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: [email protected]

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 of the most productive and diverse ecolog-
component of benthic marine ecosystems ical systems on the planet. Out of the 34
both in terms of abundance and biomass existing metazoan phyla, representatives of
and are closely related with other organ- 28 phyla have been found in coral reef
isms playing a key role in trophic webs (see communities (Adrianov 2004).
Giere 2009 for review). Knowledge of the Documenting species diversity and its
global distribution of marine free-living spatial patterns are fundamental for defin-
nematodes is overall satisfying, but many ing priorities in programs of conservation
geographical regions are still poorly and management, and this is particularly
known or even completely unknown. An urgent in tropical areas (Montiel et al.
example is the Indian Ocean, where only 2011). Here, the major causes of coral
fragmentary and mostly dated investiga- death include overfishing, frequent hurri-
tions have been carried out (see Semprucci canes, water pollution, surface water heat-
& Balsamo 2012). ing, and specific diseases. Most of these
Coral reefs occupy only a small fraction factors are due also to local environmental
of the oceans, but they are known as one conditions or global climate change (Gor-
eau et al. 2000, Barton & Casey 2005).
* Corresponding author. Despite of the increasing scientific interest
36 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

for these habitats and the problems related of 300–400 m (Ciarapica & Passeri 1993,
to their conservation, studies on meio- Gischler & Kikinger 2007).
benthic biodiversity have been overall
scarce.
The present work is part of a research Materials and Methods
project on the biodiversity and ecology of
the Maldivian Archipelago (Semprucci et Samples were collected from the central
al. 2010, 2011, 2013). This area is peculiar atolls of Felidu, South Malé, and Ari, the
for appearing as a double series of peripheral islands Rasdhoo and Thoddoo,
numerous atolls close to each other. Here, and the southern atoll of Suvadiva. The
the coral bleaching episode of 1998 caused central atolls were investigated during
widespread coral mortality, up to 90% in scientific cruises carried out in May 2005,
shallow coral reefs (Wilkinson et al. 1999, 2007, and 2008, which mainly focused on
Bianchi et al. 2006, Lasagna et al. 2010), back-reef platforms, lagoons, and outer
and a consequent remarkable change in reefs (Semprucci et al. 2010, 2011, 2013).
the composition of coral communities at Samples from Suvadiva were collected in a
the genus level (Tkachenko 2012). All of lagoon habitat in May 2009.
these changes may have had, and still have, Samples for the study of meiofauna and
a notable influence on the associated nematodes were treated using standard
meiofauna, which occur in a great variety techniques (for details see Semprucci et
of micro-niches in reef sediments. Biodi- al. 2010). Nematode specimens were iso-
versity estimates and the compilation of lated, transferred in glycerine, and mount-
species lists are crucial for the development ed as permanent slides (Seinhorst 1959).
of conservation initiatives aimed at pre-
Identification at the species level was
serving local species richness. The faunistic
performed using a light microscope
knowledge of free-living marine nematodes
equipped with Nomarski optics (Nikon
of the Maldives is based on the data
Optiphot-2) and was aided by the pictorial
collected during the Xarifa-Expedition
keys of Platt & Warwick (1983, 1988),
1957/1958 and published by Gerlach
Warwick et al. (1998), and the NeMys
(1962, 1963a, 1963b, 1964).
The present paper aims to provide online identification key (Deprez et al.
additional information on the presence 2004), and the literature by Gerlach (1962,
and distribution of marine nematode 1963a, 1963b, 1964).
species in the Maldivian archipelago. 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
Study Area species known so far from the area. They
The Maldives Islands are situated in the were found in the northern atolls of
Indian Ocean, in the central part of the Miladummadulu, Fadiffolu, and the pe-
Chagos-Maldives-Laccadive Ridge. With ripheral island of Gaha Faro, in the central
21 atolls and some 1200 islands, Maldives atoll of Ari, and in the southern atoll of
extend in a north-south direction from Addu.
about 7806 0 N and 0842 0 S in latitude, rising The taxonomic status of the species
steeply from the abyssal plain, 2500 m reported has been checked by consulting
deep eastward and 4000 m deep westward the following literature: Gerlach & Rie-
(Fig. 1). The sea floor of the lagoons inside mann (1973/1974), Verschelde et al. (1998),
the atolls is 50–60 m deep, whereas the Deprez et al. (2004), and Tchesunov
channels between the atolls reach a depth (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- respectively), followed by Enoplida (8 and
tion, at the University of Urbino. 22), and Monhysterida (6 and 18) (Table
1). The highest number of species is
recorded in the families Desmodoridae
Results (36 species in 19 genera), Cyatholaimidae
(9 in 9), Chromadoridae (8 in 6), Oxy-
Sixty-five species were found overall in stominidae (6 in 5), and Xyalidae (5 in 3).
the present survey. A comparison with the The most widespread families are Desmo-
species list from the Xarifa-Expedition doridae (9 localities from northern to
1957/1958 (Gerlach 1962, 1963a, 1963b, southern areas of the archipelago), Cya-
1964) reveals that 46 of these species, in 36 tholaimidae (7), Axonolaimidae (6), Oxy-
genera, and 16 families, are recorded in the stominidae (5), and Monoposthiidae (5).
archipelago for the first time (Table 1). The most widespread genera are the
Most of the new records are from Suvadi- Desmodoridae Desmodora (7 localities),
va (25 species), the others from Felidu (13), Desmodorella (6), Eubostrichus (6), Zalo-
and South Malé (7). In general, the new nema (6), Laxus (5), and the Xyalidae
records are mainly species belonging to the Theristus (4). The most widespread species
families Desmodoridae, Chromadoridae, are Desmodorella tenuispiculum, Eubostri-
Xyalidae, Cyatholaimidae, and Dracone- chus parasitiferus, Zalonema megalosoma,
matidae. The Desmodoridae and Chroma- Laxus gerlachi, Desmodora varioannulata,
doridae are found in sediments with a wide and T. denticulatus. Most of these wide-
range of grain size, from coarse to fine spread species are found in shallow sub-
sands but especially in medium sands, tidal habitats (mainly back-reef platforms)
whereas the Xyalidae occur mostly in fine and sporadically in deeper subtidal habi-
and medium sands, and Cyatholaimidae tats (lagoons or outer reefs).
occur in a variety of substrates (see Table The nematode assemblages of the Suva-
1). diva atoll appear to be the richest ones,
Nineteen species, that are here newly with the highest number of families (14),
recorded, appear particularly associated genera (34), and species (39), followed by
with medium sands, 16 species with fine those of the atolls of Addu (11 families, 33
sands, and 6 species with coarse sands genera, and 36 species), Felidu (11, 21, and
(Table 1). Five species are present in 26), and Ari (10, 18, and 17). Overall, the
sediments with various grain sizes, from southern and central regions of the archi-
coarse to medium, such as Chromadorella pelago show higher numbers of species
macris, Dracognomus annae, Dracognomus compared to northern areas, where only 11
notohalensis, Odontophora villoti, and species were collected (9 species in Fadif-
Theristus denticulatus (Table 1). folu and 2 in Miladummadulu) (Table 2).
With the present update, 111 species of
free-living marine nematodes belonging to
79 genera and 25 families are now recorded Discussion
from the Maldivian Archipelago. Of these,
23 species and 2 genera (Maldivea and Desmodoridae, Chromadoridae, Cya-
Squanema) were first described from the tholaimidae, and Xyalidae are the richest
Maldives by Gerlach (1962, 1963a, b, nematode families in the Maldives. Quan-
1964). titative data collected reveal that they are
Considering the composition of the also the dominant meiobenthic compo-
nematode fauna, the order Chromadorida nent in Maldivian coral sediments (Sem-
appears to be the richest, with the highest prucci et al. 2010, 2011, 2013), which
number of families and genera (9 and 68, agrees with the data reported for coral
VOLUME 127, NUMBER 1 39

sediments of other geographical areas, Desmodoridae, the most abundant and

also named coral degradation zones richest family, includes 6 taxa of the
(CDZ) (Renaud-Mornant & Gourbault subfamily Stilbonematinae: Eubostrichus
1984, Boucher & Gourbault 1990, Gour- cf. hopperi, E. parasitiferus, L. gerlachi,
bault & Renaud-Mornant 1990, Tietjen L. sigma, Robbea tenax, and Stilbonema
1991, Boucher 1997, Ndaro & Ólafsson annulatum. In particular, E. parasitiferus
1999, Kotta & Boucher 2001, Raes et al. and L. gerlachi are dominant in medium
2007). CDZs are very dynamic habitats and coarse sands of the Maldives (Sem-
made up of sediments associated with the prucci et al. 2010). Stilbonematinae are
coral reef and can be found in all the well known as sediment dwellers often
functional zones of the reef. In this zone, associated with anoxic sediments in which
coral material coming from the main reef they are able to exploit the high concen-
is progressively degraded into smaller trations of sulfide, present around the
pieces (coral gravel) until it becomes chemocline, by means of their ectosym-
coralline sediment. Some authors have biotic bacteria (Hentschel et al. 1999, Ott
suggested that the nematode composition et al. 2004). However, this subfamily has
of the CDZ is more related to the grain been frequently reported as abundant in
size of the sediment than to its carbonatic subtropical and tropical sediments and
nature (see for review Raes et al. 2007). especially in coralline sands of different
Indeed, the richest and dominant families geographical regions (Boucher 1997, Rie-
in coral sediments are typical of medium- mann et al. 2003, Raes et al. 2007). It has
coarse sands (Heip et al. 1985) such as been suggested that its high abundance
those found in Maldives. Here, the and richness in coralline sediments may be
seafloor receives a continuous supply of related to the presence of large grains in
originally biogenic sediments of different generally sheltered habitats, a condition
size, but waves and currents are not able that represents an unusual combination of
to rework and sort them effectively wide interstitial spaces and sulfide condi-
(Semprucci et al. 2010) so that the most tions with an associated rich thiobios
common sediments are poorly sorted (Riemann et al. 2003, Giere 2009).
medium and coarse sands. The high level Different types of epibionts (the Sucto-
of angularity of the coralline sediment ria Thecacineta calyx, T. halacari, and
particles creates numerous interstitial Limnoricus sp.) are observed on the cuticle
spaces, representing ideal micro-habitats of some Desmodoridae, mainly Desmodora
suitable for well-diversified assemblages pontica, Croconema sphaericum, and D.
of meiofauna and nematodes (Semprucci tenuispiculum. Thecacineta calyx is well
et al. 2010). Indeed, these areas provide known as an epibiont of nematodes (for
sheltering as well as settling of biofilms review see Ingole et al. 2010), whereas T.
and microalgae on the grain surfaces, a halacari and Limnoricus sp. have been
basic food source also for nematodes (see reported as epibionts of Halacarida, Co-
Giere 2009 for an overview). pepoda Harpacticoida, and Kinorhyncha,
A high level of diversity and richness are but not Nematoda (Dovgal et al. 2008,
observed even in the deeper subtidal 2009).
habitats that are often characterized by The richest and most abundant families
fine coralline sediments (Semprucci et al. are characterized by various cuticular
2013). An overall high complexity of these ornamentations, probably representing an
micro-habitats has been inferred, which adaptation to the respective micro-habitat
may be related again to the poor selective and are mainly epistrate feeders. This
action of the transport processes (Sem- trophic guild is commonly found in
prucci et al. 2013). 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 licauda was moved to the genus
tropical habitats, a high benthic primary Stygodesmodora by Blome (1982). Nanno-
production, a great abundance of diatoms, laimus armatus was transferred to the
and numerous wide surfaces colonized by genus Nannolaimoides by Ott (1972).
algal and bacterial biofilms are easily Worthy of mention is the finding of
found (Boucher 1997, Raes et al. 2007). some species reported first in the Maldives
Differences in the number of nematode by Gerlach, whose distribution is still
species found from the sampled localities limited to this area: Maldivea xarifae,
of the Maldives clearly are the result of a Craspodema reflectans, Longicyatholaimus
heterogeneous effort in faunistic and tax- maldivarum, Chromaspirina indica, Zalone-
onomic research in the various areas but ma maldivensis, and Stygodesmodora ba-
also suggest potential differences between cillicauda.
local environmental conditions. For in-
stance, in the southern part of the archi-
pelago, the anthropogenic pressure is still Conclusions
limited.
It is worthwhile noting that few taxa (5 Information on the distribution of free-
of 65) described from the Maldives by living marine nematodes in the Indian
Gerlach (1962, 1963a, 1963b, 1964) have Ocean is fragmentary and often dated.
changed their taxonomic status since the Knowledge for the Maldives was limited to
first record. In particular, the genus the species collected during the Xarifa-
Desmodora de Man, 1889 (sensu Lorenzen, Expedition 1957/1958 and reported by
1981) originally comprised six sub-genera, Gerlach in the (1962, 1963a, 1963b,
of which Croconema Cobb, 1920 and 1964). Here we provide an updated syn-
Zalonema Cobb, 1920 were then consid- thesis of the faunistic and taxonomic data
ered as genera by Gourbault & Vincx available so far for the Maldives Islands:
(1990) and Verschelde & Vincx (1996), we integrate the global information on
respectively. Of the Maldivian species free-living marine nematodes, allowing for
Desmodora (Xenodesmodora) torquens was future comparisons from zoogeographical
moved to the genus Croconema by Ver- point of view. The present checklist
schelde et al. (1998), who also transferred comprises 111 species belonging to 79
Desmodora (Desmodora) maldivensis to the genera and 25 families: 46 species, in 36
genus Zalonema. Leptonemella sigma was genera and 16 families are newly recorded
transferred to the genus Laxus by Tchesu- for the Maldives. The distribution range of
nov (2013), whereas Metadesmodora bacil- the species has been updated, and a greater
44 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

faunistic richness in the southern archipel- Cobb, N. A. 1920. One hundred new nemas (type
ago is recorded. The presence of six species species of 100 new genera). Contributions to a
Science of Nematology 9:217–343.
described first in the Maldives by Gerlach de Man, J.-G. 1889. Troisi ème note sur les
and still known only for this region is nématodes libres de la Mer du Nord et de la
confirmed. Manche. Mémoires de la Société Zoologique
de France 2:182–216.
Deprez, T., E. Vanden Berghe, & M. Vincx. 2004.
Acknowledgments NeMys: a multidisciplinary biological infor-
mation system. Pp. 57–63 in E. Vanden
We are grateful to Professor Paolo Berghe, M. Brown, M. J. Costello, C. Heip,
Colantoni and Dr. Giuseppe Baldelli for S. Levitus & P. Pissierssens, eds., Proceedings
the sample collection and the sedimento- of ‘the colour of ocean data’: international
symposium on oceanographic data and infor-
logical analysis. We would like also to mation management with special attention to
thank Dr. Silvia Tazioli for the taxonomic biological data Brussels, Belgium, November
identification of the specimens of Suctoria. 25–27, 2002. IOC Workshop Report 188/
VLIZ Special Publication 16. UNESCO/
IOC, VLIZ, Paris, France, 308 pp.
Literature Cited Dovgal, I., T. Chatterjee, B. Ingole, & M. Nanajkar.
2008. First report of Limnoricus ponticus
Adrianov, A. V. 2004. Current problems in marine Dovgal & Lozowskiy (Ciliophora: Suctorea)
biodiversity studies. Russian Journal of Ma- as epibionts on Pycnophyes (Kinorhyncha)
rine Biology 30(Suppl. 1):S1–S16. from the Indian Ocean with key to species of
Alongi, D. M. 1986. Population structure and the genus Limnoricus. Cahiers de Biologie
trophic composition of the free-living nema- Marine 49:381–385.
todes inhabiting carbonate sands of Davies Dovgal, I., T. Chatterjee, D. V. Subba Rao, B. K. K.
Reef, Great Barrier Reef, Australia. Austra- Chan, & M. De Troch. 2009. New records of
lian Journal of Marine and Freshwater Praethecacineta halacari (Schulz) (Suctorea:
Research 37:609–619. Ciliophora) from Taiwan, Tanzania and
Barton, A. D., & K. S. Casey. 2005. Climatological Canada. Marine Biodiversity Records 2:1–3.
context for large-scale coral bleaching. Coral Gerlach, S. 1962. Freilebende Meeresnematoden von
Reefs 24:536–554. den Malediven. Kieler Meeresforschungen
Bianchi, C. N., C. Morri, M. Pichon, F. Benzoni, P. 18:81–108.
Colantoni, G. Baldelli, & M. Sandrini. 2006. Gerlach, S. 1963a. Freilebende Meeresnematoden
Dynamics and pattern of coral recolonization von den Malediven II. Kieler Meeresforschun-
following the 1998 bleaching event in the reefs gen 19:67–103.
of the Maldives. Pp. 30–37 in Y. Suzuki, T. Gerlach, S. 1963b. Robbea tenax sp. n., ein
Nakamori, M. Hidaka, H. Kayanne, B. E. merkwürdiger mariner Nematode von den
Casareto, K. Nadaoka, H. Yamano, & M. Malediven. Internationale Revue der Gesamt-
Tsuchiya, eds., Proceedings of 10th Interna- en Hydrobiologie und Hydrographie 48:153–
158.
tional Coral Reef Symposium, Okinawa,
Gerlach, S. 1964. Neue Cyatholaimidae (Nematoda,
Japan, June 28–July 2, 2004. Japanese Coral
Chromadorida) von den Malediven. Veröffen-
Reef Society, Tokyo, Japan.
tlichungen des Instituts für Meeresforschung
Blome, D. 1982. Systematik der Nematoda eines
in Bremerhaven 9:70–78.
Sandstrandes der Nordseeinsel Sylt. Mikro-
Gerlach, S., & F. Riemann. 1973/1974. The Bremer-
fauna des Meeresbodens 86:1–194. haven checklist of aquatic nematodes: a
Boucher, G. 1997. Structure and biodiversity of catalogue of Nematoda Adenophorea exclud-
nematode assemblages in the SW lagoon of ing the Dorylaimida. Veröffentlichungen des
New Caledonia. Coral Reefs 16:177–186. Instituts für Meeresforschung in Bremer-
Boucher, G., & N. Gourbault. 1990. Sublittoral haven, Supplement 4:1–734.
meiofauna and diversity of nematode assem- Giere, O. 2009. Meiobenthology: the microscopic
blages off Guadeloupe Islands (French West motile fauna of aquatic sediments. Second
Indies). Bulletin of Marine Science 47:448–463. edition. Springer-Verlag, Berlin, Germany,
Ciarapica, G., & L. Passeri. 1993. An overview of the 527 pp.
Maldivian coral reefs in Felidu and North Gischler, E., & R. Kikinger. 2007. Effects of the
Malé atoll (Indian Ocean): platform drowning tsunami of 26 December 2004 on Rasdhoo
by ecological crises. Facies 28:33–66. and Northern Ari Atolls, Maldives. Pp. 93–
VOLUME 127, NUMBER 1 45

104 in D. R. Stoddart, ed., Tsunamis and Platt, H. M., & R. M. Warwick. 1983. Free-living
coral reefs. Atoll Research Bulletin No. 544. marine nematodes. Part I: British enoplids.
Smithsonian Institution, Washington, D.C. Synopses of the British fauna (new series), No.
Goreau, T., T. McClanahan, R. Hayes, & A. Strong. 28. Cambridge University Press, Cambridge,
2000. Conservation of coral reefs after the United Kingdom, 307 pp.
1998 global bleaching event. Conservation Platt, H. M., & R. M. Warwick. 1988. Free living
Biology 14:5–15. marine nematodes. Part. II: British chroma-
Gourbault, N., & J. Renaud-Mornant. 1990. Micro- dorids. Synopses of the British fauna (new
meiofaunal community structure and nema- series), No. 38. E. J. Brill/W. Backhuys,
tode diversity in a lagoonal ecosystem (Fan- Leiden, The Netherlands, 502 pp.
gataufa, Eastern Tuamotu Archipelago). Raes, M., M. De Troch, S. G. M. Ndaro, A.
P.S.Z.N.I Marine Ecology 11:173–189. Muthumbi, K. Guilini, & A. Vanreusel. 2007.
Gourbault, N., & M. Vincx. 1990. Two new species of The structuring role of microhabitat type in
brood protecting Desmodoridae (Nematoda) coral degradation zones: a case study with
from Guadeloupe. Nematologica 36:131–143. marine nematodes from Kenya and Zanzibar.
Heip, C., M. Vincx, & G. Vranken. 1985. The ecology Coral Reefs 26:113–126.
of marine nematodes. Oceanography and Renaud-Mornant, J., & N. Gourbault. 1984. Pre-
Marine Biology Annual Review 23:399–489. mières prospections meiofaunistiques en Gua-
Hentschel, U., E. C. Berger, M. Bright, H. Felbeck, deloupe. Hydrobiologia 118:113–118.
& J. A. Ott. 1999. Metabolism of nitrogen and Riemann, F., F. Thiermann, & L. Bock. 2003.
sulfur in ectosymbiotic bacteria of marine Leptonemella species (Desmodoridae, Stilbo-
nematodes (Nematoda, Stilbonematinae). nematinae), benthic marine nematodes with
Marine Ecology Progress Series 183:149–158. ectosymbiotic bacteria, from littoral sand of
Ingole, B., R. Singh, S. Sautya, I. Dovgal, & T. the North Sea island of Sylt: taxonomy and
Chatterjee. 2010. Report of epibiont Thecaci- ecological aspects. Helgoland Marine Re-
neta calix (Ciliophora: Suctorea) on deep-sea
search 57:118–131.
Desmodora (Nematoda) from the Andaman
Seinhorst, J. W. 1959. A rapid method for the
Sea, Indian Ocean. Marine Biodiversity Re-
transfer of nematodes from fixative to anhy-
cords 3:1–3.
drous glycerin. Nematologica 4:67–69.
Kotta, J., & G. Boucher. 2001. Interregional
Semprucci, F., & M. Balsamo. 2012. Key role of
variation of free-living nematode assemblages
free-living nematodes in the marine ecosys-
in tropical coral sands. Cahiers de Biologie
tem. Pp. 109–134 in F. Boeri & J. A. Chung,
Marine 42:315–326.
eds., Nematodes: morphology, functions and
Lasagna, R., G. Albertelli, P. Colantoni, C. Morri, &
C. N. Bianchi. 2010. Ecological stages of management strategies. Animal Science, Is-
Maldivian reefs after the coral mass mortality sues and Professions Series, Nova Science
of 1998. Facies 56:1–11. Publishers, Inc., Hauppauge, New York.
Lorenzen, S. 1981. Entwurf eines phylogenetischen Semprucci, F., P. Colantoni, G. Baldelli, M. Rocchi,
Systems der freilebenden Nematoden. Veröf- & M. Balsamo. 2010. The distribution of
fentlichungen des Instituts für Meeresfor- meiofauna on back-reef sandy platforms in
schung in Bremerhaven 7:1–472. the Maldives (Indian Ocean). Marine Ecology
Montiel, A., E. Quiroga, & D. Gerdes. 2011. 31:592–607.
Diversity and spatial distribution patterns of Semprucci, F., P. Colantoni, G. Baldelli, C. Sbrocca,
polychaete assemblages in the Paso Ancho, M. Rocchi, & M. Balsamo. 2013. Meiofauna
Straits of Magellan Chile. Continental Shelf associated with coral sediments in the Maldi-
Research 31:304–314. vian subtidal habitats (Indian Ocean). Marine
Ndaro, S. G. M., & E. Ólafsson. 1999. Soft-bottom Biodiversity 43:189–198.
fauna with emphasis on nematode assemblage Semprucci, F., P. Colantoni, C. Sbrocca, G. Baldelli,
structure in a tropical intertidal lagoon in M. Rocchi, & M. Balsamo. 2011. Meiofauna
Zanzibar, eastern Africa: I. spatial variability. in sandy back-reef platforms differently ex-
Hydrobiologia 405:133–148. posed to the monsoons in the Maldives
Ott, J. A. 1972. Twelve new species of nematodes (Indian Ocean). Journal of Marine Systems
from an intertidal sandflat in North Carolina. 87:208–215.
Internationale Revue der Gesamten Hydro- Tchesunov, A. V. 2013. Marine free-living nema-
biologie und Hydrographie 57:463–496. todes of the subfamily Stilbonematinae (Nem-
Ott, J., M. Bright, & S. Bulgheresi. 2004. Marine atoda, Desmodoridae): taxonomic review with
microbial thiotrophic ectosymbioses. Ocean- descriptions of a few species from the Nha
ography and Marine Biology: An Annual Trang Bay, Central Vietnam. Meiofauna
Review 42:95–118. Marina 20:71–94.
46 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Tietjen, J. H. 1991. Ecology of free-living nematodes new desmodorids (Nematoda) from hydro-
from the continental shelf of the Central thermal vents of the Pacific. Journal of the
Great Barrier Reef Province. Estuarine, Marine Biological Association of the United
Coastal and Shelf Science 32:421–438. Kingdom 78:75–112.
Tkachenko, K. S. 2012. The northernmost coral Warwick, R. M., H. M. Platt, & P. J. Somerfield.
frontier of the Maldives: the coral reefs of 1998. Free-living marine nematodes. Part III:
Ihavandippolu Atoll under long-term envi- monhysterids. Synopses of the British fauna
ronmental change. Marine Environmental
(new series), No. 53. Field Studies Council,
Research 82:40–48.
Shrewsbury, United Kingdom, 296 pp.
Verschelde, D., & M. Vincx. 1996. Four new species
of the family Desmodoridae (Nematoda, Wilkinson, C., O. Linden, H. Cesar, G. Hodgson, J.
Desmodoroidea) from Kenya. Zoologica Rubens, & A. E. Strong. 1999. Ecological and
Scripta 25:1–20. socioeconomic impacts of 1998 coral mortality
Verschelde, D., N. Gourbault, & M. Vincx. 1998. in the Indian Ocean: an ENSO impact and a
Revision of Desmodora with descriptions of warning of future change? Ambio 28:188–196.