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FIG. 1 Movement trajectories of eight female zebra Equus quagga collared on the Chobe floodplains in Botswana and Namibia from
the late dry season (September–October) 2012 until June 2013. Grey polygons indicate actual (Nxai Pan National Park) and potential
alternative (Savuti Marsh and Seloko Plains) migration destinations. The rectangle on the inset indicates the location of the main map
in southern Africa.
mid October on the Namibia side (Fig. 1). Adult female determinant of grazing potential for herbivores such as
zebras (n 5 8) were darted from the air or ground and zebras; Fynn & Bonyongo, 2011) in Botswana. Nxai Pan is
immobilized using a mixture of etorphine hydrochloride, an important wet season habitat for wildlife because of
azaperone and hyaluronidase. The age and family group size the filling of waterholes and the growth of new grass on
were estimated for each individual, and a satellite-tracking the pan (Department of Wildlife and National Parks,
collar attached. Tracking collars (Africa Wildlife Tracking, 1995). Nxai Pan soils are categorized as fertile to very fertile
Pretoria, South Africa) were programmed to record global and are therefore attractive to grazers because of their
positioning system (GPS) locations at either 4- or 5-hour high nutritional value (Lindsay et al., 1998). There are
intervals. The total number of data points per individual was only two other locations in northern Botswana that
562–1,403 (mean 1,008 ± SD 215). Animals were captured have similarly fertile soils; both are closer to the Chobe
and treated according to the protocols approved under re- River and zebras could reach them without having to
search permit 1,537/2010 from the Ministry of Environment navigate around fences or other potential anthropogenic
and Tourism in Namibia, and EWT 8/36/4 XVII (57) barriers (Fig. 1). The Seloko Plains are c. 60 km from zebra
from the Department of Wildlife and National Parks in dry season ranges along the Chobe River and have fertile
Botswana. to very fertile soils (Lindsay et al., 1998). A small section
GPS locations were used to map and animate movement of the eastern part of these grasslands has been converted
trajectories of all individuals. We calculated net squared to commercial farming but the remainder is unconverted
displacement distances from the collaring site over time to and zebras and other wildlife are present (Chase, 2013).
characterize the scale and timing of migration (Bunnefeld The Savuti Marsh/Matabe Depression (Savuti Marsh
et al., 2011). hereafter) is c. 100 km from zebra dry season ranges and
We assessed whether migration to Nxai Pan could be has moderately fertile to fertile soils. Zebras and other
driven by unique environmental conditions at the site, first ungulates are known to migrate to Savuti from the Chobe
using spatial data layers on soil fertility (a proxy for the and Linyanti (the Linyanti is the name of the Chobe River
nutrition content of forage, and therefore a critical further westwards) floodplains (Vandewalle, 2000), and the
grasslands of this marsh support large herds of wildlife water), present (1–3 waterholes) or abundant (. 3 water-
(Fynn & Bonyongo, 2011; Sianga et al., 2013). These three holes and/or the presence of a stream or river). All cells
sites (Nxai Pan, Seloko Plains and Savuti Marsh) are also were categorized by the same observer to maintain
broadly classified as tree and grass savannah (Lindsay et al., consistency. We were unable to assess whether waterholes
1998). Remaining habitats accessible to zebras moving contained standing water that was accessible to zebras at
southwards from the Chobe River all occur on moderate- or the time they were encountered, only whether cells
low-fertility soils that would have limited appeal to contained waterholes or watercourses that had the potential
migrating grazers. to be filled. Similar to the minimum convex polygon
We characterized environmental conditions at Nxai Pan environmental analyses above we also calculated mean cell
and at the two potential alternative destinations (Seloko fractions covered by woody vegetation and mean en-
Plains and Savuti Marsh); our hypothesis was that Nxai Pan hanced vegetation index values for each 5 × 5 km cell,
should appear significantly different to the other two sites separately for the southward and northward movement
if the environment was driving zebra migration there. We phases.
used available data on vegetation biomass, tree cover and We conducted four separate statistical analyses, using
rainfall, and averaged the values of these variables over all generalized linear models and multi-model inference, to
cells in each defined site area (Fig. 1). We assessed whether investigate how environmental characteristics of cells along
temporal changes in mean vegetation biomass varied across the migratory pathway affected zebra movements.
these three potential wet season migration destinations, Southward migration
using the MODIS 250-m resolution enhanced vegetation e−ui [ui ]yi
index, which is produced at 16-day intervals (Pettorelli et al., Prob(Y = yi ) = , yi = 0, 1, 2, . . . (1)
yi !
2005). We used remotely sensed data from the Tropical
Rainfall Monitoring Mission (half-degree resolution) to where
characterize daily precipitation levels and cumulative pre-
In(ui ) = a + β1 ∗ Water + β2 ∗ Woody
cipitation at Nxai Pan, Seloko Plains and Savuti Marsh
during September 2012–March 2013. Finally, we character- + β3 ∗ EVISouth + β4 ∗ EVISouth2
ized tree cover at each of these three sites by averaging the
We used a negative binomial regression model to estimate
percentage of each cell covered by woody vegetation, using
counts of zebra GPS fixes in cells ( yi) as a function of water-
250-m MODIS continuous field data from 2010 (Hansen
holes (Water), the mean cell fraction covered by woody
et al., 2003).
vegetation (Woody), and a second-order polynomial of
We conducted an aerial census of zebras at Nxai Pan,
mean enhanced vegetation index values; a is the regression
using a fixed-wing aircraft, on 21 February 2013. Strip trans-
intercept and β is the regression coefficient of the associated
ects were flown, using standard methodology (Norton-
independent variable.
Griffiths, 1978). All zebras within 200 m of either side of a
transect were counted, with 27 parallel transects flown e[a + β1 ∗ Water + β2 ∗ Woody
during 07.30–10.00. This design ensured 100% coverage of + β3 ∗ EVISouth + β4 ∗ EVISouth2 ]
Prob.StopoverSouth =
the pan and resulted in a complete census of individuals in 1 + e[a + β1 ∗ Water + β2 ∗ Woody
the area. High-resolution photographs were taken for more + β3 ∗ EVISouth + β4 ∗ EVISouth2 ]
accurate counting post-flight.
(2)
To assess the relationship between environmental
conditions and zebra movements along the main migratory We used a logistic regression model to investigate the prob-
pathway we calculated the minimum convex polygon across ability of a cell being used as a stopover location as a func-
all GPS locations of all individuals moving from the Chobe tion of waterholes (Water), the mean cell fraction covered by
River to Nxai Pan and back. We then overlaid a 5 × 5 km woody vegetation (Woody), and a second-order polynomial
grid across this polygon and calculated the number of of mean enhanced vegetation index values; a is the re-
GPS points in each cell separately for two time periods: gression intercept and β is the regression coefficient of the
the southward (4 December 2012–10 January 2013) and the associated independent variable.
northward (18 February 2013–13 June 2013) migratory Northward migration
phases. We classified grid cells as stopover areas when at e−ui [ui ]yi
least one zebra spent at least 48 consecutive hours in an Prob(Y = yi ) = , yi = 0, 1, 2, . . . (3)
yi !
encamped state in the cell (Morales et al., 2004; Sawyer &
Kauffman, 2011). where
Using high-resolution imagery from Google Earth
In(ui ) = a + β1 ∗ Water + β2 ∗ Woody
we assessed the availability of standing water to zebras in
each grid cell as absent (no visual evidence of standing + β3 ∗ EVINorth + β4 ∗ EVINorth2
TABLE 1 Summary statistics for migration movements of eight female zebras Equus quagga collared on the Chobe floodplains in Botswana
and Namibia (Fig. 1).
Zebras spent a mean of 73 days in the Nxai Pan area at least 48 hours during the migratory phases) appeared
(range 53–96), mostly on the pan itself. We counted 1,534 unrelated to waterhole abundance, enhanced vegetation
zebras in the area during our aerial survey on 21 February index or woody vegetation cover (Table 2). The number
2013, although correspondence with safari guides in the park of GPS relocations in cells along the southward migratory
indicated that in the 2 weeks prior to our survey there had pathway also appeared unrelated to waterhole abundance,
been more zebras in the vicinity of the pan. enhanced vegetation index or woody vegetation cover.
Return trajectories of zebras moving from the Nxai Pan However, cells with higher numbers of GPS fixes during
north to the Chobe River were less direct and took sig- the northward migratory phase contained higher densities
nificantly longer. The mean return time was 85 days (vs (and presence) of waterholes than those used less frequently
15 days for the southward journey; t 5 4.33, P 5 0.004) and (Table 2). The top 10 models for each of our four analyses
mean distance travelled was 653 ± SE 153 km (vs 302 km; accounted for 68–100% of the cumulative Akaike model
t 5 5.79, P 5 0.0006). The mean distance of actual move- weight, indicating that a relatively small fraction of models
ment trajectories of the seven zebras that survived to (15%) had high explanatory power (Supplementary
complete the entire return journey from the Chobe flood- Table S1).
plains to Nxai Pan was 955 km (range 735–1,170 km).
Environmental characteristics of Nxai Pan overlapped Discussion
with the two possible alternative wet season destinations for
Chobe River zebras. Woody vegetation cover, a measure of Prior to the discovery of this migration it was unclear
broad habitat type, was similar at all three potential destin- where Chobe River zebras moved to in the wet season, with
ations (Nxai Pan, 18.0 ± SE 3.7%; Seloko Plains, 17.1 ± SE local residents and wildlife professionals only aware that
2.6%; Savuti Marsh, 20.0 ± SE 5.3%), as was total rainfall animals left at the beginning of the wet season and returned
during September 2012–March 2013 (Nxai Pan, 444 mm; the following dry season (Chase, 2011). Similarly, the
Seloko Plains, 477 mm; Savuti Marsh, 470 mm). The area of appearance of zebras on the Nxai Pan in the wet season is
Nxai Pan used by zebras had a consistently lower mean an annual occurrence but there was no evidence that
vegetation biomass than Savuti Marsh. However, biomass animals originated as far away as the Chobe River. The
levels at Nxai Pan and Seloko Plains were similar from mid timing of the migration from the Chobe River appeared to
December to early February, which encompasses most of coincide with heavy, episodic rainfall events early in the wet
the wet season period that zebras were present at Nxai Pan season at Nxai Pan. Other migratory ungulate species
(Fig. 3). Biomass levels at Nxai Pan dropped below those of (Sinclair & Norton-Griffiths, 1979), as well as zebras in
Seloko Plains in mid February; at the same time the zebras the recently re-established migration from the Okavango
began to move back north towards the Chobe River Delta to the Makgadikgadi Pans (Bartlam-Brooks et al.,
floodplains (Table 1). 2011), also appear to time migration to rainfall events,
As suggested by initial analyses, averaged coefficients and zebras are able to orient long-range movements to
across all model combinations showed that spatial autocor- environmental conditions outside their perceptory range
relation in zebra location data was strong, with coefficients (Brooks & Harris, 2008). However, as we have only observed
on the first-order neighbourhood variable significantly two southward migration onsets we are unable to assess
greater than zero in all four analyses (Table 2). Controlling statistically the nature of the relationship between timing
for this autocorrelation the probability of a 5 × 5 km cell of rainfall and migration without a substantially longer
being used as a stopover point (i.e. where zebras stopped for time series.
Our aerial census showed that there were at least 1,500 unable to quantify access to drinking water at any of the
zebras on Nxai Pan in the wet season of 2012–2013, with three locations but at the time of migration Savuti Marsh
anecdotal information indicating this number may have held water for the first time in 28 years and was therefore
been substantially higher a few weeks before our census. much wetter than either Seloko Plains or Nxai Pan. Access
This figure is similar to dry season estimates of zebra to water would therefore not have been a problem at that
numbers (1,558–3,593) on the Chobe floodplains during site, although it is possible that having been dry for so long
2007–2012, which were also estimated from strip-transect the marsh is still an unfavoured destination that will need to
surveys (MC, unpubl. data). Combined with the facts that all be rediscovered by migratory ungulates. Also, we could not
eight of the female zebras we collared in 2012 moved from quantify predation risk, which is an important driver of
the Chobe River to Nxai Pan, the six surviving zebras re- migration in ungulates (Fryxell & Sinclair, 1988), although
peated this southward movement to Nxai Pan in December lions Panthera leo, the main predators of zebras in this
2013, and zebra numbers on the Chobe floodplains during region, are present in sufficient numbers to draw tourists in
the wet season (255) are an order of magnitude lower than both Savuti Marsh and Nxai Pan (Power & Compion, 2009;
during the dry season (Chase, 2013), this suggests that most Johnson, 2010).
of the zebras on the Chobe floodplains may be moving to With no obvious environmental differences between
Nxai Pan in the wet season. Further censuses and satellite Nxai Pan and the other two possible destinations for zebras
tracking of more individuals over time could confirm the from Chobe River, this migration may represent an ancient,
annual periodicity of the migration, as well as the fraction of conserved phenomenon that has a genetic basis and/or is an
Chobe animals that move to Nxai Pan vs those that remain outcome of learned behaviour and cultural transmission
or migrate elsewhere. Additional studies are also needed to between individuals and social groups (Alerstam, 2006;
determine whether other ungulates such as the wildebeest, Bolger et al., 2008). Research has shown that the Okavango–
long-distance migrants that move in tandem with zebras in Makgadikgadi zebra migration in Botswana, itself a long-
other ecosystems (e.g. Serengeti, Liuwa Plains—Mussuma distance event involving c. 15,000 zebras, reoccurred
Transfrontier Conservation area; Harris et al., 2009), during 2008–2009 for the first time following the removal of
perform the same Chobe–Nxai Pan migration. a veterinary fence that had blocked the route during
The reasons for such a long-distance migration, with 1968–2004. This completely separate migration suggests a
closer and apparently suitable alternative destinations by- conserved memory of an ancient route that may be
passed, remain unknown. Our analyses of potential environ- genetically encoded (Bartlam-Brooks et al., 2011), and raises
mental drivers of the migration are preliminary and our the question of why zebras from Chobe River move to Nxai
hypothesis that the migration is a fixed behaviour rather Pan rather than the nearby Makgadikgadi Pans (and vice
than a variable response to environmental conditions needs versa). This fixed-route phenomenon has also been ob-
further testing. Seloko Plains and Savuti Marsh have similar served in pronghorn Antilocapra americana in the USA,
levels of woody vegetation, soil fertility and rainfall to Nxai where individuals in the Grand Teton National Park area
Pan, and wet season vegetation biomass was similar for have used the same seasonal migration pathway, as narrow
Seloko Plains and Nxai Pan. Vegetation biomass was higher as tens of metres in some places, since the Holocene era
at Savuti Marsh than Nxai Pan, which, if anything, suggests (Berger et al., 2006).
greater grazing availability and therefore higher suitability Our analyses imply that zebra movement and stopovers
for zebras migrating from the Chobe floodplains. We were along the migratory pathway between dry and wet season
,0.001
,0.001
,0.001
0.01
0.61
0.98
0.80
0.16
0.99
0.99
0.99
0.90
0.75
0.48
ation in vegetation, and cells with waterholes are not used
P preferentially as stopover points, although cells with water-
holes had higher numbers of zebra GPS counts than those
4.64
5.28
2.78
4.82
0.51
0.03
0.25
1.42
0.01
0.01
0.01
0.13
0.31
0.71
without for the northward (but not southward) migration.
model) and to the probability of a cell being a stopover point on the migration route (logistic regression model) during both northward and southward movement phases.
−4.36E-06–3.16E-06
environmental conditions along the migratory pathway.
Confidence limits
−1.10E-06–8.9E-07
0.035–0.087
1.149–2.506
0.280–1.617
−4.300–7.294
−0.006–0.006
−0.120–0.019
−0.021–0.024
−1943–1922
−1966–1983
−2824–2853
Comparing our data with those compiled in a review
−0.28–0.13
4.05–9.6
(Harris et al., 2009) shows that the newly discovered zebra
migration ranks as the longest large-mammal migration in
Africa. The round-trip length of this migration is longer
than long-distance movements of zebras and wildebeest
in the Serengeti ecosystem (A.R.E. Sinclair, pers. comm.;
−1.30E-07 ± SE 5.16E-07
−6.01E-07 ± SE 1.91E-06
0.0015 ± SE 0.0015 Harris et al., 2009). It is also longer than migrations of topi
0.061 ± SE 0.013
1.827 ± SE 0.345
0.948 ± SE 0.339
1.497 ± SE 2.944
−0.0001 ± SE 0.003
−0.050 ± SE 0.035
8.85 ± SE 1003
14.51 ± SE 1441
6.82 ± SE 1.41
−0.07 ± SE 0.10
−10.52 ± SE 981
,0.001
0.58
0.18
0.36
0.67
0.64
0.46
0.99
1.00
1.00
0.89
0.96
0.98
4.54
1.33
0.92
0.43
0.47
0.74
0.01
0.00
0.00
0.13
0.05
0.02
0.163–0.300
−1.824–3.270
−0.192–1.008
−0.342–0.953
−0.003–0.002
−0.046–0.103
−0.010–0.009
−0.357–0.364
−1587–1575
−2920–2931
−2977–2987
5.7–14.4
−3.92E-08 ± SE 8.23E-07
0.408 ± SE 0.305
0.305 ± SE 0.329
−0.0005 ± SE 0.001
0.028 ± SE 0.037
−0.0006 ± SE 0.005
0.004 ± SE 0.183
5.39 ± SE 1486
5.29 ± SE 1514
10.08 ± SE 2.21
−5.94 ± SE 802
along the Boteti River excluded most of the river from the
South
Waterholes present
Intercept
migrations should be a high conservation priority. The B R I E R S , R.A. (2002) Incorporating connectivity into reserve selection
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an assessment of potential threats and impediments to and yearly differences. Journal of Animal Ecology, 80, 466–476.
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We thank Hans Swartbooi, Lise Hanssen, Carl-Heinz D O B S O N , A.P., B O R N E R , M., S I N C L A I R , A.R.E., H U D S O N , P.J.,
Moeller, Michelle Kastern, Werner Kilian and Johannes A N D E R S O N , T.M., B I G U R U B E , G. et al. (2010) Road will ruin
Kapner for help in the field, Alicia Ellis, Colby Loucks, Serengeti. Nature, 467, 272–273.
Sylvia Thompson, David Wilcove, Tony Sinclair, Dave D O R M A N N , C.F., M C P H E R S O N , J.M., A R A U J O , M.B., B I VA N D , R.,
B O L L I G E R , J., C A R L , G. et al. (2007) Methods to account for spatial
Ward, Raymond Peters and two anonymous reviewers for
autocorrelation in the analysis of species distributional data: a
their constructive comments, and the Namibia Ministry of review. Ecography, 30, 609–628.
Environment and Tourism, the Botswana Department E P P S , C.W., M U T AY O B A , B.M., G W I N , L. & B R A S H A R E S , J.S. (2011)
of Wildlife and National Parks, the Eppley Foundation, An empirical evaluation of the African elephant as a focal species
the Paul G. Allen Family Foundation, and the San Diego for connectivity planning in East Africa. Diversity and Distributions,
Zoo Global Wildlife Conservancy for funding and logistical 17, 603–612.
F E R G U S O N , K. & H A N K S , J. (eds) (2010) Fencing Impacts: A Review
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Veterinary Fencing in Africa with Particular Reference to the Great
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L I N D S E Y , P.A., R O U L E T , P.A. & R O M A N AC H , S.S. (2007) Economic of the Witwatersrand, Johannesburg, South Africa.
and conservation significance of the trophy hunting industry in W I L C O V E , D.S. (2009) No Way Home: The Decline of the World’s
sub-Saharan Africa. Biological Conservation, 134, 455–469. Great Animal Migrations. Island Press, Washington, DC, USA.
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on wildlife populations in the Okavango Delta, Botswana. migration disappearing? PLoS Biology, 6(7), e188.
International Journal of Wilderness, 12, 17–41. W I L L I A M S O N , D. & W I L L I A M S O N , J. (1984) Botswana’s fences and the
M O R A L E S , J.M., H AY D O N , D.T., F R A I R , J., H O L S I N E R , K.E. & depletion of Kalahari wildlife. Oryx, 18, 218–222.
F R Y X E L L , J.M. (2004) Extracting more out of relocation data:
building movement models as mixtures of random walks.
Ecology, 85, 2436–2445. Biographical sketches
N O R T O N -G R I F F I T H S , M. (1978) Counting Animals. African Wildlife
Leadership Foundation, Nairobi, Kenya. R O B I N N A I D O O is a conservation scientist who has worked in Africa
P E R K I N S , J. (2010) Fences and landscape scale degradation. In on various issues for over a decade. M I K E C H A S E is an expert on
Fencing Impacts: A Review of the Environmental, Social elephant ecology and conservation. P I E T B E Y T E L L is a conservation
and Economic Impacts of Game and Veterinary Fencing in biologist with the Namibian government. P I E R R E D U P R E E Z is
Africa with Particular Reference to the Great Limpopo and Namibia’s chief conservation biologist and runs the country’s rhino
Kavango-Zambezi Transfrontier Conservation Areas (eds recovery programme. K E L L Y L A N D E N is based in Botswana and helps
K. Ferguson & J. Hanks), pp. 108–120. Mammal Research Institute, run Elephants Without Borders. G R E G S T U A R T - H I L L has decades of
Pretoria, South Africa. experience in community-based conservation and rangeland manage-
P E T T O R E L L I , N., V I K , J.O., M Y S T E R U D , A., G A I L L A R D , J.-M., ment in southern Africa. R U S S E L L T A Y L O R is a regional expert in the
T U C K E R , C.J. & S T E N S E T H , N.C. (2005) Using the satellite-derived conservation of large wildlife and natural resources management.