lake, Nevada, include comparisons of different rates and patterns of
during December 2008. TDS and pH increases while acclimation duration and
Parameter Hazard temperature are held constant, as well as comparisons of
Effect estimate ratio P different acclimation durations, to help determine the
Acclimation duration (versus 0 d [N = 228])
most effective acclimation protocol.
Ammonia measurements taken during acclimation
3-d acclimation (N = 228) 20.401 0.670 ,0.0001 and the challenge were spoiled by sulfide interference
8-d acclimation (N = 152) 21.503 0.223 ,0.0001 and high pH; however, the higher mortality rate and
Fork length (mm) 20.027 0.973 ,0.0001 behavioral symptoms exhibited by the 3-d acclimation
treatment during acclimation suggest ammonia concen-
tration in the water collected from Walker Lake for the 3-
Discussion d acclimation treatment may have been higher than in
Although all fish ultimately died before the end of the the lake water collected for the 8-d treatment. Acute
lake water challenge, our results indicate a moderately exposure to ammonia can result in ionoregulatory
positive effect of acclimation and fish size on time to impairment, leading to reduced Na+ influx and K+
death of Lahontan cutthroat trout challenged with water effusion in fish (Randall and Tsui 2002). The toxicity of
from Walker Lake. The benefit of the 3-d acclimation total ammonia increases exponentially with alkalinity
treatment was offset by mortality during acclimation; largely due to dissociation of ionized ammonia (NH4+) to
however, the 8-d acclimation treatment appeared to the un-ionized form (NH3; Roberts 2001). Un-ionized
impart an overall advantage. Dickerson and Vinyard ammonia can attack the central nervous system by
(1999) also observed increased survival of Lahontan crossing and modifying the blood–brain barrier and
cutthroat trout challenged with Walker Lake water reducing cerebral adenosine triphosphate, resulting in
associated with acclimation and size when TDS concen- hyperventilation, hyperexcitability, coma, convulsions,
trations were lower; however, unlike our results, 3-d and death (Randall and Tsui 2002). Ammonia levels in
acclimation produced higher survival rates than longer the lake and tanks should be accurately monitored, using
(6-d) acclimation. Our results suggest the acclimation methods effective in seawater (e.g., USEPA 1997)
method might be improved by the use of longer or older throughout future studies to determine the relative
fish, longer acclimation, and better control of the water contribution of fish excretions and lake water to
temperature, ammonia concentration, and alkalinity. ammonia concentrations. In static tanks, ammonia could
Tank water temperatures decreased at a faster rate be controlled by adding ChorAm-XH (Reed Mariculture
during acclimation in the 3-d acclimation treatment than Inc., Campbell, CA). The use of flow-through tanks would
in the 8-d acclimation treatment (Figure 1). Rapidly facilitate maintenance of challenge water that is more
decreasing temperature may have caused the higher closely representative of lake ammonia concentrations
mortality rates in the 3-d acclimation treatment or may and would improve the inference of challenge survival
have acted in conjunction with increasing TDS and pH. rates to rates achievable in the lake.
Temperature fluctuated between 5.0uC and 2.3uC during Surface ammonia concentrations in Walker Lake
the challenge, but was similar across treatments. Surface fluctuated spatially and temporally during the winter
temperatures in Walker Lake ranged from 7.6uC to 8.9uC stocking seasons of 1993, 1995, and 1996, falling within
during the challenge, and they do not commonly the range of 0.000 to 0.250 mg/L (Beutel et al. 2001).
decrease to less than 6.0uC during February and March Ammonia concentrations in Walker Lake are likely to
(Beutel et al. 2001) when hatchery fish releases occur. increase with lake desiccation because the ratio of
Temperature affects the ability of salmonids to acclimate hypolimnetic volume to hypolimnetic oxygen demand
to seawater (Saunders et al. 1975; Alexis et al. 1984; decreases with reduced lake depth, resulting in longer
Virtanen and Oikari 1984; Staurnes et al. 2001) and may periods of hypolimnetic anoxia during lake stratification
have decreased survival rates in our study. The proposed (Beutel 2001). In laboratory exposures, the acute
acclimation facility should employ more effective means tolerance limit for total ammonia decreased from
of temperature control. Static tanks should be housed 181 mg/L at pH 6.51 to 2.53 mg/L at pH 9.01 in rainbow
indoors. The use of flow through tanks for acclimation trout (O. mykiss; Thurston et al. 1981). Acute ammonia
would probably provide sufficient temperature control in tolerances in westslope cutthroat trout at pH 8.0 were
outdoor, insulated tanks. similar to rainbow trout (Thurston et al. 1978). We
Many fish (24.2%) in the 3-d acclimation treatment recommend the acute toxicity of ammonia be deter-
exhibited aberrant behavior indicative of ammonia mined for Lahontan cutthroat trout in the saline, alkaline
toxicity (Post 1987) and mortality soon after initial waters of Walker Lake so appropriate measures for
exposure to 10,000 mg/L TDS, 9.68 pH, and rapidly ammonia control can be implemented during acclima-
decreasing temperature; however, fish in the 8-d tion, and stocking can be avoided when lake ammonia
acclimation treatment tolerated TDS .14,000 mg/L, pH concentrations are too high.
.9.6, and gradually decreasing temperature for 6 d Our acclimation method produced a sudden, large
during acclimation without apparent difficulty. These increase in pH after the addition of only small amounts of
lake water to the hatchery water because of the large temperature can be controlled easily. Use of flow-
buffering capacity of Walker Lake water (Figure 1). through tanks at the lake side would allow more fish
Lahontan cutthroat trout challenged with water from to be held in the tanks during acclimation and easier
saline, alkaline Pyramid Lake appear to adapt quickly to control of ammonia concentrations (assuming ammonia
increased pH by quickly increasing the number and size concentration in Walker Lake is not too high), and would
of chloride cells in their gills, where chloride anions may facilitate inference of challenge results to the lake by
be exchanged for HCO32 (mitigated by ion uptake in the eliminating effects of fish excretions and changes in
gills) to maintain lower internal pH than the alkaline water quality resulting from stagnation.
waters (Wilkie et al. 1994). Acid–base regulation is The salinity of Walker Lake (.17,400 mg/L) currently
important in maintaining safe internal ammonia concen- exceeds the salinity found by Dickerson and Vinyard (1999)
trations. Lahontan cutthroat trout exposed to Pyramid to cause complete mortality in acclimated Lahontan
Lake water exhibit chronic elevated plasma pH and cutthroat trout (i.e., 15,467 mg/L). They concluded:
ammonia concentrations, which maintain a partial ‘‘Without adequate inflow of freshwater into the lake, we
pressure NH3 gradient important for the passive bran- estimate that 100% mortality of Lahontan cutthroat trout
chial diffusion of ammonia from their blood to the water stocked into Walker Lake within 48 h of stocking can be
(Wright et al. 1993). A gradual increase in pH resulted in expected within 20 years’’ (p. 514). Our study presents no
higher survival rates than sudden pH increases in evidence with which to refute the conclusions of Dickerson
hatchery-reared rainbow trout (Murray and Ziebell and Vinyard (1999). Our results indicate stocking of
1984). Our acclimation method may therefore be Lahontan cutthroat trout into Walker Lake has become
improved by creating a gradual increase in pH, allowing at best problematic and should not be continued until
time for Lahontan cutthroat trout to increase the size effective acclimation methods are developed or salinity
and number of their branchial chloride cells. A gradual decreases. While studies such as ours can address short-
increase in pH could be accomplished in static tanks by term survival rates, the determination of long-term survival
initially adding minute amounts of lake water. This initial rates through mark and recapture studies would provide a
pH adjustment period could then be followed by a TDS more complete assessment of the stocking program.
acclimation period during which lake water could be
added at a faster rate. A simultaneous, gradual increase Acknowledgments
of pH and TDS could be achieved in static water by using
We thank Sarah Bigelow, Derek Bloomquist, Alvin
simulated lake water for acclimation. The use of
Duncan, Scott Foott, James Hoang, Erik Horgen, Art
simulated lake water would also allow acclimation to
and Barbara Jones, Ed Kelly, Tim Loux, Corene Luton,
be performed at the hatchery without the risk of fish
Roger Peka, Kris Urquhart, Cassidy Williams, and Karie
pathogen transfer from Walker Lake, facilitating temper-
Wright for their assistance with this study. We also wish
ature control without the need for a flow-through
to thank the Journal of Fish and Wildlife Management
system. The effect of alkalinity should be examined
reviewers and Subject Editor for improving this article.
separately from salinity to help determine if the increase
This research was supported by the U.S. Fish and Wildlife
of pH and TDS should be performed separately or
Service Lahontan National Fish Hatchery Complex
simultaneously during acclimation.
Fisheries Improvement Program.
The use of longer or older fish acclimated for an
extended period of time should be investigated;
References
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Articles
Characteristics of Ponds Used by Breeding Mottled
Ducks on the Chenier Plain of the Texas Gulf Coast
David Haukos,* Stephanie Martinez, Jeannie Heltzel
D. Haukos
U.S. Fish and Wildlife Service, Texas Tech University, MS 2125, Lubbock, Texas 79409-2125
S. Martinez, J. Heltzel
U.S. Fish and Wildlife Service, Texas Chenier Plain National Wildlife Refuge Complex, P.O. Box 278, Anahuac, Texas 77514
Abstract
Mottled ducks Anas fulvigula are nonmigratory and primarily depend on habitats associated with coastal marshes of
the Gulf Coast for their entire life cycle. Much of remaining available coastal marsh habitats of the Texas Gulf Coast are
managed by state and federal agencies. The Texas Gulf Coast breeding population of mottled ducks has recently
declined dramatically. To aid in development of management plans for breeding mottled ducks, we estimated the
number of potential pair ponds available on the Texas Chenier Plain National Wildlife Refuge Complex relative to
marsh type (fresh, intermediate, brackish, and saline), estimated occupancy, compared habitats of used and unused
pair ponds, and evaluated pair pond selection relative to marsh type during spring 2004 and 2005. Greater than 18,000
ponds were recorded in .22,400 ha of coastal marsh with .74% of potential pair ponds in the intermediate marsh
type. Average pond size of 425 randomly selected ponds across marsh types was 0.09 ha. Pond occupancy was
estimated as 1.3% and 2.5% in 2004 and 2005, respectively, in 634 ponds surveyed for presence of mottled duck pairs.
Mottled ducks selected for ponds in fresh marsh with short surrounding vegetation associated with recent cattle
grazing, but avoided recently (,3 mo prior) burned areas. Pond area was greater with shallower water depth for used
compared to unused ponds. The presence and coverage of submergent and emergent vegetation did not influence
use of pair ponds by mottled ducks. Development of breeding habitat (pair ponds, nesting cover, brood habitat) in
fresh marsh will benefit mottled ducks. Our results indicate development of breeding habitat on the Texas Gulf Coast,
and vegetation management using cattle grazing timed for disturbance in fresh marsh during mid- to late winter and
early spring will further attract breeding mottled ducks.
Keywords: Anas fulvigula; Chenier Plain; mottled duck; National Wildlife Refuge; pair pond; Texas; waterfowl
Received: March 7, 2010; Accepted: September 23, 2010; Published Online Early: September 2010; Published:
November 2010
Citation: Haukos D, Martinez S, Heltzel J. 2010. Characteristics of ponds used by breeding mottled ducks on the Chenier
Plain of the Texas Gulf Coast. Journal of Fish and Wildlife Management 1(2):93–101; e1944-687X. doi: 10.3996/032010-
JFWM-005
* Corresponding author: david.haukos@ttu.edu
Introduction coastal marsh habitats critical to the life history of

mottled ducks (Gulf Coast Joint Venture, unpublished
Mottled ducks Anas fulvigula are nonmigratory water- data). Much of the remaining nonpublicly owned coastal
fowl principally dependent upon coastal marshes of the marsh habitats are being subjected to a variety of
Gulf Coast for their entire life cycle (Stutzenbaker 1988; anthropogenic impacts (e.g., saltwater intrusion, oil and
Moorman and Gray 1994). On the Texas Gulf Coast, gas exploration and processing, urbanization, and
managers of federal National Wildlife Refuges (NWRs) reduced freshwater inflows), most of which result in
and state Wildlife Management Areas conserve and continued declines in habitat quality for mottled ducks
manage approximately 25% and 43%, respectively, of the (Stutzenbaker 1988; Moorman and Gray 1994). The
available palustrine emergent and estuarine emergent recent oil spill resulting from the failure of the Deepwater
Mottled Duck Pair Ponds D. Haukos et al.
Horizon well highlights the fragility of coastal marsh and

potential for devastating consequences on the marsh
ecosystem. Therefore, mottled ducks in Texas will
increasingly depend upon wetland and associated
habitats provided and protected by NWRs and Wildlife
Management Areas (Stutzenbaker 1988).
There is not currently an operational annual range-
wide survey of breeding mottled ducks as there are for
other North American waterfowl species. The only
continuous survey (since 1986) of breeding mottled
ducks for the Western Gulf Coast Population is on NWRs
of the mid and upper Texas Gulf Coast. The Chenier Plain
NWR Complex (Complex) on the upper Texas Gulf Coast
(the region east of Houston, Texas, to the Louisiana
border) historically accounted for .50% of the mottled
ducks counted on Texas coastal NWRs (U.S. Fish and Figure 1. Estimated visibility-corrected density of breeding
Wildlife Service [USFWS], unpublished data). Estimated mottled ducks on five National Wildlife Refuges (Anahuac, San
indicated breeding pairs, based on visually corrected Bernard, Brazoria, Matagorda Island, Big Boggy) of the mid and
aerial surveys on NWRs, have declined 94% since 1986 upper Texas Gulf Coast from 1985 to 2008. The long-term linear
despite a relatively large increase in the mid-1990s and trend indicates a significant decline since the late 1980s.
remained at relatively low levels throughout the 2000s
(Figure 1). Although these trends were documented on Mottled ducks initiate breeding activities much earlier
NWRs of the upper Texas Gulf Coast, other indices reveal than other North American duck species because they
recent declines of varying magnitude for mottled ducks are nonmigratory and occupy southerly habitats. Paulus
throughout Texas (GCJV 2007). Johnson (2009) conclud- (1988) reported that 90% of females were paired by
ed that the trends represented by the breeding pair December and 100% paired by February with the peak of
survey on NWRs best corresponded with the trends of nest initiation in Texas and Louisiana during March and
estimated mottled duck population finite growth rate on April. Territorial defense typically starts in January, with
the western Gulf Coast compared to other population the peak of three-bird flights (territorial resident males
surveys. Numerous hypotheses have been advanced as chasing intruding pairs) occurring in March and April
contributing to the observed decline in mottled ducks on corresponding to the peak of the breeding season
NWRs and in Texas (GCJV 2007). These include 1) loss or (Paulus 1988; Stutzenbaker 1988). Moorman and Gray
degradation of reproductive habitats (e.g., pair ponds, (1994) indicated that mated pairs usually feed together
suitable nesting cover, and brood-rearing habitats), 2) in defended pair ponds prior to and during egg laying
increasing predator populations (Elsey et al. 2004), 3) loss until pair bonds break during incubation. Spacing of
and degradation of nonbreeding habitats (i.e., winter, paired mottled ducks in territorial ponds indicates
molt), 4) reduction of ecological disturbance (e.g., cattle foraging areas used by the pair during the breeding
and goose grazing or prescribed burning), 5) redistribu- season (Paulus 1984), but rarely includes a nest site (Allen
tion of populations (i.e., from Texas to Louisiana), and 6) 1980; Baker 1983; Paulus 1984). Therefore, investigation
continued exposure to lead from spent shot shells of defended pair ponds by mottled ducks allows for
(Merendino et al. 2005). Concern for the status and evaluation of habitat characteristics that initially attract
future of mottled ducks has resulted in the USFWS breeding pairs for reproductive activities.
declaring the species a focal species in the Migratory Bird From the perspective of a coastal marsh manager,
Program Focal Species Strategy (USFWS 2005). habitat management for breeding mottled ducks may
To attract mottled ducks for breeding and subsequent conflict with habitat management for wintering water-
recruitment on NWRs, managers should provide pairing, fowl. For example, managing for waterfowl preparing to
nesting, and brood-rearing habitats. During the early migrate usually involves providing high-energy foods
portion of the breeding season through mid-incubation, (e.g., seeds and tubers) for large flocks, whereas
most duck species are territorial with females selecting management for breeding mottled ducks focuses on
pair loafing areas in wetlands, usually referred to as pair provision of invertebrates rich in protein and minerals
ponds, with spacing influenced by food distribution, necessary for reproduction in isolated areas that provide
location of nesting cover, and predator populations nesting and brood-rearing habitat. In addition, common
(Anderson and Titman 1992). Pair ponds are defended as vegetation management practices (e.g., livestock grazing
territories because they provide access to energy and and prescribed fire) on NWRs may influence habitat use
nutrition sources necessary for reproduction, ultimately by breeding mottled ducks. Thus, thorough knowledge
influencing potential recruitment (Stutzenbaker 1988; of waterfowl–habitat interactions within coastal marsh
Anderson and Titman 1992). Unfortunately, little is habitats is essential to the development of integrated
known about the habitat characteristics of ponds that management practices for all species. We investigated
attract paired mottled ducks for breeding efforts, which the use of pair ponds by breeding mottled ducks on the
is information NWR managers need to increase recruit- Chenier Plain NWR Complex on the upper Texas Gulf
ment of mottled ducks. Coast. Our objectives were to 1) determine the number
of potentially available pair ponds on the Complex, 2) foot from 3 March to 9 April 2004, and 25 February to 6
estimate the percentage of available ponds used by April 2005, from sunrise to 3 h after sunrise or from 3 h
mottled ducks, 3) evaluate selection of ponds based on before sunset to sunset to determine the presence or
habitat attributes, and 4) contrast habitat characteristics absence of a breeding mottled duck pair. These dates
between used and unused ponds. and time periods correspond to presence of feeding by
pairs in selected territorial pair or loafing ponds (Paulus
Methods 1984; Stutzenbaker 1988). We used the area of each
marsh type on each refuge to stratify pond sampling
The Chenier Plain NWR Complex is located along the across marsh types and individual refuges. Our objective
Texas Gulf Coast between the city of Houston and state was to sample 100 fresh, 150 intermediate, 125 brackish,
of Louisiana and comprised of Anahuac, McFaddin, Texas and 100 saline marsh potential pair ponds for occupancy
Point, and Moody NWRs. The total acquired area of the across the entire Complex each year of the study (2004,
Complex was 42,762 ha; 56%, 32%, 8%, and 4% on 2005). We downloaded coordinates of each randomly
McFaddin, Anahuac, Texas Point, and Moody NWRs, selected pond (ponds were randomly selected each year)
respectively (USFWS 2006). We did not sample pair into a Global Positioning System (Garmin 76S7), which
ponds on Moody NWR for this stud because of poor we used to locate each pond (accuracy 6 3 m) in the
access. Mottled ducks are considered a priority species in field.
the Comprehensive Conservation Plan of the Complex If mottled ducks were flushed from a randomly
(USFWS 2006). selected pond, we recorded whether it was a pair or a
We used 2002 aerial photography digitized in ArcMap lone bird. We never flushed greater than two birds from
(Environmental Systems Research Institute, Redlands, CA) a selected pond. When an indicated breeding pair (pair
and field measurements of salinity gradients during that or lone male) was recorded on a random pond, the pond
same year to characterize availability of ponds in the was designated as a ‘‘random/used’’ pond, and if no
coastal marsh. We recognize that salinity gradients may birds were present, the pond was designated as
have shifted between 2002 and initiation of the study, ‘‘random/unused.’’ If a marked pond no longer existed
but all field salinity measurements during the study (i.e., was overgrown with cattail Typha spp., dried up,
corresponded to the initial 2002 classification of marsh etc.), we randomly selected another pond from those
type. To determine the number of ponds available for available within the proper stratification.
breeding mottled ducks, we used ArcMap to count all At each randomly selected pond, we measured water
open water bodies (i.e., ponds) that were $9.29 m2 in salinity, water depth, and pond area; evaluated height of
the four different coastal marsh types (fresh [0–0.5 parts surrounding vegetation; estimated the percent coverage
per thousand {ppt} salinity], intermediate [0.5–3.5 ppt], of emergent vegetation; and determined the presence or
brackish [3.5–10 ppt], and saline [.10 ppt]) on the absence of submergent plant communities to character-
Complex. Bodies of water in prairies, ditches, canals, ize general habitats the ponds provided. The presence of
bayous, and manmade ponds or reservoirs were recently (within previous 3 mo) burned or grazed
excluded from this study except when water levels of vegetation surrounding each pond was also recorded
canals or ditches exceeded banks enough to be 9.29 m2 and verified with prescribed-fire or cattle-grazing re-
in area in adjacent coastal marsh. When a pond appeared cords. Vegetation structure surrounding the pond was
to be divided by vegetation, each division was marked as categorized by estimating the height of the dominant
an individual pond when sites were visually isolated vegetation. Categories of surrounding vegetation were
(vegetation was $1 m tall and at $75% of the separated ‘‘short’’ (,0.6 m; e.g., Paspalum vaginatum, Distichlis
surface area was bordered by vegetation). For large spicata, and Sagittaria spp.), ‘‘medium’’ (0.6–1.3 m; e.g.,
marsh areas of intermixed water and vegetation without Spartina patens, Schoenoplectus americanus, and Schoe-
obvious borders, individual ponds were delineated based noplectus robustus), and ‘‘tall’’ (.1.3 m; e.g., Typha spp.,
on visually defined isolated areas of open water. Because Phragmites australis, and Schoenoplectus californicus).
there were .18,000 marked as potentially available, we We measured salinity by submersing a salinity meter
measured the area of 425 randomly selected ponds (YSI, Models 30 and 63) to the bottom of the pond in
stratified by marsh type (2% of total ponds), calculated undisturbed areas and recording two to three readings
the average area by marsh type, and multiplied those (ppt) in different areas of each pond. If there was a
averages by the number of marked ponds per marsh difference .1.0 ppt among readings, we noted the high,
type for each refuge to estimate total surface area of low, and average values. Otherwise, we only recorded
identified ponds. the average. We measured water depth (mm) at three
Once locations of all potential ponds were determined, random locations with a meter stick. The stick was held
we assigned a number to each pond with ArcMap, and vertically in the water and allowed to fall to the bottom
then used a random number generator to randomly of the pond using its own weight. If there was a
choose numbers and corresponding ponds stratified by difference .100 mm in the depth readings, we recorded
marsh type and refuge. To estimate the percentage of the high, low, and average measurements. To estimate
available ponds breeding mottled ducks used, we the surface area for small ponds (,30 m2), we used a
randomly chose ponds in each marsh type to determine meter stick or tape measure to physically measure pond
pond occupancy by pairs. Each randomly selected pond perimeter and then surface area was calculated; for
was visited once by airboat, truck, all-terrain vehicle, or medium-sized ponds (30–2,000 m2), the pond edge was
Table 1. Total area (ha) of coastal marsh categorized by marsh type available in 2002 (Available area), percent total area
represented by marsh type (%), and area sampled for presence of ponds used by breeding pairs of mottled ducks Anas fulvigula
(Sampled area) in 2004 and 2005 on Anahuac, McFaddin, and Texas Point National Wildlife Refuges comprising the Chenier Plain
National Wildlife Refuge Complex (Complex) of the upper Texas Gulf Coast.
Anahuac McFaddin Texas Point Complex
Available Sampled Available Sampled Available Sampled Available Sampled
Marsh typea area (%) area area (%) area area (%) area area (%) area
Fresh 472 (5.5) 472 2,608 (11.0) 453 0 (0.0) 0 3,081 (8.6) 925
Intermediate 5,892 (68.6) 5,892 19,762 (83.4) 10,258 552 (16.0) 551 26,204 (73.3) 16,701
Brackish 1,942 (22.6) 1,942 1,333 (5.6) 1,333 930 (27.0) 877 4,206 (11.8) 4,152
Saline 278 (3.2) 278 0 (0.0) 0 1,968 (57.0) 398 2,247 (6.3) 676
Total 8,584 8,584 23,703 12,044 3,450 1,826 35,738 22,454
a
Fresh = 0–0.5 parts per thousand (ppt); intermediate = 0.5–3.5 ppt; brackish = 3.5–10 ppt; and saline = .10 ppt.
walked using a Global Positioning System to map the ANOVA, a least significant difference test was used to
perimeter, then the area was calculated using ArcMap; compare means among marsh types.
and for large ponds (.2,000 m2), an airboat was driven
around the pond and a Global Positioning System was Results
used to map the pond perimeter, then pond area was
calculated using ArcMap. In 2002, the total area of coastal marsh on the Chenier
Only a small proportion of random ponds were used Plain NWR Complex was estimated to be 35,738 ha
by breeding mottled ducks. During the study, we (Table 1). Intermediate marsh represented the highest
recorded locations of all ponds used by pairs found percentage of marsh type available on the Complex,
during daily refuge management activities (i.e., estab- followed by brackish, fresh, and saline. Of the total marsh
lished salinity transects across all units and habitat types area, we sampled 22,454 ha for pair ponds, which
of each refuge), systematic evaluation of habitat represented 63% of the total marsh available on the
condition of the refuges using air boats throughout the Complex (Table 1). The sampled area excluded inacces-
breeding season, and while traveling to and from sible areas of intermediate and fresh marsh of McFaddin
sampled random ponds on each refuge. Pair ponds used NWR and portions of saline and brackish marsh of Texas
by breeding mottled ducks during the same daily time Point NWR. Sampled marsh area was representative of
periods as for random pond visits were considered used the entire Complex; intermediate marsh represented the
ponds and combined with the randomly used ponds to highest percentage (74%), followed by brackish (18%),
constitute the used pond data set. These ponds were fresh (4%) and saline (3%; Table 1).
revisited and habitat data were collected in the same We identified 18,830 ponds in the sampled area of the
manner as at random ponds. four marsh types on the Complex; the area of these
ponds constituted 7.4% of the available sampled marsh
Statistical analyses area on the Complex (Table 2). Anahuac NWR had the
We evaluated habitat selection based on the Neu et al. greatest number of ponds in fresh, brackish, and saline
(1974) method, which is appropriate for these data marsh types, whereas McFaddin NWR had the greatest
(Alldredge and Griswold 2006). This approach compares number of ponds in the intermediate marsh type.
the 95% confidence interval associated with the propor- Overall, McFaddin NWR had the greatest number of
tion of locations among habitat types with the propor- identified ponds by a small margin (Table 2). Ranking, by
tional availability each habitat type within the study area. marsh type, of pond numbers and area was intermediate,
We compared use versus availability of ponds among brackish, saline, and fresh (Table 2). Average (SE) pond
marsh type (fresh, intermediate, brackish, and saline) and size, by marsh type, was 0.03 (0.01) ha in fresh marsh,
surrounding vegetation height (i.e., surrounding struc- 0.13 (0.06) ha in intermediate, 0.06 (0.02) ha in brackish,
ture; short, medium, and tall). We compared percent of and 0.06 (0.03) ha in saline. Across marsh types, average
used versus available ponds containing submergent and pond size was 0.09 (0.03) ha.
emergent vegetation. Finally, we assessed the influence In 2004 and 2005, we surveyed 634 randomly selected
of cattle grazing and prescribed fire disturbances on the ponds (317 ponds/y) or 3.4% of the 18,830 identified
proportion of ponds used associated with these events ponds. Because of logistical constraints in 2004, the
versus the proportion for all available ponds. original objective of visiting 425 randomly chosen ponds
We used a 2-way factorial analysis of variance (ANOVA) was not achieved. To ensure a consistent sampling effort
to compare measured habitat features of pond area (ha), between years, we visited the same number of ponds on
water depth (mm), water salinity (ppt), and pond each refuge and in each marsh type in both 2004 and
coverage by emergent vegetation (%) between used 2005. Of these, 8.8% occurred in fresh marsh, 45.7% in
and unused ponds and marsh type (fresh, intermediate, intermediate marsh, 38.8% in brackish marsh, and 6.6%
brackish, and saline). Following a significant (P , 0.05) in saline marsh (Table 2). Sampled ponds were distrib-
Table 2. Total area (ha) and number of available ponds (Available) and total area (ha) and number of randomly selected ponds
visited (Sampled) by marsh type to determine the pond occupancy of mottled ducks Anas fulvigula breeding on the upper Texas
Gulf Coast on Anahuac, McFaddin, and Texas Point National Wildlife Refuges comprising the Chenier Plain National Wildlife Refuge
Complex (Complex) in spring 2004 and 2005.
Anahuac McFaddin Texas Point Complex
Available Sampled Available Sampled Available Sampled Available Sampled
Marsh
typea Area No. Area No. Area No. Area No. Area No. Area No. Area No. Area No.
Fresh 4.5 114 1.3 42 3.8 97 1.0 14 0.0 0 0.0 0 8.3 211 2.3 56
Intermediate 431.0 4,589 6.5 108 595.1 6,636 30.2 172 69.7 742 0.6 10 1,095.8 11,667 37.3 290
Brackish 304.5 3,783 2.8 112 190.3 2,364 1.0 78 39.9 496 11.2 56 534.7 6,643 15.0 246
Saline 17.9 184 0.4 26 0.0 0 0.0 0 12.2 125 1.9 16 30.1 309 2.3 42
Total 757.9 8,670 11.0 288 789.2 9,097 32.2 264 121.8 1,363 13.7 82 1,668.9 18,830 56.9 634
a
uted across the Complex: 45% on Anahuac, 42% on recorded (37.9% of estimated total number of pairs); 17
McFaddin, and 13% on Texas Point NWRs. In 2004, a total pairs used habitats other than delineated ponds (e.g.,
pond area of 26.64 ha was sampled on the Complex, prairies, reservoirs, and bayous/ditches). Therefore, the
representing 1.6% of the potential pond area in the final 2004 data set was based on 76 ‘‘used’’ ponds (4
sampled portion of the Complex. In 2005, a total pond from random/used ponds and 72 other recorded used
area of 30.23 ha was sampled, representing 1.8% of the ponds) and 313 ‘‘unused’’ ponds. Of these 76 pairs, 54%
potential pond area on the Complex. occurred at Anahuac NWR, 30% at McFaddin NWR, and
Mottled duck pairs were present on four randomly 16% at Texas Point. In 2005, of the estimated 471 pairs of
selected ponds (1.3% occupancy) in 2004: one brackish mottled ducks occurring in the sampled area of the
pond with short surrounding vegetation, one intermedi- Complex, locations of 8 pairs on randomly searched
ate pond with short vegetation, one intermediate pond ponds and locations of an additional 127 pairs were
with medium vegetation, and one fresh pond with recorded during the search periods (28.7% of estimated
medium vegetation (Supplemental Material, Table S1, total number of pairs). Twenty-six pairs used habitats
http://dx.doi.org/10.3996/032010-JFWM-005.S1). In 2005, other than delineated ponds; thus, 101 ponds were
mottled ducks were present on eight randomly surveyed included in the data set for habitat use–availability
ponds (2.5% occupancy): three brackish ponds with short comparisons (8 from random/used ponds and 93 other
surrounding vegetation, two brackish ponds with recorded used ponds). Of these 101 pairs, 52% occurred
medium vegetation, one intermediate pond with short at Anahuac NWR, 36% at McFaddin NWR, and 12% at
vegetation, one intermediate pond with medium vege- Texas Point NWR.
tation, and one saline pond with medium vegetation. In In 2004, pairs of mottled ducks settled most frequently
2004, using the pond occupancy rate of 1.3%, we in intermediate marsh (65.8%), followed by brackish
estimated that 245 pairs of mottled ducks were in the (23.6%), fresh (7.9%), and saline (2.6%) marsh types. In
sampled area of the Complex. In 2005, based on the 2005, intermediate marsh was again used most frequent-
pond occupancy rate of 2.5%, we estimated that 471 ly (43.6%), followed by brackish (33.7%), fresh (13.9%),
pairs of mottled ducks were on the sampled area of the and saline (8.9%) marsh. However, pond occupancy was
Complex. not in proportion to pond availability within marsh type
In 2004, of the estimated 245 pairs of mottled ducks across years (x23 = 178.02, P , 0.001). Across both years,
using the sampled area of the Complex, assuming all pair mottled ducks selected ponds in fresh marsh (1.1%
sightings were independent due to establishment of availability versus 11.3% used), but used ponds in all
territories, locations of 4 pairs in random ponds and 89 other marsh types in proportion to availability (Table 3).
additional pairs observed during the search periods were Furthermore, selection of habitat structure surrounding
Table 3. Percent occurrence by breeding mottled ducks Anas fulvigula (Used) relative to availability of ponds (Available) by
coastal marsh type on the Chenier Plain National Wildlife Refuge Complex in spring 2004 and 2005.
Available Used
a
Marsh type n % n % 95% CI
Fresh 211 1.1 20 11.3 5.4–17.3
Intermediate 11,667 61.9 94 53.1 43.7–62.5
Brackish 6,643 35.3 52 29.4 20.8–37.5
Saline 309 1.6 11 6.2 1.6–10.8
Total 18,830 177
a
Table 4. Percent occurrence by breeding mottled ducks Anas fulvigula (Used) in coastal marsh ponds relative to availability of
ponds (Available) based on surrounding vegetation structure on the Chenier Plain National Wildlife Refuge Complex in spring 2004
and 2005.
Available Used
Vegetation structure n % n % 95% CI
Short (,0.6 m) 158 25.1 105 59.3 51.5–67.2
Medium (0.6–1.3 m) 375 59.6 57 32.2 24.7–39.7
Tall (.1.3) 96 15.3 5 2.8 0.2–5.5
used pair ponds was also found (x22 = 110.09, P , average water salinity between used and unused ponds
0.0001). Mottled duck pairs exhibited a preference for (F1,812 = .011, P = 0.74; Table 6).
ponds with short surrounding habitat structure across
both years (,0.6 m; 25.1% availability versus 59.3% Discussion
used). They avoided ponds with medium and tall
surrounding vegetation (Table 4). However, used ponds Although recognized as an important part of the
containing submergent (x21 = 1.13, P = 0.29; 25.9% reproductive activity of waterfowl (Anderson and Titman
availability versus 29.7% used [95% CI on use = 22.0– 1992) there have been few descriptions of habitats used
37.4]) and emergent vegetation (x21 = 3.30, P = 0.08; by paired mottled ducks during the nesting season
45.8% availability versus 54.5% used [95% CI on use = compared to most other species of ducks. Stutzenbaker
47.5–63.3]) were used in proportion to availability. (1988) reported that male mottled ducks defended ponds
The type of surrounding disturbance influenced pair formed by temporary open water sites in prairie habitats,
pond selection by mottled ducks. Selection for ponds especially those surrounded by sienna bean Sesbania
associated with cattle grazing was apparent (x21 = drummondii. However, these sites have become relatively
60.84, P , 0.001; 12.3% availability versus 31.5% used rare since the 1980s. Durham and Afton (2006) concluded
[95% CI on use = 23.7–37.4]). However, paired mottled that flooded rice fields were important loafing and
ducks avoided ponds that were associated with recent feeding habitat for mottled ducks nesting in agricultural
prescribed burns (x21 = 5.01, P = 0.02; 19.3% availability lands of former coastal prairie in Louisiana, especially
versus 10.4% used [95% CI on use = 5.3–15.5]). during drought. White and James (1978) reported that
There was no interaction between pond use categories mottled duck foraging habitat was typically ,30 cm in
and marsh type for any of the four measured habitat depth. Weeks (1969) reported that estimated loafing area
variables (pond area: F3,802 = 1.43, P = 0.23; water depth: size ranged from 10 to 130 ha, but was usually ,40 ha.
F3,810 = 0.36, P = 0.78; water salinity: F3,812 = 1.37, P = However, there are no detailed descriptions of habitats
0.25; or percent emergent coverage: F3,811 = 2.25, P = used by paired mottled ducks in coastal marsh habitats.
0.08). Pond area increased across habitat types from fresh The sheer number of potentially available ponds and
to saline marsh (F3,802 = 2.67, P = 0.04; Table 5) and was low occupancy rate (e.g., 0.8 ponds/ha and 1.9%,
nearly three times greater for used than unused ponds respectively, on the Chenier Plain NWR Complex) likely
(F1,802 = 5.95, P = 0.02; Table 6). Water depth of ponds leads to an assumption that these habitats are not
did not differ among marsh type (F3,810 = 1.12, P = 0.34; limiting to mottled duck populations in coastal marsh.
Table 5); however, used ponds had shallower water depth Such an assumption may hinder management for
than unused ponds (F1,810 = 6.70, P = 0.01; Table 6). breeding mottled ducks because it is quite apparent
Percent emergent vegetation was greater in fresh marsh that certain features of ponds attract paired mottled
compared to the other marsh types (F3,811 = 62.89, P , ducks. Therefore, managers should include consideration
0.001; Table 5). However, percent emergent vegetation of pair habitats, in addition to nesting and brood-rearing
did not differ between used and unused ponds (F1,811 = habitats, in management plans for breeding mottled
1.59, P = 0.21; Table 6). There was no difference in ducks in coastal marsh.
Table 5. Average mean and standard error (SE) surface area, water depth, and percent cover by emergent vegetation of ponds
sampled for the presence of breeding mottled ducks Anas fulvigula among four coastal marsh types on Anahuac, McFaddin, and
Texas Point National Wildlife Refuges of the upper Texas Gulf Coast during spring 2004 and 2005.
Surface area (ha) Water depth (mm) Emergent vegetation (%)
a
Marsh type x̄ SE x̄ SE x̄ SE
Fresh 0.02Ab 0.03 114.1A 14.9 51.6A 2.7
Intermediate 0.06A 0.01 134.9A 6.9 21.2B 1.3
Brackish 0.10B 0.02 126.6A 7.9 6.2C 1.4
Saline 0.15B 0.04 98.2A 18.6 7.7C 3.4
a
b
Means with the same letter do not differ (P . 0.05) among marsh type within each habitat variable.
Table 6. Average mean and standard error (SE) pond area, water depth, emergent vegetation, and water salinity (parts per
thousand [ppt]) for ponds used and unused by breeding mottled ducks Anas fulvigula on Anahuac, McFaddin, and Texas Point
National Wildlife Refuges of the upper Texas Gulf Coast during spring 2004 and 2005.
Habitat variable
Pond area (ha) Water depth (mm) Emergent vegetation (%) Salinity (ppt)
Usage x̄ SE x̄ SE x̄ SE x̄ SE
a a
Used 0.12 0.02 99.0 9.6 19.9 1.7 2.81 0.12
Unused 0.05 0.01 137.9 5.4 23.4 0.9 2.74 0.07
a
Means with the same letter differed (P , 0.05) between used and unused ponds.
Mottled ducks selected for ponds in freshwater marsh. increasing opportunity for grazing by snow geese Chen
Although ponds were used in proportion to availability caerulescens to replicate historical natural disturbance
in the other three marsh types based on habitat selection (USFWS 2006). Moderate grazing following burns in
ratios (Manly et al. 2002), mottled ducks were 11.7, 12.2, marshes also prolongs the availability of new grass
and 2.6 times more likely to select ponds available in shoots, a valuable food for snow geese (Gosselink et al.
fresh marsh than those in intermediate, brackish, and 1979). The current cattle-grazing management program
saline marsh types, respectively, on the Chenier Plain is quite flexible to take advantage of varying quantity
NWR complex. Considering the relatively low availability and quality of forage, availability of fresh groundwater to
of freshwater marsh on the Complex, such definitive provide watering sites within all marsh types, and unit-
selection ratios indicate that it is critical to concentrate specific management objectives (USFWS 2006). Grazing
management efforts for breeding mottled ducks on pressure and timing is used to create conditions that
freshwater marsh. allow for the germination, growth, and reproduction of
The relatively smaller selection ratio of saline marsh species adapted to frequent disturbance (i.e., early
was due to inclusion of Texas Point NWR, where saline successional sere) by alternation of species composition
marsh was the predominant marsh type. Excluding Texas of plant communities (Valentine 1961; USFWS 2006).
Point NWR data, the freshwater : saline marsh type Mottled duck pair avoidance of recently burned but not
selection ratio increased to 4.6. Use of saline habitats by grazed areas indicates that pond selection includes factors
breeding mottled ducks appears to be more than in addition to the structure of surrounding vegetation
expected considering the perceived negative factors because the distribution of height of surrounding
associated with breeding attempts in these habitats (e.g., vegetation was similar between burned and grazed ponds
Moorman et al. 1991). However, breeding pairs of (short grazed = 59.9% burned = 63.8%; medium grazed =
mottled ducks are frequently observed feeding in saline 34.3% burned = 31.2%; tall grazed = 5.8% burned =
marsh habitats, especially those habitats dominated by 5.0%). Apparently, the physical disturbance from cattle
Ruppia maritima. On the Chenier Plain NWR Complex, grazing is creating additional conditions that attract paired
saline marsh is frequently separated from adjacent mottled ducks. Comparisons of invertebrate populations
intermediate marsh by salt prairie ridges, facilitating between grazed and burned ponds may provide insight
easy movements among habitat types (P. Walther and M. into the attractiveness of grazing-disturbed habitats to
Whitbeck, USFWS, personal observation). Further evalu- paired mottled ducks.
ation is needed to determine resources provided by It is unlikely that pair-pond habitats would be used by
saline marsh habitats for breeding mottled ducks and female mottled ducks for brood rearing (Rigby 2008) or
whether females preferentially move to areas of fresher by birds during molt (Stutzenbaker 1988). Therefore, it is
marsh for nesting. recommended for managers of coastal marsh to provide
Structure of the vegetation community surrounding different habitats to attract and sustain breeding mottled
ponds had a greater influence on pond use than the ducks. However, mottled duck broods have greater
presence of submergent and emergent vegetation. survival in fresh compared to saline conditions, so
Mottled duck pairs selected for ponds that were management of breeding mottled ducks should concen-
surrounded by relatively short vegetation (,0.6m) and trate on fresh marsh (Moorman et al. 1991). Use of
habitats influenced by livestock grazing. These condi- vegetation disturbance, especially cattle grazing, is useful
tions are complementary, with 60% of all ponds with for management of mottled ducks in coastal marsh.
short surrounding vegetation being grazed, whereas Further, prescribed fire can be used to focus and
28% of the ponds that were not being grazed were distribute cattle grazing within fresher areas of coastal
represented by short surrounding vegetation. For used marsh to create pair-pond habitats attractive to mottled
ponds only, 73% of grazed ponds had short surrounding ducks. Grazing management is also used to create
vegetation. nesting cover desired by mottled ducks on the Chenier
Disturbance by grazing herbivores is a natural Plain NWR Complex (USFWS 2006). Our results indicate
component of the coastal marsh ecosystem (Bhattachar- development of adaptive grazing management plans in
jee et al. 2007). Management of the Chenier Plain NWR fresh and intermediate coastal marsh will be beneficial to
Complex includes prescribed burning, cattle grazing, and breeding mottled ducks.
Supplemental Material Johnson FA. 2009. Variation in population growth rates

of mottled ducks in Texas and Louisiana. Reston,
Please note: The Journal of Fish and Wildlife Management
Virginia: Administrative Report to U.S. Fish and Wildlife
is not responsible for the content or functionality of any
supplemental material. Queries should be directed to the Service. Available: http://www.fws.gov/migratorybirds/
corresponding author. NewReportsPublications/SpecialTopics/BySpecies/Johnson.
Mottled%20Ducks%20Admin%20Report.Final.pdf
Table S1. Mottled duck presence and habitat (September 2010).
measurements of coastal marsh ponds, 2004–2005. Manly BFJ, McDonald LL, Thomas DL, McDonald TL,
Found at DOI: 10.3996/032010-JFWM-005.S1 (251 KB XLS). Erickson WP. 2002. Resource selection by animals:
statistical design and analysis for field studies. 2nd
Acknowledgments edition. Dordrecht, The Netherlands: Kluwer Academic.
Merendino MT, Lobpries DS, Neaville JE, Ortego JD,
We greatly appreciate the assistance of biologists and
Johnson WP. 2005. Regional differences and long-
staff of the Chenier Plain National Wildlife Refuge
term trends in lead exposure in mottled ducks.
Complex during the study. In particular, we would like
to thank J. Neaville, P. Walther, M. Whitbeck, A. Loranger, Wildlife Society Bulletin 33:1002–1008.
and K. McDowell for their support. P. Walther, M. Moorman AM, Moorman TE, Baldassarre GA, and Richard
Whitbeck, J. Moon, J. Haskins, and M. Brasher reviewed DR. 1991. Effects of saline water on growth and
earlier drafts of the manuscript. Comments provided by survival of mottled duck ducklings in Louisiana.
W. Conway, an anonymous reviewer and the Subject Journal of Wildlife Management 55:471–476.
Editor helped to improve this manuscript. Moorman TE, Gray PN. 1994. Mottled duck. Account 81 in
Funding for the study was provided by the U.S. Fish Poole A, Gill F, editors. The birds of North America.
and Wildlife Service, Region 2, Migratory Bird Office and Philadelphia, Pennsylvania: The Academy of Natural
administered by the Texas Cooperative Fish and Wildlife Sciences, and Washington, D.C.: American Ornitholo-
Research Unit at Texas Tech University. gists’ Union.
Neu CW, Byers CR, Peek JM. 1974. A technique for
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Articles
Woodpecker Densities in the Big Woods of Arkansas
David G. Krementz,* Jason D. Luscier
D.G. Krementz
U.S. Geological Survey Arkansas Cooperative Fish & Wildlife Research Unit, Department of Biological Sciences,
University of Arkansas, Fayetteville, Arkansas 72701
J.D. Luscier
Department of Biological Sciences, University of Arkansas, Fayetteville, Arkansas 72701
Abstract
Sightings of the now-feared-extinct ivory-billed woodpecker Campephilus principalis in 2004 in the Big Woods of
Arkansas initiated a series of studies on how to best manage habitat for this endangered species as well as all
woodpeckers in the area. Previous work suggested that densities of other woodpeckers, particularly pileated
Dryocopus pileatus and red-bellied Melanerpes carolinus woodpeckers, might be useful in characterizing habitat use by
the ivory-billed woodpecker. We estimated densities of six woodpecker species in the Big Woods during the breeding
seasons of 2006 and 2007 and also during the winter season of 2007. Our estimated densities were as high as or higher
than previously published woodpecker density estimates for the Southeastern United States. Density estimates ranged
from 9.1 to 161.3 individuals/km2 across six woodpecker species. Our data suggest that the Big Woods of Arkansas is
attractive to all woodpeckers using the region, including ivory-billed woodpeckers.
Keywords: woodpeckers; density; Big Woods; Arkansas; bottomland hardwood forests
Received: March 23, 2010; Accepted: June 9, 2010; Published Online Early: June 2010; Published: November 2010
Citation: Krementz DJ, Luscier JD. 2010. Woodpecker densities in the Big Woods of Arkansas. Journal of Fish and Wildlife
Management 1(2):102–110; e1944-687X. doi: 10.3996/032010-JFWM-006
* Corresponding author: krementz@uark.edu
Introduction cies on how best to manage bottomland hardwoods in

the LMAV. In response to these concerns, the Lower
Bottomland hardwood forests of the Lower Mississippi Mississippi Valley Joint Venture (LMVJV) Forest Resource
Alluvial Valley (LMAV) have experienced habitat loss due Conservation Working Group developed a set of habitat
to fragmentation and changes in hydrology (Twedt and management guidelines and monitoring schemes for
Loesch 1999; King et al. 2005). However, the Big Woods forest-interior birds, emphasizing woodpeckers and with
(see study site description and Figure 1) of eastern a focus on the ivory-billed woodpecker (Wilson et al.
Arkansas has remained relatively intact and is one of the 2007). One of their main objectives was to estimate
largest stands of bottomland hardwoods in the South- densities of forest-interior birds, with an emphasis on
eastern United States (The Nature Conservancy 2009). woodpeckers.
Therefore, this component of the LMAV provides a In order to develop habitat guidelines for ivory-billed
valuable model system for evaluating current densities woodpeckers and woodpeckers in general, the LMVJV
and habitat use patterns of wildlife in this largely needs a better understanding of their habitat use.
fragmented ecosystem. The large-scale loss of bottom- Unfortunately, habitat needs of ivory-billed woodpeckers
land hardwood forests throughout the LMAV, in con- are not well-known and current recommendations are
junction with the development of a recovery plan for the primarily based on information collected approximately
ivory-billed woodpecker Campephilus principalis, sighted 65 y ago (Jackson 2002) by Tanner (1942). Under the
in 2004 in the Big Woods (Fitzpatrick et al. 2005) by the ivory-billed woodpecker Recovery Plan (USFWS 2006),
U.S. Fish and Wildlife Service (USFWS 2006), has raised the USFWS requested assumption-driven research re-
concerns by federal, state, and nongovernmental agen- garding ivory-billed woodpecker habitat use. Tanner’s
Woodpeckers in the Big Woods of Arkansas D.G. Krementz and J.D. Luscier
Figure 1. Public lands in the Big Woods of Arkansas. National Wildlife Refuge (NWR); Wildlife Management Area (WMA).
(1942) study, albeit old, provides the only baseline Q. lyrata, water hickory Carya aquatic, see forest type 2
information from which to make assumptions directing below). These habitat affinities probably met both
research and management for this species. Tanner (1942) foraging needs, and nesting or roosting tree require-
suggested that pileated woodpeckers Dryocopus pileatus ments. Whether any one, or some, combination of these
and red-bellied woodpeckers Melanerpes carolinus could variables met some limiting requirement is unknown, but
be used as surrogates to study ivory-billed woodpecker Tanner (1942) suggested that forage availability was an
habitat use. Tanner believed that densities of 3.9 pileated important determinant for the presence of ivory-billed
or 19 red-bellied woodpeckers/km2 would indicate woodpeckers in a particular woodland tract.
suitable ivory-billed woodpecker habitat, based on
densities of these species in the home range of 1 ivory-
billed woodpecker. Tanner (1942) also indicated that in Study Site
addition to abundances of pileated and red-bellied The Big Woods of Arkansas encompasses about
woodpeckers, abundances of other species of wood- 222,500 ha of the Cache, Arkansas, and White River
peckers could be indicative of quality ivory-billed floodplains and Bayou DeView in the Arkansas Delta
woodpecker habitat. Based on these hypotheses by (Figure 1). Our study areas included Sheffield Nelson
Tanner (1942), if the Big Woods can support at least 1 Dagmar, Trusten Holder, Rex Hancock Black Swamp,
ivory-billed woodpecker, then there should be at least Bayou Meto, Mike Freeze Wattensaw, and Henry Gray
3.9 pileated or 19 red-bellied woodpeckers/km2. Our Hurricane Lake Wildlife Management Areas of the
objective here was to estimate densities of six wood- Arkansas Game & Fish Commission, and Cache River
pecker species in the Big Woods of Arkansas to evaluate (CRNWR) and White River (WRNWR) National Wildlife
the suitability of the region for ivory-billed woodpeckers. Refuges. There were approximately 28,000 ha in the
Before we developed our sampling scheme, it was Wildlife Management Areas, 25,000 ha in CRNWR, and
necessary to understand the habitat use of ivory-billed 65,000 ha in WRNWR, hereafter ‘‘units.’’ This is the largest
woodpeckers. Based on historical accounts (Jackson corridor of bottomland hardwood forest remaining in the
2002), we know that ivory-billed woodpeckers had some LMAV north of Louisiana’s Atchafalaya Basin. The
habitat affinities including a selection for: 1) extensive bottomland hardwood forests that occur here include a
continuous forest areas, 2) very large trees, 3) continuous wide range of species and community types that can
supply of recently dead trees, 4) an open canopy, 5) tolerate inundation or soil saturation (Wharton et al.
certain tree species (e.g., sweetgum Liquidambar styraci- 1982), resulting in a complex mosaic of community types
flua, Nuttall oak Quercus texana, see forest type 3 below), reflecting differences in the alluvial and hydrologic
and 6) avoidance of other tree species (e.g., overcup oak environment (Smith and Klimas 2002). We recognized
five hydrologic forest types (Wilson et al. 2007): 1) swamp the bole of a mature tree (Barnes et al. 1998). The LMVJV
forests—bald cypress Taxodium distichum, bald cypress– calculated these habitat metrics at the transect or stand
water tupelo Nyssa aquatica; 2) wet bottomland forest— level (R. Wilson, USFWS, personal communication). Based
overcup oak–water hickory, black willow Salix nigra, on logistical constraints, we randomly chose one out of
laurel oak Q. laurifolia–red maple Acer rubrum; 3) moist the four transects in each stand. When possible, we used
bottomland forest—sugarberry Celtis laevigata, elm the center plot of a transect line as the woodpecker
Ulmus spp., ash Fraxinus spp., oak–elm–ash, oak–sweet- survey location. Upon reviewing the habitat metrics
gum; 4) dry bottomland forest—cherrybark oak Q. along transects selected, we found that some forest
pagoda, swamp chestnut oak Q. michauxii, post oak Q. types or habitat metrics that we were interested in
stellata, blackgum N. sylvatica; and 5) levee forest— surveying were poorly or not represented. We were
cottonwood Populus deltoides, sycamore Platanus occi- particularly concerned that swamp forest stands were
dentalis, pecan Carya illinoiensis, boxelder Acer negundo. not well-represented in our initial sample because all
Forest management across the study area is difficult to sightings of ivory-billed woodpeckers in the Big Woods
characterize because ownership spans three federal in 2004–2005 were in swamp forests (Fitzpatrick et al.
agencies, two state agencies, and numerous private 2005). To address this problem, we added four swamp
landowners. Over the past 10 y, about 50% of the Big forest stands to include this missing forest type.
Woods has been harvested using partial cuts with a We used a repeated-survey distance-sampling ap-
target of retaining larger, older age-class trees of various proach based on the recommendations of Wilson et al.
sizes (J. Denman, USFWS, personal communication). On (2007) to estimate densities of each woodpecker species.
federal properties, this objective has been pursued for We attempted to survey each plot five times during three
. 20 y (Denman and Karnuth 2005). Private landowners seasons: breeding season 2006 (2 May–17 June, n = 82
tend not to manage for older trees. The partial cuts, plots), winter 2007 (16 January–25 February, n = 63
where portions of the overstory are removed, vary from plots), and breeding season 2007 (25 May–1 July, n = 87
30 to 90% removal with an overall average of 50% plots). Due to logistical constraints, we surveyed a
overstory opening (Denman and Karnuth 2005). This different number of plots during each season. We did
management approach results in a multiple-canopied not survey many of our sites in WRNWR during summer
condition that is representative of the ‘‘Desired Forest 2006 due to lack of manpower and due to high flood
Conditions’’ of the LMVJV Forest Resource Conservation levels during winter 2007. We began surveys in the
Working Group (Wilson et al. 2007). About 10% of the Big morning when it was light enough to see 200 m and
Woods undergo forest management in any single year ended them no later than 1000 hours. Surveys were not
with private lands initiating more actions than on public conducted when wind speeds exceeded Beaufort scale
lands. No. 4 (i.e., 20–28 km/h) or during heavy rain. Any
woodpecker seen or heard during the 10-min period was
marked in one of four predetermined distance bands
Methods (0–25 m, 25–50 m, 50–100 m, and . 100 m). Because
different technicians were used each survey season, we
Survey design followed the advice of Robbins and Stallcup (1981) by: 1)
The LMVJV developed an ivory-billed woodpecker training each set of observers in advance with appropri-
habitat assessment scheme based on the above five ate books and sound recordings, 2) field training to
forest types, with the objective of quantifying current familiarize observers with field conditions and local
habitat conditions in the Big Woods of Arkansas (Wilson dialects, 3) examining field records of all observers for
et al. 2007, appendix 4). The sampling frame was comparability during early phases of each survey season,
centered on those areas with evidence of ivory-billed and 4) rotating observers among plots so that surveys
woodpecker existence with additional nearby areas would be as comparable as possible and to reduce the
added to increase the chance of observing ivory-billed possibility of overlooking or misidentifying species. Also,
woodpeckers. Individual units within the area of interest observers practiced estimating distances with laser
were broken down into stands of approximately 200 ha rangefinders. As a final precaution before collecting field
each. Within each stand (n = 146), an average of four data, we conducted simultaneous surveys with all
randomly allocated point-transects (n = 587) were technicians independently field-recording woodpecker
located (transect numbers ranged from two to five detections. Any problems with species identifications
based on the size of the stand). Along each transect, five were discussed at the end of these initial surveys during
equidistant plots (80 m apart) were established. At the training period to correct future identification errors
WRNWR only, the four transects were connected and (Kepler and Scott 1981).
nonrandomly placed due to logistical constraints. At
each of these plots (0.08 ha each, n = 2,935), we selected Analyses
our woodpecker survey plots across three strata pertain- We used the Multiple Covariate Distance Sampling
ing to tree density (mean trees/ha, mean trees . 61 cm analysis engine in program DISTANCE 5.0 (Thomas et al.
diameter at breast height/ha), tree health (mean snags/ 2005; Marques et al. 2007) to model detectability and
ha, mean epicormic branching score/transect), and estimate densities of each woodpecker species (individ-
predominant tree species per stand. Epicormic branching uals/km2) during the three seasons. There were not
is disturbance-induced development of new branches on enough detections to estimate stand- or unit-level
detection probabilities for any species. We estimated a

global detection function for each species per season
from all data pooled together, allowing for more precise
density estimates (Marques et al. 2007). We used these
global detection functions to estimate densities first at
the level of CRNWR, WRNWR, and Arkansas Game and
Fish Commission Wildlife Management Areas, and then
at the level of the Big Woods. We also modeled the
effective detection radius to assess whether the detect-
ability for a species was a function of habitat or area
surveyed (Buckland et al. 2001). A larger value can be
suggestive of a considerably larger area surveyed.
The Multiple Covariate Distance Sampling analysis
engine only allows hazard-rate and half-normal key
functions for modeling detectability. Our candidate set
of models for explaining detectability for each species
included these key functions with a cosine series Figure 2. Estimated effective detection radius (m; bars
expansion and observer effects. We used Akaike’s indicate 95% confidence interval [CI]) for six woodpecker
Information Criterion corrected for small sample size species surveyed in the Big Woods of Arkansas during 2006,
2007 breeding seasons and 2007 winter season.
(AICc; Burnham and Anderson 2002) to rank these
models and we used density estimates from the top-
ranked model. We did not model average parameter
estimates because currently program DISTANCE 5.0 does varying observers in model selection results were
not allow model-averaging within the Multiple Covariate pileated woodpeckers during winter 2007, red-bellied
Distance Sampling analysis engine. woodpeckers during winter and breeding seasons 2007,
To place our woodpecker density estimates into and yellow-bellied sapsuckers during winter 2007. For all
perspective, we reviewed the literature for woodpecker other species and season combinations, detectability
density estimates derived from the Southeastern United models with effects from varying observers were at least
States, usually from bottomland hardwood habitats. 36 times more plausible than models without. Detect-
However, comparing density estimates based on differ- ability estimates (standard errors in parentheses) from
ent studies should be done with caution because top models ranged from 0.06 (0.003) to 0.49 (0.06) across
different methods for estimating densities were used all species during all survey seasons (Figure 3). While
(see Bull and Jackson 1995, p 15). most detectability estimates were around 0.10, the
pileated woodpecker had consistently higher probability
Results of detections compared to the other woodpeckers each
season.
We detected 4,047 individual woodpeckers across Although the 95% confidence intervals (CIs) over-
3,588 detections, which varied in number by species and lapped, there was a general pattern for pileated and
survey season. Thus, most detections were of a single downy woodpecker densities to be lower during the
woodpecker. We had enough detections to estimate winter season than during the breeding season (Fig-
densities of downy woodpeckers Picoides pubescens, red- ure 4). The opposite pattern was true for red-headed
bellied, and pileated woodpeckers during all three survey woodpeckers, yellow-bellied sapsuckers, and northern
seasons. Red-headed woodpeckers M. erythrocephalus, flickers when densities could only be estimated during
yellow-bellied sapsuckers Sphyrapicus varius, and north- the winter. Unlike the other woodpeckers, the red-bellied
ern flickers Colaptes auratus are primarily winter visitors woodpecker density estimates increased with each
to the Big Woods; thus, we were only able to estimate successive survey season. Downy woodpecker densities
their densities during winter 2007. In no season were were similar each season. Downy and red-bellied
we able to estimate densities for hairy woodpeckers woodpecker densities were higher than pileated wood-
P. villosus (Supplemental Material, Table S1, http://dx.doi. pecker densities each season. Pileated woodpecker
org/10.3996/032010-JFWM-006.S1; Table S2, http://dx. densities were always lower than the other estimable
doi.org/10.3996/032010-JFWM-006.S12; Table S3, http:// woodpecker densities. We detected at least 4.5 pileated
dx.doi.org/10.3996/032010-JFWM-006.S3). woodpeckers and at least 98.6 red-bellied woodpeckers/
We found the effective detection radius for downy and km2 during the winter and at least 16.6 pileated
red-bellied woodpeckers was constant across seasons woodpeckers and at least 72.9 red-bellied woodpeck-
and varied around 60 m (Figure 2) while the effective ers/km2 during the breeding seasons (Table). We found
detection radius for pileated woodpecker was higher that compared to all other estimates, our pileated and
during the winter as compared to the breeding season. red-bellied woodpecker estimates were as high as or
The pileated woodpecker detection radius was about higher than most estimates from the literature. Also, for
100 m. The best-fitting detectability models were hazard- all other comparable woodpecker density estimates that
rate cosine and hazard-rate cosine plus observer effects. we could locate, ours were as high as or higher than
The only data sets that showed weak effects from estimates from other studies (Table).
Figure 4. Density estimates (bars indicate 95% confidence

Figure 3. Estimated detectability (bars indicate 95% confi- interval [CI]) for six woodpecker species surveyed in the Big
dence interval [CI]) for six woodpecker species surveyed in the Woods of Arkansas during 2006, 2007 breeding seasons, and
Big Woods of Arkansas during 2006, 2007 breeding seasons and 2007 winter season.
2007 winter season.
including the CRNWR. They concluded that the bird

Discussion community at CRNWR was organized along ‘‘the hydro-
Density estimates of six woodpecker species in the Big logic gradient acting either directly through avoidance of
Woods of eastern Arkansas during both the breeding flooded areas during the breeding season or indirectly
(2006 and 2007) and winter (2007) seasons were higher through selection of habitats having greater structural
than estimates from other parts of the Southeast. Our development of understory and midstory vegetation’’
estimates were at least 28% and 280% higher for pileated (Wakeley et al. 2007, p 435). They implied that the altered
and red-bellied woodpeckers, respectively, than the hydrologic cycle of the rivers in the Big Woods resulted
densities Tanner (1942) suggested coincided with suitable in disturbance to the plant community at various spatial
ivory-billed woodpecker habitat. Following Tanner’s hypo- scales, which in turn was organizing the bird community.
thesis that high pileated and red-bellied woodpecker We suspect that the frequent and prolonged flooding in
densities are indicative of good ivory-billed woodpecker the Big Woods (Denman and Karnuth 2005) stressed
habitat, then it follows that the Big Woods of Arkansas trees, resulting in increased food availability for the
should be a preferred location for ivory-billed woodpeckers. woodpeckers. The great variety of foraging methods
Southern bottomland hardwood forests harbor a employed by woodpeckers (Conner et al. 1994) suggests
diverse and abundant woodpecker community. For that the altered hydrologic disturbance was not affecting
example, of the eight woodpecker species detected by a single aspect of the forest but was impacting the forest
Shackleford and Conner (1997) in eastern Texas, the red- at a variety of levels (e.g., under-, mid-, and overstory),
bellied woodpecker, northern flicker, downy woodpecker, and the variable topographic relief in the Big Woods
red-headed woodpecker, and yellow-bellied sapsucker contributed to the patchy availability of different plant
were significantly more abundant in bottomland hard- communities. One primary aspect of the prolonged
wood forests than in either nearby longleaf or mixed inundation of water in the forests was increased tree
pine–hardwood forests. However, these woodpecker mortality (Conner and Sharitz 2005; 2005a, 2005b), which
densities are not static because overall woodpecker is highly correlated with arthropod biomass, the primary
densities increased markedly during the autumn and forage item of woodpeckers (Conner et al. 1994). Also,
winter (Bock and Lepthien 1975; Shackleford and Conner the Big Woods is composed predominantly of mature
1997). These elevated woodpecker populations during the hardwoods (Denman and Karnuth 2005). These older
nonbreeding season result, in part, from migrating and growth areas are preferred by most woodpeckers not
wintering northern flicker, red-headed woodpecker, yel- only because of the available forage but also because
low-bellied sapsucker, and possibly red-bellied wood- some woodpeckers (e.g., pileated woodpecker) excavate
pecker moving into these southern forests (Shackelford their nest and roost cavities in large trees (Evans and
and Conner 1997; Shackelford et al. 2000; Leonard and Conner 1979; Renken and Wiggers 1993; Conner et al.
Strout 2006). Additionally the high use of bottomland 1994; Tanner and Hamel 2001).
hardwoods during the nonbreeding season results from P.B. Hamel (United States Forest Service, personal
the abundance of soft mast, an important food source communication) investigated Tanner’s hypothesis that
during this time of year (Leonard and Strout 2006). high pileated and red-bellied woodpecker winter densi-
Why might the Big Woods be so attractive to ties could be used to aid in searching for ivory-billed
woodpeckers, especially during the nonbreeding season? woodpecker as well as characterizing habitats used by
Wakeley et al. (2007) compared bottomland hardwood ivory-billed woodpeckers. He estimated pileated and red-
bird communities across the Southeastern United States, bellied woodpecker winter densities across the South-
Table. Woodpecker density estimates (individuals/km2) for this study and from the literature rounded to the nearest integer
(ranges are in parentheses). We included estimates principally from studies conducted in the Southeastern United States and in
bottomland hardwood habitats.
Density estimate (individuals/km2)
Species Breeding season Winter season Annual Source
Pileated woodpecker 21 (17–28) 9 (5–18) This study
(5–15) Carter 1967
(5–15) (5–17) Collins 1960
(1–6) Dennis 1951
14 Hall 1983
(10–14) (3–9) Hamel 1992
7 Hamel, P.B. (USFS, pers. comm.)
3 Keller and Range 1990
4 (2–8) Renken and Wiggers 1993
17 Shackleford and Conner 1997
4 Tanner 1942
Red-bellied woodpecker 124 (73–181) 117 (99–139) This study
29 Carter 1967
(25–30) (5–35) Collins 1960
(2–3) Dennis 1951
(3–30) (2–65) Hamel 1992
16 Tanner 1942
Downy woodpecker 111 (83–145) 82 (66–101) This study
(15–16) Carter 1967
(24–60) (12–30) Collins 1960
3 Dennis 1951
(1–15) (3–20) Hamel 1992
39 Shackelford and Conner 1997
Red-headed woodpecker 78 (52–117) This study
13 Dennis 1951
6 5 Hamel 1992
28 Morse 1970
Northern flicker 60 (50–73) This study
6 (5–32) Collins 1960
(6–9) Dennis 1951
(1–15) (1–62) Hamel 1992
Yellow-bellied sapsucker 55 (42–73) This study
(3–5) Collins 1960
2 Dennis 1951
(1–40) Hamel 1992
eastern United States in bottomland hardwoods, and might not have the same minimum criteria for nesting
usually documented densities equal to or higher than and roosting tree circumferences as the ivory-billed
those estimated by Tanner (1942). However, smaller woodpeckers, nor would the foraging needs be as high
woodpeckers, especially the red-bellied woodpeckers, or specific as the ivory-billed woodpecker. Thus, Hamel
suggested that pileated and red-bellied woodpecker Table S3. Woodpecker species survey data, Spring
densities could act as surrogates for ivory-billed wood- 2007.
pecker but that those densities may not be the most Found at DOI: 10.3996/032010-JFWM-006.S3 (214 KB XLS).
effective strategy for developing searches or managing
habitats for ivory-billed woodpeckers. Acknowledgments
We found the greatest seasonal variation in densities
in yellow-bellied sapsuckers, red-headed woodpeckers, We thank A. Clifton, K. Ercit, A. Finfera, D. Hollis, C.
and northern flickers. Seasonal differences in densities of Kovach, A. Miller, H. Pruett, P. Tidwell, and M. Strauser for
these three woodpecker species were primarily due to help collecting data. We thank M. Blaney, R. Crossett, J.
migratory patterns and varying life-history patterns. The Denman, T. Foti, E. Johnson, A. Keister, P. Hamel, R. Hines,
yellow-bellied sapsucker was only present during the C. Hunter, M. Lammertink, A. Mueller, S. Reagan, K.
winter season because it is migratory and breeds far to Ribbeck, C. Rideout, B. Uhlein, and R. Wilson for logistical
the north (Walters et al. 2002). Red-headed woodpeckers support. Thanks to P.F. Doherty, Jr, P. Hamel, M. Kissling,
occurred in relatively high densities during the winter S.E. Lehnen, P. Lukacs, T. Nudds, and four anonymous
because they are both migratory and nomadic in reviewers for comments on earlier versions of this
foraging (Smith et al. 2000). We detected red-headed manuscript. We thank the Arkansas Game and Fish
woodpeckers during both breeding seasons, but not Commission and (USFWS) Cache and White River NWRs
enough to reliably estimate densities. Lastly, the situation for providing study areas.
for the northern flicker is less clear to us. Although the Funding was provided by the USFWS and the USGS
northern flicker is described as ‘‘strongly migratory,’’ the Arkansas Cooperative Fish & Wildlife Research Unit. The
southern populations are sedentary (Moore 1995). We use of trade, product, industry or firm names or products
originally anticipated detecting sufficient numbers of is for informative purposes only and does not constitute
northern flickers during the breeding seasons to estimate an endorsement by the U.S. Government or the USGS.
densities, but we did not. We assume from our results
that northern flickers moved away from the Big Woods References
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Articles
Roost Selection by Male Indiana Myotis Following Forest
Fires in Central Appalachian Hardwoods Forests
Joshua B. Johnson,* W. Mark Ford, Jane L. Rodrigue, John W. Edwards, Catherine M. Johnson
J.B. Johnson, J.W. Edwards
Division of Forestry and Natural Resources, West Virginia University, Morgantown, West Virginia 26506
W.M. Ford
U.S. Army Engineer and Research Development Center, Vicksburg, Mississippi 39180
J.L. Rodrigue
U.S. Forest Service, Northern Research Station, Parsons, West Virginia 26287
C.M. Johnson
U.S. Forest Service, Monongahela National Forest, Elkins, West Virginia 26241
Abstract
Despite the potential for prescribed fire and natural wildfire to increase snag abundance in hardwood forests, few
studies have investigated effects of fire on bat roosting habitat, particularly that of the endangered Indiana myotis
Myotis sodalis. From 2001 to 2009, we examined roost selection of Indiana myotis in burned and unburned forests in
Tucker County, West Virginia. We radiotracked 15 male Indiana myotis to 50 roost trees; 16 in burned stands and 34 in
unburned stands. Indiana myotis roosted in stands that had initially been burned 1–3 y prior to our observations. In
burned stands, Indiana myotis roosted exclusively in fire-killed maples (Acer spp.). In unburned stands, they roosted in
live trees, predominately hickories (Carya spp.), oaks (Quercus spp.), and maples. Roost trees in burned stands were
surrounded by less basal area and by trees in advanced stages of decay, creating larger canopy gaps than at random
trees in burned stands or actual roost trees located in unburned stands. Compared to random trees in unburned
stands, roost trees in unburned stands were less decayed, had higher percent bark coverage, and were surrounded by
less basal area, also resulting in larger canopy gaps. Roost-switching frequency and distances moved by Indiana myotis
among roost trees were similar between burned and unburned stands. Our research indicates that use of fire for forest
management purposes, at minimum provoked no response from Indiana myotis in terms of roost tree selection, and
may create additional roost resources, depending on spatial context.
Keywords: Fernow Experimental Forest; forest fire; Indiana myotis; Myotis sodalis; prescribed fire; roost selection; West
Virginia
Received: April 21, 2010; Accepted: September 27, 2010; Published Online Early: October 2010; Published: November
2010
Citation: Johnson JB, Ford WM, Rodrigue JL, Edwards JW, Johnson CM. 2010. Roost selection by male Indiana myotis
following forest fires in Central Appalachian Hardwoods Forests. Journal of Fish and Wildlife Management 1(2):111–121;
e1944-687X. doi: 10.3996/042010-JFWM-007
* Corresponding author: j-johnson3@juno.com
Introduction mast–producing species such as oaks (Quercus spp.) and

hickories (Carya spp.) in the Appalachian Mountains and
Formerly discouraged as a forest management prac- Central Hardwoods forest regions (Brose et al. 1999,
tice in many hardwood forest types, the use of 2006). Although effects of fire have been examined
prescribed fire has gained increasing acceptance among mostly from a forest management perspective, including
managers as a tool to help regenerate and retain hardin the Appalachian Mountains (Keyser and Ford 2006), its
Indiana Myotis Roost Selection J.B. Johnson et al.
role is often overlooked relative to life histories and Keyser and Ford 2006). Nonrandom selection of roosts,
management strategies for wildlife species (Brennan et roost-switching frequency, and distance traveled be-
al. 1998). Depending upon season, severity, and micro- tween successive roosts are indicators of roost availabil-
site conditions, fire can result in an increased abundance ity, but have not been well-researched in the eastern
of dead and dying trees, which typically has positive United States (Wilkinson 1985; Crampton and Barclay
implications for tree-dwelling wildlife species, including 1998; Sedgeley and O’Donnell 1999; Kunz and Lumsden
bats (Tucker et al. 2004; Blake 2005; Boyles and Aubrey 2003; Chaverri et al. 2007).
2006; Hayes and Loeb 2007). Recent research indicates Our objectives here were to: 1) compare roost tree
that prescribed fires alter forest structure in a way that is selection of Indiana myotis in forest stands subjected to
favorable to bats that roost in tree cavities or under fire (wildfire and prescribed burning) to unburned stands
exfoliating bark (Boyles and Aubrey 2006; Johnson et al. in the central Appalachian Mountains based on physical
2009; Lacki et al. 2009). Whereas empirical data exist characteristics of roost trees and random trees; and 2)
indicating that fire positively affects roosting habitat of examine roost tree abundance and availability by
common species such as evening bats Nycticeius humeralis comparing roost-switching frequency and distances
and northern myotis Myotis septentrionalis, no work has traveled between successive roosts located in burned
specifically examined this condition relative to roost and unburned forest stands. We predicted that Indiana
selection and habitat of Indiana myotis M. sodalis, a myotis would select roost trees within forest gaps
species listed as endangered pursuant to the U.S. created by fire, and that roost-switching frequency
Endangered Species Act (ESA) as amended (ESA 1973; would be higher and distances travelled to new roosts
Federal Register, 11 March 1967 [MacGregor et al. 1999; shorter in forest stands subjected to fire (Boyles and
Gumbert 2001; Dickinson et al. 2009]). Typically, Indiana Aubrey 2006; Johnson et al. 2009), which is consistent
myotis roost under the exfoliating bark of live trees such as with other bat species in the region and existing
shagbark hickory C. ovata or dead trees that are taller and ecological observations of the Indiana myotis.
larger in diameter than surrounding trees (Menzel et al.
2001). During a fire event, bark of live trees tends to peel Methods
away from the sapwood from heated sap in the phloem–
cambium layer (Barnes and Van Lear 1998), potentially Study area
providing an increased number of roosts for bats. We conducted our research at the Fernow Experi-
The geographic range of Indiana myotis largely mental Forest (FEF) and Petit Farm in Tucker County,
overlaps oak-dominated forest communities in Midwest, West Virginia (Figure 1). Both areas are located in the
mid-Atlantic, and northeastern United States (Braun Unglaciated Allegheny Mountains subsection of the
1950; Thomson 1982). Recent population stabilization Appalachian Plateau Physiographic Province (Kochen-
and, in some cases, modest population increases are now derfer et al. 2007). American chestnut Castanea dentata
being reversed by the rapidly spreading enzootic white- and oak species, such as northern red oak Q. rubra, were
nose syndrome Geomyces destructans. Heretofore, suc- historically significant components of the forest over-
cessful efforts to promote species recovery have focused story at Petit Farm and FEF. Chestnut blight Cryphonec-
largely on protection of hibernacula (e.g., caves where tria parasitica and subsequent fire suppression, white-
Indiana myotis overwinter [Clawson 2002; USFWS 2007, tailed deer Odocoileus virginanus herbivory, and use of
2009]). Following population declines due to white-nose uneven-aged harvesting systems has allowed forest
syndrome, protection of critical summer habitat will also composition to shift toward shade-tolerant tree species,
be important for species recovery. such as maples and American beech Fagus grandifolia
Many studies have focused on summer roosting (Schuler and Fajvan 1999, Schuler 2004). Overall mean
habitat of Indiana myotis, but few have examined spring annual precipitation in the area is 145.8 cm, ranging
and autumn roosting habitat (Menzel et al. 2001; Brack from 9.7 cm in October to 14.4 cm in June. Mean annual
2006; Britzke et al. 2006; Carter 2006). Regardless of temperature is 9.2uC, ranging from 218.0uC in January
season, Indiana myotis generally select large-diameter to 20.6uC in July (Kochenderfer 2006). The FEF is a 1,900-
trees and snags or live trees with exfoliating bark ha experimental forest managed by the U.S. Forest
(Gardner et al. 1991; Callahan et al. 1997; MacGregor et Service, Northern Research Station. Elevations range
al. 1999; Gumbert 2001; Menzel et al. 2001; Ford et al. from 530 to 1,100 m at FEF, and nearly 1,300 m on
2002). Fire may negatively impact Indiana myotis portions of the surrounding national forest. Elklick Run,
roosting habitat by reducing roost availability or by a 2.4-km fourth-order stream, roughly bisects the FEF
creating unsuitable conditions at existing roost trees. from east to west. Approximately 5.5 km of dendritic
Alternatively, fire may provide additional roosting intermittent and permanent streams feed into Elklick
resources for Indiana myotis and disturb a forest stand Run and incise the steep slopes and plateau-like
in a way similar to that in flooded forests, where snag ridgetops, providing all possible slope aspects (Madarish
abundance increases several-fold over preexisting con- et al. 2002). Vegetation at FEF is largely a mosaic of
ditions (Kurta et al. 1996; Carter et al. 2002; Carter 2006). second- and third-growth, mixed-mesophytic and north-
Fire might be an effective tool to foster recovery of this ern hardwood forest. Forests on the FEF, largely
species, because Indiana myotis may benefit from resembling those in the surrounding region, have been
additional snags and canopy disturbances provided by managed by even (patch clearcut)- and uneven (single-
active management (Carter et al. 2002; Carter 2006; tree selection)-aged silviculture since the mid-20th
Figure 1. Indiana myotis Myotis sodalis roost tree locations within unburned areas and forest stands subjected to prescribed fire at
Fernow Experimental Forest (FEF) and forest fires at Petit Farm, Tucker County, West Virginia, 2004–2009.
century, or has been left undisturbed following initial red maple A. rubrum, yellow-poplar Liriodendron tulipi-
harvesting in the Elklick watershed from 1903 to 1911 fera, black cherry Prunus serotina, American beech,
(Schuler and Fajvan 1999). Prescribed fire has recently sweet birch Betula lenta, and basswood Tilia americana
been used to promote oak regeneration in stands (Schuler 2004). In spring (April or May) 2007–2009,
currently dominated by sugar maple Acer saccharum, prescribed fire treatments were conducted in three
management compartments (12-, 13-, and 121-ha areas) Mammalogists (ACUC 1998). We used a radio receiver
on FEF (Figure 1). The compartments were burned using and three-element Yagi antenna (Wildlife Materials, Inc.,
strip head-fires, ignited with handheld drip torches. Murphysboro, IL) to attempt to locate roost trees daily.
Actual flame heights and combustion varied from To record the geographic locations (610 m) of roost
immediately dying out to .3.5 m high in some spots, trees, we used global positioning units (various
due to variable leaf litter, slope, and aspect. Additionally, models).
48, 20-m-radius plots were randomly located in each of
three management compartments, and all overstory or
midstory trees, other than oak or hickory species, were Habitat variables
treated with herbicides or girdled (T. Schuler, U.S. Using a point-quarter sampling method, we mea-
Department of Agriculture Forest Service, personal sured physical characteristics of each roost tree and
communication). those of four trees (.10 cm diameter at breast height
Petit Farm is located on private land, approximately [dbh]) nearest to the roost tree. Measurements were
8 km southeast of FEF, and has elevations ranging from taken within a few weeks of bats roosting in a tree. For
750 to 1,000 m. Petit Farm contains several first- and each roost tree, we determined tree species, dbh (cm)
second-order streams that drain into Glady Fork. Slopes using a diameter tape, decay class (Cline et al. 1980; i.e.,
are less steep at Petit Farm than at FEF. Vegetation 1 = live, 2 = declining, 3 = recent dead, 4 = loose
composition at Petit Farm is similar to that of FEF. bark, 5 = no bark, 6 = broken top, 7 = broken bole),
However, small portions of forests at lower elevations crown class (Nyland 1996; i.e., 1 = suppressed, 2 =
were cleared for pastoral grazing and hay production, and intermediate, 3 = codominant, 4 = dominant), tree
in 2001, a diameter-limit harvesting occurred on much of height (m) with a hypsometer, roost height when
the approximately 400-ha area. Northern red oak, black known, and visually estimated percent bark remaining
cherry, and sugar maple $46 cm diameter were on the tree. For each of four trees nearest to the roost
harvested, whereas all red maple were retained. In March tree, we determined tree species, dbh (cm), distance (m)
2003, an escaped campfire burned through a portion of to the roost tree, decay class, and crown class. For each
the forest stand that had been harvested in 2001. Fire roost tree, we averaged each variable for these four
intensity was variable, but flame heights and residual nearest trees irrespective of species. We measured
overstory mortality was greater than that observed at FEF, percent canopy gap either with a densiometer at four
in large part due to abundant downed tops and slash from cardinal directions surrounding the tree, or through
the recent logging (J. Rodrigue, U.S. Department of photographic methods (Johnson et al. 2009). We
Agriculture Forest Service, unpublished data). recorded aspect with a compass, percent slope with a
From spring through autumn, the extant bat commu- clinometer at the plot (11.3-m radius centered on the
nity of both sites consists of northern myotis, little brown roost tree), and stand basal area with a 20-factor prism
myotis M. lucifugus, big brown bats Eptesicus fuscus, tri- (m2/ha).
colored bats Perimyotis subflavus, eastern red bats Within each burned stand and in unburned areas, we
Lasiurus borealis, silver-haired bats Lasionycteris noctiva- located random roost trees at random coordinates
gans, and hoary bats Lasiurus cinereus (Owen et al. 2004; generated with ArcMap (Version 9.2; Environmental
Ford et al. 2005). A small number of endangered Indiana Systems Research Institute, Redlands, CA). In burned
myotis and Virginia big-eared bats Corynorhinus town- stands and unburned areas, we located 6 and 15 random
sendii virginianus that hibernate in cave systems on or trees, respectively. We considered unburned areas to be
near both FEF and Petit Farm remain in the area during any area outside areas within FEF where prescribed fires
summer (Owen et al. 2001; Ford et al. 2002). were used or the forest stand on Petit Farm where the
wildfire occurred. Random trees were identified as any
Radiotelemetry tree (.10 cm dbh) that had exfoliating bark that possibly
We captured bats in mist nets (Avinet, Inc., Dryden, could be used by an Indiana myotis as described by
NY) erected over stream corridors, small pools and Menzel et al. (2001). We used a subset of random trees
ponds, and skidder trails, during summer 2004–2006 at meeting Indiana myotis roosting requirements (e.g.,
Petit Farm and 2008–2009 at FEF. During autumn 2007– exfoliating bark) from a separate study (Johnson et al.
2008 at FEF, we used a harp trap to capture bats during 2009). Therefore, we could not match random trees with
the autumn swarm at Big Springs Cave, an Indiana roost trees in terms of sample size. We measured the
myotis hibernaculum. For each captured bat, we same tree characteristics, with exception of roost height,
determined species, sex, age, weight, forearm length, at the random roost trees as we did at the actual roost
and reproductive condition (Menzel et al. 2002). We trees.
used Skin BondH (Smith and Nephew, Largo, FL) In ArcMap, we determined the elevation for each roost
surgical cement to affix a 0.35-g radiotransmitter and random tree and the distance of each roost and
(Model LB-2N; lifespan 1–3 wk; Holohil Systems Ltd., random tree from the closest permanent water source
Carp, ON, Canada) between the scapulae of captured (Supplemental Material, Table S1, http://dx.doi.org/
Indiana myotis. Bat capture and handling protocols 10.3996/042010-JFWM-007.S1). We also used ArcMap to
were approved by the Animal Care and Use Committee determine straight-line distances that bats moved
of West Virginia University (Protocol No. 08-0504) and among successive roost trees (Supplemental Material,
followed the guidelines of the American Society of Table S2, http://dx.doi.org/10.3996/042010-JFWM-007.S2).
Table 1. Indiana myotis Myotis sodalis roost-tree use in burned and unburned forest stands at the Fernow Experimental Forest
(FEF) and Petit Farm in Tucker County, West Virginia, 2004–2009.
Tree species used
a
Year Radiotracked bats In burned area In unburned area
2005 Petit Farm: 4 males in summer Acer rubrum (n = 8) A. rubrum (n = 2)
A. saccharum (n = 2) Carya cordiformis (n = 1)
Sassafras albidum (n = 1)
Quercus rubra (n = 1)
Q. prinus (n = 1)
Betula lenta (n = 1)
2006 Petit Farm: 3 males in summer A. rubrum (n = 1) Undetermined (n = 1) A. saccharrum (n = 2)
A. rubrum (n = 1)
B. lenta (n = 1)
Q. rubra (n = 1)
Robinia pseudoacacia (n = 1)
Undetermined (n = 1)
2007 FEF: 2 males in autumn — C. ovata (n = 4)
Fagus grandifolia (n = 2)
Q. rubra (n = 2)
Prunus serotina (n = 1)
Liriodendron tulipifera (n = 1)
2008 FEF: 1 male in summer, 3 males in autumn — Summer roosts:
C. glabra (n = 1)
F. grandifolia (n = 1)
A. rubrum (n = 1)
Autumn roosts:
C. ovata (n = 4)
L. tulipifera (n = 1)
A. saccharum (n = 1)
Ulmus rubra (n = 1)
2009 FEF: 2 males in summer A. rubrum (n = 4) —
a
We tracked 1 female Indiana myotis to a 31-cm-dbh, fire-killed red maple Acer rubrum on Petit Farm (data not shown), where it roosted with about
45 other bats (Keyser and Ford 2006).
Statistical analysis between consecutive roost trees in the burned and

We used Mann-Whitney tests (Proc Npar1way; SAS unburned areas.
Institute, Inc. 2004) to make the following comparisons
of tree characteristics: 1) roost trees located within Results
burned areas to those in unburned areas, 2) roost trees in
burned areas to random trees in burned areas, 3) roost From 2004 to 2009, we captured and radiotracked 15
trees in unburned areas to random trees in unburned male Indiana myotis, and located 50 roost trees ([mean =
areas, and 4) roost trees used in autumn to those used in 3.3, 1 SE = 0.5, range = 1–5 roosts/bat] 24 at FEF and 26 at
summer. We made the four comparisons on appropriate Petit Farm); 16 in burned areas and 34 in unburned areas.
subsets of the same data set. Although differences Ten bats were captured and tracked to roosts during
between FEF and Petit Farm probably exist in terms of summer (June–July); five bats were captured and tracked
topography and fire intensity, we combined data to roosts during autumn (September–October). All roost
collected at both sites due to small sample sizes trees of bats captured at FEF during the autumn swarm
collected at either site. We used a Fisher’s Exact Test were located on or near FEF except one roost located on
(Proc Freq; SAS Institute, Inc. 2004) to determine whether Petit Farm, approximately 9 km from Big Springs Cave
roost trees were more frequently located on any slope (Figure 1). In burned areas, Indiana myotis roosted
aspects within burned and unburned areas. To deter- exclusively in fire-killed sugar maple and red maple snags.
mine roost tree availability, we used a Mann–Whitney In unburned areas, Indiana myotis roosted in a variety of
test to compare the frequency with which Indiana myotis live tree species, predominately hickories, oaks, and
switched roosts (measured as the number of consecutive maples (Table 1). At FEF and Petit Farm, some Indiana
days spent in a roost tree, between burned and myotis roosted in fire-killed trees in stands that had initially
unburned areas) and to compare distances (m) travelled been burned 1–3 y prior to our observation (Table 1).
Table 2. Characteristics of roost trees used by Indiana myotis Myotis sodalis and random trees subjected and not subjected to
fire in Tucker County, West Virginia, 2004–2009.
Fire roost (n = 16) Fire random (n = 6)
a b b
Variable Mean SE Mean SE Sc P
DBHR (cm) 41.37 2.54 26.63 4.85 35.0 0.018
HEIGHT (m) 21.65 1.11 12.92 2.17 28.0 0.004
DECAYN 3.54 0.31 2.21 0.42 39.5 0.042
GAP (%) 28.05 3.48 11.43 0.71 34.0 0.014
BASAL (m2/ha) 11.82 2.83 26.86 3.63 91.5 0.017
SLOPE (%) 29.13 2.92 43.50 5.12 100.5 0.022
ELEV (m) 833.56 10.86 680.17 26.26 21.0 ,0.001
Fire roost (n = 16) Nonfire roost (n = 34)
a b b
BARK (%) 63.87 7.18 76.79 5.31 272.0 0.034
DBHN (cm) 24.87 1.54 30.79 1.54 257.0 0.014
DECAYN 3.54 0.31 1.54 0.14 561.5 ,0.001
GAP (%) 28.05 3.48 14.63 3.10 90.0 0.005
BASAL (m2/ha) 11.82 2.83 23.55 2.08 183.5 0.002
WATER (m) 74.88 18.81 272.97 47.70 288.5 0.013
Nonfire roost (n = 34) Nonfire random (n = 15)
a b b
DECAYR 2.95 0.29 4.27 0.36 481.5 0.010
BARK (%) 76.79 5.31 46.67 8.20 245.0 0.006
GAP (%) 14.63 3.10 8.49 0.45 167.0 ,0.001
BASAL (m2/ha) 23.55 2.08 32.52 3.29 400.0 0.041
ELEV (m) 812.59 16.66 721.27 20.58 231.5 0.002
Summer roost (n = 33) Autumn roost (n = 17)
Variablea Meanb SE Meanb SE Sc P
BARK (%) 68.55 4.96 80.41 8.11 528.5 0.016
DBHN (cm) 26.60 1.32 33.20 2.18 521.5 0.024
DECAYN 2.63 0.26 1.32 0.11 272.5 0.002
GAP (%) 27.67 7.67 11.66 0.62 145.0 0.018
2
BASAL (m /ha) 16.15 2.64 24.34 2.15 438.5 0.030
SLOPE (%) 26.30 2.20 42.00 4.88 571.5 0.005
WATER (m) 121.33 38.41 380.88 52.56 650.0 ,0.001
a
DBHR = diameter (cm) at breast height of roost or random tree; DECAYR = decay class of roost or random tree (Cline et al. 1980; 1 = live, 2 =
declining, 3 = recent dead, 4 = loose bark, 5 = no bark, 6 = broken top, 7 = broken bole); HEIGHT = roost or random tree height (m); BARK =
estimation of percent bark remaining on roost or random tree; SLOPE = percent slope; BASAL = stand basal area (m2/ha); ELEV = elevation of
roost or random tree; WATER = distance of roost tree to closest permanent water source; GAP = percent canopy gap over roost or random tree;
DBHN = average diameter (cm) at breast height of four trees nearest to roost or random tree; DECAYN = average decay class of four trees nearest
to roost or random tree.
b
Mann–Whitney tests were performed on ranked values. Actual means and standard errors (SEs) are presented.
c
Mann–Whitney test statistic.
Roost trees in burned stands were surrounded by less unburned areas, roost trees were more frequently
basal area and by trees in more advanced stages of located on west- and south-facing slopes (n = 49,
decay, resulting in larger canopy gaps than at random df = 3, x2 = 9.30, Fisher’s Exact test P = 0.011). Com-
trees in burned stands and at roost trees located in pared to summer roost trees, roost trees used in autumn
unburned stands (Table 2). Compared to random trees in had a higher percentage of bark coverage, were
unburned stands, roost trees in unburned stands were surrounded by greater basal area and by trees that
less decayed, had higher percent bark coverage, and were larger in diameter and less decayed, resulting in
were surrounded by lower basal area, resulting in larger smaller canopy gaps surrounding these trees (Table 2).
canopy gaps (Table 2). In burned areas, roost trees were Roost trees used in summer were closer to water and
not more frequently located on any slope aspects (n = on gentler slopes than roost trees used in autumn
22, df = 2, x2 = 5.37, Fisher’s Exact test P = 0.065). In (Table 2).
Roost-switching frequency was similar (Mann–Whitney roosted in fire-killed trees (Gumbert 2001). In that study,
statistic = 229.5, P = 0.945) between burned (n = 12, summer and autumn roost areas also were similar in
mean = 1.25 d, SE = 0.25, switched every 1–4 d) and terms of canopy gap, aspect, slope, and distance to
unburned areas (n = 33, mean = 1.08 d, SE = 0.06, water. We found that most differences in site character-
switched every 1–2 d). Distances that Indiana myotis istics, including slope and distance to water likely were
moved among roost trees was similar (Mann–Whitney attributable to differences between Petit Farm and FEF
statistic = 96, P = 1.000) between burned (n = 8, mean rather than selection per se, because the majority of
= 219.62 m, SE = 49.65, range = 21–435 m) and autumn roost trees were located in or near the FEF and
unburned (n = 15, mean = 476.95 m, SE = 177.23, the majority of summer roost trees were located at Petit
range = 1–2,420 m) areas. Farm. Also, fire intensity and forest harvesting (basal area
removal) at Petit Farm were greater than at FEF (J.
Discussion Rodrigue, unpublished data). Compared to autumn roost
trees, the larger canopy gaps at summer roost trees were
Our observations suggest that Indiana myotis may due to more summer roosts being located in burned
respond favorably to structural changes in forests caused areas. Although differences were probably attributable
by fire. Although it is unclear whether Indiana myotis to different study areas where roosts were found
roosted in burned areas prior to fires, the fact that between the two seasons, other studies suggest there
Indiana myotis roosted in spaces under exfoliating bark are real and tangible physiological benefits to roosting in
of fire-killed sugar and red maples, indicates that the less open sites during autumn. During autumn, when
bats may benefit from forest alterations due to fire insect availability may be unstable, male Indiana myotis
(Brose et al. 1999). Characteristics of roost trees and may select relatively shaded, cooler roosts that allow
surrounding trees in burned areas were important in them to conserve energy by lowering their body
roost selection. temperature near ambient air temperatures to a state
Within burned areas, Indiana myotis selected trees that of torpor (Henshaw and Folk 1966). Conversely, during
were taller and larger in diameter than surrounding trees, summer, insect availability may be more consistent,
similar to that reported by others (Menzel et al. 2001). allowing male Indiana myotis to maintain a higher body
Roost trees were surrounded by trees that were in more temperature during the day by selecting warm trees in
advanced stages of decay, possibly providing additional canopy gaps on favorable aspects. Therefore, the
roosting structure. Also, the forced senescence of roost benefits of fire may be limited to summer roosting
trees and surrounding trees resulted in an enlarged habitat, possibly at the detriment of autumn roosting
canopy gap, allowing more solar radiation to reach the habitat. However, prescribed fires typically create a
roost trees (Johnson et al. 2009). In burned areas, canopy heterogeneous canopy layer with various gap sizes from
gaps were larger than those in unburned areas. From our which to select in summer and autumn (Johnson et al.
observations, mean canopy gap size at roost trees in 2009).
burned areas and those used during summer were In Virginia, Brack (2006) found that males and females
.30%, but it remains unclear whether enlarged canopy sometimes shared roost trees during autumn. Because
gaps such as those in burned areas provide actual fitness our study included data from only one female Indiana
benefits (e.g., increased recruitment success) for Indiana myotis, we were unable to make any substantive male–
myotis beyond roost use in unburned areas. female roost-selection comparisons (Menzel et al. 2001).
Roost trees in unburned areas occurred in naturally Despite known differences between roost selection of
existing canopy gaps. However, roost trees were less male and female bats (Kunz and Lumsden 2003), roost
decayed than random trees in unburned areas, and were trees of males in our study are very similar to those
comprised of a larger number of species than in burned reported for Indiana myotis maternity colonies elsewhere
areas. Although we did not investigate tree species within the species’ distribution (Menzel et al. 2001;
selection, the diversity of tree species used in unburned Britzke et al. 2003; Carter and Feldhamer 2005; Carter
areas was more typical of previous observations of 2006; Watrous et al. 2006; Johnson and Gates 2010).
Indiana myotis roosts (Menzel et al. 2001). In unburned Although it is unknown whether there is a minimum
areas, Indiana myotis selected trees surrounded by less density of roost trees required for Indiana myotis
basal area than random trees, and occurred in larger occupancy of an area, our roost-switching data and that
canopy gaps. of others suggest that there may be a sufficient number
During autumn, male Indiana myotis selected roost of roost trees in burned and unburned stands (Menzel et
trees that were less decayed, and in smaller canopy gaps al. 2001; Gumbert et al. 2002; Kurta et al. 2002; Britzke et
than roost trees used during summer. Also, Indiana al. 2006). We caution that areas we studied included
myotis roosted in trees that occurred mostly on southern small patches of burned forest, and roost densities in
or southwestern slopes, augmenting solar exposure. In areas adjacent to our study areas were unknown. Fire
the mountains of western Virginia, almost half of roost may be important in creating additional roost resources.
trees used during autumn were shagbark hickory (Brack Fire-killed trees may be suitable Indiana myotis roosts for
2006). Although we documented no bats roosting in an unknown number of years. Nonetheless, the ephem-
burned areas during autumn, Indiana myotis have been eral existence of roost trees remains to be quantified,
reported roosting in burned areas in Kentucky in depending on species and site conditions (i.e., suscep-
autumn, although it is unclear whether those bats tibility to wind-throw).
Our results indicate that Indiana myotis respond to are minimal in terms of disturbance to bats roosting in
effects of fire on forest stands, roosting in fire-killed trees trees within a burned stand, but may affect species (e.g.,
1–3 y postfire. This quick response to effects of fire allows eastern red bats) that roost in forest leaf litter (Rodrigue et
the bats to maximize the time period of roost suitability al. 2001; Dickinson et al. 2009). However, further research
for the newly created roost trees, because these trees will is necessary examining the effects of wildfire on bats
eventually become unsuitable. The mechanisms behind during summer. Short-term (,10 y) effects of forest fires
the initial use of a roost tree remain unclear (Wilkinson include the increase in roost tree abundance resulting in
1992; Ruczyński et al. 2009), but our observations more trees with exfoliating bark and favorable conditions
indicate that Indiana myotis likely were using trees in such as enlarged canopy gaps at trees that may have been
or near the areas impacted by fires prior to the previously unsuitable for Indiana myotis. These conditions
disturbances (Ford et al. 2002). Although we did not will gradually change as canopy gaps close and exfoliating
observe the same individuals for multiple years, the fact bark is lost. Relative to bat-roosting habitat, effects of fire
that Indiana myotis roosted in fire-killed trees in burned on upland hardwood forest stands are perhaps function-
areas suggests that they selected these new roost trees ally similar to the disturbance impacts following flood
over those used in previous years, (i.e., roost selection damage and mortality in bottomland and riparian forests,
took precedence over roost fidelity). The exfoliating bark as many trees are killed and become suitable snags for
on fire-killed trees will gradually or quickly slough from roosts, with surrounding dead trees and canopy gaps
the trees depending on species (Hallett et al. 2001), (Kurta et al. 1996; Carter 2006). In the Midwest, for
eventually rendering the tree unsuitable for Indiana example, Indiana myotis maternity colonies often use
myotis. Therefore, the intermediate- to longer term snags within floodplains or in ponds created by beavers
(.10 y after fire event) benefits of fire are uncertain for Castor canadensis. Forest fires in upland stands may mimic
Indiana myotis. effects of floods in systems in terms of snags (i.e., roost
Roost densities may return to or decline below prefire creation and forest structure change; Carter et al. 2002;
densities in absence of additional disturbance as fire- Carter 2006). However, floods may be more spatially
killed roost trees lose their bark and, therefore, much of extensive, providing a more abundant source of snag
their Indiana myotis roost-specific suitability. However, resources than forest fires, which may be less extensive
adjacent or nearby forest stands can be burned, or over the montane landscape where our study occurred.
herbicided, or girdled to create additional roost trees in Although some aspects of Indiana myotis roost selection
the short term. Systematic rotation of fire treatments may differ among regions, disturbance regimes that
increase roost resources may evoke similar responses
among adjacent forest stands would maintain or increase
from this species.
density of roost trees for Indiana myotis populations at
stand to landscape scales. By temporally staggering
prescribed fire treatments among adjacent forest stands, Supplemental Material
potential roost trees could be maintained in an area
(Rudolph and Conner 1996). Moreover, maintaining a Please note: The Journal of Fish and Wildlife Management
heterogeneous canopy layer across the landscape would is not responsible for the content or functionality of any
provide unshaded roost trees used during summer and supplemental material. Queries should be directed to the
relatively shaded roosts that are used during autumn. corresponding author.
In the long-term, canopy gap creation and reduction
or elimination of fire-sensitive tree species due to fires Table S1. Characteristics of Indiana myotis roost trees
will favor an oak-dominated forest that is beneficial to and random trees in burned and unburned forest stands
many other wildlife species (McShea and Healy 2002). at the Fernow Experimental Forest (FEF) and Petit Farm
However, this will also aid in the regeneration and in Tucker County, West Virginia, 2004–2009.
retention of hickories, a preferred roost tree genera of Found at DOI: 10.3996/042010-JFWM-007.S1 (114 KB XLS).
Indiana myotis (Brose et al. 1999; Nowacki and Abrams Table S2. Roost-switching distances of Indiana myotis
2008). Because it could take several decades for shagbark in burned and unburned forest stands at the Fernow
hickories to reach roost tree size, burning adjacent Experimental Forest (FEF) and Petit Farm in Tucker
County, West Virginia, 2004–2009.
stands on systematic rotation will create and retain snags
Found at DOI: 10.3996/042010-JFWM-007.S2 (27 KB XLS).
and other large trees with exfoliating bark in the interim.
Certainly, additional research is needed to fully assess the
spatial and temporal dynamics of snags and burning in Acknowledgments
relation to Indiana myotis roosting habitat requirements
relative to foreseeable management regimes in the The U.S. Department of Agriculture Forest Service
central Appalachians and elsewhere. Northern Research Station Joint Venture Agreement 06-
JV-11242300-140 provided primary funding for our study
Conclusions from the National Fire Plan to West Virginia University,
Division of Forestry and Natural Resources. Funding for
We believe that efforts to manage forests with prescribed burning was provided by a grant from Florida
prescribed fire to enhance oak regeneration appear to Power and Light and BHE Environmental to the U.S.
be compatible with Indiana myotis habitat management. Forest Service Northern Research Station through
Immediate effects of prescribed fires conducted in spring Collection Agreement 06-CO-112331-034.
We thank the staff of the U.S. Department of Agriculture Carter TC. 2006. Indiana bats in the Midwest: the
Forest Service Monongahela National Forest and the U.S. importance of hydric habitats. Journal of Wildlife
Department of Agriculture Forest Service Northern Management 70:1185–1190.
Research Station Timber and Watershed Laboratory for Carter TC, Carroll SK, Hofmann JE, Gardner JE, Feldhamer
conducting the prescribed burns. Field assistance was GA. 2002. Landscape analysis of roosting habitat in
graciously provided by J. Boyle, K. Boyle, C. McDaniels, A. Illinois. Pages 160–164 in Kurta A, Kennedy J, editors.
Stump, and M. Walsh. We thank T. Carter, R. Brooks, and J. The Indiana bat: biology and management of an
Stanovick, two anonymous reviewers, and the Subject endangered species. Austin, Texas: Bat Conservation
Editor for helpful critiques regarding this manuscript. International.
Carter TC, Feldhamer GA. 2005. Roost tree use by
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Articles
Summer and Winter Spatial Habitat Use by the Lake
Erie Watersnake
Kristin M. Stanford, Richard B. King,* Doug Wynn
K.M. Stanford, R.B. King
Department of Biological Sciences, Northern Illinois University, DeKalb, Illinois 60115
D. Wynn
Westerville North High School, Westerville, Ohio 43081
Abstract
In an effort to provide information to guide habitat management for the Lake Erie watersnake Nerodia sipedon
insularum, a federally threatened and Ohio state endangered species, we used radiotelemetry to obtain spatial habitat
data for adult snakes during the summer active season and during winter hibernation. During the summer active
season, terrestrial habitat use was limited to a narrow band of shoreline. Among individuals, maximum distance inland
from shore ranged from 1 to 50 m (mean = 8 m) and linear extent of shoreline ranged from 30 to 1,360 m (mean =
261 m). Winter hibernation occurred at varying distances inland with individual hibernation sites ranging from 1 to
580 m (mean = 29 m) from shore. Habitat use did not differ between males and females. Existing U.S. Fish and Wildlife
Service habitat management guidelines suggest that ground-disturbing activities within potential hibernation areas
(defined as terrestrial habitat within 161 m of shore) should be avoided in winter to prevent harm to hibernating
snakes. They suggest further that excavation and removal of shrubs, standing or downed trees, root masses, animal
burrows, piled rocks, cliffs, or bedrock within 21 m of shore should be avoided in summer to prevent harm to active
snakes. Given that Lake Erie watersnakes have recovered to the point where delisting is being proposed, these habitat
guidelines appear to be sufficient. However, maintaining voluntary compliance with habitat guidelines and meeting
the need for continued public outreach will be vital to ensure long-term persistence.
Keywords: conservation; habitat use; hibernation; management; snake

Received: May 26, 2010; Accepted: September 27, 2010; Published Online Early: October 2010; Published: November
2010
Citation: Stanford KM, King RB, Wynn D. 2010. Summer and winter spatial habitat use by the Lake Erie watersnake.
Journal of Fish and Wildlife Management 1(2):122–130; e1944-687X. doi: 10.3996/052010-JFWM-013
* Corresponding author: rbking@niu.edu
Introduction snake restricted to a cluster of islands in the Ohio and

Ontario waters of Lake Erie. The subspecific status of
Recovery of threatened and endangered species often island populations derives from color pattern differences
involves habitat management strategies designed to from mainland populations of the northern watersnake
provide for stable or increasing populations. Ideally, such N. s. sipedon (Conant and Clay 1937), a difference
strategies provide accommodations for key activities maintained by a dynamic balance between natural
(foraging, predator avoidance, reproduction) and are selection favoring a reduction in color pattern on
inclusive of all life-stages. In reality, knowledge of habitat exposed rocky islands shorelines and gene flow from
use for some species falls short of that needed for the nearby mainland (King 1993a, 1993b; King and
effective management and fundamental questions re- Lawson 1995, 1997). Although the Lake Erie watersnake
garding the size and location of key habitat elements spends much of its time on land, it forages for fish and
may be unanswered. amphibians in the near-shore waters of Lake Erie. A
The Lake Erie watersnake Nerodia sipedon insularum is dramatic shift in watersnake diet occurred in the 1990s
a medium-sized, nonvenomous, live-bearing colubrid with the arrival of the invasive round goby Apollonia
Watersnake Summer and Winter Spatial Habitat Use K.M. Stanford et al.
melanostomus, the abundance of which has contributed Bass (2001–2003), North Bass (2001–2002), and Gibraltar
to more rapid growth and greater reproductive output (2001–2003; figure 2 in King et al. 2006a). Following
among watersnakes (King et al. 2006b, 2008). Legal release, we located the snakes during daylight hours by
protection was conferred on this snake when it was searching outward from the release site or previous
declared threatened pursuant to the U.S. Endangered location using an AVM Instrument Co. LA12-Q receiver
Species Act (ESA) as amended (ESA 1973; Fazio and and Telonics RA-14K antenna. Depending on transmitter
Szymanski 1999) and endangered under the State of size and snake location, detection distance ranged from
Ohio Administrative Code, Province of Ontario Endan- about 200 to 1,000 m. Snakes were located on average
gered Species Act, and Committee on the Status of once every 8 d during the active season (from
Endangered Wildlife in Canada (summarized in USFWS implantation or emergence from hibernation until death,
2009a). Restricted geographic distribution and declining transmitter failure, or entry into hibernation) and at
population size were primary factors contributing to longer intervals during winter (King RB et al. in review).
legal protection (Fazio and Szymanski 1999; King et al. Intervals between location dates were variable because
2006a). Recovery efforts involving population monitor- of weather conditions and the logistics of interisland
ing, habitat management, and public outreach, together transportation.
with beneficial effects of abundant round gobies as food, Each time we located a snake, we recorded GPS
have resulted in rapid population recovery (R.B. King and coordinates using a handheld Garmin eTrex GPS receiver.
K.M. Stanford, Northern Illinois University, personal In addition, we measured distance inland from shore in
observation), leading the U.S. Fish and Wildlife Service meters using a tape measure or distance-measuring
(USFWS) to initiate actions to delist the Lake Erie wheel. We plotted locations onto Digital Orthophoto
watersnake (USFWS 2009a; USFWS 2010). Because Quarter Quadrangles using ArcView Geographic Infor-
delisting will eventually result in reduced legal protec- mation Systems software. We estimated the amount of
tion, having appropriate habitat management recom- terrestrial habitat used during summer active season in
mendations in place is vital to the long-term persistence two ways. First, following Pattishall and Cundall (2008),
of the snake. we estimated the linear extent of shoreline used as the
Here we describe spatial habitat use by the Lake Erie length of shoreline between the two most distant
watersnake as determined using radiotelemetry. We focus locations (Figure 1). Second, we estimated the amount
on the location and extent of habitat used during the of terrestrial habitat by determining the maximum
summer active season and the location of winter distance inland from shore at which each snake was
hibernation sites because these represent areas necessary observed. For snakes monitored during multiple sum-
for successful reproduction and overwinter survival, mers, both extent of shoreline and amount of terrestrial
respectively. We combine this with published information habitat were pooled across years because these snakes
on Lake Erie watersnake foraging habitat (Jones et al. used approximately the same areas each season.
2009) to evaluate USFWS habitat management recom- We measured the distance to shore from hibernation
mendations (USFWS 2003 [appendix D]; USFWS 2009b) sites (Figure 1) as described above. For those snakes that
and the ‘‘Habitat Protection and Management’’ criterion of hibernated inland from shoreline areas not used during
the Lake Erie Watersnake Recovery Plan (USFWS 2003). the summer, the distance between the shoreline point
closest to the hibernation site and the nearest shoreline
Methods location used during summer was estimated using
ArcView (Figure 1).
Radiotransmitters (Holohil, Inc.; model SI-2T) ranging in Distributions of distance measures (extent of shoreline,
size from 5.2 to 13.8 g with an expected battery life of 12– maximum distance inland from shore, and distance to
36 mo were implanted into adult Lake Erie watersnakes in shore from hibernation site) were all significantly right-
2000, 2001, and 2002. We conducted surgery under skewed. For analysis, we normalized these variables by
aseptic conditions using either isofluorane or halothane adding one and taking natural logarithms. Means and
anesthetic following methods modified from Reinert and other descriptive statistics were back-transformed for
Cundall (1982) and Brown and Weatherhead (2000). We clarity of presentation.
positioned transmitters within the body cavity about one-
third anterior to the vent and loosely sutured them to the
Results
body wall. We inserted transmitter antennas subcutane-
ously in an anterior direction. We monitored the snakes Transitions between summer habitat and hibernation
continuously until they recovered from anesthesia and sites were identified based on qualitative changes in
gave them daily injections of antibiotic (10 mg/kg Baytril) watersnake behavior. During the active season, terrestrial
until we released them at their site of capture 2–5 d later. habitat use was restricted to a narrow continuous band
Adult males (n = 22) averaged 210 g (range = 118–347 g) of shoreline habitat. In autumn, watersnakes typically
and adult females (n = 32) averaged 525 g (range = 306– made longer distance movements to hibernation sites
954 g) at the time of surgery. Transmitter mass averaged further inland, after which they were found repeatedly
2.4% of adult female mass (range = 1.4–3.7%) and 4.1% of within highly localized areas (i.e., within 10 m). In some
adult male mass (range = 2.7–7.1%). cases, hibernation sites were directly inland and adjacent
Telemetry occurred on five Lake Erie islands, including to shoreline areas used during summer. However, in
Kelleys (2000–2003), South Bass (2001–2003), Middle other cases, watersnakes moved to hibernation sites that
Figure 1. Movements of a representative Lake Erie watersnake, illustrating extent of shoreline used during summer, distance
outside summer range to the shoreline point nearest hibernation site, and distance to shore from hibernation site. Open circles are
locations at which this snake was found between 19 June 2001 and 12 October 2001 (active-season locations) and span 260 m. The
‘bullseye’ represents the hibernation site where this snake remained October–April. Before entering hibernation, this snake moved
about 135 m along the shore beyond where it spent the active season and 280 m inland to hibernate.
were inland from shoreline areas not used during maximum distance inland from shore: t = 0.60, P =
summer (Figure 1). Entry into hibernation occurred 0.55). Therefore, sexes were pooled to generate a
between 8 September and 18 October. Spring emer- population-wide description of active-season move-
gence was invariably followed by movements back to ments. Extent of shoreline ranged from 30 to 1,360 m
shoreline areas used the previous summer, and occurred among individuals and averaged 261 m (Figure 2).
between 20 April and 31 May. Features of summer habitat were not quantified but
watersnakes made frequent use of on-shore retreat sites
Summer habitat use consisting of crevices within exposed bedrock, among
Information on habitat use during summer was loose rocks overlaying bedrock, and within human-made
obtained from 54 watersnakes, 20 that were tracked structures (e.g., crib docks, shoreline protection).
during a single summer, 27 during two summers, 5 Given that snakes were not monitored continuously
during three summers, and 2 during four summers and some individuals were monitored for only a portion
(Table 1). Locations used by individual watersnakes of the active season, distances reported above may be
during summer were recorded 5–41 times throughout underestimates of the true extent of shoreline used
the study. For individuals tracked in multiple years, the during the active season. In fact, estimated extent of
number of locations in later years was sometimes low shoreline increased as the number of times snakes were
because mortality and battery life limited data collection. located increased (Pearson correlation, P = 0.005).
On average, individual snakes were monitored for a span However, the amount of variation explained by number
of 137 active-season days (range = 36–360 d). of locations was small (r2 = 0.14), and distributional
Neither extent of shoreline nor maximum distance characteristics of extent of shoreline (i.e., mean, median,
inland from shore differed significantly between males SD, data not shown) changed little when analyses were
and females (extent of shoreline: t = 20.92, P = 0.366; repeated using only the 40 snakes located $10 times
Table 1. Characteristics of adult Lake Erie watersnakes (n = 54) monitored on five Lake Erie islands in 2000–2003 using
radiotelemetry, including island, sex, initial snout–vent length (SVL) and Mass, Dates tracked with number of relocations in
parentheses, total relocations (Total), extent of shoreline used (Extent) and Maximum inland distance during summer, and distance
from Hibernation to shore.
Dates tracked (relocations)
Maximum Hibernation
SVL Mass Extent inland to shore
ID Islanda Sex (mm) (g) 2000 2001 2002 2003 Total (m) (m) (m)
123 KI m 753 289 8/9–9/18 (9) 5/9–6/18 (7) 16 155 13.7 20
128 KI f 928 819 7/24–9/21 (15) 15 85 10.0
129 KI f 1,000 954 7/24–9/21 (17) 5/9–11/9 (16) 33 180 10.0 190
157 KI f 1,079 652 7/24–9/16 (11) 5/10–8/15 (11) 6/4–9/6 (8) 30 725 9.1 580
159 KI f 977 778 7/24–9/8 (12) 12 110 22.9
165 KI m 752 275 7/24–9/25 (14) 14 1,360 20.0 35
168 KI f 910 651 7/24–9/22 (17) 5/13–9/25 (16) 6/2–10/1 (5) 5/1–6/4 (3) 41 1,180 20.0 24
173 KI f 794 408 7/24–9/18 (13) 5/22–9/24 (14) 6/2–9/24 (9) 6/4 (1) 37 1,090 20.0 24
184 KI m 647 221 7/24–1014 (17) 5/10–9/3 (14) 5/17–6/22 (4) 35 140 50.0 15
186 KI m 723 234 7/24–9/19 (14) 14 665 7.6 10
193 KI f 864 577 7/29–9/24 (16) 16 160 4.6
196 KI m 640 158 7/24–9/24 (13) 5/9–7/9 (9) 22 360 1.2 20
213 KI m 745 268 7/20–8/22 (8) 5/13–5/16 (2) 10 160 30.0 40
215 KI f 946 829 7/29–9/18 (11) 5/9–9/24 (16) 27 155 15.2 6
216 KI m 647 203 7/30–9/25 (12) 5/10–9/25 (13) 25 580 15.2 100
218 KI f 933 607 7/30–9/16 (12) 5/22–7/19 (7) 19 230 18.3 50
221 KI m 754 347 7/29–9/25 (15) 5/9–9/3 (15) 6/2–8/12 (7) 37 755 7.6 12
047 MBI m 700 191 7/12–9/11 (8) 5/23–8/27 (13) 21 420 1.5 20
111 NBI f 795 461 7/11–11/13 (11) 5/1–8/27 (15) 25 250 7.6 47
139 MBI f 830 418 7/12–9/11 (7) 7 180 6.1
16A KI f 910 553 7/6–10/9 (9) 9 180 9.1 21
22E NBI f 865 565 7/11–9/18 (9) 9 260 9.1
255 MBI f 890 653 7/12–9/11 (8) 5/1–7/31 (10) 18 490 7.0 6
26C NBI m 645 150 7/11–9/11 (8) 5/1–8/17 (9) 17 440 9.1 141
301 KI m 770 252 7/9–9/24 (6) 6 725 9.1 18
309 MBI m 605 131 7/12–9/11 (8) 8 70 4.6 45
35C SBI m 590 118 6/21–9/17 (11) 11 720 4.6 17
477 MBI m 725 237 7/12–9/11 (8) 5/23–9/27 (16) 24 500 3.0 1
507 SBI f 790 384 6/21–9/17 (11) 6/10–9/6 (12) 23 250 15.0 280
551 MBI m 645 137 7/12–9/11 (8) 8 120 0.9 180
628 SBI f 850 397 6/21–9/17 (11) 11 490 6.1 12
67D KI f 920 564 7/9–8/20 (6) 6 260 10.7 40
708 KI m 760 246 7/9–9/25 (8) 4/27–10/1 (14) 22 480 12.2 21
730 MBI m 675 164 7/12–9/11 (8) 8 485 12.2 395
80A NBI f 850 485 7/11–9/11 (8) 8 190 4.6 6
83E Gib f 930 606 7/12–9/18 (8) 5/21–10/3 (15) 4/15–5/13 (2) 25 360 5.0 11
A54 MBI f 740 330 7/12–9/11 (8) 5/1–10/1 (18) 5/26 (1) 27 150 2.0 30
A77 SBI m 730 214 6/21–9/6 (9) 6/28–9/19 (11) 20 85 10.0 220
B15 SBI m 680 169 6/21–9/24 (8) 8 105 3.0 162
B1E KI f 970 478 7/9–9/24 (8) 8 140 7.6 15
B35 KI f 695 519 6/23–9/25 (2) 6/2–6/27 (3) 5 230 9.1 40
B69 SBI f 895 322 7/11–10/21 (11) 4/29–5/31 (5) 16 290 7.6 12
D1C SBI m 735 192 7/11–10/12 (10) 4/29–7/23 (12) 22 340 9.1 15
D60 Gib m 655 170 7/12–9/5 (6) 6 30 3.0 6
D64 SBI f 820 360 6/21–10/12 (12) 6/10–6/28 (4) 12 100 6.1 9
E66 MBI f 690 328 7/12–9/11 (7) 5/23–9/5 (14) 21 250 12.2 180
Table 1. Continued.
Dates tracked (relocations)
Maximum Hibernation
SVL Mass Extent inland to shore
a
ID Island Sex (mm) (g) 2000 2001 2002 2003 Total (m) (m) (m)
E71 MBI f 725 351 7/12–9/7 (8) 5/1–7/19 (8) 16 90 6.0 7
F0F NBI f 965 550 7/11–9/18 (9) 9 350 4.6 105
F7D SBI f 805 431 6/21–10/12 (12) 4/30–5/2 (2) 14 260 12.2 280
467 MBI f 825 345 6/12–10/11 (15) 5/13–7/12 (3) 18 380 6.0 20
50C SBI m 745 248 6/18–9/19 (13) 5/13–8/21 (5) 18 210 6.0 11
629 SBI f 785 306 6/10–9/19 (14) 6/24–8/21 (3) 17 315 5.0 6
712 MBI f 960 591 6/10–9/23 (13) 5/13–10/16 (3) 16 105 12.0 3
D76 SBI f 965 580 6/10–9/19 (14) 4/15–5/13 (2) 16 490 30.0 9
a
Island abbreviations: KI = Kelleys Island, SBI = South Bass Island, MBI = Middle Bass Island, NBI = North Bass Island, Gib = Gibraltar Island.
(mean = 296 m) or the 33 snakes located $15 times during summer (Figure 1). Whether snakes hibernated
(mean = 279 m). Maximum distance inland from shore directly inland from summer habitat was independent of
ranged from 1 to 50 m among individuals and averaged sex (x2 = 0.002, P = 0.961). To reach hibernation sites
8 m (Figure 3). located inland from shoreline areas not used in summer,
snakes were estimated to have moved between 35 and
Hibernation sites 1,410 m (mean = 216) along the shore prior to moving
Information on hibernation sites was based on 49 inland. Snakes moved through unused shoreline areas
individual Lake Erie water snakes (27 females, 22 males; quickly (e.g., in ca. 4–14 d between two successive
Table 1). Hibernation sites were located in a single year tracking dates), suggesting that they were not an
for 36 snakes, in two successive years for 10 snakes and undetected part of summer habitat.
in three successive years for 2 snakes. Because snakes Distance to shore from hibernation sites did not differ
observed to hibernate, more than once used the same significantly between males and females (t = 0.30, P =
hibernation location (within 10 m), information on 0.762). Therefore, sexes were pooled to generate a
hibernation sites was pooled across years. population-wide description of this variable. Distance to
Of the 49 hibernation sites, 31 were located directly shore from hibernation sites ranged from 1 to 580 m
inland from shoreline areas used during the summer by among individuals and averaged 29 m (Figure 4). The
those individuals. The other 18 hibernation sites were elevation above lake level of the ground surface over
located inland from shoreline areas outside of that used hibernating snakes ranged from 1 to 10 m.
Figure 3. Maximum distance inland from shore during

Figure 2. Extent of shoreline used during summer active summer active seasons by adult Lake Erie watersnakes (n =
seasons by adult Lake Erie watersnakes (n = 54). Vertical 54). Vertical dashed lines show the estimated maximum
dashed lines show the estimated extent of shoreline used by, distance inland for, from left to right, 50% (#8 m), 75%
from left to right, 50% (#261 m), 75% (#444 m), 90% (#715 m), (#13 m), 90% (#21 m), 95% (#26 m), and 99% (#42 m) of the
95% (#952 m), and 99% (#1,622 m) of the population (based population (based on mean and variance of log-
on mean and variance of log-transformed values). transformed values).
predator was a fox, raccoon, or domestic dog. Cause of

death could not be inferred for two males and four females
that could not be retrieved from under large rocks.
Discussion
In describing the habitat and habits of the Lake Erie
watersnake, Conant and Clay (1937) noted that
The water snakes, for the most part, appear to be restricted

to the edges of the islands. They are not numerous on sand
and gravel beaches but where rocks are strewn upon the
shores, where low cliffs are close to the water’s edge, or
where docks extend into the water they are abundant
(pp 8–9).
The suggestion that the Lake Erie watersnake is largely

restricted to the island peripheries is found in subsequent
studies (e.g., Camin and Ehrlich 1958; King 1986) although
some authors note that it ‘‘does occasionally inhabit more
inland environs’’ (p 38; Kraus and Schuett 1982). Data presented
Figure 4. Distance to shore from hibernation sites for adult here largely confirm this observation with respect to the
Lake Erie watersnakes (n = 49). Vertical dashed lines show the summer active season; on average, the maximum distance
estimated distance inland to hibernation sites for, from left to inland from shore at which telemetered watersnakes were
right, 50% (#29 m), 75% (#71 m), 90% (#158 m), 95% observed was just 8 m. However, the results of this study go
(#255 m), and 99% (#621 m) of the population (based on further in documenting the linear extent of shoreline used by
mean and variance of log-transformed values). Lake Erie watersnakes and by providing information on the
distance inland to suitable hibernation sites. Extent of habitat
Characteristics of hibernation sites were variable. Most used during summer is highly variable, with some individuals
had soil and rock substrates and appeared to consist of restricting their activity to stretches of shoreline encompassing
natural openings or fissures. Access holes to some just tens of meters and others ranging over a kilometer.
hibernation sites were apparent and may have been Furthermore, hibernation can occur well inland, which has
the result of burrowing by small mammals, shoreline significant conservation implications. Although on average,
erosion, or tree falls. Eleven hibernation sites were found Lake Erie watersnakes hibernated within 29 m of shore,
in or near human-made structures including abandoned individual hibernation sites occurred as far inland as 580 m
building foundations, drainage tile, sewage line, concrete and 25% of the adult population was estimated to hibernate
shoreline protection material, the foundation of a 71 m or further inland.
cottage, and an unused wine cellar. Vegetation around Fine-scale characteristics (e.g., depth, substrate) of
hibernation sites was also variable and included mature hibernation sites used by Lake Erie watersnakes remain
forest, woody scrub, and grass and herbaceous vegeta- unknown, but given the shallow soils of the islands
tion (sometimes including mown lawns). Usually only generally (Core 1948), hibernation sites likely occur in
one telemetered snake was observed to hibernate at a crevices within the bedrock or among loose rocks and
given location although other untelemetered animals other materials immediately above the bedrock. Further-
were sometimes observed nearby. In three cases, two more, because the substrate over hibernation sites was
telemetered snakes hibernated within 10 m of each sometimes as little as 1 m above lake level, it is possible
other. that Lake Erie watersnakes are partly or fully submerged
during hibernation. Submergence appears to reduce
Mortality energy use and risk of desiccation and leads to increased
Of the 54 watersnakes included in this study, 29 were overwinter survival in the common gartersnake Thamno-
alive when transmitters or transmitter batteries failed. phis sirtalis (Costanzo 1986, 1989a, 1989b), but the
Eleven of these survivors were recaptured during annual degree to which submergence occurs in Lake Erie
population censuses in subsequent years (King et al. watersnakes is not known.
2006a). The other 25 watersnakes were known or Lake Erie watersnakes forage for fish and amphibians
presumed to have died over the course of the study; 22 (primarily round gobies and mudpuppies Necturus
apparently of natural causes and 3 as a consequence of maculosus), in waters adjacent to shoreline areas used
mowing, brush burning, or hibernation site disruption. in summer (Jones et al. 2009). Observations made during
Those that died of natural causes included four males and systematic boat surveys indicate that foraging by Lake
nine females that died during hibernation or shortly after Erie watersnakes can extend $200 m from shore
spring emergence. Two females died prior to entry into (Figure 5) in water up to 7.5 m deep (Jones et al.
hibernation. These females had recently given birth as 2009). Thus, while on-shore activities are restricted to a
evidenced by their thin appearance, and this may have narrow band of land close to the water’s edge, foraging
been a contributor to their deaths. One female apparently activities extend to a wider band of the near-shore
died as the result of predation; bite marks suggest the waters of Lake Erie.
transformed means = 153 vs. 261 m; note that Pattishall

and Cundall [2008] did not transform their data and so
report mean stream length as 280 m). Like Lake Erie
watersnakes, Northern watersnakes remained close to
water, with 57% of relocations occurring within 5 m, 88%
within 20 m, and 97% within 50 m of the stream
(Pattishall and Cundall 2008; cf. figure 3). These
similarities between stream-dwelling and lakeshore-
dwelling populations are striking and contrast markedly
with spatial habitat use by marsh and wetland dwelling
N. sipedon (reviewed by Pattishall and Cundall 2008).
Conservation implications
Spatial habitat use by Lake Erie watersnakes differs
markedly between the summer active season and winter
hibernation, and successful management should address
habitat use during both time periods. In summer, habitat
use is restricted to a narrow band of terrestrial shoreline
Figure 5. Distance offshore at which Lake Erie watersnakes and a somewhat wider band of near-shore aquatic
were observed foraging (n = 130). Vertical dashed lines show habitat. In contrast, winter hibernation occurs in both
the estimated distance offshore within which foraging occurred shoreline and inland habitats. Because there do not seem
for, from left to right, 50% (#74 m), 75% (#106 m), 90% to be habitat features strongly associated with areas
(#147 m), 95% (#178 m), and 99% (#257 m) of the population used in summer or for hibernation, general guidelines for
(based on mean and variance of log-transformed values).
human activities within these areas may represent the
best way to minimize watersnake mortality.
Spatial habitat use can be difficult to compare among As part of the Lake Erie watersnake recovery plan, the
telemetry studies because authors analyze data in USFWS developed a set of Guidelines for Construction,
different ways and only infrequently report detailed Development, and Land Management Activities (USFWS
tracking information on individuals (Pattishall and 2003 [appendix D]; USFWS 2009b). Similar guidelines
Cundall 2008). Comparisons are further complicated by address habitat management at Middle Bass Island State
differences in the dimensionality of habitat use, even Park (USFWS 2003 [appendix B]) and within conservation
within species. For example, terrestrial habitat use by easements in the region (M. Seymour, USFWS, personal
Lake Erie watersnakes in summer is essentially linear communication). To avoid harming watersnakes during
(one-dimensional) and similar to that of Northern water- hibernation, these guidelines recommend that no
snakes Nerodia sipedon sipedon dwelling in a stream excavation occur within potential hibernation areas
habitat in Pennsylvania (figure 6 in Pattishall and Cundall (defined as terrestrial habitat up to 161 m inland from
2009). In contrast, N. sipedon inhabiting marshes and shore) between 15 October and 15 April. Furthermore,
wetlands exhibit planar two-dimensional habitat use during periods of hibernation egress (defined as 16
(figures 2–4 in Tiebout and Cary [1987]; figure 1 in Brown April–31 May) and ingress (defined as 15 September–14
and Weatherhead [1999]; figure 1 in Roe et al. [2004]; October), excavation within potential hibernation areas
Roth and Greene [2006]). Even greater dimensionality in should occur only when air temperatures exceed 15.5uC.
habitat use is seen in species living in high-relief Presumably, during warm weather, surface-active snakes
environments where elevation (or depth) provides can move quickly enough to avoid harm from excavation
another habitat dimension. equipment. Because root masses may provide hiberna-
The detailed data presentation of habitat use by tion sites, tree removal should be limited to the above-
stream-dwelling Northern watersnakes in Pennsylvania ground portion of the tree. To avoid harming water-
(Pattishall and Cundall 2008) allows direct comparison snakes during the active season, these guidelines
with this study. Much like the extent of shoreline used recommend that mowing within potential summer
during summer by Lake Erie watersnakes, distances habitat (defined as terrestrial habitat within 21 m of
between the most upstream and downstream locations shore) between 15 April and 15 September occur at dusk
of Northern watersnakes were right-skewed (table 2 in with mower blades set high above the ground. By
Pattishall and Cundall 2008). In both studies, habitat use waiting until dusk, it is assumed that surface-active
varied widely among individuals, ranging from 30 to snakes have sought cover and are less likely to be
1,360 m in Lake Erie watersnakes (this study) and 10 to harmed by mowing equipment. Furthermore, these
1,425 m in Northern watersnakes (Pattishall and Cundall guidelines specify that excavation, removal of shrubs,
2008). Northern watersnakes were more variable in standing or downed trees, root masses, animal burrows,
habitat use (Levene’s test, F = 9.95, P = 0.002, SD of piled rocks, cliffs, or bedrock within 21 m of shore should
natural log–transformed data = 1.8 vs. 0.78) but on be avoided and use of heavy machinery should be
average used shorter lengths of habitat than did Lake limited to paved areas. Because Lake Erie watersnakes
Erie watersnakes (t-test using separate variance esti- make frequent use of docks and shoreline erosion-
mates: t = 22.58, df = 73.86, P = 0.012, back- control structures, it is recommended that maintenance
and construction of these structures should use designs Lonz Winery, Paramount Distillers, Saunders Cottages, M.
that provide watersnakes with access to retreat sites and B. Barnhill, D. Buris, G. and S. Cooper, the Dress
(steel or timber cribs, riprap) rather than impenetrable family, J. and R. Eaton, S. and K. Hahn, P. and L. Hays, D.
materials (concrete, sheet steel). Finally, these guidelines Honor, the Jones family, T. Kranyak, T. Marsh, the
specify that construction sites should be actively Minshall family, T. Pangrace, C. and K. Schneider, and
monitored for watersnakes before and during construc- other landowners for access to field study sites. We thank
tion activities. The low rate of mortality among M. Thomas, J. Hageman, J. Reutter, and the F.T. Stone
telemetered snakes attributed to human activities in this Laboratory for logistical support, and M. Andre, J.
study (3 of 54 animals) suggests that these management Schofield, and K. Bekker for assistance with telemetry.
guidelines have been effective. Comments from the Subject Editor and two anonymous
Definitions of potential hibernation areas (terrestrial reviewers substantially improved the manuscript.
habitat up to 161 m from shore) and summer habitat Funding was provided by Northern Illinois University,
(terrestrial habitat up to 21 m from shore) used in these the Ohio State University F.T. Stone Laboratory, Ohio
guidelines were based in part on prior analyses of the Department of Natural Resources Division of Wildlife, and
telemetry data presented here (King 2003) and were USFWS. Permits were provided by the Ohio Department
selected to encompass the estimated area used by 90% of Natural Resources Division of Parks and Recreation,
of the adult population. These definitions, together with Ohio Department of Natural Resources Division of
other analyses presented here, might be used to further Wildlife, and USFWS. Protocols were approved by
characterize an ‘ideal’ Lake Erie watersnake habitat patch institutional animal care and use committees at Northern
as also encompassing approximately 715 m of shoreline Illinois University and the Ohio State University.
(Figure 2) and extending approximately 147 m offshore
(Figure 5). Guidelines could be expanded based on
habitat use by a larger proportion of adult watersnakes
References
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ed by federal and state agencies. Delisting of the Lake Erie Costanzo JP. 1986. Influences of hibernaculum microen-
watersnake, as is being proposed by the USFWS (USFWS vironment on the winter life history of the garter
2010), will create additional challenges for managers snake (Thamnophis sirtalis). Ohio Journal of Science
charged with ensuring population persistence within a 86:199–204.
less restrictive legal framework. Adherence to the habitat Costanzo JP. 1989a. A physiological basis for prolonged
management guidelines outlined above will become submergence in hibernating garter snakes Thamno-
largely voluntary, even for projects that currently require phis sirtalis: evidence for an energy-sparing adapta-
federal consultation. We contend that continued public tion. Physiological Zoology 62:580–592.
outreach will be helpful because Lake Erie islands are a Costanzo JP. 1989b. Effects of humidity, temperature,
destination for large numbers of seasonal visitors, many of and submergence behavior on survivorship and
whom have limited knowledge regarding snakes gener- energy use in hibernating garter snakes, Tham-
ally (Burghardt et al. 2009) and Lake Erie watersnakes in nophis sirtalis. Canadian Journal of Zoology 67:2486–
particular (e.g., that they are a unique feature of the 2492.
regional biota, are protected, and are nonvenomous). [ESA] Endangered Species Act of 1973, Pub. L. No. 93-
205, 87 Stat. 884 (Dec. 28, 1973).
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We thank Camp Patmos, Cleveland Museum of Natural wildlife and plants; threatened status for Lake Erie
History, Craft’s Lakeview Lane, Long Point Owners Group, water snakes (Nerodia sipedon insularum) on the
offshore islands of western Lake Erie. Federal Register Kraus F, Schuett GW. 1982. A herpetological survey of the
64:47126–47134. coastal zone of northwest Ohio. Kirtlandia 36:21–54.
Jones PC, King RB, Stanford KM, Lawson TD, Thomas M. Pattishall A, Cundall D. 2008. Spatial biology of northern
2009. Frequent consumption and rapid digestion and watersnakes (Nerodia sipedon) living along an urban
prey by the Lake Erie watersnake with implications for stream. Copeia 2008:752–762.
an invasive prey species. Copeia 2009:437–445. Pattishall A, Cundall D. 2009. Habitat use by synurbic
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snake, Nerodia sipedon insularum. Copeia 1986:757– 198.
772. Reinert HK, Cundall D. 1982. An improved surgical
King RB. 1993a. Color pattern variation in Lake Erie implantation method for radio-tracking snakes. Co-
water snakes: inheritance. Canadian Journal of Zool- peia 1982:702–705.
ogy 71:1985–1990. Roe JH, Kingsbury BA, Herbert NR. 2004. Comparative
King RB. 1993b. Color-pattern variation in Lake Erie water water snake ecology: conservation of mobile animals
snakes: prediction and measurement of natural that use temporally dynamic resources. Biological
selection. Evolution 47:1819–1833. Conservation 118:79–89.
King RB. 2003. Hibernation, seasonal activity, movement Roth TC, Greene BD. 2006. Movement patterns and home
patterns, and foraging behavior of adult Lake Erie range use in the northern watersnake (Nerodia
water snakes. Annual Report to the Ohio Division of sipedon). Copeia 2006:797–802.
Wildlife and U.S. Fish and Wildlife Service. Available: Tiebout HM III, Cary JR. 1987. Dynamic spatial ecology of
http://respectthesnake.com/files/Annual_Report_III.pdf the water snake Nerodia sipedon. Copeia 1987:1–18.
(September 2010). [USFWS] U.S. Fish and Wildlife Service. 2003. Lake Erie
King RB, Lawson R. 1995. Color pattern variation in Lake watersnake (Nerodia sipedon insularum) recovery. Fort
Erie water snakes: the role of gene flow. Evolution 49: Snelling, Minnesota: U.S. Fish and Wildlife Service.
885–896. Available: http://www.fws.gov/midwest/endangered/
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snakes. BioScience 47:279–286. [USFWS] U.S. Fish and Wildlife Service. 2009a. Lake Erie
King RB, Queral-Regil A, Stanford KM. 2006a. Population watersnake (Nerodia sipedon insularum) 5-year review:
size and recovery criteria of the threatened Lake Erie summary and evaluation. Available: http://www.fws.
watersnake: integrating multiple methods of popula- gov/midwest/endangered/recovery/5yr_rev/pdf/lews5Yr.
tion estimation. Herpetological Monographs 20:83–104. pdf (September 2010).
King RB, Ray JM, Stanford KM. 2006b. Gorging on gobies: [USFWS] U.S. Fish and Wildlife Service. 2009b. Lake Erie
beneficial effects of alien prey on a threatened watersnake management guidelines for construc-
vertebrate. Canadian Journal of Zoology 84:108–115. tion, development, and land management activities.
King RB, Stanford KM, Ray JM. 2008. Reproductive Available: www.fws.gov/midwest/ohio/documents/
consequences of a changing prey base in island endangered_lewsguide_2009.pdf (May 2010).
watersnakes. South American Journal of Herpetology [USFWS] U.S. Fish and Wildlife Service. 2010. Endan-
3:155–161. gered and threatened wildlife and plants; proposed
King RB, Stanford KM, Wynn D, Bekker KA. In review. rule to remove the Lake Erie watersnake (Nerodia
Hibernation phenology and over-winter body tem- sipedon insularum) from the Federal list of endan-
perature of the Lake Erie Watersnake (Nerodia sipedon gered and threatened wildlife. Federal Register 75:
insularum). Submitted to Journal of Herpetology. 30319–30338.
Notes
Assessing Wetland Changes in the Prairie Pothole Region
of Minnesota From 1980 to 2007
Fred T. Oslund,* Rex R. Johnson, Dan R. Hertel
U.S. Fish and Wildlife Service, Habitat and Population Evaluation Team, 18965 County Highway 82, Fergus Falls,
Minnesota 56537
Abstract
Wetlands in the Minnesota Prairie Pothole Region are critical landscape elements because of their unmatched
importance to breeding waterfowl, and other wildlife. They provide vast benefits to store runoff or act as nutrient sinks
and offer other environmental and socio-economic returns. Data on location, extent and types of wetlands collected by
the U.S. Fish and Wildlife Service National Wetlands Inventory is used for developing conservation strategies and
evaluating net landscape changes affecting fish and wildlife populations. Minnesota wetlands were mapped 27 y ago
by the National Wetlands Inventory. We examined 176 10.2-km2 (4-mi2) sample plots in the Minnesota Prairie Pothole
Region, using aerial photo interpretation techniques, to determine the current accuracy of the National Wetlands
Inventory data used in the eastern Prairie Pothole Region for conservation planning and evaluation. We stratified our
analysis by Bailey’s (1995) Ecological Subsections. We estimated that across the entire Minnesota Prairie Pothole Region
4.3% of wetland area has been lost since 1980 with losses varying from 0 to 15% among Ecological Subsections.
Implications of these findings suggest that National Wetlands Inventory data should be regularly updated in areas
subject to rapid wetland change.
Keywords: National Wetlands Inventory; Minnesota; Prairie Pothole Region; wetlands

Received: December 18, 2009; Accepted: June 11, 2010; Published Online Early: July 2010; Published: November
2010
Citation: Oslund FT, Johnson RR, Hertel DR. 2010. Assessing wetland changes in the Prairie Pothole Region of
Minnesota from 1980 to 2007. Journal of Fish and Wildlife Management 1(2):131–135; e1944-687X. doi: 10.3996/122009-
JFWM-027
* Corresponding author: fred_oslund@fws.gov
Introduction North American harvest (Batt et al. 1989). As well as

being essential habitat, potholes provide nutrient and
The Prairie Pothole Region (PPR) covers portions of sediment sinks from surrounding agricultural and urban
Minnesota, Iowa, North Dakota, South Dakota, Montana, lands, retain runoff (which reduces flooding), promote
and the prairie provinces of Canada that were historically groundwater recharge, sequester large amounts of
grassland and covered by the last advance of Pleistocene organic carbon, and have other environmental and
glaciers. The PPR was once one of the largest grassland– socio-economic values (Johnson et al. 2008).
wetland ecosystems on earth before crop-based agricul- Prairie potholes historically covered up to 20% of the
ture transformed the landscape (Prairie Pothole Joint PPR and often exceeded 40 basins/km2 ([100/mi2]
Venture 2005). The region is composed of millions of Kantrud et al. 1989; Johnson and Higgins 1997; R.R.
small depressional wetlands, called potholes, which were Johnson, U.S. Fish and Wildlife Service, unpublished
left in the wake of retreating glaciers (Johnson et al. data). A five-county area in west-central Minnesota
2008). Prairie potholes provide crucial habitat for historically estimated to support approximately 300,000
breeding waterfowl and other migratory birds. In wet breeding dabbling-duck pairs is able to support less than
years, the PPR may contribute over 70% of the total duck 59,000 breeding pairs today (280%; Johnson et al. 2008).
production of species that comprise the majority of the In this same area, 81% of historic wetlands and 49% of
Assessing Wetland Changes in the Prairie Pothole Region F.T. Oslund et al.
wetland area have been converted to agriculture, data set to be compared to NWI data (Figure 2). For
illustrating the propensity of farmers to drain small, estimating waterfowl population estimates, plots were
shallow wetlands. stratified based on land ownership: Federal—the plot
Although prairie pothole drainage began as early as contained at least 65 ha (160 ac) of USFWS Waterfowl
the last quarter of the 19th century at the time of Production Area; Easement—the plot contained 65 ha
European settlement (Wooten and Jones 1955; Dahl and (160 ac) of USFWS wetland easements; Refuge—the plot
Allord 1996), it reached its zenith from the 1950s through contained any amount of land in a National Wildlife
the 1970s as more powerful farming implements became Refuge; or Private—the plot contained .94% private
commonplace (McManis 1964). Most Minnesota prairie land and contained no National Wildlife Refuge. Plots
wetlands (.75%) had been drained by 1980 (R. Johnson, were assigned to Bailey (1995) subsections for this
unpublished). assessment (Figure 1) and we acknowledge that the
The National Wetland Inventory (NWI) was established sample design was not designed for an assessment of
in 1974 to develop and provide information on the estimated wetland change.
location, extent, and types of wetlands (U.S. Fish and A subset of aerial photographs of each sample plot
Wildlife Service [USFWS] NWI 2002). The NWI delineated collected from 1991 to 2007 was used to determine
wetlands in the Minnesota PPR using circa 1980 high- wetland change. We overlaid NWI data layer onto
altitude color-infrared photographs at a 1:63,000-scale aerial photographs, which allowed for detection of
(Johnson and Higgins 1997; who also provide a wetland alteration (Figure 2). We used TNT Map and
description of basic NWI mapping conventions in the Image Processing System software (MicroImages, Inc.),
PPR, B. Huberty, USFWS, personal communication). By version 7.3, to compare and map changes in wetland
about 1980, the estimated statewide number of wetland condition and add or delete wetlands on aerial photo-
acres in Minnesota had been reduced by 42% to graphs by comparing them to NWI data (circa 1980).
3,521,000 ha (8,700,000 ac; Dahl 1990), which primarily National Wetland Inventory wetland delineations were
occurred within Minnesota PPR. assumed to be accurate at that time. Data on presence,
Our objective of this study was to assess the current absence, classification and size were recorded for every
accuracy of NWI data in Minnesota, which were mapped wetland that occurred in the NWI data base or on aerial
using color-infrared photography collected 27 y ago. photos for each plot. Wetlands that were significantly
Evaluation of post–NWI-mapping wetland gains and altered in extent, including those that were completely
losses was necessary to quantify the amount, type, and drained or restored, were recorded as losses (Supplemen-
distribution of wetland change. It is important to tal Material, Table S1, http://dx.doi.org/10.3996/122009-
determine the current reliability of NWI data because JFWM-027.S1).
they are used extensively for conservation planning in Wetlands were recorded as completely or partially
the Minnesota PPR. drained if they showed altered hydrology (e.g., altered size
or duration of ponding) in four consecutive years of recent
Study Area aerial photography from 2004 to 2007 or we saw evidence
that indicated tiling, ditching, filling, or intensive cultiva-
We evaluated wetland change in five Ecological
tion without crop damage that usually results from short-
Subsections ([ES] Bailey 1995; Figure 1) within the
duration ponding. If partially drained, the remaining
Minnesota PPR. The Prairie Coteau, Minnesota River
wetland area was delineated and compared to the NWI
Prairie, and Red River Prairie ES comprise the Prairie
delineation to determine wetland area loss.
Parkland Province (Bailey 1995). This province was
‘‘Restored’’ wetlands resulted when a wetland was
historically dominated by tall grass prairie but now is
purposely restored or a drainage system fell into
intensely farmed. The Aspen Parklands ES lies within the
disrepair. Wetlands were classified as restored when
Tall Grass Aspen Parklands Province, which is a transi-
water or emergent vegetation was visible on aerial
tional area between prairie and semihumid mixed
photographs in at least two consecutive years between
conifer–deciduous forests, and although frequently
2004 and 2007. Deliberately restored wetlands were
farmed in the early 20th century retains large areas of
usually easily identified because there was a dike, water
grassland pasture, hayland, and aspen-dominated Popu-
control structure, or ditch plug visible on aerial photos.
lus tremuloides deciduous forest. The Hardwood Hills ES
lies in the Eastern Broadleaf Forest, an area regarded as a Wetlands that were affected by drainage (i.e., generally
transitional zone from open grassland to forest and partially drained) were assigned by Cowardin a d modifier
woodland (Minnesota Department of Natural Resources (e.g., partially drained) in circa-1980 NWI data (Cowardin et
2008; Figure 1). al. 1979). Altered wetlands with d modifiers were
compared with those without d modifiers to determine
whether drainage rates differed between altered wetlands
Methods
and wetlands unmodified by drainage (circa 1980).
We used existing 10.2-km2 (4-mi2) breeding waterfowl Finally, data were compiled by ES and the plot-based
and wetland survey plots, which were randomly selected estimates were expanded to each ES based on the ratio
in 1987, for this assessment. Existing plots were used of plots sampled to total number of plots in each
because color-infrared aerial photography of each plot ownership stratum within the ES. This, combined with
has been acquired every spring (1991–present) as part of NWI data, enabled us to estimate the overall change in
the 4-mi2 waterfowl survey, which allowed this long-term wetland area and number in each ES.
Figure 1. The Prairie Pothole Region (PPR) of western Minnesota showing locations of sample plots and percent change by
Ecological Subsections (ES) evaluated (1980–2007).
Results Wetlands with d modifiers in the NWI data base

accounted for 72% of post-1980 losses. Wetlands without
Our 176 sample plots contained 22,497 ha of wet- prior drainage history or with no drainage evidence
lands. A total of 1,028 wetland ha were lost to accounted for 28% of all losses.
drainage and 479 total wetland ha were restored
(Table 1). A total wetland area net loss of 549 ha or Discussion
approximately 2.3% occurred on sample plots since 1980
(Table 1). Estimates increased to a loss of 4.3% wetland A routine assumption of conservation planning is that
areas when stratified samples were expanded to the geospatial data are accurate. Accuracy assessment is
entire Minnesota PPR (Table 2). Total wetland losses in crucial if we are to have confidence in the outcomes of
the Minnesota PPR were estimated at 52,097 ha and an conservation planning and evaluation. Accuracy for land
estimated 13,011 ha were restored (Table 2). The cover and wetlands data may be expected to degrade
difference between the two estimates indicated a net over time. Periodic accuracy checks and remapping
loss of 39,086 ha since 1980. Estimates of net change by when warranted is essential to developing credible
ES are provided (Table 2). conservation strategies.
Figure 2. National Wetlands Inventory (NWI) data overlaid on aerial photographs allowed for detection of altered wetlands. Lines
displayed show where wetlands were on the landscape during the mapping process in 1980 (circa 2006).
Although this assessment indicated that wetland enhancing preexisting drainage systems. Regardless of
changes throughout the entire Minnesota PPR were the legality of this drainage, it represents a significant
relatively small (24.3% of total area) over nearly a 30-y loss of wetland functions in these areas. Finally, much of
span, data for some regions are more outdated than the new drainage (post-1980) can be attributed to the
others, which should be a consideration when prioritiz- loss of small temporary wetlands (temporarily flooded,
ing remapping efforts. For example, net change ranged palustrine, emergent wetlands; Cowardin et al. 1979),
from 8 to 15% for ES in west-central and southern which accounted for 61.2% of wetland losses observed.
Minnesota. Remapping wetlands in these regions may be Temporary wetlands are routinely cultivated, altering
a high priority for conservation partnerships in the state. their depth due to redistribution of soil to the extent that
We believe that most losses can be attributed to fixing or they become incapable of retaining water in wet years.
Table 1. Wetland change data from 1980 through 2007 collected from sample plots within each Ecological Subsection.
Negatives represent a loss in wetland hectares.
Total wetland hectares Wetland hectares Wetland hectares Net change
Ecological region in sample drained restored (ha) % Change
Aspen Parklands 3,718.7 78.5 0.7 277.8 22.4
Prairie Coteau 529.9 55.4 10.1 245.4 28.6
Hardwood Hills 4,962.8 45.8 42.1 23.6 20.1
Minnesota River Prairie 11,592.9 769.7 333.3 2436.4 23.8
Red River Prairie 3,295.8 78.2 92.6 14.4 0.4
Total 24,100.1 1,027.7 478.8 2548.9 22.3
Table 2. Estimates of wetland change from 1980 through 2007 expanded to Ecological Subsection and the entire Minnesota
Prairie Pothole Region (MN PPR). Negatives represent a loss in wetland hectares.
Total wetland hectares Wetland hectares Wetland hectares Net change for PPR
Ecological region in MN PPR drained restored (ha) % Change
Aspen Parklands 345,256.3 8,181.1 72.9 28,108.1 22.4
Prairie Coteau 19,675.5 3,022.1 73.1 22,949.0 215.0
Hardwood Hills 54,907.2 1,958.2 1,703.5 2254.7 20.5
Minnesota River Prairie 353,883.2 35,204.1 7,382.7 227,821.4 27.9
Red River Prairie 127,781.3 3,732.0 3,778.6 46.6 0.0
Total 901,503.5 52,097.4 13,010.8 239,086.6 24.3
Improvements in drainage technology, including pattern www.npwrc.usgs.gov/resource/wetlands/wetloss/index.

tiling fields around wetlands (which intercepts runoff), htm (Version 16JUL97) (January 2008).
are likely to exacerbate these losses in the future. Dahl TE, Allord GJ. 1996. History of wetlands in the
Finally, it is difficult to overstate the importance of conterminous United States. Pages 19–26 in Fretwell
reliable up-to-date NWI data for conservation planning JD, Williams JS, Redman PJ, compilers. National water
and evaluating progress toward objectives. For example, summary on wetland resources. Washington, D.C.: U.S.
conservation partnerships need to periodically assess
Geological Survey Water-Supply Paper 2425.
changes in wetland status to assess net progress toward
Johnson RR. 2010. Drained wetland data for Minnesota.
objectives for wetland goods and services. We recom-
mend that the NWI make regular wetland mapping Unpublished. Fergus Falls, Minnesota: U.S. Fish and
updates a high priority in regions of the United States Wildlife Service. Available: http://prairie.ducks.org/
experiencing rapid changes in wetland abundance, type, index.cfm?&page = minnesota/restorablewetlands/home.
or distribution. htm& (August 2010).
Johnson RR, Higgins KF. 1997. Wetland resources of
Supplemental Material eastern South Dakota. Brookings: South Dakota State
University.
Johnson RR, Oslund FT, Hertel DR. 2008. The past, present,
supplemental material. Queries should be directed to the and future of prairie potholes in the United States.
corresponding author. Journal of Soil and Water Conservation 63:84–87.
Kantrud HA, Krapu GL, Swanson GA. 1989. Prairie basin
Table S1. Minnesota Prairie Pothole Region wetland wetlands of the Dakotas: community profile. Wash-
data. Observations on wetland alterations are recorded in ington, D.C.: U.S. Fish and Wildlife Service Biological
the Status column of each tab. Report 85(7.28).
Found at DOI: 10.3996/122009-JFWM-027.S1 (5 MB XLS). McManis DR. 1964. The initial evaluation and utilization
of the Illinois prairies, 1815–1840. Chicago: University
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Notes
Effects of Orientation and Weatherproofing on the
Detection of Bat Echolocation Calls
Eric R. Britzke,* Brooke A. Slack, Mike P. Armstrong, Susan C. Loeb
E.R. Britzke
U.S. Army Engineer Research and Development Center, Vicksburg, Mississippi 39180
B.A. Slack
Kentucky Department of Fish and Wildlife Resources, Frankfort, Kentucky 40601
M.P. Armstrong
U.S. Fish and Wildlife Service, Kentucky Ecological Services Field Office, Frankfort, Kentucky 40601
S.C. Loeb
U.S. Forest Service, Southern Research Station, Clemson, South Carolina 29634
Abstract
Ultrasonic detectors are powerful tools for the study of bat ecology. Many options are available for deploying acoustic
detectors including various weatherproofing designs and microphone orientations, but the impacts of these options
on the quantity and quality of the bat calls that are recorded are unknown. We compared the impacts of three
microphone orientations (horizontal, 45u, and vertical) and two weatherproofing designs (polyvinyl chloride tubes and
the BatHat) on the number of calls detected, call quality, and species detected by the Anabat II bat detector system at
17 sites in central Kentucky in May and June 2008. Detectors with BatHat weatherproofing recorded significantly fewer
call sequences, pulses per file, species per site, and lower quality calls. Detectors in the horizontal position also tended
to record fewer files, fewer species, and lower quality calls. These results illustrate potential impacts of deployment
method on quality and quantity of data obtained. Because weatherproofing and orientation impacted the quality and
quantity of data recorded, comparison of results using different methodologies should be made with caution.
Keywords: activity; Anabat; bats; orientation; weatherproofing

Received: July 27, 2010; Accepted: October 4, 2010; Published Online Early: October 2010; Published: November 2010
Citation: Britzke ER, Slack BA, Armstrong MP, Loeb SC. 2010. Effects of orientation and weatherproofing on the
detection of bat echolocation calls. Journal of Fish and Wildlife Management 1(2):136–141; e1944-687X. doi: 10.3996/
072010-JFWM-025
* Corresponding author: eric.r.britzke@usace.army.mil
Introduction bats without an observer present, thereby allowing a

small crew to sample multiple sites simultaneously and
Monitoring of bat echolocation calls has greatly for long periods of time (e.g., Gorresen et al. 2008).
expanded our knowledge of bat ecology. Ultrasonic The Anabat II (Titley Electronics, Ballina, NSW, Aus-
detectors permit nonintrusive sampling of the bat tralia) is an ultrasonic detection system that is widely
community and can be used to sample bats in habitats used for the study of bats. Although the system allows
that are ineffectively sampled using traditional capture for long periods of automated recording, the equipment
techniques (e.g., open fields, large rivers). Additionally, is susceptible to damage from rain. To protect the
bat detectors can detect more species at a site than equipment, researchers have developed two primary
capture techniques (Murray et al. 1999; O’Farrell and types of weatherproofing. The first protective measure
Gannon 1999). Further, ultrasonic detectors can be involves a detector placed in a polyvinyl chloride (PVC)
deployed to passively record the echolocation calls of tube in a waterproof box so that sound enters the PVC
Deployment of Ultrasonic Detectors E.R. Britzke et al.
tube, while the water drains out through a small hole in hoarybat.com). We placed data from each unit in a
the bottom of the tube (O’Farrell 1998). The second separate directory and scanned the data with a
system is comprised of the microphone enclosed in a customized filter in Analook (Version 4.9j) to delete
PVC housing pointed down at an acrylic-glass plate used extraneous noise (Britzke 2003), with subsequent visual
for call deflection (BatHat; EME Systems Inc., www. examination to ensure that only files with echolocation
emesystems.com; Arnett et al. 2006). The ability to detect calls remained.
echolocation calls of bats can also be affected by the We used the scanFiles option in Analook to determine
orientation angle of the detector. Depending on the the number of files and the Countscan option to
height of the detector, detector orientation can signif- calculate the total number of pulses for each sequence.
icantly affect the number of bat detections (Weller and We then used a customized filter (modified from Britzke
Zabel 2002). and Murray 2000) to extract parameters for those
While different weatherproofing methods and orien- echolocation call sequences that were of sufficient
tations have been extensively used by researchers, there quality (e.g., no broken pulses) to permit species
is a lack of published data examining the effectiveness of identification. We saved the parameters to a text file
the two most commonly used weatherproofing options, and identified them by comparison to a known call
despite the importance of the potential impacts on the library using a mixed Discriminant Function Analysis
results and subsequent interpretation of the data. These model in the statistical program R (v. 2.2.1; http://
impacts could affect the quantity of activity data www.r-project.org; Duchamp 2006). Because species
collected or quality of the data, which may render identification is probabilistic, we determined species
species identification impossible in some circumstances. presence using the methodology described in Britzke et
The objective of this study was to determine how al. (2002).
common detector orientations and weatherproofing We compared the total number of files, number
options affect the quantity and quality of bat calls of pulses per file, and percentage of files surviving
recorded. the identification filter among the five treatments using
a randomized block (by site) analysis of variance.
Methods We then compared means using a Tukey test, and
species richness using a Median test. For all tests alpha =
We conducted this study in two areas, one each in 0.05.
Franklin and Spencer counties, Kentucky. Both areas
contained small woodlots dominated by oaks (Quercus
Results
spp.), elms (Ulmus spp.), and hickories (Carya spp.)
interspersed with grazed pasture. We chose sampling We surveyed 17 sites in which sampling equipment
sites within each area to represent the range of suitable functioned properly for all five treatments. Mean number
sampling sites based on our experience with recording of files recorded per night varied by treatment (F = 4.02;
echolocation calls and included ponds, linear corridors P = 0.006). The mean number of files recorded per night
(e.g., streams and roads), canopy gaps, and open fields. by the BatHat was significantly lower than all other
We used Anabat II detectors connected to a compact treatments; the units with PVC protection recorded the
flash-storage Zero-Crossings Analysis Interface Module, highest number of files per night (Figure 2A). Mean
as well as the SD1 units, in which the detector and the number of pulses per file varied among treatments (F =
storage Zero-Crossings Analysis Interface Module are 8.02; P , 0.001) and the BatHat had significantly fewer
contained in one unit (Titley Scientific; www.titley.com. pulses per file than all other treatments (Figure 2B).
au). Before sampling, we calibrated the detectors by Percentage of files that passed through the identification
following methods used by Larson and Hayes (2000). At filter were significantly different among treatments (F =
each sampling site, we deployed five Anabat II systems 15.37; P , 0.001), with the BatHat recording calls of
side by side on tripods at 1.5 m. Each day of sampling we significantly lower quality than the other treatments
assigned a specific detector for each treatment. The five (Figure 2C).
treatments included three detector orientations (0u Six bat species were detected: big brown Eptesicus
[horizontal], 45u, and 90u [vertical]) and two weather- fuscus, eastern red Lasiurus borealis, northern long-eared
proofing options (a PVC tube and the BatHat). A Myotis septentrionalis, Indiana M. sodalis, little brown M.
schematic of the two weatherproofing options is shown lucifugus, and tri-colored Perimyotis subflavus. Species
in Figure 1. We randomly determined the relative richness varied significantly among treatments (P =
position of each of the five treatments for each site. 0.003): the mean number of species was 1.7 for the
Sampling occurred in May and June 2008. We sampled horizontal orientation, 2.4 for the 45u orientation, 2.4 for
each site for one night. the vertical orientation, 2.3 for the PVC weatherproofing,
We set up detectors before dark and recorded the and 1.1 for the BatHat weatherproofing. In general, the
echolocation calls throughout the night. We only pattern of higher detections by the PVC and 45uand 90u
conducted sampling on nights with no heavy precipita- orientations held for each of the species, although
tion and when winds were light. We placed detectors to northern long-eared bats were only detected by the
maximize detection at the recording site. The following vertically oriented detectors and Indiana bats were not as
morning, we gathered units and uploaded data to a readily detected by the detectors with PVC or BatHats
laptop computer using the CFCread program (www. (Figure 3).
Figure 1. Schematics for the two weatherproofing options tested in this study. (A) the PVC weatherproofing option, and (B) the
BatHat weatherproofing option. Both views are lateral views.
Discussion to conduct species identification or compare their results

to studies where other weatherproofing designs or
Significant differences among the treatment groups orientations were used. Future studies using passive
suggest that weatherproofing and detector orientation sampling with the Anabat system should include
may impact the detection of bat echolocation calls. In discussion of the method of deployment and the
particular, these data suggest that studies employing the potential impact on the results.
BatHat system we used may detect lower activity and We measured a variety of parameters to test the
species richness than are present at a site. If researchers effects of orientation and weatherproofing on detector
are only interested in relative activity levels among sites, performance. The total number of call sequences or files
any weatherproofing or orientation may be acceptable is often used as a measure of overall activity (e.g., Hayes
as long as detectors are deployed in a similar way among 1997), while the number of pulses per sequence can
all sites. However, problems may arise if researchers want provide a measure of the intensity of activity (Gorresen et
Figure 2. Echolocation sequences and pulses recorded by Anabat II detectors at three orientations and two weatherproofing
types in Kentucky, May and June 2008. (A) the total number of files, (B) average number of pulses per sequence, and (C) percent of
bat calls surviving identification (ID) filter. Bars with different letters above them were found to be significantly different from each
other using the Tukey pair-wise comparisons.
Figure 3. Proportion of sites at which each bat species was detected by Anabats at three orientations and two weatherproofing
designs. PVC = polyvinyl chloride tube. Big brown Eptesicus fuscus, eastern red Lasiurus borealis, tri-colored Perimyotis subflavus,
northern long-eared Myotis septentrionalis, Indiana M. sodalis, little brown M. lucifugus.
al. 2008), and the percentage of calls surviving the detected at as far a distance by the detectors as higher
identification filter is a measure of the quality of the intensity calls). Knowledge of the impacts of weather-
recordings. Factors affecting the detection, quality, or proofing and orientation on detector performance allows
length of sequences all have large impacts on the results researchers the opportunity to deploy equipment in such
of studies using Anabat detectors or other acoustic a way as to maximize success or to deploy additional
sampling equipment. The consistent pattern of differ- equipment to gain a more accurate representation of
ences across all parameters suggests that the differences species present (Duchamp et al. 2006).
among treatments are real. When conducting echolocation surveys for bats,
Weller and Zabel (2002) found no difference between multiple sampling sites may be required. Many detectors
detectors oriented at 30u or 45u if they were on 1.4-m- are often deployed throughout the project area to record
high stands. However, we found that horizontal deploy- simultaneously. Similarly, studies designed to test the
ment tended to record lower (not statistically significant) effects of habitat type or management activities on bat
activity levels (number of files) and species richness than habitat use and activity often set detectors simulta-
the other orientations. The appropriate orientation may neously in each habitat type or treatment to control for
depend on the question of interest. For example, if a the effects of temporal variation on activity (e.g., Loeb
researcher is interested in recording bats that forage near and Waldrop 2008). These situations require the use of
the water surface, such as gray bats M. grisescens, weatherproofing equipment because detectors are
horizontal deployment can be expected to be better widely scattered across the landscape and sudden
than other orientations. storms are possible. Our results suggest that detections
Six species were detected across the five treatments, obtained from detectors at two common orientations
but no treatment detected all six species. The PVC, 45u, without weatherproofing (45u and 90u) are similar to
and vertical (90u) treatments consistently recorded those from detectors with the PVC weatherproofing,
almost twice as many species as the BatHat. Further, indicating that data can be compared across studies
the number of species recorded by detectors in the using any of these methods.
horizontal position tended to be lower than the This investigation was not meant to test all possible
detectors at 45uand 90u, even though the number of configurations, but was instead focused on the compar-
calls recorded by detectors in the horizontal orientation ison of the most common orientations and weather-
was just slightly below the number of calls recorded by proofing options. The impacts of the orientation and
detectors in the vertical (90u) position. This may be due weatherproofing options likely vary with local site
to species use of the habitats or the intensity of the conditions and the bat community present. For example,
echolocation calls (e.g., low-intensity calls are not Gruver et al. (2009) found that the number of bat passes
recorded by the BatHat was greater than the number Kennedy J, editors. The Indiana bat: biology and
recorded by a PVC setup in one area but found no management of an endangered species. Austin, Texas:
differences in the number of passes recorded by the two Bat Conservational.
systems in another area. Results of this study should not Duchamp JE. 2006. Modeling bat community structure
necessarily be applied to other modifications of these and species distribution across fragmented land-
weatherproofing options, but instead should illustrate scapes within the upper Wabash River basin.
the potential impacts of orientation and weatherproof-
Doctoral dissertation. West Lafayette, Indiana: Pur-
ing options on the quality and quantity of data obtained.
due University.
Knowledge of the impacts or weatherproofing and
orientation should assist in appropriate use of data Duchamp JE, Yates M, Muzika R-M, Swihart RK. 2006.
obtained by maximizing the quality of data obtained Estimating probabilities of detection for bat echolo-
through studies involving acoustic surveys, thereby cation calls: an application of the double-observer
improving our knowledge of the impacts of manage- method. Wildlife Society Bulletin 34:408–412.
ment activities on bats. Gorresen PM, Miles AC, Todd CM, Bonaccorso FJ, Weller
TJ. 2008. Assessing bat detectability and occupancy
Acknowledgments with multiple automated echolocation detectors.
Journal of Mammalogy 89:11–17.
This study was supported by the Kentucky Department
of Fish and Wildlife Resources and the U.S. Fish and Gruver JC, Solick DI, Sonnenberg MK. 2009. Comparison
Wildlife Service, Kentucky Ecological Service Field Office. of bat activity recorded using different acoustic
We thank Aric Payne and Corey Knight, who helped with sampling equipment: implications for study design
data collection, and the numerous landowners who at wind-energy facilities. Bat Research News 50:109.
provided access to their lands. Jim Barnard provided the Hayes JP. 1997. Temporal variation in activity of bats and
illustrations of the two weatherproofing options. Erin Gillam, the design of echolocation-monitoring studies. Jour-
two anonymous reviewers, and the Subject Editor improved nal of Mammalogy 78:514–524.
the quality of the manuscript through their comments. Larson DJ, Hayes JP. 2000. Variability in sensitivity of
Anabat II bat detectors and a method of calibration.
References Acta Chiropterologica 2:209–213.
Arnett EB, Hayes JP, Huso MMP. 2006. An evaluation of Loeb SC, Waldrop TA. 2008. Bat activity in relation to fire
the use of acoustic monitoring to predict bat fatality and fire surrogate treatments in southern pine stands.
at a proposed wind facility in south-central Pennsyl- Forest Ecology and Management 255:3185–3192.
vania. An annual report submitted to the Bats and Murray KL, Britzke ER, Hadley BM, Robbins LW. 1999.
Wind Energy Cooperative. Austin, Texas: Bat Conser- Surveying bat communities: a comparison between
vation International. Available: http://batsandwind. mist nets and the Anabat II bat detector system. Acta
org/pdf/ar2005.pdf (October 2010). Chiropterologica 1:105–112.
Britzke ER. 2003. Use of ultrasonic detectors for acoustic O’Farrell MJ. 1998. A passive monitoring system for
identification and study of bat ecology in the eastern Anabat II using a laptop computer. Bat Research News
United States. Doctoral dissertation. Cookeville: Ten- 39:147–150.
nessee Technological University. O’Farrell MJ, Gannon WL. 1999. A comparison of acoustic
Britzke ER, Murray KL. 2000. A quantitative method for versus capture techniques for the inventory of bats.
selection of identifiable search-phase calls using the Journal of Mammalogy 80:24–30.
Anabat system. Bat Research News 41:33–36. Weller TJ, Zabel CJ. 2002. Variation in bat detections due
Britzke ER, Murray KL, Heywood JS, Robbins LW. 2002. to detector orientation in a forest. Wildlife Society
Acoustic identification. Pages 220–224 in Kurta A, Bulletin 30:922–930.
Notes
A Comparison of Mercury Contamination in Mussel and
Ammocoete Filter Feeders
Jamie B. Bettaso, Damon H. Goodman*
U.S. Fish and Wildlife Service, Arcata Field Office, 1655 Heindon Road, Arcata, California 95521-4573
Abstract
The Trinity River, California, has an extensive history of gold and mercury mining, and recent studies demonstrated
uptake of legacy mercury contamination by teleost fishes. We investigated mercury concentrations of larval lampreys
(ammocoetes; Entosphenus spp.) and western pearlshell mussels (Margaritifera falcata) in the Trinity River to determine
whether these two long-lived and sedentary filter feeders show site-specific differences in uptake of this contaminant.
We analyzed ammocoetes from four sites in the Trinity River and one reference site in the Mad River for total mercury
and mussels from three of the Trinity River sites for total and methyl mercury. We identified longitudinal gradients in
ammocoete total mercury levels and methyl mercury in mussels. We found a 70% increase in total mercury in
ammocoetes between two of the sample sites, suggesting a potential point source of contamination. Ammocoetes
contained levels of mercury 12 to 25 times those of mussels from the same site. Our data indicate that ammocoetes
may be a preferred organism to sample for mercury contamination and ecological effects compared with mussels in
the Trinity River.
Keywords: mercury; Trinity River; lamprey; ammocoete; western perlshell mussel; mussel
Received: November 7, 2009; Accepted: August 2, 2010; Published Online Early: August 2010; Published: November
2010
Citation: Bettaso JB, Goodman DH. 2010. A comparison of mercury contamination in mussel and ammocoete filter
feeders. Journal of Fish and Wildlife Management 1(2):142–145; e1944-687X. doi:10.3996/112009-JFWM-019
* Corresponding author: damon_goodman@fws.gov
Introduction issue a health advisory for consumption of Trinity River

fish upstream of Lewiston Dam (Klasing et al. 2005).
The Trinity River, California, has an extensive history of Focusing on taxa within a specific feeding guild
gold and mercury mining dating from the 1850s through provides the opportunity to elucidate the impacts of
the 1950s (Holmes 1965; Alpers et al. 2005). The mercury heavy metal contamination on biota in relation to life
introduced into the Trinity River system has led to history strategies such as filter feeding. In this study, we
varying concentrations of mercury in sediments and evaluate the levels of mercury in two filter feeders,
biota. Reconnaissance surveys of mercury concentrations western pearlshell mussels (Margaritifera falcata; hereaf-
in game fish by May et al. (2005) demonstrated elevated ter mussels) and larval lampreys (Entosphenus spp.;
levels in the watershed. Mercury can pose a threat to hereafter ammocoetes), at collocated sample sites.
individual organisms via lethal and sublethal impacts Several studies have demonstrated that mussels and
such as immunosuppression, teratogenesis, and endo- ammocoetes uptake mercury in areas of environmental
crine disruption (summary in Wiener and Spry 1996). At exposure (Mallatt et al. 1986; Malley et al. 1996; Renaud
an ecosystem level, bioaccumulation of mercury in et al. 1998; Haas and Ichikawa 2004), but these filter
aquatic biota can result in biomagnification in higher feeders have never been evaluated in a comparative
order predators and is of particular human health study. The objectives of this study were to 1) assess
concern when such organisms are used as a food source. mercury concentrations in mussels and ammocoetes in
Information on mercury contamination collected by the the Trinity River and 2) examine the distribution of
U.S. Geological Survey has led the state of California to mercury concentration in mussels and ammocoetes at
Mercury in Filter Feeders J.B. Bettaso and D.H. Goodman
Figure 1. Map of the Trinity River study area and collection Figure 2. Total mercury (THg) concentrations in ammocoetes
locations. Ammocoetes were collected at sites 1–5; mussels and mussels and methyl mercury (MeHg) concentrations in
were collected at sites 2–4. mussels by river mile in the Trinity River. THg is reported as mg/
g wet weight and MeHg is reported as ng/g wet weight. River
mile represents the river distance from the confluence of the
four sites within the upper Trinity River Basin and one Trinity and Klamath rivers at Weitchpec. Lewiston Dam is
reference site in the Mad River. located at river mile 110 and the confluence of Canyon Creek
The use of filter feeders in this study has two and the upper extent of Hocker Flat restoration site is at river
advantages over the previously studied salmonids, mile 78. Arrow represents direction of river flow towards the
centrarchids, and macroinvertebrates in the basin (May Pacific Ocean.
et al. 2005): 1) mussels and ammocoetes have extended
freshwater residencies, with life spans of ,60–100 and wet weight in mg/g (ppm) for total mercury or ng/g (ppb)
,4–6 y, respectively, providing a longer period of time for methyl mercury.
for bioaccumulation (Close 2002; Nedeau et al. 2005);
and 2) mussels feed primarily from the water column
Results
attached to the surface substrate, whereas ammocoetes
feed near the water–sediment interface from burrows in Concentrations of total mercury in ammocoetes col-
soft sediments, providing two different mediums of lected from the four sites in the Trinity River ranged from
potential mercury exposure on a site specific basis 0.379 to 0.882 mg/g wet weight (Table 1). All Trinity River
(Hardisty and Potter 1971). ammocoete samples had greater concentrations of total
mercury than the 0.291 mg/g measured at the reference
Methods site on the Mad River. Total mercury concentrations in
mussels collected at the three sites on the mainstem
We collected ammocoetes at three sites distributed Trinity River below Lewiston Dam ranged from 0.030 to
below Lewiston Dam, one site above the Trinity 0.036 mg/g. Methyl mercury levels in mussel samples
Reservoir, and one reference site on the Mad River ranged from 8.0 to 10.0 ng/g. The percentage of methyl
(Figure 1). In the study area, identification of ammo- mercury in mussels was 22% of total mercury at site 2, 31%
coetes to species is not possible using morphological at site 3, and 29% at site 4. Total mercury concentrations in
characters; therefore, ammocoetes were identified to ammocoete samples varied longitudinally in the Trinity
genus following Goodman et al. (2009). Mussel samples River (Figure 2). Concentrations increased in all down-
were collected at the three sites on the Trinity River stream samples, particularly between sample sites 3 and 4,
below Lewiston Dam. We collected mussels by snorkel- where a 70% increase was observed. An increasing
ing and ammocoetes by using a backpack electroshock- downstream trend was also apparent in the methyl
ing unit (Smith-Root, Inc., Vancouver, WA). Sample sizes mercury concentrations in mussels, but this relationship
were 50 mussels and 5 ammocoetes at each site. All was not mirrored in the total mercury concentrations in
samples were collected in March 2007 and composited mussel samples, which were similar among the three sites.
by site. We submitted samples to the California Statistical significance of these differences or trends was
Department of Fish and Game, at the Moss Landing not tested due to small sample sizes.
Marine Pollution Studies Laboratory. All samples were
analyzed for total mercury by cold-vapor atomic
Discussion
absorption spectrometry following EPA Method 7473
(EPA 1998). We assumed that total mercury in ammo- Overall, total mercury concentrations were more than an
coetes is .95% methyl mercury (Bloom 1992); however, order of magnitude higher in ammocoetes than in mussels
mussels were also analyzed for methyl mercury by EPA collected at the three mainstem locations on the Trinity
Method 1630 (EPA 2001). Analyses included appropriate River below Lewiston Dam. Similar differences in mercury
blanks, duplicates, and spikes for quality assurance and accumulation were found between ammocoetes and
quality control purposes, and all results are reported as mussels in the Châteauguay River in Quebec, Canada,
Table 1. River, sample site, river mile, date of collection, coordinates, sample size (n), and total mercury (THg), methyl mercury
(MeHg) concentrations, and percentage of moisture in ammocoetes and mussels. Sample site numbers correspond to numbers on
Figure 1. River mile represents the river distance from the confluence of the Trinity and Klamath rivers at Weitchpec. THg is reported
as mg/g wet weight and MeHg is reported as ng/g wet weight.a
Ammocoetes Mussels
UTM (NAD 83)
River % % %
River Site mile Date N E n Moisture THg n Moisture THg Moisture MeHg
Trinity 1. Above Trinity Reservoir 135 March 12, 4544870 525432 7 67.8 0.379
2007
2. Rush Creek 108 March 13, 4507758 514504 9 77.9 0.466 50 86.4 0.036 88.6 9.0
2007
3. Junction City Hole 79 March 13, 4508605 494769 6 70.8 0.520 50 90.4 0.030 86.3 9.3
2007
4. Bagdad 72 March 14, 4513175 489465 5 80.6 0.882 50 88.0 0.035 87.5 10.0
2007
Mad 5. Near water treatment NA March 19, 4529094 411244 7 66.3 0.291
facility 2007
a
UTM (NAD 83), Universal Transverse Mercator North American Datum 1983; NA, not applicable.
where ammocoetes had 5 times higher concentrations sampled, with the furthest downstream sample showing
compared with mussels (Renaud et al. 1998). In an a markedly higher concentration. This trend may be
ontogenetic study of sea lamprey (Petromyzon marinus) indicative of chronic mercury contamination in the
in the Connecticut River, New England, ammocoetes were system. A 70% increase in contamination between the
found to have high mercury concentrations for their two most downstream sample locations was observed
trophic level compared with teleosts (Drevnick et al. 2006). (approximately 7 miles), indicating a possible point
Similarly, increased mercury concentrations in ammo- source of contamination. Our data cannot resolve the
coetes compared with other fish species surveyed have source of this contamination. However, several potential
been detected in several watersheds in California where sources exist between the sample sites, including a major
they exhibited order of magnitude higher levels of total tributary (Canyon Creek) with a history of mining, historic
mercury at several sites (Haas and Ichikawa 2004; Haas and mining along the Trinity River main channel, and the
Morrison 2004). Concentrations of total mercury in Hocker Flat Rehabilitation site (constructed in 2005),
rainbow trout (Oncorhynchus mykiss) on the Trinity River which includes large dredge pilings (Rytuba et al. 2005).
averaged 0.0327 mg/g (n = 10) at Big Flat and 0.0499 mg/g These results indicate that further localized sampling is
(n = 10) at Hayden Flat; therefore, concentrations in our warranted to determine the source of the contamination.
ammocoete samples were an order of magnitude higher Elevated concentrations of total mercury in ammo-
than those in salmonids collected at nearby locations by coetes pose two potential threats: 1) adverse health
May et al. (2005). Our results are similar those reported in effects to the individual ammocoetes and 2) ecosystem
the foregoing studies; however, differences in concentra- effects on ammocoete predators through bioaccumula-
tions were more extreme in that ammocoetes had 12 to 25 tion. All of the ammocoete samples evaluated in this
times higher levels of total mercury compared with study had mercury concentrations considered detrimen-
mussels from the same site, indicating that mussels may tal to early life stages of fish (0.2 mg/g; Beckvar et al.
accumulate less mercury than ammocoetes in the same 2005). In addition, mercury is known to biomagnify
location. across trophic levels in aquatic systems (Watras and
The difference in total mercury concentrations be- Bloom 1992). Our data indicate that ammocoetes are
tween mussels and ammocoetes may be related to better bioindicators of mercury contamination than
differences in life history strategies because mercury mussels in the Trinity River and elsewhere when the
more readily accumulates in sediments compared with goal is to evaluate food web biomagnification and
the water column (Boudou and Ribeyre 1997). Freshwa- differentiate sources of contamination. We recommend
ter mussels reside primarily above sediments where they further investigation of mercury contamination in the
Trinity River system by using ammocoetes. Additional
filter feed from the water column, whereas ammocoetes
sampling could further refine the apparent longitudinal
typically reside in subsurface burrows in direct contact
trend, as well as temporal trends of mercury contamina-
with the substrate and filter feeder microflora and fauna
tion in the system by repeating the current sample sites,
near the sediment interface (Hardisty and Potter 1971;
adding additional sites, and sampling seasonally.
Potter 1980).
Increased accumulation of total mercury levels at
Acknowledgments
successive downriver sites was apparent in ammocoetes;
a similar pattern occurred in methyl mercury levels in the We thank Charles Chamberlain, Jim Haas, Jason May,
mussels. The ammocoete samples indicated a gradual Toby McBride, and Joe Polos for reviewing the report and
increase of total mercury concentrations from above the providing many useful comments. This project was
Lewiston and Trinity dams to the downstream sites funded through the Arcata Fish and Wildlife Service
Office in California. We also thank the Subject Editor and MW, Potter IC, editors. Biology of lampreys, vol. 1. New
anonymous reviewers for thoughtful comments and York: Academic Press.
contributions to this manuscript. Holmes GW Jr. 1965. Mercury in California. Pages 87–206
in U.S. Bureau of Mines, editor. Mercury potential of the
References United States. Washington, D.C: U.S. Department of the
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Mercury contamination from historical gold mining in Klasing S, Brodberg R, Gassel M, Roberts S. 2005. Health
California. U.S. Geologic Survey fact sheet 2005-3014 advisory: safe eating guidelines for fish from Trinity
version 1.1. Available: http://pubs.usgs.gov/fs/2005/ Lake, Lewiston Lake, Carrville Pond, the Trinity River
3014/. upstream from Trinity Lake and the East Fork Trinity
Beckvar N, Dillon TM, Read LB. 2005. Approaches for River (Trinity County). Sacramento, California: California
linking whole-body fish tissue residues of mercury or Environmental Protection Agency. Available: www.
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Toxicology and Chemistry 24:2094–2105. pdf.
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fish and marine invertebrate tissue. Canadian Journal of tions in fishes from selected water bodies in Trinity
Fisheries and Aquatic Sciences 49:1010–1017. County, California, 2000–2002. Sacramento, California:
Boudou A, Ribeyre F. 1997. Mercury in the food web: U.S. Geological Survey. Open-file report 2005-1321.
accumulation and transfer mechanisms. Pages 289– Available from the corresponding author upon request.
319 in Sigel A, Sigel H, editors. Metal ions in biological Mallatt J, Barron MG, McDonough C. 1986. Acute toxicity
systems: mercury and its effects on environment and of methyl mercury to the larval lamprey, Petromyzon
biology. New York: Marcel Dekker. marinus. Bulletin of Environmental Contamination and
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Notes
Long-Term Effects of Large Woody Debris Addition on
Stream Habitat and Brook Trout Populations
John A. Sweka,* Kyle J. Hartman, Jonathan M. Niles
J.A. Sweka
U.S. Fish and Wildlife Service, Northeast Fishery Center, P.O. Box 75, 308 Washington Avenue, Lamar,
Pennsylvania 16848
K.J. Hartman, J.M. Niles
West Virginia University, Division of Forestry, 320 Percival Hall, Morgantown, West Virginia 26506-6125
Abstract
In this study, we resurveyed stream habitat and sampled brook trout Salvelinus fontinalis populations 6 y after large
woody debris additions to determine long-term changes in habitat and brook trout populations. In a previous study,
we added large woody debris to eight streams in the central Appalachians of West Virginia to determine whether
stream habitat could be enhanced and brook trout populations increased following habitat manipulation. The large
woody debris additions had no overall effect on stream habitat and brook trout populations by 6 y after the additions.
The assumption that a lack of large woody debris is limiting stream habitat and brook trout populations was not
supported by our results. In high-gradient streams, habitat complexity may be governed more by the abundance of
boulders and large woody debris may have a lesser influence on trout populations.
Keywords: brook trout; large woody debris; habitat; stream

Received: January 20, 2010; Accepted: May 24, 2010; Published Online Early: June 2010; Published: November 2010
Citation: Sweka JA, Hartman KJ, Niles JM. 2010. Long-term effects of large woody debris addition on stream habitat and
brook trout populations. Journal of Fish and Wildlife Management 1(2):146–151; e1944-687X. doi: 10.3996/012010-
JFWM-002
* Corresponding author: john_sweka@fws.gov
Introduction habitat manipulation and found no significant changes

following the addition of LWD. Brook trout populations
The abundance of stream-dwelling salmonids is often fluctuated from year to year, but there was no net
correlated with the abundance of large woody debris increase in density. In this paper, we compare data
(LWD [Fausch and Northcote 1992; Berg et al. 1998; collected in 2006 (6 y postmanipulation) to that collected
Neumann and Wildman 2002]). However, manipulative in 2003 (3 y postmanipulation), the last year of our
studies designed to test such correlations have often previous study, to determine long-term changes to
shown mixed results (Solazzi et al. 2000; Roni and Quinn stream habitat and brook trout populations due to
2001; Warren and Kraft 2003; Sweka and Hartman 2006). LWD additions. Monitoring of habitat manipulations on
The lack of a significant influence of LWD manipulations this temporal scale is rare in the literature (except see
on salmonid populations may be due to pseudo- Gowan and Fausch 1996a), but is needed to evaluate the
replicated studies (e.g., Cederholm et al. 1997), relatively effectiveness of habitat manipulations in achieving
small spatial scales of habitat manipulation, or post- desired habitat and fish population responses.
manipulation monitoring that is not long enough to
detect a population change (Hunt 1976, 1988; Sweka and Materials and Methods
Hartman 2006).
In a previous study, Sweka and Hartman (2006) We added LWD to eight tributaries of the Middle Fork
assessed stream habitat and brook trout Salvelinus River, Randolph County, West Virginia (see Sweka and
fontinalis populations for 1 y prior to, and 3 y after, Hartman 2006 for complete details). The streams were
Brook Trout and Large Woody Debris J.A. Sweka et al.
first- and second-order with temperatures rarely exceed- years. We also classified the function of each added piece
ing 20uC with brook trout as the dominant fish species. in the same manner as naturally occurring LWD.
We divided the study into two experiments: a within- Brook trout populations were estimated by multiple-
stream experiment and an among-stream experiment. pass removal electrofishing in a randomly chosen 100-m
In the within-stream experiment, four streams had LWD section of each 300-m reach (both control and treat-
added to a single 300-m stream reach with additional ment reaches). Electrofishing occurred in the autumn
300-m reaches, upstream and downstream of the (October–November) of 2006. Population estimates were
manipulated reach, serving as controls. Treatment generated using the program CAPTURE (White et al.
reaches were separated from control reaches by 1982) and density was estimated as the population
100 m. We compared stream habitat and brook trout estimate divided by the area of the electrofishing site.
densities between controls and manipulated reaches. We compared our 2006 habitat results to those from
The among-stream experiment compared habitat and 2003 to determine whether any changes had occurred
brook trout densities from the four streams used in the since our previous study (Sweka and Hartman 2006).
within-stream experiment to four additional streams Brook trout density data were compared to those found in
where LWD was added to three 300-m reaches the autumn 2002, because our previous study ended in
(separated by 100 m). The purpose of the among- the spring 2003. Total number of pieces of LWD, total
stream experiment was to determine the effect of the number of pools, total pool area, and brook trout density
scale of habitat manipulation upon stream habitat and were analyzed as a repeated-measures analysis of variance
brook trout density. using the PROC MIXED procedure in SASH version 9.1. We
A logging crew added LWD by felling trees into the were specifically interested in the treatment 6 time
streambed at a rate of approximately 15 trees/300-m interaction in these tests to determine whether any
stream length. Each added piece of LWD was individually changes had occurred between treatments since 2003.
tagged to assess movement and functionality of The function of the added LWD was compared between
individual pieces between years. The felled trees had a 2003 and 2006 using a chi-square test.
minimum diameter of 10 cm and median diameter of
22 cm (Sweka and Hartman 2006). We assessed stream Results
habitat once prior to the addition of LWD (summer 2000)
and three times following LWD addition (summers 2001– The majority of the added LWD remained within the
2003). Brook trout populations were assessed in the study areas and continued to serve some function by 2006
autumn and spring of each year. Pretreatment brook (Supplemental Material, Table S1, http://dx.doi.org/
trout population assessment occurred in the autumn 10.3996/012010-JFWM-002.S1). Three hundred fifty-four
1999 and spring 2000, and posttreatment brook trout pieces of LWD were added to all the streams in 2000
population assessment occurred from autumn 2000 (Sweka and Hartman 2006). By 2006, 304 (86%) were
through spring 2003. found, although 107 (30%) pieces moved . 5 m from their
We used the same methodology in 2006 for assessing original location in 2000. Of the added pieces, 11% were
stream habitat and brook trout populations as in our involved in pool formation, 38% stored organic material,
previous study (Sweka and Hartman 2006). Briefly, 7% stored sediment, 9% provided overhead cover for
habitat was surveyed under summer base-flow condi- brook trout, 2% were involved in bank stabilization, and
tions and individual habitat units were classified as riffles, 33% did not serve any apparent function under summer
runs, or pools. We measured the length of each habitat base-flow conditions. The frequencies of these functions
unit along the thalweg, and wetted and bank-full widths were not significantly different from those in 2003 (x2 =
were visually estimated at transects of 0.25, 0.50, and 5.84, df = 5, P = 0.32). The frequency of the added LWD
0.75 of the thalweg length. The widths of 20% of the involved in pool formation continued to be greater than
habitat units were estimated and then verified by that of naturally occurring LWD (11% versus 2%), which is
measuring to the nearest 0.1 m. The area of each habitat likely due to the large size of the added pieces compared
unit was calculated as the mean width multiplied by the to naturally occurring pieces (Sweka and Hartman 2006).
length. Verified area was regressed on estimated area Although the majority of added LWD was retained,
and predicted values of the regression equation were stream habitat did not significantly change between 2003
used for the area of estimated habitat units. and 2006 in the within-stream experiment. Treatment
We considered LWD to be any piece of wood within reaches in the within-stream experiment continued to
the bank-full channel with a minimum diameter of 10 cm have an overall greater amount of total LWD (natural +
and minimum length of 1 m. Pieces were classified added pieces [Figure 1a; F1,7 = 19.08, p , 0.01]) than
according to the size criteria proposed by Flebbe and control reaches, but there was no significant interaction
Dolloff (1995) and according to their function within the with time (F1,10 = 0.63, P = 0.44). Reaches where LWD was
stream channel (pool formation, sediment storage, added tended to have a greater number of pools
organic storage, overhead cover, bank stabilization). (Figure 2a; F1,7 = 4.64, P = 0.07) and pool area/300 m
The location of each added piece of LWD was (Figure 2b; F1,7 = 5.09, P = 0.06) compared to control
determined by measuring the distance of each piece in reaches, but these differences were not significant, and
relation to the upstream or downstream end of each there was no time-dependent treatment effect for the
300-m study reach. Movement of added pieces of LWD number of pools/300 m (F1,10 = 2.72, P = 0.13) or pool
was estimated as the difference in locations between area/300 m (F1,10 = 0.15, P = 0.71).
Figure 1. Total abundance of large woody debris (LWD) in

the within-stream experiment conducted in tributaries of the
Middle Fork River, Randolph County, West Virginia (2003 and Figure 2. Number (a) and area (b) of pools in the within-
2006; Supplemental Material, Table S2, http://dx.doi.org/ stream experiment conducted in tributaries of the Middle Fork
10.3996/012010-JFWM-002.S2). Panel (a) corresponds to total River, Randolph County, West Virginia (2003 and 2006;
amounts of LWD (both natural and added), and panel (b) Supplemental Material, Table S3, http://dx.doi.org/10.3996/
corresponds to total amounts of LWD in the largest size classes 012010-JFWM-002.S3). Open bars correspond to control reach-
(. 10 cm diameter and . 5 m length; Flebbe and Dolloff 1995). es and shaded bars correspond to reaches where large woody
Open bars correspond to control reaches and shaded bars debris was added to 300 m of stream. Error bars represent
correspond to reaches where LWD was added to 300 m of standard errors.
stream. Error bars represent standard errors.
The added LWD continued to have no effect on age 1+

Stream habitat also did not significantly change in the
brook trout density by 2006. In the within-stream
among-stream experiment. Streams that had LWD added
experiment, age 1+ brook trout density showed an
to three 300-m reaches did not have significantly more
overall decrease between 2002 and 2006 (Figure 5; F1,10
total LWD (total LWD = natural LWD + added LWD) than
= 13.35, P , 0.01), but there was no overall difference
those where LWD was added to a single 300-m reach
between reaches where LWD was added and control
(Figure 3a; F = 1.86, P = 0.22). However, when
reaches (F1,7 = 4.50, P = 0.07) and no differential change
considering the largest size classes of LWD (diameter .
between treatments with time (F1,10 = 0.08, P = 0.78).
10 cm and length . 5 m; Flebbe and Dolloff 1995), the
Likewise, age 1+ brook trout density decreased between
streams where LWD was added to three 300-m reaches
2002 and 2006 in the among-stream experiment
continued to have significantly more of this large size
(Figure 6; F1,6 = 9.26, P = 0.02), but there was no
class of LWD than streams where LWD was added to only
overall effect of the scale of habitat manipulation (F1,6 =
a single 300-m reach (Figure 3b; F1,6 = 7.93, P = 0.03).
1.07, P = 0.34) or a differing effect of scale with time (F1,6
There was also no differential change between treat-
= 0.14, P = 0.72). The overall mean age 1+ brook trout
ments through time in the among-stream experiment for
density in all eight streams in 2006 was 2.96 6 1.38
total LWD (F1,6 = 0.25, P = 0.63) or the largest size
fish/100 m2 (6 95% CI) while it was 5.85 6 3.83 fish/
classes of LWD (F1,6 = 0.55, P = 0.49). In the among-
100 m2 in 1999 when our previous study began (Sweka
stream experiment, pool number tended to decrease
and Hartman 2006). Thus, the addition of LWD did not
between 2003 and 2006, but this was not significant
increase brook trout populations in these streams.
(Figure 4a; F1,6 = 4.44, P = 0.08), and there was no
overall effect of the scale of habitat manipulation (F1,6 =
Discussion
0.14, P = 0.72) or a differing effect of scale with time (F1,6
= 3.60, P = 0.11). Pool area also decreased significantly Long-term monitoring of trout population response to
between 2003 and 2006 in all streams (Figure 4b; F1,6 = habitat manipulation is rare in the literature. Hunt (1976,
12.82, P = 0.01), but there was no overall effect of the 1988) suggested that population response may take
scale of habitat manipulation (F1,6 = 0.11, P = 0.75) or a greater than 5 y postmanipulation and we also partly
differing effect of scale with time (F1,6 = 3.30, P = 0.12). attributed the lack of a sustained increase in brook trout
Figure 3. Total abundance of large woody debris (LWD) in

the among-stream experiment conducted in tributaries of the
Middle Fork River, Randolph County, West Virginia (2003 and Figure 4. Number (a) and area (b) of pools in the among-
2006; Supplemental Material, Table S2, http://dx.doi.org/ stream experiment conducted in tributaries of the Middle Fork
10.3996/012010-JFWM-002.S2). Panel (a) corresponds to total River, Randolph County, West Virginia (2003 and 2006;
amounts of LWD (both natural and added), and panel (b) Supplemental Material, Table S3, http://dx.doi.org/10.3996/
corresponds to total amounts of LWD in the largest size classes 012010-JFWM-002.S3). Open bars correspond to streams where
(. 10 cm diameter and . 5 m length; Flebbe and Dolloff 1995). large woody debris (LWD) was added to a single 300-m reach
Open bars correspond to streams where LWD was added to a and shaded bars correspond to streams where LWD was added
single 300-m reach and shaded bars correspond to streams to three 300-m reaches. Error bars represent standard errors.
where LWD was added to three 300-m reaches. Error bars
represent standard errors.
addition of log structures in Colorado streams, but
density in our original study (Sweka and Hartman 2006) Gowan and Fausch (1996a) attributed this increase to
to the lack of time since the LWD was added. Even with immigration from outside their study reaches rather than
monitoring of populations for 6 y following habitat an increase in recruitment and survival. Likewise, Warren
manipulation, we still did not observe an increase in and Kraft (2003) did not observe a decrease in brook
brook trout density. Brook trout densities were actually
lower in 2006 than in 2002. Pool habitat in the among-
stream experiment decreased in both scales of habitat
manipulation from 2003 to 2006, which may be one
potential explanation for decreasing brook trout densi-
ties in the among-stream experiment. However, pool
area remained the same in the treatment and control
reaches in the within-stream experiment and yet brook
trout densities still declined. Thus, the cause of the
decrease in brook trout density from 2002 to 2006
appears independent of the LWD manipulations and
resulting effects on stream physical habitat. Brook trout
populations of the Appalachian region commonly show
much inter-annual variability (Petty et al. 2005; Sweka
and Hartman 2006), making it difficult to differentiate
variation due to habitat manipulation from variation due Figure 5. Mean brook trout density (#/100 m2) in the
autumn 2002 and 2006 in the within-stream experiment
to other factors.
conducted in tributaries of the Middle Fork River, Randolph
The importance of large woody debris to structuring County, West Virginia (Supplemental Material, Table S4, http://
stream habitat remains a strong tenet of stream dx.doi.org/10.3996/012010-JFWM-002.S4). Open bars corre-
restoration, although manipulative studies have had spond to control reaches and shaded bars correspond to
difficulty showing a population-level effect. Riley and reaches where large woody debris was added. Error bars
Fausch (1995) found trout abundance increased after represent standard errors.
considering such habitat manipulations. For example,

the streams of this study had an average gradient of
. 3% (Sweka and Hartman 2006) with abundant
boulders. In such systems, stream habitat complexity
may be governed more by boulders than by LWD
(Warren and Kraft 2003). Also, the significant correlations
seen in the literature between salmonid abundance and
LWD do not necessarily mean that LWD is a limiting
factor for species such as brook trout and increasing
LWD does not always increase fish abundance.
Supplemental Material
Figure 6. Mean brook trout density (#/100 m2) in the
autumn 2002 and 2006 in the among-stream experiment is not responsible for the content or functionality of any
conducted in tributaries of the Middle Fork River, Randolph supplemental material. Queries should be directed to the
County, West Virginia (Supplemental Material, Table S4, http:// corresponding author.
dx.doi.org/10.3996/012010-JFWM-002.S4). Open bars corre-
spond to streams where large woody debris (LWD) was added Table S1. Functions of added LWD in tributaries of
to a single 300-m reach and shaded bars correspond to streams Middle Fork River, Randolph County, West Virginia in
where LWD was added to three 300-m reaches. Error bars 2003 and 2006.
represent standard errors. Found at DOI: 10.3996/012010-JFWM-002.S1 (9.6 KB
XLSX).
trout relative abundance following the removal of LWD Table S2. Amounts of LWD in tributaries of Middle
from Adirondack mountain streams. Fork River, Randolph County, West Virginia in 2003 and
The underlying assumption in manipulative studies 2006.
with LWD, and in stream restoration projects utilizing Found at DOI: 10.3996/012010-JFWM-002.S2 (11.7 KB
LWD, is that LWD is a limiting factor for stream habitat XLSX).
complexity and stream fish populations. Perhaps the Table S3. Number of pools and area of pool habitats
significant correlations seen between the abundance of in tributaries of Middle Fork River, Randolph County,
LWD and salmonid populations (Fausch and Northcote West Virginia in 2003 and 2006.
1992; Fleebe and Dolloff 1995; Neumann and Wildman Found at DOI: 10.3996/012010-JFWM-002.S3 (9.96 KB
2002) are an artifact of overall past land use within a XLSX).
watershed. For example, disturbed watersheds may have Table S4. Density of brook trout age 1+ in tributaries
had instream structure removed, increased sedimenta- of Middle Fork River, Randolph County, West Virginia in
tion, decreased habitat connectivity, removal of riparian autumn 2002 and 2006.
vegetation, and degraded water chemistry. Conversely, Found at DOI: 10.3996/012010-JFWM-002.S4 (8.94 KB
in undisturbed watersheds all these factors may still be in XLSX).
pristine condition, with accompanying high abundances
of LWD resulting in positive correlations between Acknowledgments
salmonid abundance and LWD abundance. Although We thank the U.S. Forest Service Monongahela National
LWD can increase habitat complexity, the underlying Forest and Northern Research Station, the MeadWest-
geology, stream-channel type (i.e., Rosgen Channel vaco Corporation, West Virginia Division of Natural
types), and abundance of boulders may have more of Resources, and West Virginia University Division of
an influence on stream habitat than LWD in relatively Forestry for funding this research.
high-gradient streams (Warrren and Kraft 2003). Also, We also thank those individuals who aided in data
stream-dwelling salmonids move much more than is collection during our previous study and J. Studinski, J.
typically acknowledged by resource managers (Gowan et Howell, and J. Stolarski for aiding in data collection in
al. 1994; Gowan and Fausch 1996b; Logan 2003) and 2006. This manuscript was improved with preliminary
utilize differing habitats seasonally (Petty et al. 2005). reviews by J. Mohler, M. Millard, and three anonymous
Thus, it is not surprising that manipulative studies that reviewers.
modify stream habitat over a relatively small spatial scale
(i.e., a few hundred meters of stream length) often fail to References
show a population-level effect. Successful habitat resto-
ration needs to consider watershed-scale processes and Berg N, Carlson A, Azuma D. 1998. Function and
the mobility of target species, such as brook trout (Logan dynamics of woody debris in stream reaches in the
2003; Petty et al. 2005), in order to result in population- central Sierra Nevada California. Canadian Journal of
level effects on stream fish. Fisheries and Aquatic Sciences 55:1807–1820.
Although our LWD additions failed to significantly Cederholm CJ, Bilby RE, Bisson PA, Bumstead TW,
modify stream habitat or increase brook trout popula- Fransen BR, Scarlett WJ, Ward JW. 1997. Responses
tions, our results provide valuable information to others of Coho salmon and steelhead to placement of large
woody debris in a coastal Washington stream. North Neumann RM, Wildman TL. 2002. Relationships between
American Journal of Fisheries Management 17:947– trout habitat use and woody debris in two southern
964. New England streams. Ecology of Freshwater Fish 11:
Fausch KD, Northcote TG. 1992. Large woody debris and 240–250.
salmonid habitat in a small coastal British Columbia Petty JT, Lamothe PJ, Mazik PM. 2005. Spatial and
stream. Canadian Journal of Fisheries and Aquatic seasonal dynamic of brook trout populations inhab-
Sciences 49:682–693. iting a central Appalachian watershed. Transactions of
Flebbe PA, Dolloff CA. 1995. Trout use of woody debris the American Fisheries Society 134:572–587.
and habitat in Appalachian Wilderness streams of Riley SC, Fausch KD. 1995. Trout population response to
North Carolina. North American Journal of Fisheries habitat manipulations in six northern Colorado
Management 15:579–590. streams. Canadian Journal of Fisheries and Aquatic
Gowan C, Fausch KD. 1996a. Long-term demographic Sciences 52:34–53.
responses of trout populations to habitat manipula- Roni P, Quinn TP. 2001. Density and size of juvenile
tion in six Colorado streams. Ecological Applications 6: salmonids in response to placement of large woody
931–946. debris in western Oregon and Washington streams.
Gowan C, Fausch KD. 1996b. Mobile brook trout in two Canadian Journal of Fisheries and Aquatic Sciences 58:
high-elevation Colorado streams: re-evaluation of the 282–292.
concept of restricted movement. Canadian Journal of Solazzi MF, Nickelson TE, Johnson SL, Rodgers JD. 2000.
Fisheries and Aquatic Sciences 53:1370–1381. Effects of increasing winter rearing habitat on abun-
Gowan C, Young MK, Fausch KD, Riley SC. 1994. dance of salmonids in two coastal Oregon streams.
Restricted movement in resident stream salmonids: a Canadian Journal of Fisheries and Aquatic Sciences 57:
paradigm lost? Canadian Journal of Fisheries and 906–914.
Aquatic Sciences 51:2626–2637. Sweka JA, Hartman KJ. 2006. Effects of large woody
Hunt RL. 1976. A long-term evaluation of trout habitat debris addition on stream habitat and brook trout
development and its relation to improving manage- populations in Appalachian streams. Hydrobiologia
ment-related research. Transactions of the American 559:363–378.
Fisheries Society 105:361–364. Warren DR, Kraft CE. 2003. Brook trout (Salvelinus
Hunt RL. 1988. A compendium of 45 trout stream habitat fontinalis) response to wood removal from high-
development evaluations in Wisconsin during 1953– gradient streams of the Adirondack Mountains (N.Y.,
1985. Madison: Wisconsin Department of Natural U.S.A.). Canadian Journal of Fisheries and Aquatic
Resources Technical Bulletin No. 162. Available from Sciences 60:379–389.
corresponding author. White GC, Anderson DR, Burnham KP, Otis DL. 1982.
Logan MN. 2003. Brook trout (Salvelinus fontinalis) Capture–recapture and removal methods for sam-
movement and habitat use in a headwater stream of pling closed populations. New Mexico: Los Alamos
the Central Appalachian Mountains of West Virginia. National Laboratory LA-8787-NERP. 235 pp. Available:
Master’s thesis. Morgantown: West Virginia University. http://warnercnr.colostate.edu/,gwhite/software.html;
Available from corresponding author. accessed August 2010.
Surveys
Population Demographics and Dynamics of Colonizing
Elk in a Desert Grassland–Scrubland
Louis C. Bender,* Jessica R. Piasecke
L.C. Bender
U.S. Geological Survey, New Mexico State University, P.O. Box 30003 MSC 4901, Las Cruces, New Mexico 88003
Present address: Extension Animal Sciences and Natural Resources, New Mexico State University, P.O. Box 30003 MSC
3AE, Las Cruces, New Mexico 88003
J.R. Piasecke
Department of Fishery and Wildlife Sciences, New Mexico State University, P.O. Box 30003 MSC 4901, Las Cruces,
New Mexico 88003
Abstract
Elk Cervus elaphus are rare in Southwestern desert grassland and desert-scrub habitats, and these habitats are anecdotally
considered unsuitable for elk. We studied a colonizing herd in a Southwestern desert grassland–scrubland in
northwestern New Mexico to determine the condition and population dynamics of elk in this habitat type. We
radiotracked # 24 adult (1.5 y and older) cows and # 8 calves annually, 2003–2006; the sample of radiocollared cows in
this study was very close to a census of the entire population for 2004–2006 based on repeated surveys and monitoring.
Mean body fat of lactating cows in autumn was 10.6–13.2% in 2003–2004, indicating that lactating elk were acquiring
high moderate–low good nutrition, but dropped to 5.6% in 2005 and 6.8% in 2006, indicating poor nutrition for these
years of low precipitation. We found adult female survival of 0.94–1.00 and calf survival of 0.38–1.00. Calf survival was
related to maternal size, maternal condition, and cumulative annual precipitation through parturition; all calf mortality
occurred at or near parturition. Pregnancy rates averaged 0.33 for yearling cows and 0.86 for $ 2.5-y-olds, 2003–2006,
and pregnancy was positively related to body mass for $ 2.5-y-old adults. Lactation rates of $ 2.5-y-olds averaged 0.59,
2003–2005, but dropped to 0.30 in 2006, due to poor maternal condition and low precipitation. Since colonization, elk
have increased from about 20 to $ 53 individuals, a mean rate of increase of 18%/y prior to 2006, but decreasing to # 4%
in 2006. The high sustained rate of increase (with few exceptions) indicates that desert grassland–scrubland habitats are
suitable for elk and merit consideration in elk management plans of Southwestern agencies.
Keywords: elk; desert Southwest; population dynamics; condition; survival; productivity

Received: October 7, 2009; Accepted: May 20, 2010; Published Online Early: June 2010; Published: November 2010
Citation: Bender LC, Piasecke JR. 2010. Population demographics and dynamics of colonizing elk in a desert grassland–
scrubland. Journal of Fish and Wildlife Management 1(2):152–160; e1944-687X. doi: 10.3996/102009-JFWM-013
* Corresponding author: lbender@nmsu.edu
Introduction Skovlin et al. 2002). These limitations would result in

lower individual condition and consequently reduced
Desert grassland and desert-scrub (hereafter, grass- demographic vigor and population performance
land–scrubland) habitats are anecdotally considered (Hanks 1981; Gaillard et al. 2000; Piasecke 2006;
marginal habitat for elk Cervus elaphus (Skovlin et al. Bender et al. 2008). Despite this, elk have colonized
2002). Presumed detriments of arid habitats include or been introduced into arid grassland or scrubland
scarcity of water, low forage quantity and quality, lack habitats in several areas, and many of these popula-
of vegetative cover, and thermal stress; this potentially tions subsequently grew at rates comparable to elk
results in nutritive deficiencies, dehydration, high populations in montane habitats (McCorquodale et al.
thermoregulatory costs, and increased risk of preda- 1988; Carpenter and Silvy 1991; Strohmeyer and Peek
tion and harvest (Young 1988; Jiang 1993; Sowell 2001; 1996).
Elk Population Dynamics L.C. Bender and J.R. Piasecke
Around January 2000, about 20 elk colonized the Chaco

Culture National Historical Park (CCNHP) area, an arid
grassland–scrubland in northwestern New Mexico. These
elk have subsequently established a productive population
in and around CCNHP, increasing to $ 53 individuals by
2006 (see below). Because Southwestern desert grassland–
scrublands have generally been considered unsuitable for
elk, no current information was available on actual
suitability of these habitats for elk or the potential for elk
populations to increase in these habitats. Consequently,
our goal was to determine population demographics
and growth potential of elk in the Southwestern desert
habitats in and around CCNHP. Specifically, our objectives
were to: 1) determine nutritional condition, 2) determine
population size, composition, and trend, 3) determine
survival and productivity and the environmental factors
influencing them, and 4) model population growth for elk
in CCNHP.
Study Area
Our study area covered about 308 km2 on and
adjacent to CCNHP in northwestern New Mexico,
centered at about 36u009N, 108u009W (Figure 1). Eleva-
tions ranged from 1,670 to 2,079 m, and topography was
characterized by rolling plains and mesas interspersed Figure 1. Location and topographic relief of study area on
with steep ravines. Average high temperature in July was and adjacent to Chaco Culture National Historical Park,
32uC and average low temperature in January and northwest New Mexico.
December was 211uC. Average annual precipitation
was 23 cm, with 52% falling July–October, and average
annual snowfall was 37 cm. using carfentanil citrate (3.0 mg/elk) and xylazine
The dominant plant community on mesas consisted of hydrochloride (100 mg/elk) each April and November,
fourwing saltbush Atriplex canescens, rubber rabbitbrush 2003–2006. We similarly captured a limited number of
Chrysothamnus nauseosus, mountain mahogany Cerco- bulls to aid in population estimation (see below). We
carpus intricatus, winterfat Krascheninnikovia lanata, captured calves by hand if found less than about 5 d
galleta Pleuraphis jamesii, blue grama Bouteloua gracilis, after birth, 2003–2006. We captured calves . 5 d old by
Indian ricegrass Achnatherum hymenoides, and bigelow net-gunning or aerial darting from a Bell 206B Jet Ranger
sage Artemisia bigelovii, with scattered pinyon pine Pinus helicopter, 2004–2006. We blindfolded all captured elk to
edulis and oneseed juniper Juniperus monosperma. minimize stress during handling and administered
Characteristic vegetation of ravines descending from penicillin, vitamin B, vitamin E–selenium (MUSE), and an
mesas included mound saltbush Atriplex obovata, galleta, eight-way Clostridium bacterin to alleviate capture stress.
blue grama, alkali sacaton Sporobolus airoides, bigelow We fit elk with radiocollars (adults, 1.5 y and older) or ear-
sage, winterfat, Mormon tea Ephedra spp., and oneseed tag transmitters (calves; Advanced Telemetry Solutions,
juniper. Vegetation in large washes included fourwing Asanti, MN).
saltbush, galleta, giant dropseed S. giganteus, and sand We collected blood by venal puncture, checked
dropseed S. cryptandrus. Only about 15% of the type of lactation status, and measured chest girth at each
ecosystem represented by CCNHP remains intact in capture. We tested serum for pregnancy-specific placen-
North America, with most perturbation due to grazing tal protein B to determine pregnancy status (Piasecke
(Ricketts et al. 1999). 2006). We checked lactation status of each cow to assess
Ungulates other than elk and mule deer Odocoileus calf survival up to the point of capture (November).
hemionus were rare in CCNHP. Potential predators of elk Lactation indicated successful nursing by a calf within 3–
present on CCNHP include coyotes Canis latrans, pumas 11 d (Bender et al. 2002). We used spring scales or girth
Puma concolor, and bobcats Lynx rufus. Domestic to estimate body mass (kg; Cook et al. 2003), where
livestock including cattle and horses were common on calves = 0.0003901 6 G2.6430078; yearlings = 2.038 6 G
adjacent Tribal, State of New Mexico, and Bureau of Land 2 97.533; and adults = 2.549 6G 2 156.760, where G =
Management holdings. chest girth (cm).
We estimated a rump body condition score (rBCS) by
Methods palpation of the soft tissue of the rump near the base of
the tail and a withers body condition score (wBCS) by
Elk capture and condition palpation of the top of the withers just posterior to the
We captured (and recaptured previously radiocollared) shoulder hump (Cook 2000). We scored results of rBCS
cow elk by darting from a Bell 206B Jet Ranger helicopter and wBCS measures from standards that ranged from
one (emaciated) to five (obese) in intervals of 0.25 June) and lactation (January–September). We used odds
(Cook 2000). We used a SonoVet 2000 ultrasound ratios to interpret results of logistic analyses.
(Medison, Seoul, South Korea) with a 5-mHz probe to
measure subcutaneous fat thickness along a straight Pregnancy and lactation
line midway between the spine, at its closest point to We determined proportions pregnant and propor-
the coxal tuber (hip bone), and the ischial tuber (pin tions lactating by age class (yearlings, $ 2.5-y-olds) by
bone; MAXFAT; Cook 2000). We estimated percent body dividing the number of pregnant cows by the total
fat (BF) by combining rBCS and MAXFAT into an index numbers tested, determined standard errors of esti-
(rLIVINDEX) using: rLIVINDEX = rBCS when MAXFAT , mates using the normal approximation (Zar 1996), and
0.3 cm; and rLIVINDEX = (MAXFAT 2 0.3) + rBCS when used Fisher’s exact tests (Zar 1996) to compare
MAXFAT $ 0.3 cm (Cook et al. 2001). We then proportions among years. We used logistic regression
calculated BF from rLIVINDEX using BF = 27.1527185 (Hosmer and Lemeshow 1989) to model variables
+ 7.323081 6 L 2 0.98980456 6 L2 + 0.057445567 6 L3, related to the probability of an individual elk being
where L = rLIVINDEX (Cook et al. 2001). If MAXFAT was pregnant. Our dichotomous outcome variable was
not measured, we calculated BF directly from rBCS pregnant or not pregnant and our predictors were
using BF = 4.478 6 rBCS 2 4.618. We also used an age, mass, girth, MAXFAT, rBCS, wBCS, loin, and BF, with
ultrasound to measure the thickness of the loin all variables collected from cows during the autumn
(longissimus dorsi) muscle between the 12th and 13th when pregnancy status was assessed.
ribs near the spine to index lean tissue (muscle)
catabolism (Cook et al. 2001). Population size and dynamics
We visually located all radiocollared individuals a We estimated minimum population size from herd
minimum of once per week. When located, we composition surveys during November 2004–2006 cap-
recorded survival status (live or dead) of each elk and ture events. Surveys covered the entire study area and
determined causes of death for all mortalities following were flown two to three times during each capture
Bender et al. (2004). We considered deaths that event. We recorded numbers of bulls, cows, and calves
occurred within 30 d of capture to be capture-related during surveys, and determined variance around ratios
and censored these from survival analyses (Berringer et (bulls/100 cows, calves/100 cows) following Czaplewski
al. 1996). et al. (1983). We compared ratios among years using
parametric bootstrapping (Bender et al. 1996).
Survival We estimated the minimum number of cows in the
We calculated annual survival rates of cows and population by summing the numbers of radiocollared
calves (for pooled radiotagged calves and calves not cows and the highest number of uncollared cows seen
radiotagged but whose dam was radiocollared [i.e., during surveys. Because each individual survey was
calves-at-heel]) using monthly intervals with the stag- accomplished in , 1 d, two or three complete surveys
gered-entry Kaplan–Meier estimator and used Z-tests to occurred during each capture event, groups of elk were
compare survival among years (Pollock et al. 1989). We usually separated by . 5 km, uncollared elk were generally
also used pregnancy status of individual cows and their associated with collared elk, and all collared elk were
lactation status the following autumn to determine calf observed multiple times, we were confident that there was
survival (Bender et al. 2002; i.e., Ŝcalf = Lactatingy / little or no intermixing during surveys and, hence, little
Pregnanty21). This estimate of calf survival incorporates potential for duplication of counts. In addition, we also
calves that may have died shortly after parturition but tallied the number of uncollared cows seen throughout
before we were able to either capture them or observe the summer to corroborate their minimum numbers.
them with their dam, and thus would not have been Minimum calf and bull numbers were determined
included in the radiotagged or calf-at-heel samples. identically. We also determined calving success for each
However, this estimate is potentially confounded by radiocollared cow to corroborate minimum calf numbers.
fetuses that are lost in utero. Because observed in utero Last, we correlated minimum population estimates of all
losses are , 3% for elk (Piasecke 2006), this potential individuals and only cows and calves with pregnancy rate,
positive bias is low. lactation rates, calf survival rates, and calf : cow ratios to
We used logistic regression (Hosmer and Lemeshow detect any trends in population demographics as popu-
1989) to model the dichotomous outcome of calf survival lation size or density increased.
(i.e., live or die) as a function of its mother’s age, girth, We estimated maximum potential finite rate of
mass, MAXFAT, rBCS, wBCS, loin, and BF the previous population increase (l) annually as l = ŜF + 0.5 6 (C/C),
autumn and late winter, which represent the approximate where ŜF = annual survival rate of adult cows and C/C =
seasonal peak (autumn) and low (late winter) in condition, observed calf : cow ratio (White and Bartmann 1997). This
respectively. We also modeled calf survival as a function of was a maximum estimate for l, because our observed
precipitation during several biologically relevant periods, calf : cow ratios (collected in November) did not include
including: winter prior to birth (December–March); late calf mortality from December through the following
gestation period (April–June); lactation period (June– September, when calves would be recruited into the
October); annually based on calendar (January–Decem- adult population. We also estimated mean annual l since
ber) and biological (June–May) years; and cumulative colonization using l = (Ny / Ninitial)1/y, where y = number
annual precipitation through parturition (i.e., January– of years since colonization.
Table 1. Mean percent body fat (x̄), standard error (SE), and Table 2. Mean body mass (x̄; in kg), standard error (SE), and
sample sizes (n) for lactating and dry cow elk in the Chaco sample sizes (n) of calf (both sexes), yearling, and $ 2.5-y-old
Culture National Historical Park area, New Mexico, April 2003– cow elk in the Chaco Culture National Historical Park area, New
November 2006. Means followed by the same letter within a Mexico, 2003–2006.
column (ABC) or row (XY) do not differ (P . 0.10).
Calf Yearling $ 2.5-y-old
Lactating Dry Pooled
Year x̄ SE n x̄ SE n x̄ SE n
Month x̄ SE x̄ SE x̄ SE n
2003 — — — 178.6 4.3 2 213.5 6.7 4
April 2003 — — — — 5.8 A 0.8 4 2004 99.8 13.5 6 181.7 0.0 1 214.2 27.3 13
December 2005 97.2 24.9 4 169.9 12.0 5 213.0 14.1 15
2003 13.2 AX 1.5 12.8 AX 0.6 — — 9
2006 — — — 175.6 5.8 4 211.4 14.7 20
April 2004 — — — — 4.1 B 0.4 14
November
2004 11.3 AY 0.5 13.2 AX 0.4 — — 18
April 2005 — — — — 4.0 B 0.3 21 illegal harvest (2006); three females from unknown
November causes for which we could not exclude capture as the
2005 5.6 CY 0.4 7.5 CX 1.2 — — 20 ultimate cause of mortality; and three that were capture-
April 2006 — — — — 3.7 B 0.3 22 related. In addition, two calves (one each in 2004 and
November 2005) and one adult female lost their collars and one
2006 6.8 BY 0.5 10.1 BX 0.6 — — 24 adult female was never redetected using telemetry. The
latter 10 cases were censored from survival analyses.
Survival of adult females for the biological year (June–
Results May) was similar among years (Z , 1.0; P $ 0.318) and
ranged from 0.94 (SE = 0.06) to 1.00 (SE = 0.00; Table 3).
We captured and processed 11 elk (4 adult cows, 7 None of the individually identifiable calves (those
adult bulls), 14 adult cows (7 new captures, 7 recaptures), radiocollared or associated with a collared cow) died in
21 adult cows (3 new captures, 18 recaptures), and 23 2004–2005, 2005–2006, or 2006–2007, resulting in a
adult cows (all recaptures), April 2003–2006, respectively. maximum overall calf survival rate of 1.00 (SE = 0.00) for
In addition, we captured and processed 10 (8 new each year. Using pregnancy and lactation status from
captures, 2 recaptures), 18 (4 new captures, 14 recap- individual adult cows, estimated preweaning survival
tures), 20 (two new captures, 18 recaptures), and 24 adult was lowest (Z $ 3.4; P # 0.0006) in 2006–2007 (0.38 vs.
cows (all recaptures), late November–early December, 0.80 and 1.00 for 2004–2005 and 2005–2006, respec-
2003–2006. Last, we captured and processed seven tively) except for 2003–2004 (0.50), which was similar
calves each summer, 2004–2005, and an additional four (Z = 0.4; P = 0.719; Table 3). All radiocollared calves or
calves in December 2005. calves observed at heel that were alive at weaning were
verified by lactation in the mother, and all subsequently
Nutritional condition survived until the following June, resulting in postwean-
Body fat varied by year (F3,128 = 29.7; P , 0.001), ing calf survival of 1.00 (SE = 0) for each year. Thus,
season (late winter v. late autumn [F1,128 = 20.1; P , annual survival rates were identical to preweaning
0.001]), and lactation status (F1,128 = 13.7; P , 0.001 (Table 3).
[Table 1]). Body fat levels were higher in November than Calf survival was positively related to maternal
April regardless of lactation status (P # 0.009). Among MAXFAT, girth, mass, rBCS, and BF the previous autumn
years, BF of lactating cows was lower (P , 0.001) in 2005 (Table 4). Maximum subcutaneous fat thickness was the
and 2006 than in 2003 and 2004 (Table 1). Similarly, dry best individual predictor of maternal attributes on calf
cows were in better condition in 2003 and 2004 (P # survival and correctly classified the fate of 71.6% of
0.001). Condition levels did not vary by year at the low of calves. Odds ratios (3.48; 95% CI = 1.37–8.86) indicated
condition in early April except for April 2003 (P # 0.052),
when sample sizes were small (4 vs. 14–22; Table 1). Table 3. Annual survival (Ŝ), standard error (SE), and sample
Mean body mass of $ 2.5-y-old cows (H3 = 0.2; P = sizes (n) of adult cow and calf elk in the Chaco Culture National
0.983), yearling cows (H3 = 2.7; P = 0.436), or calves Historical Park area, New Mexico, June 2003–May 2007. Means
(U = 20.7; P = 0.219) did not differ among years followed by the same letter (ABC) within a column do not
(Table 2). Because of small sample sizes in 2003 and differ (P . 0.10).
2004, we pooled yearling cows from 2003 and 2004, Adult Calf
when condition of elk was significantly higher than in
2005 and 2006, and compared these years against 2005 Year Ŝ SE n Ŝ SE n
and 2006, but yearling mass still did not differ (H2 = 2.3; 2003–2004 1.00 A 0.00 16 0.50 ABC 0.34 4
P = 0.311). 2004–2005 0.94 A 0.06 22 0.80 B 0.12 10
2005–2006 1.00 A 0.00 24 1.00 A 0.00 14
Survival 2006–2007 0.96 A 0.04 24 0.38 C 0.03 16
We documented eight mortalities during this project:
x̄ 0.975 0.015 — 0.670 0.141 —
one female from harvesting (2004); one female from
Table 4. Relationships between probability of survival of individual calves and maternal age, size, and indices of condition by
season in the Chaco Culture National Historical Park area, north-central New Mexico, 2003–2006. Presented are Wald Chi-square test
statistic (x2), probability of test statistic (P), coefficient (b) and standard error (SE) of the variable in the logit, and sample sizes (n).
Autumn Late winter
a 2 2
Index x P b SE n x P b SE n
Age 2.1 0.143 — — 28 3.1 0.077 0.397 0.225 27
Girth 3.5 0.061 0.155 0.083 27 4.7 0.031 0.211 0.097 32
Mass 3.7 0.054 0.060 0.031 27 5.1 0.024 0.081 0.036 32
Loin 1.8 0.177 — — 27 4.2 0.042 4.812 2.360 34
wBCS , 0.1 0.964 — — 28 0.5 0.477 — — 34
rBCS 4.7 0.030 1.651 0.760 28 0.2 0.680 — — 34
MAXFAT 6.8 0.009 1.247 0.477 27 , 0.1 0.957 — — 34
BF 5.0 0.025 0.322 0.122 28 0.4 0.548 — — 34
a
wBCS, withers body condition score; rBCS, rump body condition score; MAXFAT, maximum subcutaneous fat thickness at the rump; BF, body fat.
that probability of survival for a calf increased about 3.5 (1.08; 95% CI = 1.01–1.16) indicated that probability of
times for each 1-cm increase in MAXFAT of their mother. survival for a calf increased about 1.1 times for each 1-kg
Among precipitation periods, calf survival was related increase in maternal body mass. For this subset of calves,
(x2 = 6.9, P = 0.008; b = 0.402 [SE = 0.153]) only to survival was also only related (x2 = 9.1; P = 0.003; b =
cumulative annual precipitation through parturition (i.e., 0.354 [SE = 0.118]) to cumulative annual precipitation
January–June). Odds ratios (1.50; 95% CI = 1.11–2.02]) through parturition. Odds ratios (1.42; 95% CI = 1.13–
indicated that probability of a calf surviving increased 1.79) indicated that probability of a calf surviving
about 1.5 times for each 1-cm increase in cumulative increased about 1.4 times for each 1-cm increase in
precipitation through June. When MAXFAT and cumula- cumulative precipitation through June. When maternal
tive precipitation through parturition were modeled mass and cumulative precipitation through parturition
together, MAXFAT was dropped from the final model were modeled together, both maternal mass (x2 = 3.7;
(x2 = 0.1; P = 0.712) while cumulative precipitation was P = 0.054) and cumulative precipitation (x2 = 5.9; P =
retained (x2 = 6.5; P = 0.011) indicating that cumulative 0.015) were retained in the final model, but odds ratios
precipitation through parturition had the dominant (1.078; 95% CI = 0.999–1.163) that included one
effect on calf survival (Figure 2). suggested that the effect of maternal mass was weaker
For calves whose mothers were assessed for size and than the effect of cumulative precipitation (odds ratio =
condition in late winter, calf survival was positively 1.42; 95% CI = 1.07–1.90).
related to maternal age, girth, mass, and depth of the
loin muscle in late winter (early April; Table 4). The Pregnancy and lactation
strongest relationship of maternal attributes during late Pregnancy rates ranged from 0.00 to 0.50 for yearling
winter on calf survival was maternal body mass, which and 0.69 to 1.00 for $ 2.5-y-old cows, respectively, and
correctly classified the fate of 74.4% of calves. Odds ratios were similar within age classes (Fisher’s exact P = 1.000
and 0.251 for yearlings and $ 2.5-y-old cows, respec-
tively) among years (Table 5). Lactating cows (including
yearlings) comprised 22–56% of cows and proportions
were similar (Fisher’s exact P = 0.109) among years
(Table 5). For $ 2.5-y-olds only, lactation rates were 0.29,
0.63, 0.67, and 0.30 for 2003–2006, respectively (Table 5).
Logistic models including age, girth, and rBCS were
positively related to the probability of a $ 2.5-y-old cow
conceiving (Table 6). Probability of pregnancy of $ 2.5-y-
old cows was best predicted by body mass, which
correctly classified the pregnancy status of 70.0% of the
elk. Odds ratios indicated that probability of pregnancy
increased about 1.06 times (95% CI = 1.008–1.111) for
each 1 kg increase in body mass. Pregnancy in yearling
cows was unrelated to any measure of size or condition.
Figure 2. Relationship between cumulative annual precipita- Population size and growth
tion (cm) through gestation (June) and calf survival in the We classified and documented a minimum of 43, 51,
Chaco Culture National Historical Park area, New Mexico. and 53 elk during autumn composition surveys, 2004–
Annual calf survival rates are presented immediately below 2006, respectively. Age and sex composition (SE) of elk
the line for that year’s precipitation. was 60 (4) bulls and 55 (4) calves, 50 (4) bulls and 44 (4)
Table 5. Mean (x̄), standard error (SE), and sample sizes (n) for pregnancy rate, lactation rate, and calves/100 cows in late autumn
for elk in the Chaco Culture Historical Park area, north-central New Mexico, 2003–2006. Means not sharing a letter (ABC) within a
column differ (P , 0.10).
Pregnant
Yearling $ 2.5-y-old Lactating Calf ratio
Year x̄ SE n x̄ SE n x̄ SE n x̄ SE n
2003 0.50 0.31 2 1.00 0.0 5 0.22 0.11 9 — — —
2004 0.00 0.00 1 0.69 0.07 16 0.56 0.06 18 55A 4 43
2005 0.40 0.13 5 0.93 0.04 15 0.50 0.06 20 44B 4 51
2006 0.00 0.00 1 0.80 0.04 20 0.25 0.03 24 24C 2 53
x̄ 0.33 0.13 — 0.86 0.07 — 0.38 0.09 — — — —
calves, and 59 (3) bulls and 24 (2) calves/100 cows, 2004– (2005 and 2006) to high marginal–low good (2003 and
2006, respectively. Bull ratios were similar (P $ 0.035 at 2004) nutrition, based on thresholds from nutritional
aexp = 0.10) but calf ratios varied (P # 0.030) among studies of penned elk (Cook et al. 2004). For free-
years (Table 5). The maximum estimate of l was 1.22 ranging elk, nutrition effectively ceases to be limiting
( = 0.94 + 0.5 60.55), 1.22 ( = 1.00 + 0.5 60.44), and 1.08 when lactating cows are able to accrue about 13.7% BF
( = 0.96 + 0.5 6 0.24) for 2004–2006, respectively. If we (Piasecke and Bender 2009). Lactating cows in CCNHP
substituted observed lactation rates for calf : cow ratios, never achieved . 13.2% BF, indicating that habitat
estimates of l were 1.11, 1.22, 1.25, and 1.09 for 2003– quality was always limiting to some degree. This was
2006, respectively. further evidenced by lactating cows being unable to
Minimum population size in 2006 was $ 53 elk for an accrue condition levels comparable to dry cows
approximate elk density of 0.15 elk/km2 (Table 7). Similarly (Table 1). However, strong effects on individual and
derived estimates for 2004 and 2005 were $ 43 (0.13 population performance are unlikely to be seen in elk
elk/km2) and $ 51 (0.15 elk/km2), respectively. Assuming populations until BF levels of lactating cows drop
that the initial January 2000 population estimate of 20 below 7.9% (Piasecke and Bender 2009). Lactating cows
elk was correct, the population has grown at a mean rate were below this threshold only in 2005 and 2006
of 15%/y (l = 1.15) from colonization in 2000 through (Table 1), and this eventually resulted in decreased calf
December 2006. Realized rate of population increase survival (0.38 vs. 0.77 for 2003–2005) and consequently
based on minimum population estimates declined from lowered (l = 1.04 vs. 1.18 for 2000–2004) population
l = 1.18 (2000–2005) to l = 1.04 (2005–2006). rates-of-increase.
Neither minimum population size (Z # 1.4; P $ 0.318) In CCNHP, accrual of BF in cows was most strongly
or number of cows and calves (Z # 1.4; P $ 0.303) were positively related to cumulative precipitation from
related to cow : calf ratios, $ 2.5-y-old cow pregnancy September to November (i.e., from the end of the
rates, yearling pregnancy rate, adult survival, calf survival, primary lactation to forage senescence; Bender 2007).
proportions of adult cows lactating, or proportions of all Declines in nutritional condition were, thus, a result of
cows lactating. density-independent effects (i.e., low precipitation),
highlighting the sensitivity of elk in our arid study area
Discussion
to pronounced annual variation in habitat quality related
Body fat level of lactating cows indicated that cows in to spatially and temporally patchy precipitation charac-
CCNHP were able to acquire diets ranging from poor teristic of arid environments (Sowell 2001; Bender 2007).
Table 6. Relationships between probability of pregnancy of age $ 2.5-y-old and yearling cow elk with age, size, and condition in
the Chaco Culture National Historical Park area, north-central New Mexico, 2003–2006. Presented are Wald Chi-square test statistic
(x2), probability of test statistic (P), coefficient (b) and standard error (SE) of the variable in the logit, and sample sizes (n).
$ 2.5-y-old Yearling
a
Index x2 P b SE n x2 P b SE n
Age 3.6 0.057 0.581 0.306 52 — ---- — ---- 13
Girth 6.3 0.012 0.179 0.071 58 0.6 0.441 — — 13
Mass 5.2 0.022 0.057 0.025 52 0.5 0.462 — — 13
Loin 0.3 0.587 — — 49 2.5 0.113 — — 13
wBCS 1.8 0.174 — — 58 1.7 0.193 — — 13
rBCS 3.3 0.069 1.307 0.719 58 0.5 0.467 — — 13
MAXFAT 0.7 0.391 — — 49 0.3 0.560 — — 13
BF 2.3 0.133 — — 58 0.2 0.624 — — 13
a
wBCS, withers body condition score; rBCS, rump body condition score; MAXFAT, maximum subcutaneous fat thickness at the rump; BF, body fat.
Table 7. Minimum numbers of elk in the Chaco Culture National Historical Park area, New Mexico, 2004–2006. The 2005 estimate
excludes a radiocollared cow and her calf that emigrated from the study area. Known individuals include individuals associated with
radiocollared cows or bulls or seen on survey flights that covered the entire study area (bulls and cows) and unmarked calves of
radiocollared cows.
Cows Bulls Calves
Year Collared Known Collared Known Collared Known Total
2004 18 2 4 8 6 5 43
2005 23 3 4 9 7 5 51
2006 25 4 8 9 0 7 53
Production and survival of calves is the most variable survival through summer was 0.95 for a population in
and influential component of elk and other ungulate Michigan (Bender et al. 2002), and elk in CCNHP
population dynamics (Clutton-Brock et al. 1982; Gaillard approached that level (0.80–1.0) in 2 of 4 y. This high
et al. 2000; Eberhardt 2002). Past research has shown that productivity was reflected in the mean historic growth
conception, calf survival, and recruitment were depen- rate (l = 1.18) seen prior to 2006, despite inclusion of
dent on condition of adult cows (Clutton-Brock et al. the lower growth potential seen in 2003 (l = 1.11). In
1982; Cook et al. 2004), results that we also observed in contrast, habitat limitations associated with unfavorable
CCNHP. Population density also can affect population precipitation patterns in 2005 and 2006 resulted in lower
productivity because of competition for resources, and (l = 1.04) growth rates in 2006.
productivity can also be limited by density-independent Maternal condition positively affected calf survival in
factors such as poor forage quality and precipitation CCNHP, similar to responses found elsewhere (Wegge
(Clutton-Brock et al. 1987; Smith and Anderson 1998; 1975; Thorne et al. 1976; Guinness et al. 1978; Clutton-
Piasecke and Bender 2009). We found that productivity in Brock et al. 1982). However, most of the causes of
CCNHP was affected by each of these factors except for preweaning mortality these studies found were classed
density dependence, because neither maternal condition as density independent (i.e., drowning, accidents, preda-
nor population demographics were related to density. tion), suggesting that chance also played a strong role in
Body mass most strongly affected pregnancy in $ 2.5- preweaning survival in addition to maternal influences.
y-old cows in CCNHP, similar to results seen in other free- We also saw this in CCNHP, where cumulative annual
ranging elk populations (Piasecke 2006). In contrast, precipitation through gestation, a density-independent
pregnancy in yearlings was unrelated to size or effect, was also positively related to calf survival.
condition, perhaps because small sample sizes (n = 13) Additionally, all calf mortality occurred at or near
precluded a sensitive analysis of factors affecting yearling parturition, a period primarily associated with density-
pregnancy. Both absolute (Grier 1968; Cook et al. 2004) independent mortality (Harper et al. 1967; Arman et al.
and relative (Haigh and Hudson 1993) body mass have 1978; Clutton-Brock et al. 1987; Gogan and Barrett 1987;
been implicated as important variables for yearling Smith and Anderson 1998) and when calves typically
pregnancy in elk. Yearling cows in CCNHP were able to experience their greatest mortality (Guinness et al. 1978;
achieve about 80–85% of the mean $ 2.5-y-old cow Clutton-Brock et al. 1988). Because pregnancy was also
body mass (Table 2), a greater proportion than repro- related to cow condition, most aspects of productivity in
ductive thresholds delineated for penned (70%; Haigh CCNHP (i.e., conception, preweaning calf survival) were,
and Hudson 1993) and free-ranging elk (50%; Piasecke thus, influenced to some degree by both maternal
2006), but pregnancy rates were still low (x̄ = 0.33). In condition and density-independent effects (i.e., precipi-
terms of absolute mass, yearling pregnancy exceeded tation). The only exception was postweaning calf
25% only when body mass exceeded 163–169 kg in survival, which was uniformly high (Table 3).
Yellowstone National Park (Greer 1968) and about 170 kg The importance of cumulative precipitation through
for multiple free-ranging elk populations (Piasecke 2006). gestation in CCNHP was likely due to effects on spring
Yearling mass in CCNHP ranged from 170 to 180 kg green-up and, hence, quantity and quality of forage.
(Table 2); thus, pregnancy in yearling cows may have Successfully raising a calf came at a strong cost to cows;
been related to a threshold in absolute body mass, rather lactating cows were unable to accrue condition at the
than relative mass. rate of dry cows, and consequently entered winter in
Calf survival ranged from 0.38 to 1.00 in CCNHP, and poorer condition (Table 1). Because condition is largely a
the calf survival rate of 0.38, calf : cow ratio of 24:100, and result of nutrition (Cook et al. 2004), and condition also
lactation rate of 0.25 in 2006 all reflected the decreased affected calf survival, the influence of precipitation, a
and low productivity of elk following the decline in density-independent effect, was likely at least partially
condition first seen in 2005. From 2003 to 2005, mediated through maternal condition, which is most
productivity in CCNHP was comparable to the highest generally associated with density dependence. Produc-
productivity rates seen (51–56/100 for elk in Michigan; tivity of elk can, thus, be influenced by factors other than
Bender et al. 2002) and with the fastest growing elk herd resource competition (i.e., density dependence), high-
documented (l = 1.22; Eberhardt et al. 1996; Bender et lighting the importance of density-independent effects
al. 2002). Similarly, the highest documented elk calf on elk population dynamics.
Adult female survival (0.94–1.00) in CCNHP was the journal editors for their efforts to improve this
comparable to the highest rates seen for free-ranging manuscript. All activities were in accordance with NMSU
elk (Bender et al. 2008) even in 2005 and 2006, when elk IACUC Permit NM-2002-027.
condition was extremely low. This indicates that even
given conditions such as elk faced in 2005–2006, habitat
conditions were still suitable to allow high adult female References
survival, the last demographic affected by resource stress Arman P, Hamilton WJ, Sharman GAM. 1978. Observa-
(Gaillard et al. 2000). Elk are protected from harvest while tions of the calving of free-ranging tame red deer
on CCNHP, but off the Park they are subject to regulated (Cervus elaphus). Journal of Reproductive Fertility 37:
tribal harvest during the autumn and early winter. Thus, 87–90.
although the population is lightly hunted (the only non– Bender LC, compiler. 2007. Growth, health, and impacts
capture-related mortalities for 2003–2006 we observed of a pioneering elk herd on and adjacent to Chaco
were one cow harvested in 2004 and one cow illegally
Culture National Historical Park, New Mexico. Final
killed in 2006), adult survival rate was high even
Report to National Park Service. Las Cruces: New
compared to unhunted populations and was at or near
the expected maximum for adult cows (x̄ = 0.98, 2003– Mexico Cooperative Fish and Wildlife Research Unit.
2006 [Eberhardt 2002; Bender et al. 2008]). Available from the corresponding author.
Because of the low number of mortalities during this Bender LC, Carlson E, Schmitt SM, Haufler JB. 2002.
study, we were unable to model survival and, thus, Production and survival of elk (Cervus elaphus) calves in
identify whether condition or other factors influenced Michigan. American Midland Naturalist 148:163–171.
adult cow vulnerability. Given the extremely high survival Bender LC, Cook JG, Cook RC, Hall PB. 2008. Relations
seen, however, it is unlikely that any factor was between nutritional condition and survival of elk in
significantly affecting adult cow survival in CCNHP. the Pacific Northwest. Wildlife Biology 14:70–80.
However, the relative lack of disturbance associated with Bender LC, Roloff GJ, Haufler JB. 1996. Evaluating
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Surveys
The Aquatic Turtle Assemblage Inhabiting a Highly
Altered Landscape in Southeast Missouri
Brad M. Glorioso,* Allison J. Vaughn, J. Hardin Waddle
B.M. Glorioso
IAP World Services, Inc., U.S. Geological Survey, National Wetlands Research Center, Lafayette, Louisiana 70506
A.J. Vaughn
Division of State Parks, Missouri Department of Natural Resources, Jefferson City, Missouri 65102
J.H. Waddle
U.S. Geological Survey, National Wetlands Research Center, Lafayette, Louisiana 70506
Abstract
Turtles are linked to energetic food webs as both consumers of plants and animals and prey for many species. Turtle
biomass in freshwater systems can be an order of magnitude greater than that of endotherms. Therefore, declines in
freshwater turtle populations can change energy transfer in freshwater systems. Here we report on a mark–recapture
study at a lake and adjacent borrow pit in a relict tract of bottomland hardwood forest in the Mississippi River
floodplain in southeast Missouri, which was designed to gather baseline data, including sex ratio, size structure, and
population size, density, and biomass, for the freshwater turtle population. Using a variety of capture methods, we
captured seven species of freshwater turtles (snapping turtle Chelydra serpentina; red-eared slider Trachemys scripta;
southern painted turtle Chrysemys dorsalis; river cooter Pseudemys concinna; false map turtle Graptemys
pseudogeographica; eastern musk turtle Sternotherus odoratus; spiny softshell Apalone spinifera) comprising four
families (Chelydridae, Emydidae, Kinosternidae, Trinoychidae). With the exception of red-eared sliders, nearly all
individuals captured were adults. Most turtles were captured by baited hoop-nets, and this was the only capture
method that caught all seven species. The unbaited fyke net was very successful in the borrow pit, but only captured
four of the seven species. Basking traps and deep-water crawfish nets had minimal success. Red-eared sliders had the
greatest population estimate (2,675), density (205/ha), and biomass (178 kg/ha). Two species exhibited a sex-ratio bias:
snapping turtles C. serpentina in favor of males, and spiny softshells A. spinifera in favor of females.
Keywords: agriculture; Big Oak Tree State Park; hydrology; population demography; red-eared slider; Trachemys scripta
Received: July 6, 2010; Accepted: September 29, 2010; Published Online Early: October 2010; Published: November
2010
Citation: Glorioso BM, Vaughn AJ, Waddle JH. 2010. The aquatic turtle assemblage inhabiting a highly altered
landscape in southeast Missouri. Journal of Fish and Wildlife Management 1(2):161–168; e1944-687X. doi: 10.3996/
072010-JFWM-020
* Corresponding author: gloriosob@usgs.gov
Introduction 1940s ,50% of the floodplain remained forested (Twedt

and Loesch 1999). Since that time, an additional 2–3
Historically, bottomland hardwood forests occurred million ha have been converted to agriculture (Forsythe
nearly contiguously over an estimated 10 million ha 1985), facilitated by flood control projects that alter the
throughout the Lower Mississippi Alluvial Valley (Putnam natural hydrology of the landscape (Galloway 1980). An
et al. 1960; Hefner and Brown 1985). The nutrient-rich estimated 96% of bottomland hardwood forest losses
alluvial soil within the floodplain of large rivers like the can be attributed to conversion into agricultural
Mississippi has encouraged people to convert native production (MacDonald et al. 1979). The remaining
forests to agriculture (Twedt and Best 2004). By the late bottomland forest (25%) are most often small patches,
Turtle Population in a Southeast Missouri State Park B.M. Glorioso et al.
Figure 1. Aerial view of Big Oak Tree State Park, Missouri, showing the approximately 400-m-diameter manmade circular lake,
which is surrounded by a levee over which the borrow pit lies.
#1,012 ha, and highly fragmented, separated from each In union with these efforts, we examined the
other by nonforest land cover (Rudis 1995; Twedt and freshwater turtle assemblage at BOTSP. Turtles play a
Loesch 1999). One such example is Big Oak Tree State very important role in ecosystems because they are
Park (BOTSP), located in the northernmost extent of the linked to energetic food webs both as consumers of
Lower Mississippi Alluvial Valley in the boot-heel of plants and animals and as prey species for a variety of
southeast Missouri (Figure 1). organisms (Mitchell and Buhlmann 2003). Iverson (1982)
A far-ranging effort by local citizens to protect an showed turtle biomass in populations can be quite large,
exceptionally large bur oak Quercus macrocarpa culmi- typically at least an order of magnitude greater than that
nated in the purchase of approximately 408 ha of land of endotherms. Declines in a turtle population will,
dedicated in 1938 as BOTSP. In 1977, nearly the entire therefore, affect ecosystem health in terms of changes in
park (ca. 380 ha) was designated as a Missouri Natural energy transfer. In this mark–recapture study, we
Area, including the approximately 32-ha virgin wet– examined sex ratio and size structure, and estimated
mesic bottomland forest surrounding the aforemen- population size, density, and biomass for turtles captured
tioned bur oak. Trees in the park today are unequaled in at BOTSP.
the state for their size, with a canopy averaging 36.5 m
and several trees .40 m tall. Due to the small size of the Study Area
park and land management practices that negatively
altered the park’s forest and hydrology, regeneration of Big Oak Tree State Park is located in the northernmost
oaks is nearly nonexistent and many of the trees are old section of the Mississippi Alluvial Valley in the St. James
and dying. The park is currently in the planning process Bayou Clayey Lowland land type association (Figure 1).
of hydrological restoration, and in conjunction with this The park’s elevation ranges from 87 to 88.5 m above sea
effort, studies have commenced to gather baseline data level, with the lowest elevation occurring in the shrub
on the natural history of the park, including surveys and swamp. Due to a small land acquisition (16 ha) since the
studies of fungi, plants, insects, crawfish, fish, and birds. original purchase, BOTSP now encompasses 423.5 ha,
The data acquired from these investigations will allow for and is representative of the northern boundary of the
posthydrological restoration comparisons, and provide Mississippi Embayment, a land formation created when
insight into restoration effectiveness. the ancestral seas retreated for the last time during the
Cretaceous Period. The park rests in an early meander used deep-water crawfish nets baited with chicken leg
belt of the Mississippi River, which is significant to the quarters to capture turtles on three occasions (Glorioso
hydrology. The topography is characterized as a ridge and Niemiller 2006). Lastly, a few turtles were captured
and swale complex with higher ground inundated only by hand in the shallows or on land. The trapping success
during significant river flooding, and lower elevations of the hoop-nets and fyke net was measured using catch
holding water for most of the growing season. The per unit effort (CPUE), where the total number of turtles
climate, topography, floral, and faunal associations at captured was divided by the total number of trap-nights
BOTSP more closely resemble those of the southeastern for each capture method.
states than anywhere else in Missouri. We identified each captured turtle to species and
Big Oak Tree State Park is essentially an island of trees, individually marked them with unique three-letter
separated from any other wooded corridors by thou- identification by filing the marginal scutes (Cagle 1939;
sands of ha of agriculture (Figure 1). The park’s Dorcas 2005). We modified the marking scheme for
hydrology is directly compromised by an elaborate ditch eastern musk turtles Sternotherus odoratus to compen-
system built to expedite water removal from agricultural sate for the reduced number of marginal scutes in
lands. Access to the Mississippi River has been closed by kinosternid turtles. We marked spiny softshells Apalone
a levee system, but during extreme high-water events, spinifera by cutting shallow pie-shaped pieces at the
floodwaters occasionally back through the ditches, margins of the epidermis. We also examined red-eared
bringing water and agricultural pesticides into the park. slider Trachemys scripta males for the presence of
Backwater flooding did not occur during the duration of melanism on the shell and skin.
our turtle survey. With the exception of the manmade We determined the sex of turtles by examining
lake, which is pumped full with groundwater each spring, secondary sexual characteristics such as elongated
rainwater is the primary source of the park’s water. Yearly foreclaws and tails in males. We determined the sex of
rainfall ranges from 68 to 203 cm (mean = 130 cm), with snapping turtles Chelydra serpentina by taking a ratio of
most rainfall occurring in the early autumn and spring the distance from the cloaca to the posterior tip of the
months. The mean monthly temperature is 29.4uC in July, plastron and the distance from the posterior tip of the
though temperatures during the survey reached 37.8uC plastron to the anterior edge of the femoral scute
in August. (Dorcas 2005). If the ratio was .0.86 we considered it
The park contains a bottomland hardwood forest and male, and if it was ,0.86 we considered it female. In
the remains of a bald cypress (Taxodium spp.) swamp, addition, we attempted to extract the penis of all
formally categorized as a shrub swamp. A 9-ha manmade snapping turtles. Most individuals that we determined
recreational lake built in the 1960s on the historic Grassy to be males by the above ratio were confirmed by
Pond (a large swamp system dominated by sedges and presence of the penis, and no individuals that we
cypress trees) is located in the center of the park. determined to be female by the above ratio were shown
Surrounding the circular lake, which is choked with to be male by the presence of a penis. We performed G-
coontail Ceratophyllum demersum, is a borrow pit from tests in Program R (R Development Core Team 2008) to
which the levee around the lake was built. The borrow detect differences in sex ratios. The level of significance
pit connects to the shrub swamp and water levels in for all analyses was set at a = 0.05.
both of these areas varied during the survey. During a 7- We took measurements of straight-line carapace and
wk drought in August and September, when ,2.5 cm of plastron lengths at the midline using digital calipers
rain fell, water levels in the borrow pit fell to about 45 cm (60.1 mm) for smaller turtles (,160 mm), tree calipers
in the center, from a high of at least 122 cm at the start (61 mm) for larger turtles (.160 mm), and a measuring
of the survey. The shrub swamp completely dried during tape (61 mm) for softshell turtles. We measured body
this drought. mass with hanging spring scales of various sizes. We
performed a two-tailed Kolmogorov–Smirnov test in
Methods Program R (R Development Core Team 2008) to test the
null hypothesis that the samples of male and female
In this survey we used 76-cm and 91-cm three-ring carapace lengths came from the same distribution.
hoop-nets with 2.5-cm mesh baited with sardines. At any We estimated the population abundance of turtle
given time during the study, between 7 and 17 hoop- species in the assemblage at BOTSP using closed-
nets were set out in the park for 1–3-d intervals. We population capture–recapture models that assume full
checked the hoop-nets on 37 occasions from 10 June to demographic and geographic closure of the population
27 September 2007. There were three other days not during the study period (i.e., no births, deaths, immigra-
coincident with hoop-net trapping (22 April, 2 June, and tion, or emigration). We deemed it reasonable to assume
23 June) where we used other capture methods closure for the duration of this short-term study due to
(described below), giving 40 total trap-sampling events the isolated nature of the location, lack of aquatic
from 22 April to 27 September. migration routes outside the park, and the commence-
We used an unbaited seven-ring fyke net with 7.6-m ment of trapping occurring after peak hatching times.
leads and 6.4-cm mesh in both the borrow pit and lake. Models based on those described by Otis et al. (1978)
We also used two basking traps, one small (ca. 117 6 91 were fit using maximum-likelihood estimation in Pro-
6 30 cm) and one large (ca. 152 6 152 6 51 cm), to gram MARK (White and Burnham 1999). The four models
capture turtles in both the borrow pit and the lake. We analyzed for each species were: 1) constant capture
Table 1. Number of total individuals captured, total recaptures, and individuals recaptured (%) for seven turtle species captured
at Big Oak Tree State Park, Missouri, from 22 April to 27 September 2007.
Total Total Individuals
Species individuals recaptures recaptured (%)
Red-eared slider (Trachemys scripta) 785 172 131 (17)
Snapping turtle (Chelydra serpentina) 69 23 13 (19)
Southern painted turtle (Chrysemys dorsalis) 53 12 9 (17)
Eastern musk turtle (Sternotherus odoratus) 31 3 1 (3)
Spiny softshell (Apalone spinifera) 20 12 6 (30)
River cooter (Pseudemys concinna) 2 0 0 (0)
False map turtle (Graptemys pseudogeographica) 1 3 1 (100)
Totals 961 225 161 (17)
probability (M0), 2) capture probability varying with time sliders comprised 92% of fyke net captures in the borrow
(Mt), 3) behavioral response where probability or pit for a CPUE of 8.50. Species also captured by fyke net
recapture is different from probability of first capture included snapping turtles (10), southern painted turtles
(Mb), and 4) a combination of the effects of time and (4), and false map turtles (1). The false map turtle was
behavior (Mtb). escaping when the net was being checked because it
Capture data on individuals from the 40 total trap- could fit through the mesh, as could all eastern musk
sampling events were summarized into four sampling turtles and smaller size classes of other species.
occasions for analysis in MARK. We conducted model Basking-trap success was minimal, with only 19 total
selection using information-theoretic techniques based captures of two species (red-eared sliders and southern
on Akaike’s Information Criterion adjusted for small painted turtles). Both basking traps did poorly in the
sample sizes, and all abundance estimates reported are borrow pit, with only two southern painted turtle
model-averaged to account for model selection uncer- captures. In the lake, there were 13 red-eared sliders
tainty (Burnham and Anderson 1998). To compute and 4 southern painted turtles captured, with both traps
density, the abundance of each species was divided by having similar success. Deep-water crawfish nets were
13 ha, the approximate combined aquatic area of the used to capture turtles on three occasions, but had very
lake and borrow pit. We calculated the biomass by little success overall and were not further pursued as a
multiplying the overall mean body mass (in kg) of all capture method.
individuals at first capture by the density. The three most frequently captured turtle species
were recaptured in similar percentages of overall
Results individuals (Table 1). Excluding the lone false map turtle
individual that was captured multiple times, the species
We captured 961 individual turtles of 7 species at with the largest return rate was the spiny softshell at
BOTSP in 2007 (Table 1; Supplemental Material, Data 30%. Across all species, the majority of individuals were
S1, http://dx.doi.org/10.3996/072010-JFWM-020.S1). The recaptured in the same primary water body as the initial
most abundant turtle species, the red-eared slider, capture. With the exception of three snapping turtles
represented nearly 82% of all individuals captured. Due and one spiny softshell, all overland movements were
to few river cooter Pseudemys concinna and false map made by red-eared sliders. About one in four recaptured
turtle Graptemys pseudogeographica captures these two red-eared sliders was found to have moved between
species were excluded from all statistical analyses. primary water bodies. Of the 36 red-eared sliders that
There were 975 captures with hoop-nets of seven moved, 34 (94%) moved from the borrow pit to the lake.
species, with 634 captures in the borrow pit and 341 The majority of this movement coincided with the
captures in the lake. The overall CPUE of hoop-nets in drought that brought water levels down in the borrow
both water bodies was 0.95, with the borrow pit having pit to extremely low levels.
twice the capture success of the lake. The CPUE for the Of the five most numerous species captured, sex ratios
borrow pit was nearly twice the CPUE of the lake for red- did not differ significantly from 1:1 for red-eared sliders,
eared sliders, and three times the CPUE for snapping southern painted turtles, and eastern musk turtles. There
turtles. The largest differences in CPUE between the two was a significant male-biased sex ratio for snapping
trapped areas involved southern painted turtles Chrys- turtles, and a significant female-biased sex ratio for spiny
emys dorsalis and eastern musk turtles, with the borrow softshells (Table 2).
pit having approximately 10 and 6 times the CPUE of the The red-eared slider population was well-represented
lake, respectively. Capture success of spiny softshells was by adult and subadult size classes (Figure 2). Carapace
equal between the borrow pit and lake. length distribution in red-eared sliders appears bimodal;
The fyke net was only successful in the borrow pit; one there were many adult turtles in the sample, with
red-eared slider was captured in the lake in 14 d of younger individuals between 110 and 140 mm also
trapping. In the borrow pit, the fyke net captured 185 being well-represented (Figure 2). A Kolmogorov–Smir-
turtles of four species in 20 trapping days. Red-eared nov Z-test indicated that the carapace lengths of males
Table 2. Number (n) of males (m) and females (f), results of (Figure 2), and both species were sexually dimorphic in
G-tests (G-stat) for equal sex ratios in the five most abundant carapace length (Kolmogorov–Smirnov D = 0.61, P ,
turtle species captured at Big Oak Tree State Park, Missouri, 0.001 and D = 1.0, P , 0.001, respectively). The southern
from 22 April to 27 September 2007. Asterisk denotes painted turtle population showed a near-uniform distri-
significantly different from 1:1 ratio at a = 0.05. bution of carapace lengths in males, but adults between
Species n (m:f) G-stat P 135 and 150 mm dominated the distribution in females
(Figure 2), and only a few juveniles of southern painted
Red-eared slider 387:392 0.03 0.8578
(Trachemys scripta) turtle were captured. Carapace length was sexually
dimorphic in southern painted turtles (Kolmogorov–
Snapping turtle 46:23 7.82 0.0052*
(Chelydra serpentina) Smirnov D = 0.74, P , 0.001). The eastern musk turtle
Southern painted turtle 20:33 3.22 0.0727 population was represented by nearly all adults, but
(Chrysemys dorsalis) many smaller size class captures would have been
Eastern musk turtle 19:12 1.59 0.2067 precluded by the larger mesh size of the nets (Figure 2).
(Sternotherus odoratus) There was no evidence of sexual dimorphism in carapace
Spiny softshell 5:15 5.23 0.0222* length in eastern musk turtles (Kolmogorov–Smirnov D
(Apalone spinifera) = 0.34, P = 0.371).
Male red-eared sliders exhibited melanism on their
shells as well as their skin, whereas females sometimes
and females were sexually dimorphic (i.e., did not come showed melanism on their shells, but never on the skin.
from the same distribution; Kolmogorov–Smirnov D = The smallest male captured that showed signs of
0.37, P , 0.001). The population of both snapping turtles melanism was 172 mm carapace length. There were
and spiny softshells consisted of nearly all adults 201 male red-eared sliders individuals captured with
Figure 2. Size-class distributions by carapace length of the five most abundant turtle species captured at Big Oak Tree State Park,
Missouri, from 22 April to 27 September 2007.
Table 3. Estimates of population size (N), standard error (SE), lower and upper 95% confidence interval (CI), density, and biomass
for the five most abundant turtle species captured at Big Oak Tree State Park, Missouri, from 22 April to 27 September 2007.
Lower Upper Density Biomass
Species N SE 95% CI 95% CI (per ha) (kg/ha)
Red-eared slider (Trachemys scripta) 2,675 233.10 2,218 3,132 205.8 178.5
Snapping turtle (Chelydra serpentina) 121 22.93 76 166 9.3 57.4
Southern painted turtle (Chrysemys dorsalis) 103 28.96 46 160 7.9 2.3
Eastern musk turtle (Sternotherus odoratus) 326 314.66 0 942 25.1 4.2
Spiny softshell (Apalone spinifera) 24 4.69 14 33 1.9 7.1
$172 mm carapace length, and 81% of those individuals communication, 29 September 2007). It is certainly
were melanistic. Nearly 90% of individual males with possible that this species may be present at the park,
$185 mm carapace lengths were melanistic. but no explanation can be given as to our failure to
We obtained population size estimates for five turtle detect them in our sampling.
species (Table 3). The results of all four closed-popula- Ream and Ream (1966) found that different turtle
tion models were used for model-averaging of the trapping methods introduced considerable bias in sex
abundance estimate of each species with the exception ratios and size-class distribution due to differences in
of model Mtb on red-eared sliders and eastern musk individual behavior related to size and sex. Therefore,
turtles where that model failed to converge. The several trapping techniques were used during this study
estimated standard error and the associated 95% in an effort to maximize overall catch and diversity of
confidence interval were large for those species with species while reducing bias. The fyke net in the borrow
relatively few captures or recaptures. pit had the greatest CPUE of any capture technique, even
The red-eared slider was by far the most abundant while precluding capture of smaller turtles, but the hoop-
turtle species at BOTSP and, therefore, had the highest nets performed best with respect to overall catch and
density of the turtle species (ca. 205 individuals/ha), with diversity.
all other species considerably lower (Table 3). Due to the The two primary bodies of water in this study are quite
moderate mean mass and the high abundance, red- different and it was not surprising to see that some
eared sliders also had the largest biomass of the turtle species appear to prefer one over the other. Most
species at 178 kg/ha, followed by snapping turtles at southern painted turtles were captured early in the study
57 kg/ha, with the other three species combined at in the ditch that empties into the borrow pit. This ditch
,14 kg/ha (Table 3). gets more shallow the further one gets from the borrow
pit, and at the commencement of this study it retained
Discussion water for only about 50 m from the borrow pit. The ditch
In addition to the seven turtle species captured, there became completely dry halfway through the study. The
are four other aquatic turtle species whose range lack of southern painted turtles captured at BOTSP and
overlaps BOTSP, which were not found in this study: the location of most of their captures was surprising. At
smooth softshell Apalone mutica, alligator snapping Reelfoot Lake, approximately 20 km to the south, the
turtle Macrochelys temminckii, western chicken turtle southern painted turtle was the codominant emydid
Deirochelys reticularia and Mississippi mud turtle Kinos- species with the red-eared slider, and was captured most
ternon subrubrum. Not capturing smooth softshells at often in the main bodies of the lake, and seldom in
BOTSP is understandable because its typical habitat is backwater areas (B. Glorioso, unpublished data). South-
riverine, but alligator snapping turtles and western ern painted turtles and eastern musk turtles at BOTSP
chicken turtles were probably historically abundant in especially preferred the murky shallow waters of the
the area (Briggler and Johnson 2006). Unfortunately, borrow pit to the clear vegetation-choked water of the
these two species are now listed as rare in Missouri lake.
primarily due to habitat loss, but also due to over- Sex ratios were found to be significantly biased in
collection for the food and pet trade in the case of snapping turtles and spiny softshells. Sex ratio is an
alligator snapping turtles. It is unlikely that a viable important demographic measurement because of the
population of either occurs in the highly altered aquatic influence that it has on certain aspects of population
habitats of the park. dynamics (Gibbons 1990). Whereas a single male can
More perplexing is the nondetection of Mississippi fertilize many females, females are much more limited in
mud turtles from the waters of BOTSP. This species is their reproductive potential and, thus, a skewed sex ratio
typically found in or near swamps, sloughs, and canals. It in favor of males is cause for concern. An unequal sex
can also be found in habitats that have been highly ratio can occur naturally, but many times it can be
modified, including highly urbanized areas. Missouri attributed to sampling biases and differential catchability
Department of Conservation state herpetologist Jeff of the sexes (Gibbons 1990). Ream and Ream (1966)
Briggler stated the Mississippi mud turtle to be one of found differences in sex ratio within a population with
the most commonly trapped turtle species at conserva- respect to capture technique. However, more recent
tion areas in the boot-heel of Missouri (personal studies have found no change in sex ratios across
different capture techniques in the same population success, future research efforts at BOTSP could incorpo-
(Lovich and Gibbons 1990; Smith and Iverson 2002). Also, rate this aspect of turtle ecology. A better understanding
short-term studies may be ineffective at truly represent- of age-specific fecundity and survival, and clutch size as it
ing sex ratios within a population (Gibbons 1990). For relates to age and size of adult females, would be
example, in this study, only female spiny softshells were beneficial. Also, future studies could use radiotelemetry
captured through mid-July. Five males and one female to track gravid females to their nesting sites, gaining
were captured after this date. Thus, we could have insight into nest survival in the park and among the
observed an even greater skewed sex ratio for this agricultural lands. In conjunction, recaptures from this
species had we not sampled for the duration we did. It is study could be used in future investigations to deter-
possible, however, that the significantly different sex mine growth rates and survivorship as well as derive
ratios observed in snapping turtles and spiny softshells in better population estimates.
this study are an artifact of sampling-method bias, small
sample sizes, and the limited duration of our study.
Several nests were observed in early June that Supplemental Material
appeared to have been depredated. Most of these nests Please note: The Journal of Fish and Wildlife Management
were found on the levee around the lake, particularly near is not responsible for the content or functionality of any
the parking lot where the nests were located in areas that supplemental material. Queries should be directed to the
receive extended sunlight. At BOTSP, in addition to the corresponding author.
limited open area around the levee and parking lot,
turtles most likely use the surrounding agricultural fields Data S1. BOTSP species spreadsheets: Apalone
as nesting sites, much as they do at Reelfoot Lake in spinifera, Chelydra serpentina, Chrysemys dorsalis, Grapte-
northwest Tennessee (B. Glorioso, unpublished data). In mys pseudogeographica, Pseudemys concinna, Ster-
these areas, in addition to natural predators, nests may be notherus odoratus, and Trachemys scripta.
physically destroyed by agricultural practices, and may be Found at DOI: 10.3996/072010-JFWM-020.S1 (98.6 KB
impacted negatively by pesticides that are commonly XLSX).
sprayed upon crops adjacent to the park. There was little
evidence of successful reproduction in that only three Acknowledgments
hatchlings (red-eared sliders) were observed during the
study. Apparent below-normal egg or hatchling survival We thank K. McCarty, Chief of Natural History for the
rates at the park may be explained by a lack of suitable Missouri Department of Natural Resources, for his
nesting sites, heavier than usual predation rates, or a support of this project by allocating the necessary funds.
combination of these and other factors. We also thank M. Comer, Natural Resource Manager I of
The occasional backwater flooding that has been the Bootheel Management Unit of the Missouri Depart-
responsible for bringing water into the borrow pit, the ment of Natural Resources, for his support in retaining
preferred habitat of most turtle species in this study, is a the necessary supplies for this project. For field help on
result of the Mississippi River flowing backwards in high- one or more days we thank the following: R.R. Layton, C.
water events through an approximately 457-m gap in the Hanson, M.F. Glorioso, M. McNeary, G.R. Wyckoff, and C.F.
levee. This gap will be closed by the U.S. Army Corps of Glorioso. Thanks to C.R. Schwalbe, G.M. Fellers, the
Engineers as the main component of the St. John’s Bayou Subject Editor, and three anonymous reviewers who
and New Madrid Floodway Project (http://www.mvm. made helpful criticisms of earlier versions of this
usace.army.mil/stjohns/default.asp) aimed at protecting manuscript.
agricultural land and small towns near BOTSP from This research was conducted under Missouri Depart-
flooding. As mitigation for the project, the U.S. Army Corps ment of Conservation Wildlife Collector’s Permit No.
of Engineers has a water management plan in place for 13899. Any use of trade, product, or firm names is for
southeast Missouri, as well as a proposal to create a descriptive purposes only and does not imply endorse-
vegetative corridor between BOTSP and a nearby conser- ment by the U.S. Government.
vation area. In addition, the Missouri Department of Natural
Resources is planning to construct a $1.2 million water
retention project designed to maintain a self-sustaining References
water system at the park, thus promoting regeneration of Briggler JT, Johnson TR. 2006. Missouri’s turtles. Jefferson
the existing dominant species while inhibiting invading
City, Missouri: Missouri Department of Conservation.
understory species. Without these actions, the park would
likely transition from a wet bottomland hardwood to a Available: http://mdc4.mdc.mo.gov/Documents/8273.
much drier forest type, jeopardizing the survival of the pdf (August 2009).
turtle assemblage at BOTSP. Burnham KP, Anderson DR. 1998. Model selection and
Future studies comparable to this one could be used inference: a practical information-theoretic approach.
to evaluate the impact of the Missouri Department of New York: Springer-Verlag.
Natural Resources and U.S. Army Corps of Engineers Cagle FR. 1939. A system for marking turtles for future
planned actions on the turtle assemblage at BOTSP, identification. Copeia 1939:170–173.
using the results of this study as a baseline. Because this Dorcas ME. 2005. Herpetology laboratory guidelines and
study indicated potential problems with reproductive protocols. Davidson, North Carolina: Davidson Col-
lege, Department of Biology. Available: http://www. sion, Publication Number 420–529. Available: www.ext.
bio.davidson.edu/people/midorcas/research/Contribute/ vt.edu/pubs/fisheries/420-529/420-529.pdf (August 2009).
HERP%20LAB%20PROTOCOLS-2005-8-25.pdf (August Otis DL, Burnham KP, White GC, Anderson DR. 1978.
2009). Statistical inference from capture data on closed
Forsythe SW. 1985. The protection of bottomland animal populations. Wildlife Monographs 62.
hardwoods along the Lower Mississippi Valley. Trans- Putnam JA, Furnival GM, McKnight JS. 1960. Manage-
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Mississippi Delta. Alexandria, Virginia: U.S. Army Engi- environment for statistical computing. Vienna, Austria:
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Gibbons JW. 1990. Sex ratios and their significance www.R-project.org (August 2009).
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JW, editor. Life history and ecology of the slider turtle. methods on the estimation of population structure
Washington, D.C.: Smithsonian Institution Press. in painted turtles. American Midland Naturalist 75:
Glorioso BM, Niemiller ML. 2006. Using deep-water 325–338.
crawfish nets to capture aquatic turtles. Herpetolog- Rudis VA. 1995. Regional forest fragmentation effects on
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Hefner JM, Brown JD. 1985. Wetland trends in the values. Landscape Ecology 10:291–307.
southeastern United States. Wetlands 4:1–11. Smith GR, Iverson JB. 2002. Sex ratio of common musk
Iverson JB. 1982. Biomass in turtle populations: a turtles (Sternotherus odoratus) in a north-central
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adult sex ratio in the turtle Malaclemys terrapin. Oikos Twedt DJ, Best C. 2004. Restoration of floodplain forests
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Surveys
Bobcat Population Status and Management in North
America: Evidence of Large-Scale Population Increase
Nathan M. Roberts,* Shawn M. Crimmins
N.M. Roberts
Department of Natural Resources, Cornell University, Ithaca, New York 14853
S.M. Crimmins
Department of Forest Management, University of Montana, Missoula, Montana 59812
Abstract
Bobcat Lynx rufus populations are thought to be increasing in North America; however, little information exists
on their current population status. In the United States, management and monitoring of bobcat populations
is the responsibility of state wildlife management agencies. We surveyed state wildlife management agencies in
each of the 48 contiguous states regarding the current population status, distribution, and monitoring protocols
of bobcats within each respective jurisdiction. We also surveyed the governments of Mexico and Canada regarding
bobcat population status within their jurisdictions. We received responses from 47 U.S. states, Mexico, and 7
Canadian provinces. Responses indicate that bobcats occur in each of the contiguous states except for Delaware.
Populations were reported to be stable or increasing in 40 states, with 6 states unable to report population trends
and only 1 state (Florida) reporting decreases in bobcat populations. Of the 47 states in which bobcats occur, 41
employ some form of population monitoring. Population density estimates were available for 2,011,518 km2 (33.6%) of
the estimated bobcat range in the United States, with population estimates between 1,419,333 and 2,638,738
individuals for this portion of their range and an estimated 2,352,276 to 3,571,681 individuals for the entire United
States. These results indicate that bobcat populations have increased throughout the majority of their range in North
America since the late 1990s and that populations within the United States are much higher than previously
suggested.
Keywords: bobcat; Lynx rufus; monitoring; North America; population status

Received: December 13, 2009; Accepted: June 10, 2010; Published Online Early: June 2010; Published: November
2010
Citation: Roberts NM, Crimmins SM. 2010. Bobcat population status and management in North America: evidence of
large-scale population increase. Journal of Fish and Wildlife Management 1(2):169–174; e1944-687X. doi: 10.3996/
122009-JFWM-026
* Corresponding author: nmr25@cornell.edu
Introduction hispidus (Schnell 1968), and pronghorn Antilocapra

americana (Beale and Smith 1973). Thus, in many
Bobcats Lynx rufus are one of the most widely spread situations, proper management of bobcat populations
and adaptable carnivores in North America (Anderson may be an important component of managing for
1987). Bobcats serve as a top predator in many ecosystem function and biodiversity.
ecosystems (Conner et al. 2001) and can have significant In the United States, management of bobcat popula-
effects on ecosystem function. For example, bobcats tions is the responsibility of state wildlife management
have been identified as a significant source of predation, agencies. However, management of bobcats, as with
in specific systems, for white-tailed deer Odocoileus many other carnivores, can be difficult due to their
virginianus (Nelms et al. 2001), cotton rats Sigmodon elusive nature and the lack of information on demo-
Bobcat Status and Management in North America N.M. Roberts and S.M. Crimmins
Table 1. Bobcat monitoring methods (Monitoring), population estimates (N), range extent (Habitat, in km2), and population
status (Status; 1981–present) for selected jurisdictions in North America. NR indicates data were not reported by jurisdiction.
Monitoring methods were public sightings (PS), harvest analysis (HA), hunter surveys (HS), population models (PM), scent or sign
stations (SS), snow track surveys (TS), incidental harvest (IH), vehicle collisions analysis (VC), or other (OT).
Jurisdiction Monitoring N Habitat Status
United States
Alabama HA 68,625 116,550 Increasing
Arizona HA,HS 62,395–65,909 294,315 Increasing
Arkansas HS 14,049 134,874 Unknown
California NR 69,429–72,735 330,614 Stable
Colorado NR NR NR NR
Connecticut PS,VC Unknown 11,655 Increasing
Delaware - 0 - Absent
Florida NR 303,338 151,669 Decreasing
Georgia HA,SS 209,870–249,845 149,907 Increasing
Idaho NR 1,705–13,640 170,500 Increasing
Illinois HS 2,252 143,960 Increasing
Indiana PS,IH Unknown 92,434 Increasing
Iowa PM,OT 1,155–2,331 34,153 Increasing
Kansas HA,HS,SS 29,666–31,785 211,900 Increasing
Kentucky HA 14,000 102,896 Increasing
Louisiana HA,HS 66,431 112,825 Increasing
Maine HA Unknown 40,000 Stable
Maryland SS Unknown Unknown Increasing
Massachusetts NR Unknown 12,051 Unknown
Michigan HS,SS Unknown 116,550 Increasing
Minnesota HA,PM,SS,TS 2,857 103,600 Increasing
Mississippi HA 52,436 90,753 Increasing
Missouri HA,HS,SS 18,000–20,000 121,489 Increasing
Montana HA,TS 7,641 375,550 Increasing
Nebraska NR Unknown 171,000 Increasing
Nevada HA,PM Unknown 284,449 Stable/Increasing
New Hampshire HS,VC,IH Unknown 20,098 Increasing
New Jersey OT 106 1,600 Increasing
New Mexico HA,SS 36,249–54,373 181,243 Stable
New York PS,HA,HS 5,078 103,600 Increasing
North Carolina HA 90,115 126,161 Increasing
North Dakota HA Unknown 36,838 Unknown
Ohio PS Unknown 22,286 Increasing
Oklahoma HA Unknown 178,202 Increasing
Oregon None Unknown 248,630 Unknown
Pennsylvania HA,VC,IH,PS,PM 18,766 85,300 Increasing
Rhode Island PS,VC Unknown Unknown Unknown
South Carolina HA,SS 40,552–94,622 56,773 Stable
South Dakota HA,PM Unknown 100,000 Increasing
Tennessee HA Unknown 106,739 Stable
Texas HA 287,444–1,357,928 495,594 Stable
Utah HA Unknown 202,020 Stable
Vermont HA 2,100–3,500 23,310 Increasing
Virginia HA Unknown 64,118 Increasing
Washington NR Unknown 170,000 Stable
West Virginia HA 8,743 56,656 Increasing
Wisconsin HA,HS,PM,TS 2,850 46,620 Increasing
Table 1. Continued.
Jurisdiction Monitoring N Habitat Status
Wyoming HA,SS 2,481–13,783 253,601 Increasing
Canada
British Columbia HA Unknown NR Stable
Manitoba NR Unknown NR Stable
New Brunswick HA,TS,OT Unknown 72,784 Stable
Nova Scotia HA,OT 5,170–9,910 43,086 Fluctuating
Ontario HA Unknown 400,015 Stable
Quebec PS,IH Unknown 55,000 Stable/Increasing
Saskatchewan HA,HS Unknown 248,639 Stable
Mexico NR Unknown 1,702,545 Unknown
graphic rates. Changes in bobcat management in with previous studies and population estimates (USFWS
response to population trends are common (Rolley et 1982; Woolf and Hubert 1998) and for brevity.
al. 2001); however, most population assessments or
studies of bobcat range have been limited in geographic Data analysis
scope (e.g., Woolf et al. 2002). It is generally assumed by We compared the proportion of states reporting
managers that a harvest rate of 20% is sustainable (Knick increasing or stable populations between 1996 and
1990), but that variability in environmental factors may 2008 (Woolf and Hubert 1998) using chi-square tests. We
confound this (Anderson and Lovallo 2003). The vast only included states that reported population trends in
geographic range of bobcats in North America highlights this analysis. States that reported populations as ‘‘stable/
the need for large-scale population assessments. Bluett increasing’’ were assigned a status of ‘‘stable’’ for this
et al. (2001) reported that state agencies considered analysis. We estimated minimum population abundance
distribution and relative abundance to be among the and total suitable habitat as the sum of population or
most important research topics for bobcats. habitat estimates across all jurisdictions. Because not all
Woolf and Hubert (1998) provided the most recent states reported estimated population abundance (n =
assessment of bobcat population distribution and 26) or area of suitable habitat (n = 5), we considered
management in North America, with the most recent these minimum estimates to be very conservative. To
range-wide population assessment occurring in 1981 estimate total population abundance across the contig-
(USFS 1981). However a more contemporary assess- uous United States, we calculated average bobcat
ment is needed because human populations and density for those states reporting both population and
development have increased greatly during the past area of habitat estimates and then multiplied this
decades. Although numerous regional or local studies average by the geographic range estimates for all states
have suggested that bobcat populations are increasing that reported range estimates but no population
(e.g., Woolf et al. 2000), no empirical evidence of range- estimates (n = 17) and added this value to the minimum
wide increases in population abundance exists. Our count. This method provided a general estimate of
objective was to document monitoring efforts, spatial population abundance across the entire United States
distribution, and population trends of bobcats in North that facilitates comparison with historic estimates that
America. used similar methods (USFWS 1982).
Methods Results
Agency surveys We received responses from 47 of the 48 states, 7
We contacted wildlife management agencies from all Canadian provinces, and Mexico. Responses indicated
of the 48 contiguous states of the United States, 7 total area of suitable bobcat habitat of 8,708,888 km2,
Canadian provinces (British Columbia, Manitoba, New with 71% (6,186,819 km2) in the United States, 20% in
Brunswick, Nova Scotia, Ontario, Quebec, Saskatchewan), Mexico (1,702,545 km2), and 9% in Canada (819,524 km2).
and the country of Mexico in 2008. Each jurisdiction was Twenty-eight (50.9%) of the jurisdictions reported
asked to report: 1) what methods are employed to population estimates or densities (Table 1). Of the 48
monitor bobcat status, 2) population estimates and jurisdictions that reported population trends, 31 (64.6%)
average densities (if available), 3) the area (reported here reported increasing populations while 15 (31.3%) report-
in km2) of suitable bobcat habitat within the jurisdiction, ed stable populations, and 1 reported fluctuating
and 4) post-1981 population trends. Monitoring methods populations. Only one jurisdiction, Florida, reported
were classified as public sightings, harvest analysis, decreasing bobcat populations (Figure 1). The minimum
hunter surveys, population models, scent or sign population estimate in the United States was between
stations, snow track surveys, incidental harvest, vehicle 1,418,333 and 2,637,738 individuals, representing 68.4%
collisions analysis, or other. We primarily present of the total estimated bobcat range in the United States
information from U.S. states to facilitate comparison (Table 1). Using average density, the total population
Table 2. Methods used to monitor bobcat populations in

the lower 48 states and 7 Canadian provinces in 2008 (45
jurisdictions reporting). Twenty-five jurisdictions used . 1
method.
Method No. of jurisdictions
Harvest data analysis 33
Public sightings 7
Hunter surveys 11
Population models 6
Scent or sign stations 9
Snow track surveys 4
Figure 1. Bobcat population trends in the contiguous United Incidental harvest 4
States, 2004. Vehicle collision analysis 4
Other 4
abundance in the United States was estimated to be None 1
between 2,352,276 and 3,571,681 individuals. A signifi-
cantly higher (x2 = 5.64, P = 0.018, df = 1) proportion of
states reported increasing populations in 2008 than in mately 68% of the reported bobcat range. In 1981 similar
1996. methods were used to estimate bobcat populations; at
Of the 55 jurisdictions surveyed, 45 (81.2%) reported that time it was estimated there were 725,000–1,017,000
methods used to monitor bobcat populations, 25 (55.6%) bobcats in the United States (USFWS 1982). Using these
of which used . 1 method. Harvest data analysis and methods, our data indicate that current bobcat popula-
hunter surveys were the most commonly used methods tion abundance in the entire United States is 2,352,276–
(Table 2). No jurisdiction used . 5 methods to monitor 3,571,681. This growth is likely the result of many factors
bobcats. Only one jurisdiction (Oregon) reported having including changing agricultural and land-use practices,
bobcat populations but no monitoring method. range expansion, and habitat improvement programs
such as the Conservation Reserve Program (FSA 1985, 16
Discussion U.S.C. 3831). In addition, advances in wildlife manage-
Bobcats are highly adaptable carnivores and are the ment and monitoring have allowed states to greatly
most abundant of North America wild felids (Cowan improve wildlife management programs in recent
1971; Anderson and Lovallo 2003). Their ecological role decades, particularly furbearer management. As recently
and economic value as furbearers are the primary as 1971, 40 of the 48 contiguous United States had no
motivations behind research and management of protection or formal management of bobcats (Faulkner
bobcats (Woolf and Nielsen 2001). Woolf and Hubert 1971). In New York, for example, bobcats were desig-
(1998) reported that less than half of the states in the nated as an unprotected species and take was unregu-
contiguous United States (n = 20) had increasing bobcat lated through 1976, only 5 y prior to the 1981 population
populations as of 1996; however, we found that 32 states estimate (USFWS 1982). After 1976, bobcats were
now report increasing populations (Table 1). All other afforded legal status as a furbearer species to be
states that reported bobcat population trends, with the conserved, and harvest regulations were developed
exception of Florida, reported at least stable populations. and implemented. Similar shifts in management para-
Although the magnitude of population increase likely digms have occurred in many jurisdictions; these have
varies considerably among jurisdictions, the general trend clearly benefited bobcat populations. Bobcat harvests
supports the hypothesis that bobcat populations are did increase during 1976–1984, but recent harvest levels
expanding across much of their geographic range. The are comparable to those three decades earlier (36,674
reason for declining populations in Florida is unknown, harvested during 1997–1998 verses 37,026 during 1975–
although extensive human development may be partly 1976), despite this increase in population size (Associa-
responsible. Given the stability of bobcat populations tion of Fish and Wildlife Agencies, unpublished data).
elsewhere in the United States, the decreasing population This is likely due to decreased harvest effort.
in Florida does not pose any immediate threat to the Because population survey methods were not stan-
large-scale persistence of bobcat populations in the dardized across jurisdictions, there is likely substantial
United States. However, research into the vital rates and variability in the accuracy of the estimates provided by
demographic patterns of the Florida bobcat population management agencies. For example, only six states used
could provide useful information to wildlife managers if any form of population model in their monitoring of
populations elsewhere begin declining. bobcat populations, likely because of the paucity of
Minimum population estimates provided by jurisdic- accurate data available for developing such models.
tions indicated that the United States’ bobcat population Similarly, 19 states failed to report population estimates.
is . 1.4 million. This estimate is likely a substantial Although this variability may be substantial, it does not
underestimation of total population abundance, because preclude comparison to previous reports (USFWS 1982)
it only represented population estimates for approxi- or interpretation of jurisdiction-specific population
trends. Agency surveys such as this provide the only References

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Surveys
Recreational Carrying Capacity of Lake Umbagog
National Wildlife Refuge
Robert Manning,* William Valliere, Jeffrey Hallo
R. Manning, W. Valliere
Rubenstein School of Environment and Natural Resources, George D. Aiken Center, University of Vermont, Burlington,
Vermont 05405
Present address of R. Manning: 81 Carrigan Drive, Burlington, Vermont 05405
J. Hallo
Department of Parks, Recreation and Tourism Management, Clemson University, 280B Lehotsky Hall, Clemson, South
Carolina 29634
Abstract
Recreational carrying capacity addresses the issue of how much and what types of recreation can be accommodated in
parks and related areas without unacceptable impacts. Contemporary approaches to carrying capacity rely on
formulation, monitoring, and management of indicators and standards of quality. Recreational carrying capacity of
Lake Umbagog National Wildlife Refuge, located in northern New Hampshire and Maine, was analyzed using visitor
surveys that employed normative theory and methods and visual simulations of a range of recreation-related
conditions. Study findings suggest that indicators of quality for the visitor experience include the number of boats
seen on Lake Umbagog and associated rivers, the size of boating groups, the minimum acceptable chance of seeing
selected types of wildlife, and the minimum acceptable chance of catching selected types of fish. Study findings also
suggest a range of standards of quality for these indicators. Recreational carrying capacity of Lake Umbagog National
Wildlife Refuge can be managed by using study findings to guide formulation of indicators and standards of quality,
monitoring indicator variables, and taking management actions to ensure that standards of quality are maintained.
Keywords: carrying capacity; Lake Umbagog; normative research

Received: May 5, 2010; Accepted: August 27, 2010; Published Online Early: August 2010; Published: November 2010
Citation: Manning R, Valliere W, Hallo J. 2010. Recreational carrying capacity of Lake Umbagog National Wildlife Refuge.
Journal of Fish and Wildlife Management 1(2):175–182; e1944-687X. doi: 10.3996/052010-JFWM-011
* Corresponding author: robert.manning@uvm.edu
Introduction These initial scientific applications suggested that the

concept was more complex in this new management
How much recreational use can be accommodated in context. At first, as might be expected, the focus was
parks, wildlife refuges, and related protected areas? This placed on the impacts that recreational use can have on
long-standing and increasingly urgent question is often natural resources, including trampling of vegetation, soil
addressed under the rubric of carrying capacity (Manning compaction and erosion, water pollution, and wildlife
2007). The underlying concept of carrying capacity has a disturbance (Hammitt and Cole 1998; Leung and Marion
rich history in the natural resource professions. In 2000). However, it soon became apparent that there was
particular, it has been applied in wildlife and range another critical dimension of carrying capacity dealing
management where it refers to the number of animals with the quality of the visitor experience. Wagar (1964),
that can be maintained in a given habitat (Dasmann 1964). for example, in his early and important monograph on
Carrying capacity has obvious parallels and intuitive appeal the application of carrying capacity to outdoor recrea-
in the field of outdoor recreation. However, the first tion, reported that his study ‘‘was initiated with the view
rigorous applications of carrying capacity to management that carrying capacity of recreation lands could be
of parks and related areas did not occur until the 1960s. determined primarily in terms of ecology and the
Recreational Carrying Capacity of Lake Umbagog R. Manning et al.
deterioration of areas. However, it soon became obvious issue of crowding, but it also has been expanded to
that the resource-oriented point of view must be include other potential indicators of quality, including
augmented by consideration of human values.’’ Wagar’s ecological impacts of outdoor recreation (Manning et al.
point was that as more people visit a park or related area, 2004; Manning 2007).
not only can the natural resources of the area be affected Traditionally, norms have been measured through a
but the visitor experience can also be affected. For numerical approach. For example, respondents are asked
example, increasing recreational use can lead to per- to evaluate the acceptability of alternative use levels,
ceived crowding, conflict among visitors, and aesthetic such as a range of groups encountered per day along
degradation caused by natural resource impacts (Man- trails. Resulting data are aggregated to determine social
ning 2010). Thus, as applied to parks and protected norms. More recently, visual approaches to measuring
areas, carrying capacity has two primary components: norms have been developed (Manning et al. 1996;
resource and social. Manning and Freimund 2004). In this technique,
A growing body of research has identified a range of computer software is used to develop a set of visual
resource and social impacts caused by outdoor recrea- simulations of a range of recreation use levels and
tion (Hammitt and Cole 1998; Manning 2010). However, associated resource and social impacts.
the operative question posed by the issue of carrying The approach to carrying capacity described above
capacity is how much impact and associated use should has been used in an increasing number and variety of
be allowed (Manning 2007). To help answer this parks and protected areas with special emphasis on the
question, several carrying capacity frameworks have national park system (Manning 2007). In this article, this
been developed based on a management-by-objectives approach to carrying capacity is extended to Lake
approach. Prominent examples of these frameworks Umbagog National Wildlife Refuge (Lake Umbagog
include limits of acceptable change (Stankey et al. NWR), located in Coos County, New Hampshire, and
1985) and visitor experience and resource protection Oxford County, Maine, by identifying a suite of indicators
(Manning 2001). These frameworks rely on formulation of and standards of quality for the visitor experience. Lake
management objectives and associated indicators and Umbagog NWR is located in northern New Hampshire
standards of quality. and Maine and contains significant natural and recrea-
Management objectives are broad, narrative state- tional resources, including 2,833-ha Lake Umbagog,
ments that define the degree of resource protection and portions of three rivers flowing in and out of the lake,
the type of visitor experience to be provided. Indicators large areas of wetlands, and upland areas surrounding
of quality are measurable, manageable variables that the lake. It is estimated that the refuge accommodates
reflect the essence or meaning of management objec- approximately 50,000 recreation visits annually (U.S. Fish
tives; they are quantifiable proxies or measures of and Wildlife Service 2008).
management objectives. Indicators of quality may
include elements of both the natural and social Methods
environments. Standards of quality define the minimum
acceptable condition of indicator variables. Once man- Two surveys of visitors to Lake Umbagog NWR were
agement objectives and associated indicators and conducted to support formulation of management
standards of quality have been formulated, indicators objectives and associated indicators and standards of
of quality are monitored and management actions are quality for the visitor experience at Lake Umbagog NWR.
taken when and where needed to maintain standards of A phase 1 survey was conducted in summer 2006 to
quality. identify potential indicators of quality of the visitor
Research on standards of quality has relied on experience. The survey used both open- and close-ended
normative theory and methods developed in sociology questions. An open-ended question asked respondents
and extended to parks and outdoor recreation (Manning what they enjoyed most about their visit to the refuge.
1999; Vaske and Whitaker 2004; Manning 2007). Norms in Responses were coded verbatim and then combined into
parks and outdoor recreation are generally defined as several broad categories. A follow-up, close-ended
standards that individuals and groups use for evaluating question presented respondents with a list of 22
environmental and social conditions (Shelby and Vaske potential issues at Lake Umbagog NWR related to the
1991; Vaske et al. 1986). If visitors have normative quality of the visitor experience and respondents were
standards concerning relevant aspects of resource asked to rate the degree to which these issues were
conditions and recreation experiences, then such norms thought to be a problem, by using a three-point
can be studied and used as a basis for formulating response scale: ‘‘not a problem’’, ‘‘small problem,’’ and
standards of quality. ‘‘big problem.’’ Respondents also were asked to report
Application of norms to standards of quality has the recreation activities in which they engaged. Baseline
followed the work of Jackson (1965), who developed a characteristics of visitors and use patterns also were
methodology to measure norms (Vaske and Whitaker collected. Sampling was conducted at the four primary
2004; Manning 2007). Using these methods, the personal access points to the refuge. Interviewers were stationed
norms of individuals are aggregated to derive social at these access points on 10 randomly selected days in
norms. These social norms are often displayed graphi- July and August and asked all visitors to participate in
cally in the form of social norm curves. Normative the survey. The sampling days were spread over the days
research in outdoor recreation has focused largely on the of the week with sampling occurring during daylight
hours. Within each sampling period, a trained surveyor even) distribution of motorboats and nonmotorboats.
approached each group exiting the site and asked them Respondents were asked the same series of questions as
to participate in the survey. Visitors completed the on- described above. A series of six study photographs
site questionnaire in the presence of the surveyor, who (mounted on 61 cm 6 91 cm poster boards) also were
answered any questions that arose and collected the used to present a range of fishing boats in a generic cove
questionnaires upon completion. In total, 256 visitors on Lake Umbagog (Figure S3, http://dx.doi.org/10.3996/
were approached, and this yielded 197 completed 052010-JFWM-011.S1). The number of boats ranged from
questionnaires representing a 77% response rate. The 0 to 20. Respondents were asked the same series of
high response rate minimizes the possibility of nonre- questions as described above.
sponse bias (Vaske 2008; Dillman et al. 2009). The sample Standards of quality for group size of boaters (number
size of approximately 200 yields a margin of error of 7% of boats in a group of visitors) were measured by asking
or less at the 95% level of confidence (Lohr 1999). respondents to rate the acceptability (using the 9-point
A phase 2 survey was conducted in 2007 to identify response scale described above) of groups that ranged
potential standards of quality for indicator variables from 1 to 10 boats. Standards of quality for seeing
identified in the phase 1 survey. These indicators were wildlife were measured by asking respondents in an
number of boats seen at one time on the lake, number of open-ended series of questions to report the minimum
boats seen at one time on rivers, number of boats seen acceptable percentage of visitors who should be able to
at one time in fishing coves, size of boating groups, see each of several types of wildlife while visiting Lake
minimum acceptable chance of seeing selected types of Umbagog NWR. Standards of quality for catching fish
wildlife, and minimum acceptable chance of catching were measured in a similar way: respondents were asked
selected types of fish. The same survey and sampling to report the minimum acceptable percentage of visitor
methods as described above were used. In total, 254 who should be able to catch several species of fish.
visitors were approached, and this yielded 193 complet-
ed questionnaires representing a 76% response rate. Results
Once again, the high response rate minimizes the
possibility of nonresponse bias. Data from the phase 2 Baseline characteristics of visitors and use patterns
survey were used to derive a series of norm curves were quite similar across the two surveys (Supplemental
(described below). The level of agreement (or ‘‘crystal- Material, Table S1, http://dx.doi.org/10.3996/052010-
lization’’) about the points that describe these norm JFWM-011.S2). In the phase 1 survey, the majority of
curves can be measured using Van der Eyk’s A (Van der visitors (92.1%) were from the New England states, most
Eyk 2001; Krymkowski et al. 2009). The A statistic is (60.9%) were male, and the average group size was 3.6
always between 21 and +1, with 21 indicating complete people. Most visitors (67.2%) had been to Lake Umbagog
disagreement, +1 complete agreement, and 0 uniformly NWR before, and repeat visitors have been visiting the
distributed responses across all values of the response area for an average of 11 y and have visited an average
scale. Baseline characteristics of visitors and use patterns of six times in the past 12 mo. On average, visitors spend
also were collected. nearly 5 h/d in Lake Umbagog NWR, and approximately
Three series of visual simulations were used to one-third of visitors camp at either the state park
measure standards of quality for the number of boats campground at the southern end of Lake Umbagog or
seen in three contexts: on the lake, on rivers, and in at one of the remote campsites on the lake shore. The
fishing coves. For the lake, six study photographs most popular recreation activities were viewing wildlife
(mounted on 61 cm 6 91 cm poster boards) were used (85.8%); paddling (63.5%); swimming (52.8%); photogra-
showing a range of 0 to 20 boats, each with an even phy (52.3%); camping (49.7%); motorboating (41.1%);
distribution of motorboats and nonmotorboats (Figure and fishing for bass (47.2%), trout and salmon (26.4%),
S1, http://dx.doi.org/10.3996/052010-JFWM-011.S1). Re- and other fish (27.4%).
spondents were asked to rate the ‘‘acceptability’’ of each Four categories of variables were mentioned by more
photograph on a 9-point scale bounded by 24 (‘‘very than 10% of visitors for what they enjoyed most about
unacceptable’’) and 4 (‘‘very acceptable’’), with a neutral their visit: seeing wildlife and catching fish (22.4%);
point of 0. Respondents also were asked to report the enjoying a specific recreation activity (e.g., boating,
study photograph that showed the number of boats fishing; 20.9%); enjoying the peaceful, uncrowded
they 1) would prefer to see (‘‘preference’’), 2) would find character of the area (15.8%); and enjoying the natural
so unacceptable they would no longer visit Lake beauty of the area (12.6%). The follow-up, close-ended
Umbagog NWR (‘‘displacement’’), 3) felt should be question found that the three most problematic issues all
allowed to use the lake before visitors were restricted related to motor boats—their number, noise, and speed
from using the area (‘‘management action’’), and 4) on the lake and on rivers.
thought looked most like the number of boats they Visitor responses to the series of normative questions
typically saw on the lake on the day they were surveyed are shown in Figures 1–4 and Tables 1 and 2. The values
(‘‘typically seen’’). A series of six photographs (mounted for ‘‘acceptability’’ in Table 1 are the point at which the
on 61 cm 6 91 cm poster boards) also were used to social norm curves cross the neutral point of the
present a range of boats on refuge rivers (Figure S2, http:// acceptability scale (i.e., the point at which the aggregate
dx.doi.org/10.3996/052010-JFWM-011.S1). The number of ratings of acceptability fall out of the acceptable range
boats ranged from 0 to 15, each with and even (or near and into the unacceptable range). Levels of agreement
Figure 1. Social norm curve for number of boats seen at one time on Lake Umbagog. Photos illustrating a range of boats on Lake
Umbagog are shown in Figure S1 (http://dx.doi.org/10.3996/052010-JFWM-011.S1). The social norm curve traces the mean
acceptability rating for each of the study photographs.
for boating-related standards of quality (as measured by open-ended comments of visitors noting that seeing
Van der Eyk’s A) are moderate to high, and most wildlife and catching fish were the most enjoyable
respondents reported that they saw fewer boats than the aspects of visiting Lake Umbagog NWR, suggests that
maximum number acceptable suggesting that recrea- opportunities to see wildlife and catch fish are poten-
tional carrying capacity has not been exceeded. tially good indicators of quality for this area. Moreover,
the number of boats on the lake and on rivers and
Discussion associated issues of speed and noise of motorboats were
reported by visitors as the most problematic issues,
Study findings provide empirical guidance to help suggesting that the number of boats and size of boating
formulate a suite of indicators and standards of quality groups using Lake Umbagog also may be important
for analyzing and managing recreational carrying capac- indicators of quality.
ity at Lake Umbagog NWR. The popularity of wildlife A range of standards of quality were identified for
viewing and fishing as recreational activities, along with these indicator variables. Using visual simulations of a
Figure 2. Social norm curve for the number of boats seen at one time on refuge rivers. Photographs illustrating a range of boats
on rivers are shown in Figure S2 (http://dx.doi.org/10.3996/052010-JFWM-011.S1). The social norm curve traces the mean
acceptability rating for each of the study photographs.
Figure 3. Social norm curve for number of fishing boats seen at one time in a generic cove of Lake Umbagog. Photographs
illustrating a range of fishing boats in fishing coves on Lake Umbagog are shown in Figure S3 (http://dx.doi.org/10.3996/052010-
JFWM-011.S1). The social norm curve traces the mean acceptability rating for each of the study photographs.
range of boats in three contexts (on the lake, on rivers, question asked respondents: ‘‘Which photograph shows
and in fishing coves), respondents reported a series of the highest level of use you think should be allowed on
standards of quality that ranged from the number of the lake? In other words, at what point should visitors be
boats they preferred to see to the number of boats that restricted from using the lake?’’
would cause them to stop visiting the area (Table 1). A However, other points along the range of potential
standard of quality in the low end of these ranges would standards of quality also might be selected, depending
provide a very high-quality visitor experience but may upon management objectives for the area and other
ultimately limit the number of visitors who can use the considerations. For example, the relatively remote
area. A standard of quality in the high end of the ranges location of Lake Umbagog NWR may suggest that
would allow substantial public access, but the quality of management emphasize opportunities for solitude and
the visitor experience may be relatively low. The intimate contact with nature, thus selecting standards of
‘‘management action’’-based standard has some special quality that will provide a very high quality of experience
appeal in that it explicitly asks respondents to consider for these values. Decisions about standards of quality
the inherent trade-offs between quality and access. The also might consider the objective of providing a diversity
Figure 4. Social norm curve for boating group size on Lake Umbagog and associated rivers. The social norm curve traces the mean
acceptability rating for each of the boat group sizes.
Table 1. Summary table for standards of quality for boating on Lake Umbagog.
Mean Median Agreementa
No. of boats seen at one time on Lake Umbagog
Acceptability 13.1 0.9–0.85
Preference 5.7 4.0 0.45
Displacement 16.8 20.0 0.68
Management action 11.5 12.0 0.41
Typically seen 8.0 8.0 0.51
No. of boats seen at one time on refuge rivers
No. of fishing boats seen at one time in a generic cove of Lake Umbagog
a
Van der Eyk’s A statistic.
of visitor experiences at Lake Umbagog NWR or even indicators of quality be monitored and management
throughout wildlife-oriented recreation areas in the actions taken when needed to help ensure that
broad geographic area. This spatial zoning approach standards of quality are maintained. The indicators of
may be effective in meeting potentially wide-ranging quality identified in this study could be monitored in
demands for outdoor recreation among the public by several ways. Periodic visitor surveys could be conducted
providing a spectrum of recreation opportunities from asking respondents to report the types of wildlife seen
which people can choose. This approach is in keeping and catch rates for selected species of fish. The study
with the Recreation Opportunity Spectrum framework photographs illustrating a range of boats on the lake, on
designed to encourage diverse recreation opportunities rivers, and in fishing coves also could be used by simply
(Clark and Stankey 1979) and the goal of providing for asking visitors to report the photograph that looks most
the needs of a diverse public (Manning 2010). Formula- like what they experienced on the day of the survey. This
tion of standards of quality also should consider approach was used in the phase 2 survey, and resulting
opportunities and constraints posed by natural resource data suggest that current use levels are relatively low
considerations, such as the sensitivity of wildlife species compared with standards of quality, as reported here.
and other environmental resources. Alternatively, refuge staff could monitor the number of
The contemporary carrying capacity frameworks de- boats directly through observation or photography.
scribed at the beginning of this article require that A diverse array of management actions might be used
to help ensure that standards of quality are maintained.
The professional literature in outdoor recreation sug-
Table 2. Summary table for minimum acceptable gests four basic strategies for managing outdoor
percentage chance of seeing wildlife and catching fish at recreation: increase the supply of opportunities, reduce
Lake Umbagog National Wildlife Refuge. the demand, reduce the impact of use, and increase the
durability of the resource (Manning 2010). A wide variety
Minimum % chance of tactics or specific management practices are available
Mean Median within each of these strategies. Examples include visitor
information and education programs, providing hard-
See loon 61.7 70.0
ened facilities such as trails and campsites, and
See eagle 51.5 50.0 establishing a visitor permit system. Managing the
See osprey 50.6 50.0 percentage chance of seeing wildlife and catching fish
See waterfowl 67.8 75.0 may be especially challenging given the mobility of
See moose 43.8 30.0 wildlife, changing environmental conditions, and varia-
Catch bass 61.9 50.0 tion in visitor use levels and behavior (Anderson et al.
Catch trout and salmon 43.2 30.0 2010). However, advances in research on the relation-
Catch other fish 56.2 50.0
ships between visitor use levels and behavior and the
presence of wildlife may enhance the ability of managers
to address this issue (Klein et al. 1994; Monz et al. 2004). Acknowledgments
Managers also might increase chances of encountering
wildlife by educating visitors about optimum locations Appreciation for assistance in conducting this study is
and times for wildlife viewing and fishing. expressed to Lake Umbagog National Wildlife Refuge
Recreational carrying capacity is a challenging but staff Paul Casey and Laurie Wunder and to Carol Foss of
New Hampshire Audubon. The Subject Editor and two
increasingly important issue in management of parks and
anonymous reviewers provided helpful comments on
protected areas. The information developed in this study
this article.
offers an empirical basis for analyzing and managing
recreational carrying capacity at Lake Umbagog NWR and
other units of the NWR System. Specifically, it identifies
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Issues and Perspectives
Arguments for Using Population Models in Incidental
Take Assessments for Endangered Species
Conor P. McGowan,* Mark R. Ryan
Department of Fisheries and Wildlife Sciences, University of Missouri, 302 Anheuser-Busch Natural Resources Building,
Columbia, Missouri 65211
Present address of C.P. McGowan: U.S. Geological Survey, Alabama Cooperative Fish and Wildlife Research Unit, School
of Forestry and Wildlife Sciences, Auburn University, Auburn, Alabama 36849
Abstract
Population models can be useful tools for evaluating management strategies and risks for a given species. A major, but
often overlooked, component of endangered or threatened species management and recovery is the incidental take
allowance of many endangered species laws. In the United States population models are seldom applied to address
specific incidental take scenarios. We believe it is prudent for wildlife management agencies like the U.S. Fish and
Wildlife Service to embrace explicit predictive tools to assess the possible effects of incidental take actions and to set
standards for what constitutes unacceptable levels of incidental take in terms of predicted effect on population
viability, recovery, and extinction. We briefly give recommendations for incorporating simulation models into jeopardy
evaluations in ways that would dovetail with legislative language and provide a simple example model. Using explicit
predictive models to support jeopardy determinations and incidental take decision-making would lead to transparent
decisions rooted in measurable quantities such as changes in extinction probability or abundance projections.
Keywords: biological opinions; endangered species; incidental take; jeopardy; population models, section 7
consultations
Received: June 3, 2010; Accepted: September 9, 2010; Published Online Early: September 2010; Published: November
2010
Citation: McGowan CP, Ryan MR. 2010. Arguments for using population models in incidental take assessments for
endangered species. Journal of Fish and Wildlife Management 1(2):183–188; e1944-687X. doi: 10.3996/062010-JFWM-
014
* Corresponding author: cmcgowan@usgs.gov
Introduction and endangered species (Boyce 1992; Beissinger and

McCullough 2002; Morris and Doak 2002).
Predicting the future of an endangered or threatened A major, but often overlooked, component of endan-
species is a key part of planning management actions gered or threatened species management and recovery
and recovery (National Research Council 1995). In 1995 is incidental take (hereafter, ‘‘take’’) allowances, which
the National Research Council concluded that predictive are defined as exceptions to the prohibitions and
population models were extremely useful tools for protections under endangered species laws. For exam-
evaluating listing and delisting of an endangered ple, Section 10 of the U.S. Endangered Species Act (ESA),
species, assessing management strategies, and evaluat- as amended, states ‘‘The Secretary may permit…any
ing risks to the species. The National Research Council taking otherwise prohibited… if such taking is incidental
(1995) recommended that population models be used to, and not the purpose of, the carrying out of an
wherever possible for those purposes in endangered otherwise lawful activity,’’ and if ‘‘the taking will not
species management and policy decisions. Population reduce appreciably the likelihood of the survival and
models and population viability analyses have a long recovery of the species in the wild’’ (ESA 1973). Similar
history and have grown in importance for managing rare exceptions exist in endangered species legislation of
Population Models in Take Assessments C.P. McGowan and M.R. Ryan
other countries. For example, the Bern Convention on take effects on endangered or threatened populations
the conservation of European wildlife and natural (McGowan 2008; McGowan and Ryan 2009).
habitats prohibits destruction of protected species but As we see it, the benefits of models for the type of
permits exceptions ‘‘to prevent serious damage to crops, assessments related to endangered species take deci-
livestock, forest, fisheries, water and other forms of sion-making are three-fold. First, the resulting decisions
property…’’ as long as ‘‘the exception will not be are based on mathematical and probabilistic predictions.
detrimental to the survival of the population concerned’’ A model can tell the decision maker what is likely to
(E.U. Convention on the Conservation of European happen in future and how some proposed take is likely
Wildlife and Natural Habitats, 1981, Bern, 19.IX.1979, to affect the endangered population. Second, a formal-
Article 9). The National Research Council’s book, ‘‘Science ized model is explicit and open for review from other
and the Endangered Species Act’’ (1995), has entire scientists. Decisions based on a model (if well-crafted
chapters and sections dedicated to listing decisions, and reasonable) are immediately scientifically defendable
critical habitat designations, and habitat conservation because there are no hidden assumptions. In an
planning, but only addresses take within the context of appropriately documented explicit model, the assump-
other regulatory or management issues. The National tions used to make the predictions can be clearly
Research Council (1995) did, however, call for the use of articulated so that all involved with and affected by the
population viability analyses when evaluating take decision can see and understand the basis on which the
decisions wherever possible. The language of the decision was made. All models contain some assump-
legislative documents (i.e., ‘‘probability of survival…in tions about the system or process represented by the
the wild’’) is well-suited for use with population model and all management decisions involve some
projection models that produce predictions regarding model or method for making predictions about how
extinction probability and future population growth and the management actions will affect the system (Williams
abundance. et al. 2002). We are advocating for a process that
The legislative phrase ‘‘reduce appreciably the likeli- transparently articulates the methods used to make
hood of survival and recovery of the species in the wild’’ those predictions and the assumptions embedded within
is often referred to as ‘‘causing jeopardy’’ in U.S. that predictive model. Lastly, models can account for
legislative and government documents (U.S. Fish and uncertainty in predictions and incorporate that uncer-
Wildlife Service [USFWS] 1998). Jeopardy determina- tainty into the decision-making process. All decisions in
tions come under the Section 7 Consultation process, natural resource management are subject to uncertainty,
which (in theory) is a straight forward process. If, whether it is environmental stochasticity (e.g., unpre-
during the planning stages for any action, a Federal dictable weather events), partial controllability (e.g., a
agency determines that the actions ‘‘may affect’’ a listed prescribed fire gone awry), partial observability (e.g.,
species, the agency requests the initiation of a Section sampling or parameter estimation errors), or ecological
7 consultation. If the action agency determines that uncertainty (e.g., lack of knowledge about system
the action ‘‘may adversely affect’’ a listed species or function or population dynamics; Williams et al. 2002).
designated critical habitat, it requests ‘‘formal consulta- Appropriately crafted models can mimic some or all of
tion.’’ To analyze an action and the effects on any listed that uncertainty and make probabilistic predictions
species and critical habitat, the USFWS writes and about the outcomes of take proposals and the probable
issues a ‘‘biological opinion.’’ A biological opinion effects of take on population abundance, extinction
consists of a review of relevant literature and data to probability, or recovery probability. Objective-driven
assess the status of the species in question and to assess decision-making (e.g., maximizing species recovery
the effects of the proposed actions and whether they will probability, or minimizing species extinction probability)
cause jeopardy for the listed species. Biological opinions supported by appropriate population models is trans-
conclude with an ‘‘incidental take statement’’ that parent, defendable, and robust to uncertainty. Models
articulates what levels of take are anticipated and how that acknowledge and incorporate uncertainty can also
much take is permissible (USFWS 1998). Section 7 of the allow decision-making to proceed under the uncertainty,
ESA and the pursuant determination of what actions and can provide opportunities to learn about system
amount to jeopardy can have major implications for function to reduce uncertainty in the future. Effective
management and recovery of a listed species (Rohlf monitoring of the effects of take on an endangered
2001). population and comparisons to model predictions can
Jeopardy decisions for any take situation provide lead to insights on population dynamics and model
opportunities to apply predictive quantitative models accuracy. We argue that projection models should be
that could evaluate the effects of the proposed actions in used wherever possible for take decisions and that a
terms of extinction probability or population growth of standardized approach for the use of those models
the protected species (National Research Council 1995). should be adopted.
The phrasing ‘‘reduce the probability of survival…in the
wild’’ implies that some evaluation of future extinction An Example—Piping Plover
risk and the changes in extinction risk due to the Biological Opinions
proposed actions is conducted in making jeopardy
decisions. But, thus far, to our knowledge there have McGowan (2008) looked at 11 biological opinions for
been few attempts to use population models to evaluate piping plovers Charadrius melodus from throughout the
species’ range and examined and evaluated the use of (McGowan and Ryan 2009). We argue that models are
population models in those opinions. Piping plovers are the only tools available to thoroughly evaluate the
small migratory shorebirds that breed on beaches of the effects of management actions and take allowances that
Atlantic Coast of North America, on beaches in the Great can account for and incorporate the variety of sources of
Lakes, and on rivers, lakes, and wetlands in the northern uncertainty that factor into a jeopardy decision. In fact,
Great Plains (Elliott-Smith and Haig 2004). The piping making any prediction about the effect of some action
plover was listed as threatened in the Atlantic Coast and on an ecosystem or species requires a model of the
the Great Plains and endangered in the Great Lakes in system or population dynamics (Williams et al. 2002).
mid-1980s (USFWS 1985). The piping plover is a
representative species for this evaluation because its Developing Models
ecology and population dynamics have been studied
extensively. Furthermore, it has been listed under the Developing predictive models of system dynamics does
ESA for over 20 y and there have been numerous not require tremendous data sets and detailed knowledge
biological opinions filed on many different federal of system dynamics (Starfield 1997), but rather some
projects throughout the species’ range (Sidle et al. notion of system processes and the ecology and
1991). There have also been several attempts to model population dynamics of the species concerned. Models
piping plover populations, each with varying methodol- need to be only as complex as necessary to make
ogies and results for the USFWS to evaluate and chose adequate predictions about the issue being modeled.
among. Some situations require very complex models, such as the
The use of piping plover population models and the loggerhead sea turtle Caretta caretta model developed by
results of the models varied across opinions, time, and Crouse et al. (1987) and Crowder et al. (1994). Those
geographic range. For the Atlantic Coast breeding studies used stage-based matrix models to simulate turtle
population, the 1996 Recovery Plan (USFWS 1996) populations and their models required significant com-
included a population model that set target fledge ratios plexity because the issue being studied was the effect of
(the number of chicks successfully produced per fishing bycatch mortality of juvenile turtles on population
breeding pair) to guide management. All the opinions viability. The models required isolating the juvenile age
from that region examined the proposed take in terms of class from other stages of development and applying
the target fledge ratios, and determined permissibility bycatch to that stage specifically. Alternatively, models
based on some assessment of whether those target could be as simple as linear regression of a time series of
ratios would be reached when the action took place. abundance data (e.g., Dennis et al. 1991; Morris and Doak
For the Great Plains population there have been three 2002) where the regression slope parameter is equivalent
published models since 1993 (Ryan et al. 1993; Plissner to the finite population growth rate. Predictions about
and Haig 2000; and Larson et al. 2002), which vary in future population size and extinction risk would be as
mathematical structure and complexity. McGowan (2008) simple as multiplying the population size by the estimated
reported that opinions from that region primarily growth rate to return the expected population size in the
determined jeopardy by determining whether or not future. Applying variability to the growth rate parameter
action would result in meeting the desired fledging and replicating the projection numerous times could
target from the first published model, which had the represent uncertainty about future environmental varia-
lowest required fledge ratios to maintain a stable tion and growth-rate estimation uncertainty. However,
population size. McGowan (2008) reported that only even deterministic simulations (explicit predictions with
one of the examined biological opinions discussed the no variability) would be a more transparent approach than
effects of the proposed action on the extinction intuition or undefined unarticulated processes for evalu-
probability of the population. All others discussed the ating take. These models can easily be implemented in
effects of the proposed actions in terms of achieving statistical programs such as R (R Development Core Team
target fledge ratios that the population models conclud- 2009), but could also be developed in spreadsheet-type
ed were necessary for maintaining stable population programs such as MicrosoftH Excel, a program widely used
growth. The recovery criteria for this population are for data management in the field of natural resources.
stated in terms of regional population size targets and
not in terms of fledge ratios. In those opinions, McGowan A Simple Population Model Example
(2008) found no discussion of how not achieving the To exemplify the benefits of modeling we have crafted
prescribed fledge ratios would decrease or change the a very simple population model to represent a threat-
probability of survival in the wild or of reaching the ened or endangered species similar to the count-based
proposed delisting criteria. population viability analysis presented in Morris and
From this limited and brief examination of biological Doak (2002). The equation to predict future population
opinions written as part of Section 7 Consultation in U.S. size simply multiples current population size by the
law to evaluate the effects and permissibility of take, we estimated population growth rate to predict the
conclude that wider and more standardized use of population size at the next time step:
population models should be used. Population viability
analysis–type population models are well-suited to Ntz1 ~Nt l ð1Þ
evaluate extinction risk (Morris and Doak 2002) and,
potentially, changes in extinction risk due to take where N is the population abundance, t is time, and l is
the population growth rate. Initial N is estimated from parameters to see how the population responds to the
ongoing monitoring, counting surveys, or mark–recap- proposed take under different conditions.
ture studies, and l is estimated by examining a time Using Microsoft Excel and beginning with a population
series of population counts. size of 500 individuals, modeling l as a normally
distributed random variable with mean 0.995 (SD =
lt ~Nt =Nt{1 ð2Þ 0.02; NORMINV(RAND(), 0.995, 0.02), code for a normally
distributed random variable in Microsoft Excel) and
And after a number of successive population counts the applying annual take as a uniform random variable
geometric mean of l can be estimated: varying between 5 and 10 individuals/y (RANDBETWEEN
qffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffi (5,10), Microsoft Excel code for a uniform random
l~ y lt~1 |lt~2 | . . . |lt~y ð3Þ variable between the upper and lower value), we
simulated the population 50 y into the future and
where y is the number of years for which l is estimated. replicated the simulation 5,000 times (see Supplemental
Our approach assumes that past population growth rate Material, Table S1, http://dx.doi.org/10.3996/062010-
will continue into the future and that population size can JFWM-014.S1). The model predicted that median abun-
be estimated with some accuracy. Variability and dance at 50 y for the population with the wind farm take
uncertainty about future population growth can be will be , 65 individuals compared to , 385 individuals
incorporated by simply calculating the variance or the for the simulations with no take (Figure 1). The model
standard deviation of the mean (a simple task in also predicted that extinction probability will increase
Microsoft Excel), and replicating the population projec- from 0.00 to 0.03 when the proposed take is applied to
tion numerous times with new values of l for each year the system. With this modeling approach it is straight-
in each replication. forward to see the effects that the proposed take will
have on the system, given the assumptions and the
Ntz1 ~Nt lt ð4Þ uncertainty about population growth rates in the future.
Again, this approach to incorporating variation and Even if uncertainty in model parameters is high we can
uncertainty assumes that past variability is a reasonable still examine the relative abundance across the simula-
approximation of future variability. Calculating extinction tion scenarios and see that the proposed take is likely to
from this simulation is simply a matter of counting the have some negative impact on the population. We can
number of replications where N falls below 1 and also look at population growth predictions and see that
dividing by the total number of replications. The model allowing the proposed take leads to a decrease in
must have some restrictions applied so that the predicted population growth from a nearly stable
populations cannot recover from an abundance of less population (l̄ = 0.995) to a rapidly declining population
than two individuals. It is also important to not simulate (l̄ = 0.956). McGowan and Ryan (2009) showed that
the population too far into the future because a model simulations can be conducted to compare projected
like this has no limits on growth and could, theoretically, population growth and extinction probability with and
grow to infinity if given enough time, or conversely the without the proposed actions. Subsequently managers
probability of extinction also increases over time in a can evaluate the predicted change in extinction proba-
stochastic model. There is no rule of thumb for bility and the changes in the probability of a population
determining good simulation duration, but (depending reaching its recovery criteria to determine whether an
on the species) simulating on the order of a few decades action appreciably affects those probabilities.
is likely sufficient to avoid the problems of unimpeded
growth and make model predictions relevant to take Conclusions
decision-making.
We believe there is a dovetail between legislative
Consider a case where a wind power company wants
language and well-established population ecology mod-
to build a wind farm facility in an area where an
eling tools in the case of evaluating jeopardy in relation to
endangered bird species lives. Based on other wind farm
take permitting. We cannot envision good reasons for not
projects, they estimate that their actions will likely take
using an explicit model to make predictions and evaluate
between 5 and 10 adult breeding birds of the
take proposals. We realize that time and budgets are
endangered species each year. Using our simple model
limited and that take assessments are numerous. Howev-
described above, applying this take is simply a matter of
er, even simple models will make predictions as accurate
subtracting the annual take from the projection equa-
tion: as (and often more accurate than) intuition, and a
modeling approach is more transparent and scientifically
Ntz1 ~ðNt {Tt Þlt ð5Þ defendable. All decisions in natural resource management
involve some prediction about the future and how some
Simultaneous simulations with equations (4 and 5) can action will affect the managed system (Williams et al.
show the expected difference in abundance between a 2002). A take assessment and decision, with or without an
take and a no-take scenario, and extinction probability explicit model, must make some prediction about how the
could easily be derived and compared for these two sets proposed take will affect the protected population. An
of simulations. If no estimates of l or N are available, it explicitly articulated model allows the decision maker, the
may be appropriate to simulate multiple values of these stakeholders, and other scientists to understand the basis
Figure 1. Comparison of the simulated median population abundance trajectories for a hypothetical endangered species with
(dotted line) and without (solid line) incidental take 50 y into the future.
for the predictions that informed the decision. Models do Table S1. Sample simulation model.
not need to be complex to assist with decision-making, as Found at DOI: 10.3996/062010-JFWM-014.S1 (6423 KB
was demonstrated herein with a simple Excel spreadsheet XLS).
model, which took only a few hours to construct and
error-check. A quantitative model only needs to suit the Acknowledgments
decision at hand and fit the available data. If data on
abundance and population trend are not available, We thank the U.S. Geological Survey, the U.S. Fish and
models can still be constructed though expert opinion Wildlife Service, and the department of Fisheries and
on model parameters. Models can be constructed to Wildlife Sciences at the University of Missouri for financial
account for uncertainty in population size, population and logistical support of this work.
growth rate, or any model parameter due to lack of data, We are grateful to M.A. Larson, J.J. Millspaugh, F.R.
and sensitivity analyses or multiple scenarios could be Thompson III, E. Zipkin, two anonymous referees, and the
simulated to understand how that parameter uncertainty editors of the Journal of Fish and Wildlife Management for
affects the decision outcome. reviewing and helping us to improve this manuscript. We
Putting the criteria for jeopardy in some form of a are also grateful to J.S. Rikoon, M.C. Runge, J. Szymanski,
population viability analysis context related to extinction and J. Cochrane for many lengthy discussions on this
probability will avoid arbitrary definitions of the term subject of incidental take and endangered species policy.
‘‘appreciable.’’ We conclude that using predictive popula-
tion models to make take management decisions in References
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Issues and Perspectives
Evolution of Avian Conservation Breeding With Insights
for Addressing the Current Extinction Crisis
Jesse D’Elia*
J. D’Elia
U.S. Fish and Wildlife Service, Pacific Regional Office, 911 NE 11th Avenue, Portland, Oregon 97232
Department of Fisheries and Wildlife, Oregon State University, Corvallis, Oregon 97331
Abstract
Birds have been bred in captivity since the dawn of civilization. In the nineteenth century, breeding rare and exotic
animals for the menageries of royalty and aristocrats transitioned to the formation of public zoological gardens and
natural history museums. With industrialization and human population expansion, some bird species became rare or
disappeared entirely. Once the magnitude of the destructive forces of humans became evident, concerted efforts were
initiated to restore depleted bird populations by using, among other techniques, captive breeding. In this article, I
explore the evolution of avian conservation breeding programs and evaluate how this historical review informs our
outlook for addressing the current extinction crisis.
Keywords: birds; conservation; extinction; zoos

Received: June 20, 2010; Accepted: October 14, 2010; Published Online Early: October 2010; Published: November 2010
Citation: D’Elia J. 2010. Evolution of avian conservation breeding with insights for addressing the current extinction
crisis. Journal of Fish and Wildlife Management 1(2):189–210; e1944-687X. doi: 10.3996/062010-JFWM-017
* Corresponding author: jesse_delia@fws.gov
The charm of history and its enigmatic lesson consist in falcons [Falco peregrinus] and saker falcons [Falco
the fact that, from age to age, nothing changes and yet cherrug] and other kinds of birds in very great
everything is completely different. Huxley (1952) abundance’’ (Bergreen 2008). Other aristocrats and
noblemen of that time such as Holy Roman emperor
Frederick II of Hohenstaufen, author of the falconer’s
Menageries and Zoological Gardens
bible De arte venandi cum avibus (The Art of Hunting with
Birds have been bred in captivity since the dawn of Birds) also maintained extensive menageries and en-
civilization, as a source of food, for entertainment and gaged in trading birds and other animals (Bergreen
sport, and as status symbols, spiritual totems, or personal 2008). Social evolution in Europe during the nineteenth
companions. Breeding rare and exotic birds in captivity century resulted in a shift from these elaborate private
originated in elaborate royal menageries, first in China menageries of monarchs to public institutions (Hoage
around 3,000 B.C. and later in ancient Egypt, Rome, and and Deiss 1996). Concurrent with this shift, a scientific
Greece (Jennison 1937; Schafer 1968; Lamberton and approach to explaining and organizing life was emerging
Rotroff 1985; Cooper 1995; Graham 1998; Benson 1997; from Enlightenment thinkers, and by the nineteenth
Birkhead and van Balen 2008), and by the early medieval century, natural history studies were in vogue in Europe,
period, an extensive trade in birds had been established leading to the expansion of public natural history
throughout the known world (Graham 1998). In the museums and zoological gardens (Farber 2000).
thirteenth century, Marco Polo marveled at the menag- Zoological gardens provided a dimension to natural
erie of Kublai Khan, the fifth Great Khan of the Mongolian history studies that could not be gained by looking at
Empire and grandson of Genghis Khan, who was study skins in natural history museums. They opened up
accompanied by ‘‘ten thousand falconers … and five the possibility of studying behavior, comparative physiol-
hundred gerfalcons [sic] [Falco rusticolus], and peregrine ogy, and evolutionary relationships of exotic animals, some
Evolution of Avian Conservation Breeding J. D’Elia
of which were from opposite ends of the Earth, but which [to] any more definite conclusion, than that captivity has
could now be viewed, living and breathing, side by side an especially injurious influence on the reproductive
(Burkhardt 1999, 2001). Following his adventures on the system; & more injurious in some orders than in others,
H.M.S. Beagle, Charles Darwin studied the breeding habits but with many exceptions in every case …. We can
and hybridization of birds in the London Zoo and relied on attribute this deficient action [i.e., inability to breed in
private breeders and zoological institutions to obtain captivity] only to general constitutional derangement (C.
pigeons (Columbidae) for breeding experiments (Darwin Darwin, as cited in Stauffer 1975, pp. 80–81).
1868; Stauffer 1975; Steinheimer 2004). His study of
pigeons later helped form the basis for his opening salvo As the natural history field evolved in the early 1800s,
in On the Origin of Species (1859) as a model of how those in charge of collections assembled catalogs of
variation and selection could give rise to new species captive animals, including descriptions of morphology
(Secord 1981; Burkhardt 1999; Montgomerie 2009). Italian and behavior, sometimes indicating whether the animal
scholars also were using specimens in zoological gardens would breed in captivity. The first of these catalogs was
to conduct studies on living animals in the 1800s, and, prepared by Frédéric Cuvier (the younger brother of
when the animals died, they would turn the specimens Georges Cuvier) in the early 1800s (Burkhardt 2001)
over to natural history museums for further study and followed by a publication of the London Zoological Park
archiving (Gippoliti and Kitchener 2007), something that in 1835 (Bennett 1835), which set the standard for the
was also done in Paris (Burkhardt 2001) and presumably larger zoos around the world (e.g., Anderson 1883;
elsewhere. English aviculturists with private collections in Brown 1892; Sanyal 1892; Stephan 1900; Hornaday 1911).
the late nineteenth and early twentieth centuries also were In the early catalogs, it was apparent that a wide
donating specimens to natural history museums, as variety of bird species were, at least occasionally,
evidenced by records in museum catalogs. breeding in captivity, including parakeets (Psittacidae),
Despite advances in scientific knowledge facilitated by doves and pigeons (Columbidae), weaver-birds Hyphan-
captive animals, the menageries and zoological gardens of tornis spp., bluebirds Sialia spp., black kites Milvus
the late nineteenth and early twentieth centuries were migrans, sunbitterns Eurypyga helias, cranes (Gruidae),
consumers of wildlife (Frankham et al. 2002). Some birds emus Dromaius novaehollandiae, red-wing tinamous
certainly bred in these institutions, but many birds were not Rhynchotus rufescens, common rheas Rhea americana,
kept under conditions that would have facilitated breeding, white-eyebrowed wood-swallows Artamus superciliosus,
and as early as the 1830s, some were arguing how improved Australian brush-turkeys Alectura lathami, black-tailed
captive conditions might improve a bird’s ability to breed. native-hens Gallinula ventralis, cassowaries Casuarius
For example, in 1838, Swainson wrote the following: spp., pheasants and junglefowl (Phasianidae), gulls
(Laridae), and waterfowl (Anatidae; Bennett 1835; Mitch-
ell 1852; Sclater 1870, 1877). In some instances, where
Such species, again, as show the least disposition to species were especially prolific, there was a need to
breed, (and many, we have no doubt, would do this, if dispose of individuals. Sclater (1870, p. 19) noted ‘‘A
their habitations were constructed on the principle we certain number of the young of the various species of
are now advocating,) should be more particularly Pheasants bred in the Gardens are disposed of every year
supplied by natural shelter; that is, the shelter of leaves in the month of November. Priced lists of these birds
and branches: dwarf trees or shrubs, having foliage, may be obtained on application at the Society’s Office.’’
should be put in their inclosures, within which the nests In the mid-1800s, zoological parks were trading
may be constructed among the branches, and the breeding birds (Sclater 1877), but these efforts were
females screened from observation. The parrots, nearly opportunistic and their purpose was to promote the
all of which build in hollow trees, might have such interest of the zoo and not the species. Manchurian
retreats provided for incubation. The present mode of cranes Grus japonensis, which would have been a great
putting boxes and pigeon-holes into all aviaries, without rarity outside of the Orient, were noted breeding in
regard to the different nature of the birds, is but a rude captivity in Paris and attempting to breed in London, but
and inadequate substitute for more appropriate contriv- failure to maintain a captive population only resulted in
ances. In short, we consider that the art of preserving and requests for more birds to display:
rearing foreign birds in this country, is quite in its infancy;
and our only surprise is, that, with the inadequate means
The birds which were imported by M. de Montigny, on his
that are employed, any species are kept alive more than a
return from China in 1854, not only flourished at Paris in
few months after their being consigned to such
the most perfect health, but for three successive seasons
comfortless habitations (Swainson 1838, p. 154).
made a nest and hatched out their young. The pair in
possession of the [London Zoological] Society in 1860
It is interesting to note that two decades after nested twice but did not succeed in hatching their eggs.
Swainson’s writings, Darwin was at a loss to explain The following year, however, they were more successful,
why some species bred readily in captivity, but others and a young bird was hatched on the 24th of June, 1861,
did not (Stauffer 1975): to which they devoted unceasing care and attention for
many months. The Manchurian Crane is a favourite bird
Considering all the facts which I have been able to collect, among the Chinese, and it has been stated that a
most of which I have given, it seems impossible to come considerable number of them are always in captivity at
Pekin[g] …. It is much to be regretted that only one zoological education (Hornaday 1911). This charter,
individual of this fine Crane is now left in the Society’s along with Hornaday’s leadership and advocacy for
collection. No more valuable present could be made by wildlife conservation, would set the stage for later efforts
the Society’s correspondents in China than additional to establish international endangered species lists (see
specimens of this species (Sclater 1870, p. 8). Scott et al. 1987) and to redefine the role of zoos as
organizations committed to species preservation.
By the mid- to late 1800s, zoological gardens were Despite Hornaday’s belief that zoos had an integral
becoming increasingly popular in Europe (Farber 2000), role to play in conserving native species (some of which
and the template of these institutions was starting to be were in serious decline at the time he was appointed
copied in the United States, first in Philadelphia, director of the New York Zoological Society), he did not
Pennsylvania, and then in Cincinnati, Ohio. When the believe that captive breeding and restocking game
New York Zoological Society opened its park in 1899 decimated by hunters would produce any meaningful
(later called the Bronx Zoo), it was the largest zoo in the results in the United States, unless strict regulations were
world (approximately 4 times larger than the Berlin put into place and enforced. He noted the following:
Garden, 5 times larger than the Paris Garden, and 9 times
larger than the London Garden), and there was a
The game owners and game-keepers of England, Scot-
conscious effort to adapt lessons learned from other
land, Germany and Austria have not saved their game and
zoological parks (The New York Times 1897). As the first
their sport for 300 years by investigations, restocking,
director of the Society, William Temple Hornaday, stated
doctoring or breeding game in coops and pens. They
in an interview with The New York Times in 1896:
have provided shelter, food, water and protection from
enemies. But greater than any or all of those has been the
We have a splendid opportunity to make here in New protection afforded by savage regulations against exter-
York the finest zoological gardens in the world. We can minatory slaughter with guns! (Hornaday 1931, pp. 60–61).
borrow from the accumulated treasures of knowledge
acquired by the Zoological Society of London and by the Hornaday clearly understood one of the key limitations
managers of Jardin de Plantes in Paris, and we can avoid of captive breeding for conservation purposes: without
the mistakes necessarily made by these and other large removing the threats in the wild, breeding species in
owners of wild animals in the process of finding out the captivity for restocking populations would be fruitless.
best way to keep them in captivity (The New York Times
1896, p. 8). Bird Population Declines and Extinctions
Hornaday was already a towering figure in the In addition to fueling the spread of zoological gardens,
zoological community when he took the helm of the expansion of natural history as a new academic pursuit in
society (The New York Times 1896). He had traveled the 1800s had some unintended consequences. One was
extensively throughout Europe, studying their natural the desire to fill newly constructed natural history
history museums, and he participated in expeditions to museums with exotic specimens from around the world.
Cuba, South America, Egypt, and Southeast Asia, Specimens brought back from faraway lands were a
collecting and studying animals. Later, he was instru- status symbol in scientific circles—the rarer, the better.
mental in establishing the National Zoological Park in The story of the extinction of the great auk Pinguinus
Washington, D.C., under the control of the Smithsonian impennis serves as a sad example of this mentality (Fuller
Institution, and was superintendent of that park in 1890 1987, 1999, 2003; Gaskell 2000). By 1854, there was a
(The New York Times 1896). He would go on to help growing sentiment that the great auk was already
prevent the extinction of the American bison Bison bison, extinct, collectors having taken .40 specimens from
to successfully lobby state legislatures to pass bills the single island where they were known to breed in
banning trade in wild game, and he would successfully 1830 and 1831, and a dozen or more in 1833 (Gaskell
lobby Congress to pass the Migratory Bird Treaty Act and 2000). However, in 1854, an article was published in The
prohibit the importation of skins and plumage of birds Edinburgh New Philosophical Journal entitled The Great
intended for the millinery industry (Hornaday 1931). Auk Still Found in Iceland, and English ornithologists John
Under Hornaday’s leadership, the New York Zoological Wolley and Alfred Newton, fully aware of the extreme
Garden stood out from its European counterparts, not rarity of the bird, quickly set out for the island … to
only because of its size but also because it adopted the procure specimens! (Gaskell 2000). They did, and that is
approach of displaying animals in more natural settings the last time anyone ever saw a great auk alive.
(Farber 2000), something that Hornaday had brought to Apparently, the naturalists of the time were far more
natural history museums a few years earlier. Perhaps concerned with solidifying their reputations by acquiring
most importantly, he implemented a conservation- rare specimens for natural history museums than in
oriented charter that was revolutionary. Specifically, the preserving birds in nature for future generations.
New York Zoological Society had established three The extinction of the great auk did little to curb the
primary objectives when it was founded in 1895 under collectors’ appetite for specimens, perhaps because
a special charter granted by the legislature of the state of society, as a whole, saw little economic value in this
New York: 1) development of a public zoological park, 2) bird, or because some did not actually believe it was
preservation of native animals, and 3) promotion of completely gone from its remote haunts. Thus, other bird
species continued to be stalked by skin and egg early 1900s only made matters worse, as hunters fanned
collectors in the late 1800s and early 1900s, including out across the countryside in search of dwindling game.
the California condor Gymnogyps californianus (Wilbur As it became more and more evident that hunting was
1978), imperial woodpecker Campephilus imperialis taking a serious toll on game populations and the
(Nelson 1898), and ivory-billed woodpecker Campephilus millinery industry was wreaking havoc on birds around
principalis (Hasbrouke 1891a; Snyder 2007). Slaughter of the globe for their prized feathers, public sentiment
birds for museum specimens is evidenced by the fact began to shift from the malaise of the late 1800s toward
that in 1872 there were approximately 30,000 stuffed a call to action, followed by a full-blown movement to
birds and bird skins at the British Museum and by 1909 save the birds.
there were 500,000 (Allen 1909). Egg collecting for Direct proof of the ability of humans to destroy a once
personal collections and natural history museums was common bird species was finally put on full display with
also at its zenith around this time (Kiff 2005). Rarity only the extinction of the passenger pigeon Ectopistes
increased the collector’s appetite. When, at the turn of migratorius in 1914 (Schorger 1955; Fuller 1987). Stories
the century, there were thought to be only approxi- of passenger pigeons once blackening the sky are now
mately 50 heath hens Tympanuchus cupido cupido legendary. John James Audubon experienced the
remaining, all on Martha’s Vineyard, an island off of following firsthand:
Massachusetts, collectors were offering US$25–30 apiece
for specimens, because, as The New York Times pointed
out, ‘‘the ornithological collector who does not get a The air was literally filled with pigeons; the light of
Heath Hen for his collection at once will not get one at noonday was obscured as by an eclipse; the dung fell in
all’’ (The New York Times 1900). spots, not unlike melting flakes of snow; and the
The tide of public opinion toward bird conservation continued buzz of wings had a tendency to lull my
only started to turn when large numbers of economically senses to repose (Audubon 1832, p. 321).
valuable and otherwise ‘‘useful’’ bird species started to
show vulnerability around 1870, with the invention of Loss of a bird species that once numbered in the
breech-loading shotguns (Forbush 1913; Hartley 1922). billions and ranged throughout the central and entire
Pump-action and semiautomatic shotguns soon followed eastern half of North America put things into perspective
(both were in production by 1900). These advances in for many. The strength of our destructive abilities was
gun technology lead to drastic declines in gamebirds now evident, and the need to combat those who
throughout most of the civilized world (Hartley 1922; attempted to maintain the status quo was equally
Hornaday 1931). evident to conservation minded individuals. Thus, in
By the late 1800s, bird extinctions due to human 1900 the United States passed its first federal wildlife
activities were readily acknowledged by some in the protection law, The Lacey Act. This act prohibited the
ornithological community (e.g., Meyer 1889). However, transportation of illegally captured or prohibited animals
solutions to the extinction crisis were either not across state lines. Hartley (1922, p. 5) described the
contemplated or not readily offered. There seems to be impetus for the law as follows:
a sense of fatalism in the writings of early naturalists who
were torn between an untamed world teeming with Passenger pigeons were gone, the Heath Hen was gone,
wildlife and a world that had to sacrifice nature to prairie chickens [Tympanuchus cupido] were going,
accommodate the juggernaut of expanding civilization turkeys [Meleagris gallopavo] were disappearing, the quail
after the industrial revolution and the growing railroad [Colinus virginianus] had been entirely eliminated from
empire. Elliot Coues, one of the founders of the American some States. Soon all game birds would be distant
Ornithologists’ Union (AOU), perhaps best epitomized memories of the past. Non-game birds also were suffering
this malaise when he noted the sheer number of bird from the depredations of plumage hunters, and they too
deaths from telegraph wires in 1876 during a horseback would soon be effaced.
trip from Denver, Colorado, to Cheyenne, Wyoming
(Coues 1876). He came to the following rather dismal Although last ditch efforts were made to breed the
conclusion: few remaining passenger pigeons by Professor Charles O.
Whitman at the University of Chicago and by the
Usually a remedy has been or may be provided for any Cincinnati Zoo, breeding efforts (including cross-foster-
unnecessary or undesirable destruction of birds; but there ing passenger pigeon eggs with a rock dove Columba
seems to be none in this instance. Since we cannot livia) were insufficient to sustain the small captive
conveniently abolish the telegraph, we must be content population (Deane 1908, Herman 1948). Similarly, last
with fewer birds. The only moral I can discern is that larks ditch efforts to breed a few remaining Carolina parakeets
must not fly against telegraph wires (Coues 1876, p. 736). Conuropsis carolinensis would fail (Snyder 2004). Despite
the knowledge that populations were being decimated
Development of the railroads, along with perfection of and their range had drastically contracted since the early
cold storage facilities for shipping game to market, 1800s (Hasbrouke 1891b), aviculturists breeding Carolina
drastically increased the number of birds killed for food parakeets in Europe and America (including the Phila-
in America each year in the late 1800s (Palmer 1900). The delphia Zoo, Hanover Zoo, and Frankfurt Zoo) since the
invention and spread of personal automobiles in the mid-1800s (Nowotny 1898; McKinley 1978) did not
contemplate a coordinated effort to save the species joined by numerous other local bird clubs, natural history
from extinction. As Cokinos (2000, p. 48) explained, societies, and societies for the preservation of Australian
wildlife (Garnett and Crowley 2008). The Dutch Associ-
… no one could yet see the larger context in which ation for Bird Protection (Vereniging tot Bescherming
captive breeding could take place: through a widespread, van Vogels), the German Federation for Bird Protection
well-managed program in which private owners and zoos (Deutscher Bund für Vogelschutz), and the Irish Society
shared birds for breeding. Such a program might have for Bird Protection were all formed around the turn of
prevented the extinction of the Carolina Parakeet …. the century (Dalton 1993). With the creation of the
Instead, aviculturists and zoos competed against each French League for Bird Protection (Ligue pour la
other for prized individuals, knowing that the species Protection des Oiseaux) in 1912, a national group for
kept inching closer to extinction. bird protection existed in nearly every northern Europe-
an nation (Dalton 1993).
On the back of these extinctions and drastic waterfowl The British ornithologist Charles Dixon perhaps best
declines in the United States, which were about to get summed up the sentiment of conservationists at the
worse with the draining of wetlands and the continental time in his book Lost and Vanishing Birds:
drought of the 1930s (Perry 2008), people were
beginning to wonder if once wide-ranging and numer- … it requires no great strain upon the imagination to
ous species could be killed off, what was to stop bird picture a time when all the larger wild birds and beasts,
species that were considered more valuable, such as all the exceptionally helpless ones of the earth, must
geese, ducks, heath hens, and turkeys (at the time all in perish, or exist as specimens in our museums, or as
severe decline), from going extinct? phantom records in our scientific literature. This will be a
serious outlook for the biologist of the future, and the
A Call for Bird Conservation matter has long been sufficiently important to warrant
some strong steps being taken to avert as far as possible
In the late 1800s, a small group of mostly self-taught
such a vast calamity. After all, we only hold the fauna of
ornithologists organized to form the AOU and generated
the world in trust, and it is but our bare duty to posterity
a movement to change the status of wild birds in
to hand that fauna down as intact as we found it, or as
America (Orr 1992). At the AOU’s second meeting in
nearly so as the reasonable exigencies of life will admit
1884, William Brewster (a founding member of AOU and
(Dixon 1889, pp. 27–28).
curator of birds and mammals for the Boston Society of
Natural History) expressed concern over the slaughter of
Even suppliers of the millinery industry in the late
birds for the millinery industry. Apparently, the statistics
1800s realized that resources were not boundless. The
he presented ‘‘startled’’ some members, and he moved
law of diminishing returns eventually caught up with
for appointment of a committee for the protection of
them, so industrial farms were established for a few
North American Birds (Orr 1992). Brewster’s motion
species that bred readily in captivity. Dixon (1889) noted
passed unanimously, and he was named committee
the following:
chair. The committee announced its formation, offered a
draft state bird protection law, and suggested law
enforcement tactics in an article published in Forest It is gratifying to know that egrets [Egretta spp.] are now
and Stream, January 21, 1886. The article also included a being kept in captivity for the sake of their plumes. There
proposal to form, ‘‘an association for the protection of is, we believe, an establishment near Tunis [Tunisia,
wild birds and their eggs, which shall be called the Africa] where these birds are kept and allowed to breed in
Audubon Society’’ (Orr 1992). a large aviary. The plumes are shorn twice in the year, in
State chapters of the Audubon Society were formed, May and September, each bird furnishing about seven
and reports were sent to the AOU for publication in its grammes in the year, valued at thirty-five francs, a sum,
scientific journal The Auk (AOU 1898). Because the after deducting all expenses, which leaves a net gain of
reports were broken out by state, these efforts gave some twenty-two francs per bird. This shows how easily
spatial context to what was happening at the national we can preserve these beautiful birds from extinction and
scale in relation to bird declines. That same year, a yet gratify the whim of women for wearing ‘aigrette’
summary of a report by William Temple Hornaday at the [tufted-crest, or head-plumes of the egret, used for
New York Zoological Society was published in The New adorning a woman’s hat] (Dixon 1889, p. 295).
York Times (The New York Times 1898) with a bar chart
illustrating massive bird declines across 30 states. The Around the same time (in 1895), Edward Avery
article concluded that several species were likely to go McIlhenny, heir to the Tabasco sauce fortune, took a
extinct if measures were not implemented immediately. different approach to conserving egrets. Alarmed by the
Environmental degradation on other continents and loss of snowy egrets Egretta thula in the southeastern
the global plumage trade spurred efforts that paralleled United States due to plume hunters, and having heard
those in the United States. In Australia, the Royal stories of an Indian raja who built bamboo cages for
Australasian Ornithologists Union was formed in the captive birds that continued to return to his ‘‘flying
early twentieth century with the objective of creating gardens’’ long after the raja had died and his cages had
reserves and enforcing wildlife laws, and it was soon rotted away, McIlhenny took eight young birds from two
snowy egrets nests and raised them in an aviary he The project, in which up to 400 Wood duck[s] were
constructed from bamboo and poultry netting near his banded and released each year (2579 in all), probably had
home on Avery Island in coastal Iberia Parish, Louisiana a seminal effect on the restoration of the species in the
(McIlhenny 1935; Nickens 2010). In early winter, he set wild. Banding returns from the US Banding Laboratory
them loose to migrate south. By 1908, there were 10,000 records (pers. comm.) indicated that the Litchfield birds
egrets returning to the property and by 1911, the had reached 15 US states and Ontario in Canada, and had
population reached 100,000 egrets and McIlhenny’s formed the nuclei of breeding colonies from South
refuge on Avery Island was affectionately dubbed ‘‘Bird Carolina up to Massachusetts. Like the later success with
City’’ (McIlhenny 1935; Nickens 2010). the Hawaiian goose Branta sandvicensis, it would seem
In the late 1800s, ostriches Struthio camelus had also that captive rearing programme for birds could be a
become increasingly difficult to catch in Africa due to positive step towards the conservation of the species.
overharvesting for the millinery industry (Sales 2009).
This led European entrepreneurs in South Africa to begin Another important early waterfowl conservation
domesticating ostriches on farms starting in 1867 breeding venture in the early 1900s was the program
(Douglass 1881). The model of these farms soon spread, to breed the giant Canada goose Branta canadensis
and ostriches were shipped to North Africa, Europe, maxima (Hanson 1965; Lee et al. 1984).
India, Australia, South America, and the United States The origins of breeding depleted stocks of gamebirds
where they were raised to produce numerous products came at a time when the most of the civilized world was
(de Mosenthal and Harting 1877; Douglass 1881; just beginning to come to terms with the idea of
Doughty 1973). Although spurred by the notion that resource conservation. In 1910, Gifford Pinchot, the first
wild stocks were running out, breeding served mostly as chief of the U.S. Forest Service (under Theodore
a cost savings measure to those entrepreneurs who Roosevelt), wrote The Fight for Conservation (Pinchot
sought to make their living in the millinery industry. It 1910) that called on the American people to adopt a
was more profitable to run an ostrich farm than to track doctrine of wise use as a measure of national security.
down diminishing supplies in the more remote regions Around that time, many U.S. states began to establish
of Africa. game farms to help restock the countryside with
In the United States, concerted efforts to breed birds in gamebirds that had undergone massive declines in the
captivity for the purpose of their conservation began late nineteenth century. Birds that were raised included
around the turn of the twentieth century in conjunction ring-necked pheasants Phasianus colchicus, wood ducks,
with the establishment of Audubon societies and the black ducks Anas superciliosa, mallards Anas platy-
growing public concern for bird resources. Grinnell rhynchos, quail (Odontophoridae), ruffed grouse Bonasa
(1901, pp. 532–533) stated the following: umbellus, and wild turkeys. These operations were
initially considered hugely successful, and by 1919,
55,400 pheasant eggs were produced in New York state
It is only within a very few years that breeding wild geese alone, along with 9,206 half-grown birds to be released
and ducks has been attempted. At present, however, a later that year (Hartley 1922). Private conservation clubs
number of persons are very much interested in this also were initiated by those interested in buoying
pursuit, and there seems to be good reason to believe gamebird numbers to provide hunting opportunities.
that after a few more years in experiment, a number of However, these programs may only have been successful
species of our wildfowl will be so far domesticated that in boosting local numbers for hunters (Brakhage 1953),
they can be depended upon to breed in confinement. with little contribution to the populations that were now
allowed to heal due to increased habitat protection and
In the early twentieth century, the wood duck Aix more restricted hunting seasons (Hornaday 1931).
sponsa, once a common species in eastern North Commissions were formed and meetings were held to
America, was the rarest duck on the continent and on develop techniques on breeding game birds, and ‘‘game
the verge of extinction (Ripley 1959). Fortunately, this farming’’ of birds became part of the curriculum at
species was being reared in captivity in Belgium, Holland, Cornell University (The New York Times 1918), the
France, and England at that time (Ripley 1959). In 1918, leading institution in ornithology at the time (Battalio
when the wood duck was thought to be virtually extinct, 1998). One of these early meetings (arranged by the
there were probably more individuals in captivity than in American Protective and Propagation Association) in-
the wild (Ripley 1973). A small breeding program was cluded such notable figures as William T. Hornaday, head
initiated in Litchfield, Connecticut, by Alain C. White of the New York Zoological Society, professors at Cornell
under the direction of Herbert L. Job (Ripley 1973). Job University, superintendents of the game farms, repre-
had studied early rearing experiments with North sentatives from the U.S. Department of Agriculture, and
American diving ducks at Cornell University and the chief of the U.S. Biological Survey (The New York
attempted to emulate them with the wood duck. He Times 1918).
found that although it was relatively easy to maintain the Another man at the 1918 meeting of these conserva-
birds in captivity, they were hard to rear. In 1922, he tion luminaries was W. C. Adams. Adams was the
sought the assistance of the English gamekeeper Henry chairman of the Commissioners on Fisheries and Game
Bowden, who helped the program flourish. According to in Massachusetts, and the man in charge of managing
Ripley (1973, p. 56), heath hen populations on Martha’s Vineyard (The New
York Times 1918). Around that time, only a few years expediency and wise use to one of right and wrong
after the loss of the passenger pigeon and Carolina (Leopold 1933, 1949). Thus, the stage was set for
parakeet, there was growing concern over the precarious conserving birds from extinction as an ethical duty,
state of the heath hen, once a widespread species in New regardless of perceived economic value to society.
England and Mid-Atlantic region (The New York Times
1907, 1917; Gross 1923). In 1907, the Massachusetts Captive Breeding Evolves: Breeding
Game Commission had gathered US$2,000 in private Nongame Birds
donations for recovery of the bird and was seeking
additional money from the state legislature to build Ornithology was growing as a profession in the 1920s
firebreaks around breeding grounds and to maintain and 1930s, but experimental science in ornithology was
artificial incubators (The New York Times 1907). Conser- still in its infancy (Battalio 1998). The model of
vation efforts of establishing a preserve for the species in conservation clubs or state biologists raising game birds
1908 proved temporarily fruitful, increasing populations to maximize production was still the norm (although see
to .2,000, but habitat loss due to a wildfire, a series of Wood (1926) for a discussion of aviculture in private
severe winters, combined with an increase in predators, English aviaries). Furthermore, there were lingering
and disease devastated the single isolated population. By questions regarding some rare species, and whether
1924, only 50 birds remained, all on Martha’s Vineyard they were doomed to extinction, thus rendering captive
(Gross 1931). breeding efforts futile.
The state of Massachusetts, the Federation of Bird Loyle Miller (Miller 1942), who was studying New
Clubs of New England, and other organizations were World vulture (Cathartidae) fossil remains at the tar pits
involved in the struggle to preserve the heath hen. When of southern California was detecting some interesting
the birds were more numerous, they were introduced to trends in the historical abundance of vultures in the fossil
Long Island, New York, and the mainland of Massachu- record, and opined the following:
setts, but these attempts failed. In addition, attempts
were made to breed the birds in captivity, but these Were not teratorns an antiquated group that time had
attempts also failed (The New York Times 1917; Gross weathered away before we arrived? Is not the California
1931). Despite their best efforts, by 1927, there were only Condor a senile species that is far past its prime? It was
13 birds left and by winter 1928, only two birds widely distributed and numerically abundant in Pleisto-
remained. When in 1929 only a single bird remained, cene time (Florida, Mexico, New Mexico, Texas, Nevada,
requests were made to secure a prairie chicken California), but now it is restricted to one or two localities
Tympanuchus spp. or ruffed grouse as a potential mate. and a numberable roster of individuals within the two
These efforts were never realized. Californias. Is it not a species with one foot and even one
Although attempts to conserve the heath hen failed, wing in the grave? (Miller 1942, p. 213).
they showed that ornithologists had largely broken out
of the fatalistic mentality of the late nineteenth century. Those who opposed later efforts to take California
The fight to save species from extinction was on, and condors out of the wild for captive breeding would point
captive breeding was clearly a potential weapon in that back to Miller’s (1942) paper as an argument against
fight. According to Gross (1923), in a letter to The Auk, intervening to save the condor. However, Miller’s (1942)
‘‘The response to a questionnaire sent out by Mr. William statement must be put into the context of the time,
C. Adams, Director of the Division [of Massachusetts when the world was at the height of World War II, the
Fisheries and Game] has shown conclusively that public techniques for breeding raptors in captivity had not been
opinion favors saving the Heath Hen at all reasonable developed, no one knew the real cause of the recent
costs.’’ In the April 1923 issue of The Auk’s Notes and condor population decline, and expending dollars to
News section, there was a discussion of Mr. Adam’s conserve rare species would have been toward the
questionnaire, with the remark that ‘‘The saving of the bottom of a long list of global priorities.
Heath Hen is not a local or State matter but is of interest By the late 1940s, the war was over, science was being
to the whole country and scientific world’’ (AOU 1923, reinvigorated both financially and in public opinion,
p. 379). wildlife management was beginning to be seen as a
After extinction of the heath hen, efforts were already professional endeavor, and the concept of conservation
underway to prevent a similar result for the prairie ethics was spreading (Leopold 1933, 1949). In 1948, the
chicken. State game officials in Texas closed down the International Union for the Preservation of Nature (later
hunting season of the prairie chicken for 5 y, and captive to be renamed the International Union for the Conser-
breeding experiments were attempted by the State vation of Nature and Natural Resources, or simply the
Game Department. Although the captive breeding World Conservation Union) was formed by a consortium
experiments were unsuccessful (Associated Press 1937), of governments, national groups, and international
there was a growing sentiment that preservation of organizations at a United Nations Education, Scientific,
species had broad public support and was of national and Cultural Organization meeting in Paris, with a goal of
interest. Moreover, in 1933, Aldo Leopold, the father of conserving ‘‘the entire world biotic community … on
wildlife management, had introduced a novel concept in which rests the foundation of human civilization’’
his paper The Conservation Ethic that would transform (Holdgate 1999, p. 33). Fledgling experiments to breed
the debate of conservation from one of economic nongame birds in captivity were underway in this era.
For example, the AOU Committee on Bird Protection the eggs) to rapidly increase nēnē production (Smith
reported attempts to breed whooping cranes Grus 1952). Captive breeding efforts proved very successful
americana in captivity in 1949: and by 1961, Sir Peter Scott at the Severn Wildlife Trust in
England (later renamed the Wildfowl and Wetlands Trust)
Attempts have been made to breed captive birds had a flock of .100 birds and was offering to sell 20
[Whooping Cranes] in a large enclosure in Texas but the nēnē a year back to the Hawaii at a price of US$22.50 per
initial clutch proved to be infertile. Another pairing is now gosling (AOU 1962).
being encouraged with a different male bird and the The early successes of this effort led to captive
experiment will continue (AOU 1950, pp. 321–322). breeding programs for other Hawaiian waterbirds,
including the Hawaiian moorhen Gallinula chloropus
Whooping cranes had declined precipitously in the sandvicensis, Laysan duck Anas laysanensis, and Hawaiian
late 1800s and early 1900s, and by 1947, only two birds duck Anas wyvilliana. Captive propagation of Laysan
were known from Louisiana and only 25 from Texas duck was initiated in 1957 at the Honolulu Zoo with the
(Hamerstrom 1947). It was clear at this time that capture of eight ducks and then expanded with the
extraordinary measures would be necessary to save the capture of an additional 36 ducks in 1958 that were sent
species from extinction. By 1957, when it was believed to the Honolulu Zoo for acclimatization and ultimately
that 27 whooping cranes were left in the world (1 at the distributed to eight aviaries in the United States
San Antonio Zoo, 2 at the New Orleans Zoo, and 24 at (Honolulu Zoo, Philadelphia Zoological Gardens, San
the Aransas National Wildlife Refuge on the Texas’ Gulf Antonio Zoo, Dr. Jean Delacour, Dr. S. Dillon Ripley, San
Coast), plans to take several pairs into captivity for a Diego Zoo, Tracy Aviary, and the Bronx Zoo) and at the
breeding program were considered by the Whooping Severn Wildlife Trust in England (Ripley 1960; AOU 1962;
Crane Advisory Group (consisting of U.S. and Canadian Warner 1963). This collaboration of breeding amongst
authorities), but not carried out (Time 1957) pending multiple institutions, a practice that also was being carried
completion of studies on a more common surrogate out with the Hawaiian moorhen from 1953 to 1969
species. (Dibben-Young 2009), represented a new concept in
At the time of the first whooping crane breeding captive breeding, in direct contrast to the provincial and
experiments in 1949, the nēnē, or Hawaiian goose Branta proprietary attitudes of zoological gardens at the time the
sandvicensis, was also on the verge of extinction. Carolina parakeet went extinct in captivity in 1916.
Although the nēnē was numerous and well distributed Indeed, much had changed over the four decades
on the main Hawaiian Islands when Captain James Cook between the extinction of Carolina parakeets and the
landed in 1778, when the Journal of Wildlife Management captive breeding cooperation being undertaken for
published Baldwin’s (1945) estimate of only approxi- Laysan duck. Air transportation was now allowing
mately 50 birds remaining, all on the Island of Hawaii, the movement of animals across great distances in a short
gravity of the situation was apparent. As a result, a time, zoos were starting to contemplate their role and
breeding program for the nēnē to prevent its extinction responsibilities in endangered species conservation, the
began in 1949 by the State Department of Land and field of wildlife management was starting to form into a
Natural Resources Pohakuloa Breeding Facility (Smith recognized profession, natural history studies were
1952; Stone et al. 1983). giving way to experimental research, and the public
Although nēnē had bred at estates and zoological was starting to accept their ethical duty to conserve
gardens throughout Europe starting in 1823, and in the species.
U.S. mainland (Brown and Canton 1880), they were However, the focus of avian breeding efforts for
apparently gone from these collections by the time endangered species conservation in the 1950s was still
Germany invaded France in 1940 (Smith 1952). An ill- heavily rooted in the game farms of the early 1900s and
fated attempt to establish a captive flock and split it up continued to be directed toward those species, or taxa,
amongst wealthy landowners and menagerie owners on that were easily bred in captivity or that were valued as a
the Hawaiian islands for further breeding was made in food or hunting resource, primarily ducks, geese,
the 1920s and 1930s, but those who received the birds pheasants, and gallinaceous birds (Wildlife Conservation
lacked the facilities or abilities to raise the birds, and Committee of the Wilson Bulletin 1940; Newman 1945;
some were simply let loose, whereas a few are thought Sylvester and Lane 1946; Ginn 1947). Although there
to have ended up on the dinner table of the farmhands were a few examples of early efforts to breed nongame
at one of the pineapple Ananas comosus plantations on birds at this time (e.g., Swedish ornithologists had begun
Lanai (Smith 1952). a captive breeding program as early as 1954 to restore
At the outset of the breeding program initiated in eagle owls Bubo bubo, first in Sweden and later in
1949, it was apparent that unless the causes of decline Germany [Wayre 1969, 1975] and a program to breed
were addressed, captive breeding would be pointless and reintroduce white storks Ciconia ciconia had been
(Smith 1952). This program also had the foresight to undertaken in Switzerland starting in 1948 [Bloesch 1980;
spread the risk to captive birds by establishing two Schaub et al. 2004]), the value of nongame wildlife had
separate flocks—one flock on Hawaii and one flock in not yet been fully realized in most of the world.
England—and to use double clutching (i.e., removing But that was about to change. With increasing concern
eggs from the nest to induce females to lay a second over the impacts of pesticides on wildlife and humans,
clutch) and cross-fostering (i.e., having chickens incubate Rachael Carson’s Silent Spring (Carson 1962) hit like an
atom bomb in 1962. To Carson, there was an aesthetic science and ecology. Thus, under a single roof, propa-
value in nature and a connection between the health of gation, laboratory science, and ecology were comingling.
nature and our social and physical health. She had made This was no longer a game farm program to maximize
the link between the loss of bird species (i.e., a silent production. Saving endangered species through captive
spring) and the concomitant negative effects on society, breeding was now a scientific endeavor (an outgrowth of
and the public largely agreed with her argument. the controlled experiments of the Environmental Con-
Carson’s book was an instant best seller that would lead taminants Research Program at Patuxent starting in 1959
to major changes in environmental laws in the United when Dr. John L. Buckley became director) that included
States as well as overseas. The change in social experimentation on closely related species to work out
consciousness associated with Carson’s work would the best approach for critically imperiled species (Perry
spawn a new generation of conservation efforts. 2004).
Concurrent with Carson’s exposé of the pesticide Such an approach was first used with procuring,
industry, the Species Survival Commission of the incubating, and raising sandhill crane Grus canadensis
International Union for the Conservation of Nature and eggs to provide information that could be eventually be
Natural Resources (IUCN) was busy generating lists and used on endangered whooping cranes starting in 1966
distribution maps of threatened species throughout the (Erickson 1975, 1980). Erickson’s work with whooping
world (later to be formalized in its popular Red Data cranes eventually led to experiments with artificial
Books; Scott et al. 1987). These efforts were spawned by insemination and detailed studies of artificial incubation,
several publications highlighting the endangered state behavior, and diet (Erickson 1975). In particular, the
of various taxa, including Extinct and Vanishing Birds of adoption of artificial insemination techniques, including
the World (Greenway 1958). It was in this context that, in the use of human imprinted birds to obtain semen,
1966, the Endangered Species Preservation Act (Pub. L. would drastically change the playing field for the captive
No. 89-669, 111–3, 80 Stat. 926 [repealed 1973]) was breeding of endangered avifauna, especially for those
enacted. That act specifically called for the Secretary of species that were not prone to breed on their own when
the Interior to ‘‘carry out a program in the United States placed together in captivity (Berry 1972; Temple 1972;
of conserving, protecting, restoring and propagating Grier 1973; Blanco et al. 2009).
selected species of native fish and wildlife.’’ Three years In many ways, Erickson’s work at Patuxent with
later, The Endangered Species Conservation Act of 1969 whooping cranes—and bald eagles Haliaeetus leucoce-
(Pub. L. No. 91-135, 83 Stat. 275) was enacted, expanding phalus that were also being bred at Patuxent starting in
the authorities of the 1966 act and instituting a provision 1965 (Maestrelli and Wiemeyer 1975; Perry 2004)—
to allow listing of threatened and endangered wildlife changed perceptions about what was possible in captive
worldwide. Although the 1969 act prohibited the breeding. As Martin stated in the introduction to the First
importation of foreign wildlife that was endangered, it Conference on the Breeding of Endangered Species in
provided for exemptions in the case of zoological parks Captivity,
or other institutions interested and capable of rearing
endangered species in captivity. Despite the lack of It is noteworthy that the concept of captive breeding, in
regulatory might that would finally come in the form of this instance too [i.e., with Whooping Cranes], initially met
strict penalties and take prohibitions of the Endangered with resistance from various quarters, since it was
Species Act (ESA) as amended (ESA 1973, 16 U.S.C. probably felt that conservation in the field can be the
111531–1543 [1976 & Supp. V 1981]), the 1966 act was only answer. But it is apparent that successful conclusion
an important milestone as the first national legislation in of the pilot project with Sandhill Cranes will do much to
the world calling for the protection of endangered reduce this resistance, whilst at the same time providing a
wildlife. This would soon be followed by the ratification firm factual basis for the main scheme involving
of the Convention on International Trade in Endangered Whooping Cranes (Martin 1975, p. xxv).
Species of Wild Fauna and Flora (27 U.S.T. 1087, T.I.A.S.
No. 8249) in 1975, which aimed to ensure that Many studies at Patuxent followed the whooping
international trade in specimens of wild animals and crane and bald eagle work, including breeding experi-
plants did not threaten their survival. ments with the American kestrel Falco sparverius (Porter
and Wiemeyer 1970), Aleutian Canada goose Branta
Scientific Programs for Breeding hutchinsii leucopareia (renamed Aleutian cackling goose;
Endangered Species Springer et al. 1978; Erickson 1980), and peregrine falcon
Falco peregrinus (Maestrelli and Wiemeyer 1975). Despite
The same year the Endangered Species Preservation their successes with several species, Patuxent never had
Act was passed (1966), the Endangered Wildlife Research any success in propagating peregrine falcons because
Program was established at the Patuxent Wildlife they used wild-caught migrants, which almost never
Research Center near Laurel, Maryland, under the breed in captivity; something they learned by housing 10
direction of the Bureau of Sport Fisheries and Wildlife or so pairs for 15–20 y without any breeding success.
(an arm of the U.S. Fish and Wildlife Service at the time; The Patuxent group was not the sole faction interested
Erickson 1975). One of the three divisions of this program in experimentation with breeding captive birds. In
was solely dedicated to propagation of endangered August 1965, Professor J. J. Hickey convened the first
wildlife. The other two programs included laboratory international peregrine falcon conference at the Univer-
sity of Wisconsin, where attendees were sobered by the Burnham 2003). Cade’s research program proved to be a
extent and severity of the peregrine falcon population stunning success and The Peregrine Fund raised 434
decline (Hickey 1969). Some of the falconers and peregrine falcons, 40 gyrfalcons Falco rusticolus, 50
biologists in attendance were so concerned that at the lanner falcons Falco biarmicus (from a single pair), and
end of that conference they met to discuss captive 80 prairie falcons Falco mexicanus within 10 y (Cade
breeding as a means of saving the species from 1980; Cade et al. 1988). By the turn of the century,
extinction (Cade 1986). This rump session, chaired by .7,000 peregrine falcons had been produced in captivity
Donald V. Hunter, Jr, led directly to the formation of The and released into the wild by The Peregrine Fund,
Raptor Research Foundation in 1966, the original Patuxent, the Canadian Wildlife Service, and other
purpose of which was to promote the rapid develop- sources (Cade and Burnham 2003). Furthermore, the
ment of captive breeding projects for peregrine falcons techniques developed for peregrines proved useful for
and other raptors (Cade 1986, 1988). In its early years, other species, with conservation breeding and reintro-
this group played a pivotal role in rapidly disseminating ductions having been carried out on Mauritius kestrels
information on husbandry techniques amongst these Falco punctatus, lesser kestrels Falco naumanni, aplo-
various groups (Cade 1986, 1988). mado falcons Falco vespertinus, orange-breasted falcons
At the time The Raptor Research Foundation was Falco deiroleucus, bearded vultures Gypaetus barbatus,
formed, most people considered a systematic and griffon vultures Gyps fulvus, bald eagles, white-tailed sea
reliable raptor breeding program at the scale necessary eagles Haliaeetus albicilla, harpy eagles Harpia harpyja,
to recover the peregrine falcon to be unattainable (Cade and other raptors (reviewed by Cade 1986, 1988, 1997).
1986). Although Renz Waller, a German artist and What had been considered a pipe dream only a few years
falconer, had successfully bred peregrine falcons in an earlier by most had been transformed into a reality as
old silo near Dusseldorf, Germany, during World War II, The Peregrine Fund and others advanced the field of
attempts to replicate his success in the 1950s in Germany aviculture through a program that combined the
and Britain had failed (Cade 1988). With the magnitude traditional knowledge accumulated over centuries by
and extent of the peregrine falcon population declines, it falconers with modern scientific inquiry.
was clear that failure was no longer an option. Lessons learned from the raptor breeding programs at
One of the groups that began experimenting with The Peregrine Fund were exported to other avian taxa,
breeding peregrine falcons after the 1965 University of including parrots, cranes, and to a lesser extent,
Wisconsin meeting was The Peregrine Fund, a nonprofit passerines. George Archibald, one of Tom Cade’s
organization led by Tom Cade, a professor at Cornell students at Cornell University, embraced the philoso-
University and a veteran falconer (Cade 1986). The fund phies and techniques developed by The Peregrine Fund
had humble beginnings, starting with unsolicited dona- and Patuxent and applied and refined these techniques
tions to the Laboratory of Ornithology at Cornell from for use on cranes. In 1972, Archibald and Ron Sauey
private citizens hoping to help save the peregrine founded the International Crane Foundation, a nonprofit
(Burnham and Whaley 2010). The first donation (of organization dedicated to the study and preservation of
US$2) came from two boys in California that wrote to the the world’s crane species (Archibald 1974, 1996/1997).
Laboratory of Ornithology on October 5, 1970: The International Crane Foundation specifically sought to
maintain in captivity 15 unrelated breeding pairs of each
Dear Sirs: threatened crane species in captivity as a safeguard
Me and my Friend have been studying birds, especially against their extinction (Archibald 1974), although the
Peregrine Falcon[s]. One day he came over and showed primary interest of the foundation was the welfare of
me a newspaper article about you trying to help save cranes in the wild (Archibald 1996/1997). Through
Peregrine[s], so we have been going around collecting carefully designed manipulative experiments and close
money to help your project. I hope every thing works out attention to pair bonding, territory size, conspecific
alright. Sincerely Yours, Orin Starn and Jay Hart. Burnham isolation, species-specific behavior, and the physiological
and Whaley (2010, p. 2). and psychological needs of young, the International
Crane Foundation, in coordination with Patuxent, devel-
Shortly thereafter, the Massachusetts Audubon Society oped a modern scientific approach to breeding cranes
made a substantial contribution to the laboratory (Luthin et al. 1986; Ellis et al. 1996).
(Burnham and Whaley 2010). With the establishment of Wildlife programs in other parts of the world were also
the ESA in 1973, numerous additional funding sources starting to experiment with innovative approaches to
were available; and Bob Berry, Frank Bond, Jim Weaver, captive breeding in the 1960s and 1970s, most notably,
and Tom Cade officially founded The Peregrine Fund as a New Zealand and Australia. In New Zealand, captive
nonprofit organization the following year (Burnham and breeding efforts were attempted with the critically
Whaley 2010). Cade took the published knowledge on endangered and flightless kakapo parrot Strigops hab-
artificial insemination, artificial incubation, nutrition, roptilus. Although initial attempts failed because the six
hand-rearing, socialization of young, fostering and birds brought into captivity were all males (Clout and
cross-fostering, and reintroductions, and combined it Craig 1995), techniques that combined artificial incuba-
with his and his colleagues’ knowledge of falconry to tion and hand rearing of chicks with translocation to
create the most successful raptor propagation program island refugia, supplemental feeding and nest monitor-
in the world (Cade et al. 1988; Cade 2000; Cade and ing in the wild, eventually proved fruitful (Elliott et al.
2001). Captive breeding also was attempted with the longer cope with the hazards of life in the wild. This
flightless takahē Porphyrio hochstetteri in the 1960s should in no way be misconstrued (as it has been by
through the taking of eggs from the wild. As with the some conservationists) as an action that merely confined
kakapo, initial attempts failed, but through a series of these animals to captivity. The idea was that the captive
experiments, researchers eventually developed a suc- colonies should be set up merely as a safeguard against
cessful captive breeding program (Clout and Craig 1995). extinction, while the most stringent efforts should be
In Australia, the Victorian Government had established made to preserve the wild habitat and wild populations
a wildlife research station in 1959, known as Serendip of the species concerned and to release back to the wild
Sanctuary outside Melbourne to study, breed, and captive-bred animals when their habitat had been made
reintroduce species that had become rare or extinct in sage. This, it seemed to me, was a zoo’s major raison
Victoria (Ziembicki 2010). One of the species the program d’être.
began breeding in the mid-1960s was the Australian
bustard Ardeotis australis. Researchers at the station Durrell drew some of his grand vision from Jean
recreated the bustard’s natural lek mating system in Delacour, a French aviculturist who, after World War I,
captivity by constructing an elaborate enclosure that had the dream of creating a Garden of Eden on Earth
allowed females to select the mate of their choice but (Mayr 1986). According to Ersnt Mayr (1986), Delacour
that prevented the males from direct contact with each succeeded as closely as is practical by establishing the
other (Ziembicki 2010). Other captive breeding programs finest private zoological park in the world at Chateau
in Australia to prevent avifauna extinctions followed, Clères in Normandy, France, before World War II (Mayr
including conservation breeding of orange-bellied parrot 1986). It was there that he laid the foundations for
Neophema chrysogaster (Orange-Bellied Parrot Recovery current aviculture of rare and exotic birds, especially
Team 1999), helmeted honeyeater Lichenostomus mela- pheasants and waterfowl and was a major influence on
nops cassidix (Smales et al. 1992), regent honeyeater the next generation of conservation breeders, including
Xanthomyza phrygia (Menkhorst et al. 1999), and black- Durrell, who considered Delacour ‘‘the most incredible
eared miners Manorina melanotis (Baker-Gabb 2003). aviculturist and ornithologist’’ (Botting 1999).
With government and nonprofit organizations con- The year after the establishment of Durrell’s zoo,
ducting scientific evaluations of captive breeding, by the volume 1 of The International Zoo Yearbook (hereafter
end of the 1970s many of the modern-day husbandry Yearbook) was published by the Zoological Society of
methods for breeding endangered birds in captivity had London, representing the first authoritative channel for
been worked out (Erickson 1980), although more the international exchange of information about zoos
sophisticated approaches to demographic and genetic (Jarvis 1967; Olney 2003). Information on the latest
management of captive populations and methods for breeding techniques could now be easily shared with a
minimizing behavioral problems in released birds were large number of zoos in short order, meetings could be
only beginning. coordinated, and a network could be formed for sharing
breeding stock. Furthermore, the editors of the Yearbook
The Changing Role of Zoos in the 1960s began to solicit and compile information from all known
zoos on breeding successes, research, rare animals, and
While Erickson’s lab was at work experimenting with several other subjects (Jarvis 1967). By 1966, Yearbook
breeding endangered species, the public was reacting to editors were sending questionnaires to 588 zoos. The
Carson’s Silent Spring, and Congress was busy putting reach of the Yearbook was summed up by its editor
protections in place for endangered species, zoos were Caroline Jarvis in 1967:
undergoing a sea-change in culture that would be critical
to the development of coordinated captive breeding
[The Yearbook] is bought not only by zoos but by
programs.
zoologists in universities and scientific research institutes
In 1959, the first zoological park specializing in from Ulan Bator in Outer Mongolia, to Novosibirsk in
endangered species conservation, the Jersey Zoological Siberia, Anchorage in Alaska and Mount Hagan in
Park, was founded by Gerald Durrell on the small island Western New Guinea (Jarvis 1967, p. 131).
of Jersey in the English Channel. According to Durrell
(1990), all the authorities in the zoological community he In the United States, increasing interest in breeding
consulted on the project told him that his plan to endangered species in zoos resulted from the incorpo-
establish a zoo for the conservation of endangered ration of the Wild Animal Propagation Trust (a coalition
species, which required him to take out a £25,000 loan, of 19 U.S. zoos, lead by the Bronx Zoo) under the aegis of
was foolish and destined to fail. Durrell (1990, pp. 17–18) the American Association of Zoological Parks and
was undeterred and reflected on his endeavor: Aquariums in 1963 (Hunt 1964; Conway 1969). This was
followed by a series of international symposia and
What I had in mind was an almost completely new conferences to discuss the role of zoos in wildlife
concept of the motivation of a zoological garden. Its first conservation, first in London in 1964 (Fitter 1967) and
major objective should be to act as an adjunct to the then in San Diego in 1966 (Durrell 1975). It was out of
whole conservation movement by the setting up of viable these conferences that the idea of conserving endan-
breeding colonies of those endangered species whose gered species in captivity would expand beyond the
numbers had dropped so drastically that they could no small island of Jersey in the English Channel and move
toward the top of the international agenda for zoos. By the grave’’ argument. Vandenberg (1975, p. 85), after
that time (in 1963), Durrell had paid back his bank loan discussing some problems with breeding the Congo
and formed the Jersey Wildlife Preservation Trust (later peacock Afropavo congensis, asked the following:
renamed the Durrell Wildlife Conservation Trust; Durrell
1990). His trust still is extremely successful, helping to Should we not then ask ourselves whether some species,
prevent the extinction of many species through captive which have now become rare, and for the conservation of
breeding. Perhaps the trust’s most noteworthy success is which cultural and scientific zoological gardens are
the program it established on Mauritius, under the making the greatest of sacrifices with little marked
supervision of Carl Jones (Durrell and Durrell 1980; Jones success, are not doomed to disappear and whether
and Hartley 1995). Jones and colleagues, with the man’s interference in these cases is fully justified?
assistance of The Peregrine Fund, developed a scientif-
ically based captive breeding program that restored Others were expressing concern about the loss of
viable populations of several critically endangered birds, selective pressures when birds were brought into
including the Mauritius kestrel, Mauritius fody Foudia captivity. Matthews (1973) noted,
rubra, pink pigeon Columba mayeri, and echo parakeet
Psittacula eques (Jones and Hartley 1995; Jones et al.
1995; Jones 1999, 2008). There is the distinct risk of a change in the genetic
At the 1966 conference regarding the role of zoos in constitution of the population during its time in captivity.
wildlife conservation, William Conway, Director of the Many natural selection pressures, such as predators, food
New York Zoological Society, presented a paper—later shortage, inclement weather, are removed. An aviculturist
published in Oryx (the journal of the Fauna Preservation would be justly pleased if he raised a complete brood from
Society)—entitled The Opportunity for Zoos to Save hatching; yet he will, in fact, be dulling the edge of natural
Vanishing Species (Conway 1967). In this landmark paper, selection …. Few aviculturists are sufficiently ruthless to
Conway built upon the focus of the San Diego eliminate even those showing marked defects if the
Conference and argued that zoos around the world species is very rare. Few are even sufficiently methodical
had an obligation to cooperate in the conservation of to avoid breeding from such birds (Matthews 1973, p. 9).
endangered species (something that Durrell had been
arguing since the 1950s). He calculated that with such Despite these, and other, cautions (e.g., Myers 1979),
cooperation, it would have been possible to save at least by the end of the 1970s, captive breeding of endangered
58 extinct birds, including the elephant birds (Aepyor- species was an accepted method, albeit a last resort, to
nithidae), lesser moas Anomalopteryx spp., dodo Raphus conserving critically endangered wildlife and many zoos
culcullatus, great auk, and Carolina parakeet. He noted had accepted their role to act responsibly and cooper-
that most endangered species in the 1960s were not atively in this endeavor. Moreover, endangered species
even being kept in potential breeding groups (Conway were now fully protected under the law in the United
1967) and called for broad changes in how zoos interact States, and other countries would soon follow their lead,
with one another. His ideas would get greater readership facilitated by the growing role of the IUCN’s Species
outside of the zoological community a few years later Survival Commission in organizing international sympo-
when he restated his arguments in the journal Science sia and coordinating the publication of Red Data Books,
(Conway 1969). which detailed species threatened with global extinction
Although there was growing rhetoric regarding the (see Scott et al. 1987 for a history of the Red Data Books).
changing role of zoos, the hard work of actually However, the methods for deciding which rare individ-
instituting the change would take longer. As Jarvis uals to trade between zoos were not yet quantitative or
(1967) noted, ‘‘At the moment it is true to say that in the formalized.
majority of zoos rare animals breed despite the zoos It was not until 1979, when Katherine Ralls and
rather than because of them’’ (Jarvis 1967, p. 134). Jonathan Ballou, researchers at the National Zoo, in
Among the problems at the time, were the refusal to Washington, D.C., found a correlation between inbreed-
cull inferior or aged specimens, the common practice of ing and juvenile mortality rates for 16 species of captive
selling off young zoo-bred stock and keeping too few animals, that the need to keep track of individual
animals in breeding groups (Jarvis 1967), and the lack of pedigrees became apparent (Ralls et al. 1979; Ralls and
genetic information to help sex birds and discourage Ballou 1986). Follow-up studies on 44 species confirmed
inbreeding. Furthermore, few zoos were keeping suffi- the results and according to Hancocks (2001, p. 142),
cient records for individual animals (Jarvis 1967; Durrell ‘‘Intensive Population Management became the new
1990). For some species, there was little or no incentive catchphrase, and the American Association of Zoological
to breed them in captivity, especially for those species Parks and Aquariums (now known as the AZA) began to
that were difficult to maintain in breeding groups due to strenuously promote the breeding of animals in genet-
space limitations. As Wayre (1969, p. 47) noted, ‘‘It was ically regulated programs.’’
easier and cheaper to buy freshly-trapped replace- Although studbooks had been kept for a handful of
ments.’’ wild species before Ralls and Ballou’s work, starting with
Furthermore, not everyone was sold on the idea of the American bison in 1932, there were only eight
captive breeding as a viable means to prevent extinc- studbooks for wild animals by 1966, when the first
tions, reminiscent of Loyle Miller’s (1942) ‘‘one wing in studbook list appeared in the Yearbook (all for mammals),
but the number increased to 38 in 1979 and to 61 in 1984 prove to be incompatible, infertile eggs are common, and
(Glatston 1986). The increased number of wild animal males tend to interfere with egg laying and incubation. In
studbooks was a clear reflection of the need for pedigrees addition, inbreeding is apparently affecting fertility and
to help guide the demographic and genetic management reproductive outcomes (Zoological Society of San Diego,
of captive breeding programs, especially for imperiled unpublished data).
species (Haig and Ballou 2002). By the mid- to late 1970s,
there was also recognition that a single body to Currently, the ‘alalā is extinct in the wild and numbers
coordinate management of studbooks and facilitate ,60 individuals in captivity. It is too soon to tell whether
cooperation amongst zoos was necessary (Martin 1975). captive breeding will prove successful in saving the ‘alalā
Thus, in 1979, the Captive Breeding Specialist Group (later from extinction, but, at this point, it is the only hope.
renamed the Conservation Breeding Specialist Group) of
the IUCN’s Species Survival Commission was formed to Controversy in Modern Conservation
promote global partnerships and collaborations in ex situ Breeding Programs
species conservation programs. Among other things, the
Captive Breeding Specialist Group connects all the Despite the fact that captive breeding programs had
regional breeding programs, as well as universities and grown out of their infancy by the 1980s, most breeding
wildlife management institutions in a global network, efforts had, until then, involved managing a captive
facilitates assessments of the need for conservation population while at least some individuals of the species
breeding programs, and formulates global breeding persisted in the wild. By 1980, when the California
strategies (Ebenhard 1995; Byers and Seal 2003). Condor Research Center was founded in a partnership
With the expansion of studbook data, the organization between the National Audubon Society and the U.S. Fish
and Wildlife Service, it was obvious that a vigorous
of a central body to coordinate captive breeding efforts,
captive breeding program would be necessary to save
emerging methods to amplify and analyze DNA, and the
the species (Kiff 1990). Although the intent of the
invention of personal computers, population biologists
breeding program, which was initiated in 1982, was to
and geneticists began to develop quantitative models for
leave several birds in the wild to serve as mentors to
pairing animals to maximize founder contributions and
captive-released birds, between November 1984 and
minimize the risk of inbreeding (Haig et al. 1990;
April 1985, 9 of the remaining 15 birds in the wild
Ebenhard 1995; Frankham 1995). During this time, there
vanished (i.e., they were probably killed by wanton
was also a growing recognition of the need for
shooting or lead toxicosis from eating carcasses shot
behavioral, physiological, and nutritional considerations
with lead ammunition; Kiff 1990). Notwithstanding
in pairing and breeding endangered animals in captivity
internal acrimony and public outcries, the decision was
(Dunn 1986; Kear 1986; Wielebnowski 1998); and by the
finally made in December 1985 to bring all remaining
late 1990s, a modern integrated approach to captive
birds in for a captive breeding program and on April 19,
management was in place:
1987, the last wild condor was captured.
This decision to capture all wild condors fractured
Now [in 1998] the need for a broader approach to captive relationships between wilderness advocates (also know
management has been recognized (e.g., Hutchins et al. as preservationists, for which the condor had become an
1995; Maillinson 1995) and research efforts incorporating unofficial symbol in California; Phillips and Nash 1981)
behavior, nutrition, disease, physiology, genetics, popu- and scientists (also known as conservationists) who
lation biology, and various interdisciplinary studies are advocated captive breeding as the only remaining hope
seen as necessary to facilitate responsible and successful for the species’ survival (Rickleffs 1978). As Alagona
captive propagation and conservation (Wielebnowski (2004, p. 568) stated,
1998, p. 137).
[To preservationists] condors and wilderness existed in a
This modern integrated approach has resulted in mutually dependent, symbiotic relationship. Condors
consistently successful breeding programs for some bird embodied wild nature and justified the protection of
groups (e.g., gamebirds, waterfowl, cranes, parrots, wilderness areas as critical endangered species habitat; in
raptors, Australian finches [Passeridae], and fringillid return, wilderness endowed the condor with its symbolic
finches [Fringillidae]); however, some species have proven capital. Some took this argument even further, claiming
difficult to breed in captivity. For example, attempts to that real condors only lived in the wilderness and that
breed the ‘alalā or Hawaiian crow Corvus hawaiiensis in intensive scientific management by definition robbed
captivity have been ongoing since 1973 (Duckworth et al. them of their wild essence. Condors would be better off
1992; Banko et al. 2002), but success in breeding and left alone under any circumstances, even if doing so led
releasing the ‘alalā has, so far, proven elusive, despite the to their extinction.
involvement of numerous ornithologists and aviculturists.
As the latest recovery plan for the species indicated (U.S. To preservationists, captive condors would be nothing
Fish and Wildlife Service 2009, I-16–I-17), more than ‘‘feathered pigs’’ (Alagona 2004). Despite their
rally cries to leave the condors alone in the wilderness to
In general, this species does not breed well in captivity: die with dignity, or to stop wasting scarce conservation
pairs require separate aviaries, many potential mates resources on a bird that was past its time (Pitelka 1981,
1982), the conservationists won out, and an intensive quate institutional support and funding, it may be
captive breeding program successfully rescued the possible to produce enough birds to effectively combat
California condor from the very brink of extinction the declines caused by falconers, at least at a local scale
(Snyder and Snyder 2000). A similar controversy erupted (Bailey et al. 1998). Some have argued that without
shortly thereafter on the Big Island of Hawaii over bustards to hunt, traditional Arab falconry is finished, and
capturing wild Hawaiian crows for a captive breeding without Arab falconry, the houbara is finished, because
program (see Walters 2006). As William Conway, Director no other group in the world has such a passion for
of the Bronx Zoo, noted in his 1986 paper entitled The conservation of this bird (Bailey et al. 1998). Whether
practical difficulties and financial implications of endan- captive breeding efforts will eventually help recover the
gered species breeding programs (Conway 1986, p. 218), houbara throughout Asia, where populations are still in
decline and threats are increasing (largely from falconry
Thus far, the international conservation community has and poaching), remains to be seen (Tourenq et al. 2005;
not seriously faced up to the need to support the Stone 2008).
acquisition of animals in danger of extinction for captive
breeding. Although support has been given to the The Ark Paradigm
proposals for the capture of a few animals, the Kouprey
Controversy over the efficacy of captive breeding has
Bos sauveli, the Tamarau Bubalus mindorensis, the Arabian
been debated since its inception as a species conserva-
oryx Oryx leucoryx, the California condor Gymnogyps
tion tool. However, it was during the 1980s that a few
californianus and several birds recently rescued from
biologists and zoo directors were arguing that govern-
Guam (where seven of its 15 species have become extinct
ments, zoos, and scientists needed to do much more if
during the past year), in no case has this occurred until
they were going to prevent looming mass extinctions.
the species was nearly extinct, the chances of propaga-
Some of them were starting a new society, The Society
tion poor and the opportunity to obtain representative
for Conservation Biology, to figure out how best to
founder stock lost.
conserve the Earth’s biodiversity.
In 1986, Michael Soulé, one of the architects of the
Unfortunately, this paradigm has persisted (e.g.,
Society for Conservation Biology (who earned a Ph.D.
puaiohi Myadestes palmeri; Tweed et al. 2006), po’ouli
Melamprosops phaeosoma (VanderWerf et al. 2003, 2006), under Paul Ehrlich, author of The Population Bomb
and Maui parrotbill Pseudonestor xanthophrys (Becker et [1968]), along with William Conway of the New York
al. 2010). Exceptions are rare and are generally limited to Zoological Society and others wrote an apocalyptic
species where there are utilitarian motives. One such paper entitled The millennium ark: how long a voyage,
exception is the conservation breeding program for the how many staterooms, how many passengers? (Soulé et
houbara bustard Chlamydotis undulate carried out by the al. 1986). In it, they predicted a coming demographic
National Avian Research Center, Environmental Research winter lasting between 500 and 1,500 y in which mass
and Wildlife Development Agency of Abu Dhabi and its extinctions will occur as human populations expand
major satellite operation the Emirates Center for Wildlife beyond their resources. They argued that saving species
Propagation in Missour, Morocco. It is the largest, most will require that zoos devote most of the space in zoos
expensive, most expansive breeding restoration program to the captive breeding of endangered species. Howev-
for any avian species since the peregrine falcon (the er, unlike Noah’s Ark, this would entail more than just
breeding facility in Missour consists of 2,500 individual keeping two of each species; it would require managing
cages for adult breeders, an incubation building that can populations, which would stretch the resources of zoos
incubate up to 40,000 houbara eggs per year, a rearing (also see Conway 1986). They believed that with
complex, and a prerelease facility that can hold .2,000 advances in technology over the next 200 y, it would
birds; Emirates Center for Wildlife Propagation 2010). The be possible to transfer from a ‘‘living zoo’’ to a
captive breeding effort was initiated in 1986 at Taif, Saudi ‘‘suspended zoo’’ where genetic information from each
Arabia, in response to declining populations of houbara species is frozen in time until the end of the
bustards, which have significant cultural value in the demographic winter of humans (i.e., after society
Middle East, North Africa, and Asia, where they have collapses), and even the possibility of transporting these
been the favored quarry of falconers for centuries (Saint miniature zoos to the dark side of the moon for cold
Jalme et al. 1996). A program of scientific inquiry into storage.
breeding and release techniques has resulted in a Although their ideas were not entirely new—Jarvis
remarkable program of artificial insemination, artificial (1967) had suggested a ‘‘wild animal bank’’ in 1967,
incubation of eggs, imprinting captive breeders to Conway (1967) had talked about the need for someone to
humans, and careful artificial rearing of young destined play the role of a ‘‘stern Noah with little space in his Ark’’,
for release to ensure that they do not imprint on humans and Wayre (1969) had warned of massive habitat
(Saint Jalme et al. 1996). Although international guide- destruction in the wake of future population growth and
lines suggest that reintroduction programs should be the need to breed more endangered species in captivity—
considered only if the factors that caused the decline are the use of computational tools to estimate the number of
eliminated or sufficiently reduced (IUCN 1987), the animals, the amount of space, and the amount of time
apparent success of the houbara captive breeding and needed to conserve endangered species was novel. The
reintroduction program demonstrates that with ade- concept of miniaturizing zoos was also new.
Following this landmark paper, in September 1987, the with pending bird extinctions that rival mass extinctions
IUCN adopted a policy statement on captive breeding that recorded in the fossil record, with threats from habitat
had wide-ranging implications (IUCN 1987; Bennett 1990). destruction, invasive species, disease, and a changing
With the spread of personal computers and enhanced climate. In particular, the threat of climate change has led
genetic techniques, quantitative tools for genetic and some to call for increased attention to captive breeding,
demographic management became more sophisticated. in lieu of ecological gambling on novel and untested
This new technology coupled with the successes of some approaches, such as assisted colonization and assisted
breeding programs led the IUCN to suggest that migration (Ricciardi and Simberloff 2009). From this brief
approximately 3,000 species worldwide needed some survey of the history of captive breeding in avian
form of captive breeding program (Ebenhard 1995). conservation, there are five insights that are applicable
Others would soon co-opt the Ark paradigm to to our present situation and the future of all living things.
promote increasing the capacity of zoos and increasing
the number of captive breeding programs. However, not 1) A fatalistic attitude is a sure path to extinction
everyone rallied behind the call for proliferation of Inaction will not solve the problems currently facing
breeding programs, because so many had failed (Griffith threatened species. As with the passenger pigeon and
et al. 1989). In 1996, Noel Snyder, the lead scientist in Carolina parakeet, deciding to do nothing will simply add
efforts to save the California condor from extinction in more species to the long list of extinct birds (see Fuller
the 1980s, and colleagues published an article in 1987 for this list) and will ignore ‘‘our bare duty to
Conservation Biology entitled Limitations of captive posterity to hand [a] fauna down as we found it, or as
breeding in endangered species recovery (Snyder et al. nearly so as the reasonable exigencies of life will admit’’
1996) in which they noted the surge in recovery efforts (Dixon 1889). Furthermore, no species is ‘‘doomed to
that were calling for long-term breeding associated with extinction’’ unless we decide it is. As Soulé (1987, p. 181)
the Ark paradigm. Although they acknowledged the so aptly put it, ‘‘There are no hopeless cases, only people
critical role captive breeding played in saving several without hope and expensive cases.’’ Believing that
species from extinction, they expressed concern that not species have a destiny to go extinct ignores the all-
enough care was being given to fully evaluating the encompassing impacts that humans have had on our
limitations of captive breeding, especially for short-lived planet. At best it is a form of denial; at worst, obfuscation
species that could quickly lose genetic diversity and to promote a competing agenda.
become domesticated without the forces of natural
selection (see Sheppard [1995] for a discussion of space 2) Bold action, political fortitude, and
limitations for long-term captive breeding of endangered conservation leaders required
birds). They also warned of the enormous costs and Preventing the extinction of critically endangered
resources necessary to successfully implement captive birds by using captive breeding is possible, but it often
breeding programs, which are largely dependent upon requires bold action, political fortitude, and strong
economic forces and continued public support for conservation leadership. The leaders of past generations
sustenance. Snyder’s admonition of Ark paradigm (AOU founders, Audubon Society founders, William
promoters was undoubtedly influenced by his involve- Temple Hornaday, and others) that thought it necessary
ment in several conservation breeding efforts that were to stand up against the millinery industry saved tens,
having difficulty achieving recovery, including efforts for maybe hundreds, of bird species from extinction. Gerald
the California condor (Snyder and Snyder 2000), thick- Durrell pioneered a new approach to zoological gardens
billed parrot Rhynchopsitta pachyrhyncha (Snyder et al. on the small island of Jersey with a bank loan and a
1994), and the Puerto Rican parrot Amazona vittata dream. Ray Charles Erickson of the Patuxent Endangered
(Snyder et al 1987). Species Research Center, Tom Cade of The Peregrine
Despite this and other critiques (e.g., Balmford et al. Fund, and other aviculturists and ornithologists around
1995), the Ark paradigm is very much alive today (Bowkett the world boldly pressed forward with rigorous research
2009). However, even with advances in cryogenics, agendas to develop the technical means of breeding
extracting ancient DNA, artificial insemination, in vitro endangered species, even as others told them it was
fertilization, embryo transfer, and cloning technologies, impossible. Rachel Carson’s exposé of the pesticide
we are not close to realizing the miniaturized millennium industry helped catalyze the environmental movement,
ark that Soulé et al. (1986) had envisioned. This is not all culminating in a cascade of international environmental
bad because there seems to be a consensus among protections. Because of these leaders, and those they
conservation biologists that without concurrently con- inspired, we have healthy populations of bald eagles,
serving habitats and reducing or eliminating other causal peregrine falcons, and waterfowl. In addition, many bird
mechanisms responsible for species’ declines, the ark species, including nēnē, several crane species, Mauritius
might as well be put on the dark side of the moon. kestrels, kakapos, and California condors, are on the path
to recovery. However, the war is not won, and many
Insights for Addressing the Current contentious battles remain to be fought if we want to
Extinction Crisis save endangered birds. Recently, much political rhetoric
has been expended on the need for partnerships in
The chronicle of captive breeding efforts in avian conservation. Although this need is undoubtedly true, it
conservation has only just begun. We are currently faced is either naı̈ve or insincere to believe that we can
overcome the pending extinction crisis without signifi- captive breeding programs and for providing valuable
cant conflict. Strong conservation leaders will be needed comments on earlier versions of this manuscript. I also
to take bold actions and navigate this conflict to thank an anonymous reviewer, the Subject Editor, and
conserve biodiversity, ‘‘on which rests the foundation the Editor-in-Chief for their helpful comments and
of human civilization’’ (Holdgate 1999, p. 33). suggestions. Last, but not least, I thank Mike Osborne
for providing me with the motivation to delve into the
3) The longer we wait, the greater the risk history of science.
Waiting until species are almost gone precludes
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Volume 1, Number 2, November 2010 ISSN 1944-687X

Special Thanks to Reviewers and Editors

Layne Adams Erin Gillam Frank Panek

Keywords: fertilizer; fire; nutrient; Odocoileus virginianus; white-tailed deer

Introduction Regeneration methods, such as clearcutting, and

Study Areas Methods

phate/acre and 131 lb potash/acre. We collected soil

into an ‘‘other’’ category. We could not calculate a Effects on forage production

sassafras, Carolina buckthorn Rhamnus caroliniana, com- Discussion

Introduction bottlenecks, of high mortality may exist for some species

season and later shifted to cladocerans. In 2005, copepods

t0 = 0) in 4 y for yellow perch. No correlations between Discussion

occur than in marine environments because freshwater

Figure 4. Larval yellow perch Perca flavescens density (solid

bluegill larvae. We did not observe any larval bluegill

Table 6. Bivariate correlations between predator–prey

be able to account for observed recruitment variability

Acknowledgments flavescens. Journal of the Fisheries Research Board of

Introduction dissolved solids [TDS]) for other trout species (Behnke

where t1 , t2 , … , tD denoted the ordered times of

Introduction coastal marsh habitats critical to the life history of

Horizon well highlights the fragility of coastal marsh and

Supplemental Material Johnson FA. 2009. Variation in population growth rates

able: http://www.fws.gov/southwest/refuges/plan/docs/ Weeks JL. 1969. Breeding behavior of mottled ducks in

Keywords: woodpeckers; density; Big Woods; Arkansas; bottomland hardwood forests

Introduction cies on how best to manage bottomland hardwoods in

detection probabilities for any species. We estimated a

Figure 4. Density estimates (bars indicate 95% confidence

including the CRNWR. They concluded that the bird

Introduction mast–producing species such as oaks (Quercus spp.) and

Statistical analysis between consecutive roost trees in the burned and

Keywords: conservation; habitat use; hibernation; management; snake

Introduction snake restricted to a cluster of islands in the Ohio and

Figure 3. Maximum distance inland from shore during

predator was a fox, raccoon, or domestic dog. Cause of

The water snakes, for the most part, appear to be restricted

The suggestion that the Lake Erie watersnake is largely

transformed means = 153 vs. 261 m; note that Pattishall

Keywords: National Wetlands Inventory; Minnesota; Prairie Pothole Region; wetlands

Introduction North American harvest (Batt et al. 1989). As well as

Results Wetlands with d modifiers in the NWI data base

Improvements in drainage technology, including pattern www.npwrc.usgs.gov/resource/wetlands/wetloss/index.

Keywords: activity; Anabat; bats; orientation; weatherproofing

Introduction bats without an observer present, thereby allowing a

Discussion to conduct species identification or compare their results

Introduction issue a health advisory for consumption of Trinity River

Keywords: brook trout; large woody debris; habitat; stream

Introduction habitat manipulation and found no significant changes

Figure 1. Total abundance of large woody debris (LWD) in

The added LWD continued to have no effect on age 1+

Figure 3. Total abundance of large woody debris (LWD) in

considering such habitat manipulations. For example,

Keywords: elk; desert Southwest; population dynamics; condition; survival; productivity

Introduction Skovlin et al. 2002). These limitations would result in

Around January 2000, about 20 elk colonized the Chaco

Introduction 1940s ,50% of the floodplain remained forested (Twedt

Keywords: bobcat; Lynx rufus; monitoring; North America; population status

Introduction hispidus (Schnell 1968), and pronghorn Antilocapra

Table 2. Methods used to monitor bobcat populations in

trends. Agency surveys such as this provide the only References

Keywords: carrying capacity; Lake Umbagog; normative research

Introduction These initial scientific applications suggested that the

Introduction and endangered species (Boyce 1992; Beissinger and

Keywords: birds; conservation; extinction; zoos