PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

127(1):35–46. 2014.

New records and distribution of marine free-living nematodes

in the Maldivian Archipelago
Federica Semprucci* and Maria Balsamo
(FS, MB) Dipartimento di Scienze della Terra, della Vita e dell’Ambiente (DiSTeVA),
Università degli Studi di Urbino ‘Carlo Bo’, Loc. Crocicchia, 61029 Urbino, Italy,
e-mail: federica.semprucci@uniurb.it

Abstract.—Biodiversity estimations, in particular in tropical regions, are a

useful tool in research focused on the knowledge and conservation of
ecosystems. Inferences on the biogeography of marine nematodes are often
difficult due to the poor knowledge of the species distribution. In particular,
limited information on the composition of the free-living marine nematodes is
available from the Indian Ocean and Maldivian Archipelago. This study aims
at providing a checklist of the free-living marine nematodes associated with
coral sediments of the Maldives. Forty-six species, in 28 genera and 10
families, are newly recorded for the Archipelago, increasing the number of
known nematode species to 111, in 79 genera, and 25 families. The
distribution range of the species is updated, and a noticeable faunistic
richness is recorded for the southern archipelago. The presence of six species
described first in Maldives by Gerlach and still known only for this region is
confirmed. The present synthesis of the faunistic and taxonomic data available
so far for the Maldives Islands integrates the global information on the
distribution of this phylum and provides information for future interesting
comparisons from a zoogeographical perspective.

Keywords: biodiversity, conservation, coral reefs, Maldives Islands,

Nematoda

Free-living nematodes are an important
component of benthic marine ecosystems
both in terms of abundance and biomass
and are closely related with other organ-
isms playing a key role in trophic webs (see
Giere 2009 for review). Knowledge of the
global distribution of marine free-living
nematodes is overall satisfying, but many
geographical regions are still poorly
known or even completely unknown. An
example is the Indian Ocean, where only
fragmentary and mostly dated investiga-
tions have been carried out (see Semprucci
& Balsamo 2012).
Coral reefs occupy only a small fraction
of the oceans, but they are known as one
* Corresponding author.
of the most productive and diverse ecolog-
ical systems on the planet. Out of the 34
existing metazoan phyla, representatives of
28 phyla have been found in coral reef
communities (Adrianov 2004).
Documenting species diversity and its
spatial patterns are fundamental for defin-
ing priorities in programs of conservation
and management, and this is particularly
urgent in tropical areas (Montiel et al.
2011). Here, the major causes of coral
death include overfishing, frequent hurri-
canes, water pollution, surface water heat-
ing, and specific diseases. Most of these
factors are due also to local environmental
conditions or global climate change (Gor-
eau et al. 2000, Barton & Casey 2005).
Despite of the increasing scientific interest
36 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
for these habitats and the problems related
to their conservation, studies on meio-
benthic biodiversity have been overall
scarce.
The present work is part of a research
project on the biodiversity and ecology of
the Maldivian Archipelago (Semprucci et
al. 2010, 2011, 2013). This area is peculiar
for appearing as a double series of
numerous atolls close to each other. Here,
the coral bleaching episode of 1998 caused
widespread coral mortality, up to 90% in
shallow coral reefs (Wilkinson et al. 1999,
Bianchi et al. 2006, Lasagna et al. 2010),
and a consequent remarkable change in
the composition of coral communities at
the genus level (Tkachenko 2012). All of
these changes may have had, and still have,
a notable influence on the associated
meiofauna, which occur in a great variety
of micro-niches in reef sediments. Biodi-
versity estimates and the compilation of
species lists are crucial for the development
of conservation initiatives aimed at pre-
serving local species richness. The faunistic
knowledge of free-living marine nematodes
of the Maldives is based on the data
collected during the Xarifa-Expedition
1957/1958 and published by Gerlach
(1962, 1963a, 1963b, 1964).
The present paper aims to provide
additional information on the presence
and distribution of marine nematode
species in the Maldivian archipelago.
Study Area
The Maldives Islands are situated in the
Indian Ocean, in the central part of the
Chagos-Maldives-Laccadive Ridge. With
21 atolls and some 1200 islands, Maldives
extend in a north-south direction from
about 7806 0 N and 0842 0 S in latitude, rising
steeply from the abyssal plain, 2500 m
deep eastward and 4000 m deep westward
(Fig. 1). The sea floor of the lagoons inside
the atolls is 50–60 m deep, whereas the
channels between the atolls reach a depth
of 300–400 m (Ciarapica & Passeri 1993,
Gischler & Kikinger 2007).
Materials and Methods
Samples were collected from the central
atolls of Felidu, South Malé, and Ari, the
peripheral islands Rasdhoo and Thoddoo,
and the southern atoll of Suvadiva. The
central atolls were investigated during
scientific cruises carried out in May 2005,
2007, and 2008, which mainly focused on
back-reef platforms, lagoons, and outer
reefs (Semprucci et al. 2010, 2011, 2013).
Samples from Suvadiva were collected in a
lagoon habitat in May 2009.
Samples for the study of meiofauna and
nematodes were treated using standard
techniques (for details see Semprucci et
al. 2010). Nematode specimens were iso-
lated, transferred in glycerine, and mount-
ed as permanent slides (Seinhorst 1959).
Identification at the species level was
performed using a light microscope
equipped with Nomarski optics (Nikon
Optiphot-2) and was aided by the pictorial
keys of Platt & Warwick (1983, 1988),
Warwick et al. (1998), and the NeMys
online identification key (Deprez et al.
2004), and the literature by Gerlach (1962,
1963a, 1963b, 1964).
The species collected by Gerlach during
the Xarifa-Expedition 1957/1958 are in-
cluded in the present paper in order to
compile a complete list of nematode
species known so far from the area. They
were found in the northern atolls of
Miladummadulu, Fadiffolu, and the pe-
ripheral island of Gaha Faro, in the central
atoll of Ari, and in the southern atoll of
Addu.
The taxonomic status of the species
reported has been checked by consulting
the following literature: Gerlach & Rie-
mann (1973/1974), Verschelde et al. (1998),
Deprez et al. (2004), and Tchesunov
(2013). Specimens of all the species record-
VOLUME 127, NUMBER 1 37

Fig. 1. Study area.

38 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
ed are present in the first author’s collec-
tion, at the University of Urbino.
Results
Sixty-five species were found overall in
the present survey. A comparison with the
species list from the Xarifa-Expedition
1957/1958 (Gerlach 1962, 1963a, 1963b,
1964) reveals that 46 of these species, in 36
genera, and 16 families, are recorded in the
archipelago for the first time (Table 1).
Most of the new records are from Suvadi-
va (25 species), the others from Felidu (13),
and South Malé (7). In general, the new
records are mainly species belonging to the
families Desmodoridae, Chromadoridae,
Xyalidae, Cyatholaimidae, and Dracone-
matidae. The Desmodoridae and Chroma-
doridae are found in sediments with a wide
range of grain size, from coarse to fine
sands but especially in medium sands,
whereas the Xyalidae occur mostly in fine
and medium sands, and Cyatholaimidae
occur in a variety of substrates (see Table
1).
Nineteen species, that are here newly
recorded, appear particularly associated
with medium sands, 16 species with fine
sands, and 6 species with coarse sands
(Table 1). Five species are present in
sediments with various grain sizes, from
coarse to medium, such as Chromadorella
macris, Dracognomus annae, Dracognomus
notohalensis, Odontophora villoti, and
Theristus denticulatus (Table 1).
With the present update, 111 species of
free-living marine nematodes belonging to
79 genera and 25 families are now recorded
from the Maldivian Archipelago. Of these,
23 species and 2 genera (Maldivea and
Squanema) were first described from the
Maldives by Gerlach (1962, 1963a, b,
1964).
Considering the composition of the
nematode fauna, the order Chromadorida
appears to be the richest, with the highest
number of families and genera (9 and 68,
respectively), followed by Enoplida (8 and
22), and Monhysterida (6 and 18) (Table
1). The highest number of species is
recorded in the families Desmodoridae
(36 species in 19 genera), Cyatholaimidae
(9 in 9), Chromadoridae (8 in 6), Oxy-
stominidae (6 in 5), and Xyalidae (5 in 3).
The most widespread families are Desmo-
doridae (9 localities from northern to
southern areas of the archipelago), Cya-
tholaimidae (7), Axonolaimidae (6), Oxy-
stominidae (5), and Monoposthiidae (5).
The most widespread genera are the
Desmodoridae Desmodora (7 localities),
Desmodorella (6), Eubostrichus (6), Zalo-
nema (6), Laxus (5), and the Xyalidae
Theristus (4). The most widespread species
are Desmodorella tenuispiculum, Eubostri-
chus parasitiferus, Zalonema megalosoma,
Laxus gerlachi, Desmodora varioannulata,
and T. denticulatus. Most of these wide-
spread species are found in shallow sub-
tidal habitats (mainly back-reef platforms)
and sporadically in deeper subtidal habi-
tats (lagoons or outer reefs).
The nematode assemblages of the Suva-
diva atoll appear to be the richest ones,
with the highest number of families (14),
genera (34), and species (39), followed by
those of the atolls of Addu (11 families, 33
genera, and 36 species), Felidu (11, 21, and
26), and Ari (10, 18, and 17). Overall, the
southern and central regions of the archi-
pelago show higher numbers of species
compared to northern areas, where only 11
species were collected (9 species in Fadif-
folu and 2 in Miladummadulu) (Table 2).
Discussion
Desmodoridae, Chromadoridae, Cya-
tholaimidae, and Xyalidae are the richest
nematode families in the Maldives. Quan-
titative data collected reveal that they are
also the dominant meiobenthic compo-
nent in Maldivian coral sediments (Sem-
prucci et al. 2010, 2011, 2013), which
agrees with the data reported for coral
VOLUME 127, NUMBER 1
sediments of other geographical areas,
also named coral degradation zones
(CDZ) (Renaud-Mornant & Gourbault
1984, Boucher & Gourbault 1990, Gour-
bault & Renaud-Mornant 1990, Tietjen
1991, Boucher 1997, Ndaro & Ólafsson
1999, Kotta & Boucher 2001, Raes et al.
2007). CDZs are very dynamic habitats
made up of sediments associated with the
coral reef and can be found in all the
functional zones of the reef. In this zone,
coral material coming from the main reef
is progressively degraded into smaller
pieces (coral gravel) until it becomes
coralline sediment. Some authors have
suggested that the nematode composition
of the CDZ is more related to the grain
size of the sediment than to its carbonatic
nature (see for review Raes et al. 2007).
Indeed, the richest and dominant families
in coral sediments are typical of medium-
coarse sands (Heip et al. 1985) such as
those found in Maldives. Here, the
seafloor receives a continuous supply of
originally biogenic sediments of different
size, but waves and currents are not able
to rework and sort them effectively
(Semprucci et al. 2010) so that the most
common sediments are poorly sorted
medium and coarse sands. The high level
of angularity of the coralline sediment
particles creates numerous interstitial
spaces, representing ideal micro-habitats
suitable for well-diversified assemblages
of meiofauna and nematodes (Semprucci
et al. 2010). Indeed, these areas provide
sheltering as well as settling of biofilms
and microalgae on the grain surfaces, a
basic food source also for nematodes (see
Giere 2009 for an overview).
A high level of diversity and richness are
observed even in the deeper subtidal
habitats that are often characterized by
fine coralline sediments (Semprucci et al.
2013). An overall high complexity of these
micro-habitats has been inferred, which
may be related again to the poor selective
action of the transport processes (Sem-
prucci et al. 2013).
39
Desmodoridae, the most abundant and
richest family, includes 6 taxa of the
subfamily Stilbonematinae: Eubostrichus
cf. hopperi, E. parasitiferus, L. gerlachi,
L. sigma, Robbea tenax, and Stilbonema
annulatum. In particular, E. parasitiferus
and L. gerlachi are dominant in medium
and coarse sands of the Maldives (Sem-
prucci et al. 2010). Stilbonematinae are
well known as sediment dwellers often
associated with anoxic sediments in which
they are able to exploit the high concen-
trations of sulfide, present around the
chemocline, by means of their ectosym-
biotic bacteria (Hentschel et al. 1999, Ott
et al. 2004). However, this subfamily has
been frequently reported as abundant in
subtropical and tropical sediments and
especially in coralline sands of different
geographical regions (Boucher 1997, Rie-
mann et al. 2003, Raes et al. 2007). It has
been suggested that its high abundance
and richness in coralline sediments may be
related to the presence of large grains in
generally sheltered habitats, a condition
that represents an unusual combination of
wide interstitial spaces and sulfide condi-
tions with an associated rich thiobios
(Riemann et al. 2003, Giere 2009).
Different types of epibionts (the Sucto-
ria Thecacineta calyx, T. halacari, and
Limnoricus sp.) are observed on the cuticle
of some Desmodoridae, mainly Desmodora
pontica, Croconema sphaericum, and D.
tenuispiculum. Thecacineta calyx is well
known as an epibiont of nematodes (for
review see Ingole et al. 2010), whereas T.
halacari and Limnoricus sp. have been
reported as epibionts of Halacarida, Co-
pepoda Harpacticoida, and Kinorhyncha,
but not Nematoda (Dovgal et al. 2008,
2009).
The richest and most abundant families
are characterized by various cuticular
ornamentations, probably representing an
adaptation to the respective micro-habitat
and are mainly epistrate feeders. This
trophic guild is commonly found in
medium-coarse sands with a very poor
40 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Checklist of free-living marine nematodes reported for the Maldivian Archipelago. Report
abbreviations: * ¼ species collected during present study; published records by Gerlach: 1 ¼ 1962, 2 ¼ 1963a, 3
¼ 1963b, 4 ¼ 1964. Atoll abbreviations: A ¼ Addu, Ar ¼ Ari, F ¼ Fadiffolu, Fe ¼ Felidu, G ¼ Gaha Faro, M ¼
Miladummadulu, R ¼ Rasdhoo, S ¼ Suvadiva, SM ¼ South Malé, T ¼ Thoddoo. Substrate abbreviations: a ¼
algae, cor fr ¼ coral fragments, cor sed ¼ coral sediments, CS ¼ coarse sands, FS ¼ fine sands, Man ¼
mangrove, MS ¼ mixed or medium sands, PCS ¼ pikeperch among coralline sediment, S ¼ sand, Sþdetritus ¼
sand associated with detritus, sed ¼ sediments.

Taxon Report Atoll Type of substratum

Anticomidae
Anticoma acuminata (Eberth, 1863) 1 A
A. lata Cobb, 1898 1 F, R a, Favites-, Porites- and
Acropora-sed
A. trichura Cobb, 1898 1 A, G S, FS, CS
Anticomopsis typica Micoletzky, 1930 1 A Seriatopora- and Acropora-
sed
Leptosomatidae
Platycoma africana (Gerlach, 1959) 1 F MS
Oncholaimidae
Viscosia glabra (Bastian, 1865) * Ar MS
V. macramphida Chitwood, 1951 * SM MS
V. viscosia (Bastian, 1865) * Fe CS
Oxystominidae
Halalaimus filum Gerlach, 1962 1 M Rhizophora-Man
H. supercirrhatus Gerlach, 1955 1 Ar PCS
Litinium volutum Gerlach, 1962 1 Ar S
Maldivea xarifae Gerlach, 1962 1, * F, Fe CS
Oxystomina alpatovi (Filipjev, 1927) 1 G S
Thalassoalaimus impar Gerlach, 1962 1 Ar CS
Phanodermatidae
Phanoderma cocksi Bastian, 1865 1 Ar Psammocora-sed
P. ocellatum (Cobb, 1920) 1 A a
Thoracostomopsidae
Epacanthion buetschlii (Southern, 1914) * Fe CS
E. quadridiscus Shimada, Kajihara & * R MS
Mawatari, 2009
Mesacanthion hirsutum Gerlach, 1953 * Ar MS
Rhabdodemaniidae
Rhabdodemania dura Inglis, 1966 * S FS
Tripyloididae
Bathylaimus depressus Gerlach, 1962 1 A
Tripyloides undulatus Gerlach, 1962 1 A CS
Trefusiidae
Cytolaimium exile Cobb, 1920 1 Ar S
Xennellidae
Porocoma striata Cobb, 1920 1 A Lithothamnion-zone
Xennella suecica Allgén, 1935 * S MS
Ceramonematidae
Dasynemoides spinosus Gerlach, 1963 2 G FS
Metadasynemella elegans Vitiello, 1974 * S MS
Chromadoridae
Chromadorella macris (Gerlach, 1956) * SM CS, MS
Parapinnanema cf. harveyi Warwick & * Fe MS
Coles, 1975
Prochromadorella ditlevseni (De Man 1922) * Fe CS
P. macroocellata Wieser, 1951 * T MS
P. septempapillata Platt, 1973 * SM MS
Ptycholaimellus ponticus (Filipjev, 1922) * S FS
VOLUME 127, NUMBER 1 41

Table 1.—Continued.

Taxon Report Atoll Type of substratum

Spilophorella euxina Filipjev, 1918 * S MS

Trochamus complexus Boucher, 1976 * S FS
Cyatholaimidae
Acanthonchus setoi Wieser, 1955 * S CS
Craspodema reflectans Gerlach, 1964 4, * G, S MS
Longicyatholaimus maldivarum Gerlach, 1964 4, * R, SM, T PCS, CS
Marylynnia annae (Wieser & Hopper, 1967) * Fe FS
Metacyatholaimus brevicollis (Cobb, 1898) 4 A, Fe, S FS, MS, cor fr
Nannolaimoides armatus (Gerlach, 1964) 4 A S
Paracanthonchus barka Inglis, 1962 * S FS
Paracyatholaimoides cf. multispiralis Gerlach, * Fe CS
1953
Paracyatholaimus duplicatus Gerlach, 1964 4 A, S FS, CS
Desmodoridae
Acanthopharyngoides duplex Gerlach, 1963 2 A cor sed
Acanthopharynx micans (Eberth, 1863) 2 A Pocillopora-sed
A. rigida Stekhoven, 1950 * R MS
Bolbonema brevicollis (Cobb, 1920) 2, * A, R, S, T FS, CS, MS
Chromaspirina indica Gerlach, 1963 2, * G, T FS, MS
C. madagascariensis Gerlach, 1953 2 A S
Croconema cinctum Cobb, 1920 2, * A, S Lithothamnion-zone, MS
C. sphaericum (Kreis, 1928) * S MS
C. torquens (Gerlach, 1963) 2, * Fe MS
Desmodora californica Allgén, 1947 2, * F, Fe Halimeda-sed, CS
D. gerlachi Vitiello, 1971 2 A, R S, Lithothamnion-zone
D. pontica Filipjev, 1922 * T CS
D. varioannulata (Kreis, 1928) 2, * A, Fe, G, R, S, Lithothamnion-zone,
Acropora-sed, FS-CS
Desmodorella balteata Verschelde, Gourbault * S MS
& Vincx, 1998
D. tenuispiculum Allgén, 1928 2, * Ar, Fe, R, S, MS, CS
SM, T
Eubostrichus cf. hopperi Muthumbi, * SM FS
Verschelde & Vincx, 1995
E. parasitiferus Chitwood, 1936 2 A, F, Fe, S, FS, MS, CS, Millepora
SM, T
Laxus gerlachi (Hopper & Cefalu, 1973) 2, * Fe, G, R, S, MS, CS
SM
L. sigma (Gerlach, 1963) 2 R PCS
Metachromadora clavata Gerlach, 1957 2 A
M. serrata Gerlach, 1963 2 Ar Halimeda-sed
M. spectans Gerlach, 1957 * S MS
Molgolaimus allgeni Allgén, 1935 * S FS
Onyx dimorphus Gerlach, 1963 2 F CS
O. cf. perfectus Cobb, 1891 2 Ar S
Paradesmodora campbelli (Allgén, 1932) 2, * A, Fe, S Sþdetritus, CS, MS
P. punctata Gerlach, 1963 2 G MS
Robbea tenax Gerlach, 1963 3 Ar
Spirinia laevioides Gerlach, 1963 2 A, F, Fe Sþdetritus, a, CS
S. laevis (Bastian, 1865) * S FS
S. parasitifera (Bastian, 1865) 2, * F, Fe Sþdetritus, FS
Squanema articulatum Gerlach, 1961 2 Ar CS
Stilbonema annulatum Gerlach, 1963 2 R PCS
Stygodesmodora bacillicauda (Gerlach, 1963) 2, * G, S MS, CS
42 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Continued.

Taxon Report Atoll Type of substratum

Zalonema maldivensis (Gerlach, 1963) 2, * A, Fe, R Zoantharia-sed, MS

Z. megalosoma Steiner, 1918 2, * A, Fe, G, R, MS, CS
S, SM
Draconematidae
Apenodraconema spinicaudum (Gerlach, 1958) * Ar, S MS
Dracognomus annae Verschelde & Vincx, 1993 * Ar, Fe MS, CS
D. marioni Allen & Noffsinger, 1978 * Fe MS
D. notohalensis Allen & Noffsinger, 1978 * Ar, Fe, SM MS, CS
Ethmolaimidae
Filitonchus volutus (Gerlach, 1956) * S FS
Leptolaimidae
Camacolaimus prytherchi Chitwood, 1935 1 A, F CS, Sþdetritus
Cynura papillata Gerlach, 1962 1 A
Onchium ocellatum Cobb, 1920 1 A Lithothamnion-zone,
Montipora-sed
Procamacolaimus tubifer Gerlach, 1953 1 A
Monoposthiidae
Monoposthia costata (Bastian, 1865) 2, * A, Fe, S MS
Nudora thorakista Schulz, 1935 2 A, M Man
Rhinema retrorsum Cobb, 1920 2, * A, Ar, S FS, MS
Tarvaiidae
Tarvaia cf. angusta Gerlach, 1953 1 F Sþdetritus
Axonolaimidae
Axonolaimus arcuatus Stekhoven, 1950 1 A S
Odontophora furcata Wieser, 1956 1 Ar, G FS, MS
O. villoti Luc & De Coninck, 1959 * Fe, SM MS, CS
Parodontophora pacifica (Allgén, 1947) 1, * A, S Montipora-sed, FS
Comesomatidae
Dorylaimopsis mediterranea Grimaldi-De Zio, * S FS
1968
D. pellucida (Cobb, 1920) * S FS
Paracomesoma sigmoidalis Riera, Núnez & * Fe MS
Del Carmen Brito, 2006
Diplopeltidae
Araeolaimus bioculatus (De Man, 1876) 1 A a
Campylaimus gerlachi Timm, 1961 * S FS
Diplopeltis cirrhatus (Eberth, 1863) 1 A Lithothamnion-zone
Diplopeltula indica Gerlach, 1962 1 A Sþdetritus
Siphonolaimidae
Astomonema obscura (Boucher & Helléouët, * S MS
1977)
Sphaerolaimidae
Sphaerolaimus gracilis de Man, 1876 * S FS
Xyalidae
Cobbia urinator Wieser, 1959 * S FS
Elzalia poli Castillo-Fernández & Lambshead, * S FS
1990
E. tuberculata Hope & Aryuthaka, 2009 * S FS
Theristus denticulatus Warwick, 1970 * Ar, Fe, S, MS, CS
SM
T. flevensis Stekhoven, 1935 * S MS
VOLUME 127, NUMBER 1 43

Table 2.—Total number of free-living marine nematode families, genera, and species collected in atolls and
peripheral islands of the Maldivian Archipelago.

Latitude Longitude Atoll Families Genera Species

0 0
06800 N 73817 E Miladummadulu 2 2 2
05825 0 N 73837 0 E Fadiffolu 6 9 9
04816 0 N 73800 0 E Rasdhoo 4 10 12
04844 0 N 73829 0 E Gaha Faro 6 11 11
03858 0 N 72856 0 E Ari 10 18 17
03859 0 N 73830 0 E South Malé 7 11 12
03833 0 N 73829 0 E Felidu 11 21 26
04826 0 N 72857 0 E Thoddoo 3 7 7
00834 0 N 73812 0 E Suvadiva 14 34 39
00838 0 N 73812 0 E Addu 11 33 36

fine fraction (Alongi 1986); moreover, in
tropical habitats, a high benthic primary
production, a great abundance of diatoms,
and numerous wide surfaces colonized by
algal and bacterial biofilms are easily
found (Boucher 1997, Raes et al. 2007).
Differences in the number of nematode
species found from the sampled localities
of the Maldives clearly are the result of a
heterogeneous effort in faunistic and tax-
onomic research in the various areas but
also suggest potential differences between
local environmental conditions. For in-
stance, in the southern part of the archi-
pelago, the anthropogenic pressure is still
limited.
It is worthwhile noting that few taxa (5
of 65) described from the Maldives by
Gerlach (1962, 1963a, 1963b, 1964) have
changed their taxonomic status since the
first record. In particular, the genus
Desmodora de Man, 1889 (sensu Lorenzen,
1981) originally comprised six sub-genera,
of which Croconema Cobb, 1920 and
Zalonema Cobb, 1920 were then consid-
ered as genera by Gourbault & Vincx
(1990) and Verschelde & Vincx (1996),
respectively. Of the Maldivian species
Desmodora (Xenodesmodora) torquens was
moved to the genus Croconema by Ver-
schelde et al. (1998), who also transferred
Desmodora (Desmodora) maldivensis to the
genus Zalonema. Leptonemella sigma was
transferred to the genus Laxus by Tchesu-
nov (2013), whereas Metadesmodora bacil-
licauda was moved to the genus
Stygodesmodora by Blome (1982). Nanno-
laimus armatus was transferred to the
genus Nannolaimoides by Ott (1972).
Worthy of mention is the finding of
some species reported first in the Maldives
by Gerlach, whose distribution is still
limited to this area: Maldivea xarifae,
Craspodema reflectans, Longicyatholaimus
maldivarum, Chromaspirina indica, Zalone-
ma maldivensis, and Stygodesmodora ba-
cillicauda.
Conclusions
Information on the distribution of free-
living marine nematodes in the Indian
Ocean is fragmentary and often dated.
Knowledge for the Maldives was limited to
the species collected during the Xarifa-
Expedition 1957/1958 and reported by
Gerlach in the (1962, 1963a, 1963b,
1964). Here we provide an updated syn-
thesis of the faunistic and taxonomic data
available so far for the Maldives Islands:
we integrate the global information on
free-living marine nematodes, allowing for
future comparisons from zoogeographical
point of view. The present checklist
comprises 111 species belonging to 79
genera and 25 families: 46 species, in 36
genera and 16 families are newly recorded
for the Maldives. The distribution range of
the species has been updated, and a greater
44 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
faunistic richness in the southern archipel-
ago is recorded. The presence of six species
described first in the Maldives by Gerlach
and still known only for this region is
confirmed.
Acknowledgments
We are grateful to Professor Paolo
Colantoni and Dr. Giuseppe Baldelli for
the sample collection and the sedimento-
logical analysis. We would like also to
thank Dr. Silvia Tazioli for the taxonomic
identification of the specimens of Suctoria.
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