BIOGEOGRAPHIC ATLAS OF THE SOUTHERN OCEAN Census of Antarctic Marine Life

Census of Antarctic Marine Life
SCAR-Marine Biodiversity Information Network
BIOGEOGRAPHIC ATLAS
OF THE SOUTHERN OCEAN
 CHAPTER 5.3. ANTARCTIC FREE-LIVING MARINE NEMATODES.
Ingels J., Hauquier F., Raes M., Vanreusel A., 2014.
In: De Broyer C., Koubbi P., Griffiths H.J., Raymond B., Udekem d’Acoz C. d’, et al. (eds.). Biogeographic Atlas of the
Southern Ocean. Scientific Committee on Antarctic Research, Cambridge, pp. 83-87.
EDITED BY:
Claude DE BROYER & Philippe KOUBBI (chief editors)
with Huw GRIFFITHS, Ben RAYMOND, Cédric d’UDEKEM
d’ACOZ, Anton VAN DE PUTTE, Bruno DANIS, Bruno DAVID,
Susie GRANT, Julian GUTT, Christoph HELD, Graham HOSIE,
Falk HUETTMANN, Alexandra POST & Yan ROPERT-COUDERT
SCIENTIFIC COMMITTEE ON ANTARCTIC RESEARCH
THE BIOGEOGRAPHIC ATLAS OF THE SOUTHERN OCEAN
The “Biogeographic Atlas of the Southern Ocean” is a legacy of the International Polar Year 2007-2009 (www.ipy.org) and of the Census of Marine Life 2000-2010
(www.coml.org), contributed by the Census of Antarctic Marine Life (www.caml.aq) and the SCAR Marine Biodiversity Information Network (www.scarmarbin.be;
www.biodiversity.aq).
The “Biogeographic Atlas” is a contribution to the SCAR programmes Ant-ECO (State of the Antarctic Ecosystem) and AnT-ERA (Antarctic Thresholds- Ecosystem Resilience and Adaptation) (www.scar.org/science-themes/ecosystems).
Edited by:
Claude De Broyer (Royal Belgian Institute of Natural Sciences, Brussels)
Philippe Koubbi (Université Pierre et Marie Curie, Paris)
Huw Griffiths (British Antarctic Survey, Cambridge)
Ben Raymond (Australian Antarctic Division, Hobart)
Cédric d’Udekem d’Acoz (Royal Belgian Institute of Natural Sciences, Brussels)
Anton Van de Putte (Royal Belgian Institute of Natural Sciences, Brussels)
Bruno Danis (Université Libre de Bruxelles, Brussels)
Bruno David (Université de Bourgogne, Dijon)
Susie Grant (British Antarctic Survey, Cambridge)
Julian Gutt (Alfred Wegener Institute, Helmoltz Centre for Polar and Marine Research, Bremerhaven)
Christoph Held (Alfred Wegener Institute, Helmoltz Centre for Polar and Marine Research, Bremerhaven)
Graham Hosie (Australian Antarctic Division, Hobart)
Falk Huettmann (University of Alaska, Fairbanks)
Alix Post (Geoscience Australia, Canberra)
Yan Ropert-Coudert (Institut Pluridisciplinaire Hubert Currien, Strasbourg)
Published by:
The Scientific Committee on Antarctic Research, Scott Polar Research Institute, Lensfield Road, Cambridge, CB2 1ER, United Kingdom (www.scar.org).
Publication funded by:
- The Census of Antarctic Marine Life (Albert P. Sloan Foundation, New York)
- The TOTAL Foundation, Paris.
The “Biogeographic Atlas of the Southern Ocean” shared the Cosmos Prize awarded to the Census of Marine Life by the International Osaka Expo’90 Commemorative Foundation, Tokyo, Japan.
Publication supported by:
-
The Belgian Science Policy (Belspo), through the Belgian Scientific Research Programme on the Antarctic and the “biodiversity.aq” network (SCAR-MarBIN/ANTABIF)
The Royal Belgian Institute of Natural Sciences (RBINS), Brussels, Belgium
The British Antarctic Survey (BAS), Cambridge, United Kingdom
The Université Pierre et Marie Curie (UPMC), Paris, France
The Australian Antarctic Division, Hobart, Australia
The Scientific Steering Committee of CAML, Michael Stoddart (CAML Administrator) and Victoria Wadley (CAML Project Manager)
Mapping coordination and design: Huw Griffiths (BAS, Cambridge) & Anton Van de Putte (RBINS, Brussels)
Editorial assistance: Henri Robert, Xavier Loréa, Charlotte Havermans, Nicole Moortgat (RBINS, Brussels)
Printed by: Altitude Design, Rue Saint Josse, 15, B-1210, Belgium (www.altitude-design.be)
Lay out: Sigrid Camus & Amélie Blaton (Altitude Design, Brussels).
Cover design: Amélie Blaton (Altitude Design, Brussels) and the Editorial Team.
Cover pictures: amphipod crustacean (Epimeria rubrieques De Broyer & Klages, 1991), image © T. Riehl, University of Hamburg; krill (Euphausia superba
Dana, 1850), image © V. Siegel, Institute of Sea Fisheries, Hamburg; fish (Chaenocephalus sp.), image © C. d’Udekem d’Acoz, RBINS; emperor penguin
(Aptenodytes forsteri G.R. Gray, 1844), image © C. d’Udekem d’Acoz, RBINS; Humpback whale (Megaptera novaeangliae (Borowski, 1781)), image © L. Kindermann, AWI.
Online dynamic version :
A dynamic online version of the Biogeographic Atlas is available on the SCAR-MarBIN / AntaBIF portal : atlas.biodiversity.aq.
Recommended citation:
For the volume:
De Broyer C., Koubbi P., Griffiths H.J., Raymond B., Udekem d’Acoz C. d’, Van de Putte A.P., Danis B., David B., Grant S., Gutt J., Held C., Hosie G., Huettmann
F., Post A., Ropert-Coudert Y. (eds.), 2014. Biogeographic Atlas of the Southern Ocean. Scientific Committee on Antarctic Research, Cambridge, XII + 498 pp.
For individual chapter:
(e.g.) Crame A., 2014. Chapter 3.1. Evolutionary Setting. In: De Broyer C., Koubbi P., Griffiths H.J., Raymond B., Udekem d’Acoz C. d’, et al. (eds.).
Biogeographic Atlas of the Southern Ocean. Scientific Committee on Antarctic Research, Cambridge, pp. xx-yy.
ISBN: 978-0-948277-28-3.
This publication is licensed under a Creative Commons Attribution-NonCommercial 4.0 International License
2
Biogeographic Atlas of the Southern Ocean

Meiobenthos : Nematoda
5.3. Antarctic free-living marine nematodes
Jeroen Ingels1, Freija Hauquier2, Maarten Raes2 & Ann Vanreusel2
1
2
Plymouth Marine Laboratory, Plymouth, UK
Marine Biology Research group, Biology Department, Ghent University, Gent, Belgium
1. Introduction
Nematodes or roundworms are the most pervasive metazoans on the planet
(i.e. 80% of all living terrestrial metazoans and >90% in deep-sea ecosystems;
Danovaro 2012) and have successfully exploited nearly every imaginable
habitat. They can be found from high mountains down to the deepest depths
in the oceans; they have even been recovered from the deep subsurface biosphere at 3.5 km depth where they are able to exploit the available resources
(Borgonie et al. 2011); and can live as parasites in many organisms. Nematodes come in different sizes, from minute worms in sediments a few tenths
of millimeters in length, to large parasitical forms which may be a few meters
long. Most nematodes, however, don’t exceed a few millimeters in length.
About 16,000 nematode species have been described as parasites causing
numerous diseases in vertebrate organisms and plants, and these have been
studied intensively in the context of socio-economical and medical interests
(e.g. Chan 1997, Chitwood 2003). The free-living nematodes, on the other
hand, are perhaps less known despite their ubiquity and high levels of biodiversity. Free-living nematodes are amongst the most speciose marine benthic
organisms in the world (Snelgrove 1999), with nearly 7000 recognised marine
species and many more undescribed or undiscovered (Appeltans et al. 2012).
Estimates for marine nematode diversity may range from 10,000 to 1,000,000,
depending on the source and how ‘conservative’ or ‘liberal’ the estimate itself
was (Lambshead 1993, Mokievsky & Azovsky 2002, Snelgrove 1999). More
recently, Appeltans and co-authors (2012) showed that about 50,000 species
is a more accurate estimate of total expected nematode diversity, meaning
that nearly 90% of nematode species remains undescribed. Recent investigations indicated that there are 638 valid species that have been recovered
from deep-sea samples worldwide (Miljutin et al. 2010), but as for some other
under-explored habitats, many deep-sea samples contain numerous unknown
species. A study by Xu et al. (2013) showed that no fewer than 155 nematode
species were described in the journal Zootaxa alone in the period 2007–2012,
many of which are marine, and showing that nematode taxonomy is currently
a rather active field of research, although much remains to be described.
vorous or feed on small detrital particles freely available or attached to sediment grains; and even prey on small organisms, including other nematodes
(Heip et al. 1982). The intermediate position they take in marine sediment
food webs identifies them as important links, transferring energy available in
sediments under the form of dissolved organic matter, detritus, microbes and
other organisms to higher trophic levels (Bongers & Ferris 1999). Moreover,
nematodes play an important role in decomposition processes and nutrient
cycling in sediments. They are also known to interact with microbiota and
other metazoan organisms (Heip et al. 1982), and may have an important role
to play in bioturbating the sediments they live in. Moreover, nematode diversity
has been linked to ecosystem functioning which may suggest an important
functional role in sediments (Danovaro et al. 2008).
Given their small size and limited mobility, and the fact that they have a
conservative reproductive method and lack an active dispersal phase in their
life history, we might expect that species should have geographically limited
distributions. Consequently, the species turnover between areas should be
high compared to larger organisms which have a better chance of long-distance dispersal via pelagic larval phases (Lambshead 1993), leading to higher
global species diversity. However, recently, Bik et al. (2010) provided molecular evidence of low endemism, continued shallow-deep water exchanges, and
cosmopolitan species complexes within marine nematodes. Although these
molecular analyses do not necessarily pertain to the species level, posing
the question of the ‘meiofauna paradox‘ is inevitable, i.e. how is it possible
that meiofaunal organisms with limited active dispersal capacities are able to
become cosmopolitan? The most likely answer to this question relies on the
fact that it is their small size that makes them susceptible to entrainment by
currents impinging the sediment surface, causing passive dispersal (Boeckner
et al. 2009). Consequently, transport over larger distances by currents is likely
to be more widespread than thought previously.
2. Methods
2.1. Data collection and geographical scope
The data used for this review on Antarctic nematode species distribution
stems from different sources and was gathered and collated by the authors.
Many nematode species data originated from studies conducted at the Marine
Biology research group of Ghent University and data records available on
the NeMys database (Deprez et al. 2005). In addition, historic literature was
gathered based mainly on species lists in Gerlach & Riemann (1973). Subsequently, original descriptions were studied to obtain geographical locations,
i.e. Antarctic and sub-Antarctic species records were extracted and added
to the database. Taxonomic literature until 2012 was included in the database. For a more complete overview of the data sources used, we refer to the
marked literature sources.
The species distribution maps presented in this Chapter include both true
Antarctic and sub-Antarctic data, with a circum-Antarctic scope. Latitudinal
range of included nematode species extends from roughly 46°S in the Kerguelen Islands region to approximately 78°S in Discovery Inlet.
2.2. Limitations of coverage and taxonomic resolution
Photo 1 Desmodora campbelli (Allgén, 1932), South Georgia (Polarstern, ANT-XXVII/3, stn. 214-4, 255 m). Image © F. Hauquier, University of Ghent.
With 10,000s to 1,000,000s of individuals per square meter of seafloor,
marine free-living nematodes are the most abundant metazoan life form in
marine sediments and often represent 70–90 % of metazoan meiofaunal organisms (organisms ranging 32–1000 µm in size). Despite their ubiquity and
high abundance, we are only beginning to understand the role of nematodes
in benthic communities. They have been identified as key contributors to different ecosystem functions in marine environments. Nematodes are characterised by a wide range of morphological features which can be used to infer
their ecological roles. The size and morphology of their buccal cavities can
serve to identify their feeding strategies (Moens & Vincx 1997). They can feed
on microbiota such as bacteria, cyanobacteria, and algae; they may be fungi-
Unfortunately, the geographical coverage of (sub-)Antarctic data on free-living
nematode species is fragmental and is mostly based on taxonomical works
for which the locations were recovered. This brings the limitation inherent to
undersampling, and causes limited scope for biogeographical interpretation
on the species level. Much more data is available on the genus level, the
focus of many Antarctic nematode studies that address ecological questions.
Whilst such data suffers from lack of species information, it is unlikely that
information on genera occurrences in the Southern Ocean is useful to infer
biogeographical patterns. Nematode genera in general are not limited to particular areas and most common genera are found all over the globe in marine
environments, a phenomenon often referred to as the ‘meiofauna paradox’.
Nematode genera abundances are often thought to be associated with particular environmental conditions and so their occurrence is often studied in an
ecological rather than a taxonomical or biogeographical context. Nevertheless, there are rare genera that seem limited in distribution, but this is often
the result of limited sampling effort. To illustrate, a recent study showed that
a newly described and relatively rare genus, Dystomanema, was recovered
from both the Antarctic and North Atlantic (Bezerra et al. 2013).
3. Biodiversity
Nematodes are widespread in the Antarctic. On land, they are the most diverse and abundant invertebrate phylum (43 species; Wharton 2003). Antarctic nematode species numbers in the marine environment are much higher
than for their terrestrial counterparts, and until recently limited at nearly 400
accepted marine species. In the framework of the present Atlas, a taxonomic
revision was performed of all nematode species records, historical and recent
Biogeographic Atlas of the Southern Ocean
83

Meiobenthos : Nematoda
(until 2012) from marine Antarctic sediments (Appendix 1, at the end of volume). According to the latest literature sources and data records in our dataset 524 species are considered valid (see Map 1 for an overview of number
of species per sector; see data reference list for literature sources). Nematode systematics has been shrouded in uncertainty because of their minute
size (making identification more labor-intensive), taxonomic difficulties, and
the relatively slow advent of molecular studies on nematodes. In addition, a
huge number of nematode species were described in the first half of the 20th
century following early Antarctic expeditions which yielded a large number of
new taxa. The first descriptions of marine nematodes from (sub-) Antarctic
regions date back to 1891, when von Linstow (1891) described a number of
nematode species from South Georgia. Since then and with much disagreement between authors, many poor descriptions of Antarctic marine nematode
species have appeared, resulting in confused taxonomy and contradicting
descriptions. Some descriptive efforts following the Antarctic expeditions resulted in large numbers of species being described in a short space of time,
leading to several synonymies. Scientists such as Cobb, De Man, von Linstow, and Ditlevsen, amongst others, described tens of species from the marine Antarctic and sub-Antarctic regions. Undoubtedly one of the most prolific
nematode taxonomists working on Antarctic samples was Carl Allgén; in his
report on free-living marine nematodes from the Swedish Antarctic Expedition
in 1901–1903 (Allgén 1959), no less than 343 species were described, 200 of
which were new to science. His work strongly influenced later developments
in nematode taxonomy, not merely because he contributed significantly to our
knowledge on the diversity of Antarctic marine nematodes, but also because
the limited descriptive and illustrative material and sometimes doubtful diagnoses that he provided, causing species to be synonymised or considered
“species inquirendae” or “incertae sedis” in many later instances — the result
of inadequate descriptions. After several other collations of species lists, “The
Bremerhaven Checklist of Aquatic Nematodes” provided an exhaustive list of
all hitherto known nematodes worldwide (Gerlach & Riemann 1973). Since
then, several studies have added new species to the list and several genera
and family reviews have been conducted. The dataset presented here is an
updated account of valid Antarctic and sub-Antarctic marine free-living nematodes (“sp. inq.” and “sp. inc. sedis”. are excluded from this list). This data is
partially based on the NeMys database (Deprez et al. 2005) and has been
updated with geographical location data (coordinates and water depth, when
available) and taxonomical information contained within the original descriptive and ecological literature. The data is available through SCAR-MarBIN
(http://www.scarmarbin.be) and further details on the methods can be found in
the previous section “Methods”.
4. Biogeography of Antarctic and sub-Antarctic nematodes
Because most observations of nematode species have been conducted as
part of taxonomic works, geographical information on Antarctic marine nematodes is limited to the occurrences reported in species descriptions. In addition, studies focusing on the ecology of nematodes reported many valuable
distribution data, but are generally restricted to the genus level – species are
often not considered. Notwithstanding the over 2200 records of nematode
species in the (sub-)Antarctic, there are only a handful of studies which have
reported on the biogeography of selected groups of marine free-living nematodes.
Vermeeren et al. (2004) discussed the distribution of species belonging
to the genus Dichromadora, a genus that occurs regularly in the Southern
Ocean over a wide range of water depths. The authors compared samples
from the Indian, Pacific, Arctic and Atlantic continental margins for Dichromadora occurrences and noted the absence of this genus in the Indian and Pacific Oceans. A number of Dichromadora species were present in the Arctic and
Atlantic (two and three species, respectively), but the Southern Ocean samples contained eight species in addition to one previously described by Timm
(1978) (Maps 2–6). A high degree of endemism was observed, with seven of
the eight species only occurring in the Southern Ocean. To be noted, however,
is the fact that seven of the eight Southern Ocean species discovered in the
study, as well as all Arctic and Atlantic species, were new to science. This
suggests that the deep-sea nematode fauna in the Southern Ocean and possibly all other oceans are undersampled. The Dichromadora species exhibited
either a very limited distribution (e.g. D. antarctica, D. quadripapillata; Maps
2–3) or they appeared across various locations in the Southern Ocean (e.g. D.
weddellensis, D. southernis, D. parva, D. polarsternis, D. polaris; Maps 2–6),
the latter indicating that nematode species may have wide ranges over regional scales in the deep sea. The species studied did not show any bathymetric
limitations, at least for the depth range studied (1000–2000 m water depth).
The results of Vermeeren et al. (2004) were complemented with results from
Ingels et al. (2006) in that two Dichromadora species were recognised from
the Scotia Arc at about 300 m water depth (D. polaris; Map 6), reinforcing the
hypothesis that bathymetry per se does not seem to limit nematode distributions in the Southern Ocean, at least for the genus Dichromadora. This is perhaps not surprising considering the genus is also common in coastal areas,
but species comparisons are needed with shallow-water samples to confirm
this. A study conducted by De Mesel et al. (2006), on the other hand, showed
a considerable degree of species turnover for the 55 putative Acantholaimus
84
species identified in different parts of the Southern Ocean. In addition to high
species turnover as a result of the restricted species distributions and their
rarity, the number of congeneric species in assemblages was high, leading to
high local and regional biodiversity levels for this genus. Fourteen species had
a distribution extending from the shelf to the lower slope, pointing to a strong
degree of eurybathy too. The occurrence of the otherwise typical deep-sea
genus Acantholaimus in high densities and diversity on the continental shelf is
a unique feature of the Southern Ocean. (De Mesel et al. 2006).
Ingels et al. (2006), in a biodiversity and biogeographical nematode
study, focussed mainly on the genera Desmodora and Desmodorella. They
found that two species, Desmodora campbelli and Desmodorella aff. balteata
had wide geographical distributions in the Southern Ocean, while the other
eight species had very limited distributions (see Maps 7–8). In contrast to
Vermeeren et al. (2004), however, Ingels et al. (2006) found bathymetric restrictions for several species, indicating that shallow-water island chains such
as the Scotia Arc may provide the means for species that are depth-restricted
to disperse over larger geographical areas. Although the Scotia Arc islands
are surrounded by deep ocean, this does not necessarily prevent strong water
column currents to transport small animals between similar depth ranges of
the islands’ margins. Preliminary molecular results reported in the same study
suggest that certain nematode species exhibit extremely slow evolution with
conservation of certain species-specific genes or that hydrodynamic processes
and sediment disturbance may be behind the high rates of genetic exchange
observed between species populations of distant geographic locations (Ingels
et al. 2006). Whatever the case, some species seem geographically or bathymetrically restricted, whilst others have circum-Antarctic or eurybathic distributions. If we appreciate genus-level differences along bathymetric gradients in
the Antarctic, there are indications of distinct communities at different depth
zones but genera are not restricted bathymetrically (Vanhove et al. 2004). This
means that perceived bathymetric gradients are caused by changes in relative
abundance of genera rather than genus composition. Noteworthy in this context is the fact of sampling intensity; while presence observations can confirm
biogeographical distributions we have to be cautious on how to interpret the
absence of species — absence in a sample does not mean that the species is
not present, it may merely mean that the area is undersampled.
Fonseca et al. (2006) studied the occurrence of species of the deep-sea
genus Molgolaimus (Map 9) — a particularly species-rich genus in the deep
sea — in different oceans and concluded that geographical rather than environmental clustering of morphologically similar species does not support the
idea of a common origin of deep-sea species. In addition, the genus Molgolaimus seems to have many species with restricted distributions in the Southern
Ocean (compared to the western Indian Ocean for instance), making Molgolaimus species suitable for distinguishing between biogeographical provinces
in the Southern Ocean (Fonseca et al. 2007). Here, the authors propose that
evolutionary history may have shaped nematode species composition at the
ocean scale, while at local and regional scales ecological processes are promoting species co-existence and speciation (e.g. higher number of Molgolaimus species co-occurring at Peninsula tip and eastern Weddell Sea; see
Map 9). Whether this is also the case for other nematode genera needs to be
verified. Moreover, definite conclusions can’t be drawn because of the lack of
insight into the true presence or absence of species in the limited quantity of
samples that are available.
Undoubtedly, the abiotic environment has a significant impact on the
community structure of shallow and deep-sea nematode communities. Various environmental factors have been shown to influence benthic nematode
community structure although studies rarely involve species level information. Disturbance by physical, biological or biogeochemical processes, sediment grain size, food quantity and quality, and other trophic conditions have
been evoked as regulating community structure by creating conditions that
are more favourable for some nematode genera whilst unfavourable for others. Food input seems to be a major determinant in the Antarctic along with
the seasonality of its availability (Sebastian et al. 2007, Vanhove et al. 1998,
Vanhove et al. 1999, Vanhove et al. 2000). However, despite several ecological studies positing the link between environment and nematode community
structure, the role of ecology in biogeographical ranges remains unresolved.
5. Conclusion
In this review, we give an overview of the scarce information available on
nematode species distributions in the Southern Ocean. Due to taxonomic difficulties and the general lack of species information, biogeography of Antarctic
and sub-Antarctic nematodes remains rather elusive. Based on the information we could find and verify, it seems that, indeed, some species might be
limited to certain regions or depths in the Southern Ocean, while others may
have circum-Antarctic and eurybathic distributions. Faunal connections between the southernmost South America and the Antarctic Peninsula are present for some taxa but remain to be verified for others. More taxonomic studies
with distribution data at species level may help to overcome the problems of
lack of knowledge and undersampling. The advent of molecular techniques
is definitely something to welcome in the search of biogeographic patterns in
Southern Ocean nematode diversity.
Map 1
Nematode species count per sector
1 - 16
17 - 30
31 - 45
46 - 59
60 - 74
75 - 89
90 - 103
104 - 118
119 - 132
133 - 147
Map 2
zzDichromadora antarctica
zzDichromadora dissipata
zzDichromadora weddellensis
Map 3
zzDichromadora quadripapillata
zzDichromadora southernis
Map 4
zzDichromadora parva
Map 5
zzDichromadora polarsternis
Map 6
zzDichromadora polaris
Meiobenthos: Nematoda Map 1 Antarctic and sub-Antarctic nematode species counts per sector, based on latest valid taxonomy (until 2012); all depth ranges are included (0–4000 m) and
the latitudinal range extends from roughly 46°S (Crozet Islands) to 78°S (Discovery Inlet). Maps 3–5. Species distribution of the genus Dichromadora (bathymetric range 0–2285 m). Map 2.
D. antarctica Timm, 1978 (data: Cobb 1914), D. dissipata Wieser, 1954 (data: Guotong 1999) and D. weddellensis Vermeeren et al., 2004 (data: Vermeeren et al. 2004). Map 3. D. southernis
Vermeeren et al., 2004 and D. quadripapillata Vermeeren et al., 2004 (data: Vermeeren et al. 2004). Map 4. D. parva Vermeeren et al., 2004 (data: Vermeeren et al. 2004). Map 5. D. polarsternis Vermeeren et al., 2004 (data: Vermeeren et al. 2004). Map 6. D. polaris Vermeeren et al., 2004 (data: Vermeeren et al. 2004; Ingels et al. 2006).
Biogeographic Atlas of the Southern Ocean
85

Meiobenthos : Nematoda
Map 7
zzDesmodora campbelli
zzDesmodora microchaeta
zzDesmodora minuta
zzDesmodora scaldensis
Map 8
zzDesmodorella abyssorum
zzDesmodorella aff. balteata
zzDesmodorella tenuispiculum
Map 9
Number of Molgolaimus species per sector
1
2
3
4
5
6
7
8
9
Meiobenthos: Nematoda Maps 7–9 Map 7. Species distribution of the genus Desmodora (data: Allgén 1959; Ingels et al. 2006; bathymetric range: 0–502 m). Map 8. Species
distribution of the genus Desmodorella (data: Allgén 1959; Ingels et al. 2006; bathymetric range: 0–681 m). Map 4. Species distribution, expressed as number of species per sector,
of the genus Molgolaimus (data: Ditlevsen 1921; Guotong 1999; Fonseca et al. 2006; bathymetric range 79–4000 m).
Acknowledgments
Dr. Huw Griffiths (BAS, Cambridge) and Dr. Anton Van de Putte (RBINS, Brussels) prepared the maps. This is CAML contribution # 101.
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
Appendix 1 at the end of volume
Biogeographic Atlas of the Southern Ocean
87

Appendix 1 : Nematoda
Appendix 1: Nematoda (Chap. 5.3)
Table 1 Phylogenetic species list of (sub-)Antarctic marine free-living Nematoda
PHYLUM NEMATODA Potts, 1932
Class ENOPLEA Inglis, 1983
Subclass ENOPLIA Pearse, 1942
Order Enoplida Filipjev, 1929
Suborder Enoplina Chitwood & Chitwood, 1937
Family Enoplidae Dujardin, 1845
Enoplus heardensis Mawson, 1958
Enoplus michaelseni Linstow, 1896
Enoplus micrognathus Allgén, 1947
Enoplus paralittoralis Wieser, 1953
Family Thoracostomopsidae Filipjev, 1927
Enoplolaimus acanthospiculum Allgén, 1959
Enoplolaimus arcospiculum Allgén, 1959
Enoplolaimus falklandiae Allgén, 1959
Enoplolaimus filiformis (Allgén, 1935)
Enoplolaimus niger Allgén, 1959
Enoplolaimus notopropinquus Allgén, 1959
Enoplolaimus opacus Allgén, 1959
Enoplolaimus propinquus de Man, 1922
Enoplolaimus vulgaris (de Man, 1893)
Epacanthion brevispiculosum Mawson, 1958
Epacanthion brevispiculum Mawson, 1956
Epacanthion filicaudatum Mawson, 1956
Fenestrolaimus antarcticus Mawson, 1956
Mesacanthion brachycolle Allgén, 1959
Mesacanthion infantile (Ditlevsen, 1930)
Mesacanthion kerguelense Mawson, 1958
Mesacanthion pacificum Allgén, 1947
Mesacanthion virile (Ditlevsen, 1930)
Mesacanthoides caputmedusae (Ditlevsen, 1918)
Mesacanthoides latignathus (Ditlevsen, 1918)
Metenoploides alatus Wieser, 1953
Oxyonchus australis (de Man, 1904) Mawson, 1956
Oxyonchus brachysetosus Allgén, 1959
Oxyonchus crassicollis Allgén, 1959
Oxyonchus dentatus (Ditlevsen, 1918) Filipjev, 1927
Oxyonchus macrodon Allgén, 1959
Oxyonchus notodentatus Allgén, 1959
Oxyonchus parastateni Allgén, 1959
Oxyonchus stateni (Allgén, 1930)
Oxyonchus subantarcticus Mawson, 1958
Paramesacanthion allgeni Mawson, 1958
Paramesacanthion estridium Wieser, 1953
Paramesacanthion oxycephalum (Ditlevsen, 1926)
Paramesacanthion tricuspis (Schuurmans Stekhoven, 1950)
Family Anoplostomatidae Gerlach & Riemann, 1974
Anoplostoma campbelli Allgén, 1932
Anoplostoma tenuisetum Allgén, 1959
Chaetonema amphora Wieser, 1953
Chaetonema steineri (Filipjev, 1927)
Family Phanodermatidae Filipjev, 1927
Klugea longiseta Mawson, 1956
Klugea truncata Mawson, 1956
Micoletzkyia anomala Wieser, 1953
Micoletzkyia austrogeorgiae Allgén, 1954
Micoletzkyia falklandiae Allgén, 1954
Micoletzkyia nudicapitata Allgén, 1959
Phanoderma banzare Mawson, 1956
Phanoderma campbelli Allgén, 1927
Phanoderma cocksi Bastian, 1865
Phanoderma laticolle (Marion, 1870)
Phanoderma paracampbelli Allgén, 1958
Phanoderma parasiticum Ditlevsen, 1926
Phanoderma speculum Schuurmans Stekhoven & Mawson, 1955
Phanoderma tuberculatum (Eberth, 1863) Bastian, 1865
Phanoderma wieseri Mawson, 1956
Phanodermopsis ingrami Mawson, 1958
Family Anticomidae Filipjev, 1918
Anticoma acuminata (Eberth, 1863) Stekhoven, 1950
Anticoma allgeni Platonova, 1968
Anticoma campbelli Allgén, 1932
Anticoma columba Wieser, 1953
Anticoma curticauda Platonova, 1968
Anticoma filicauda Mawson, 1956
Anticoma graciliceps Platonova, 1968
Anticoma kerguelensis Mawson, 1958
Anticoma longissima Allgén, 1958
Anticoma major Mawson, 1956
Anticoma pellucida Bastian, 1865
Anticoma pushkini Platonova, Belogurov & Sheenko, 1979
Anticoma subsimilis Cobb, 1914
Anticoma tenuis Allgén, 1930
Anticoma trichura Cobb, 1898
Anticoma wieseri Mawson, 1958
Anticomopsis typica Micoletzky, 1930
Antopus serialis (Baylis, 1916)
Paranticoma antarctica Mawson, 1956
Paranticoma odhneri Allgén, 1959
Paranticoma tubuliphora Wieser, 1953
Suborder Trefusiina Siddiqi, 1983
Family Simpliconematidae Blome & Schrage, 1985
Simpliconema aenigmatodes Blome & Schrage, 1985
Family Trefusiidae Gerlach, 1966
Trefusia axonolaimoides Allgén, 1953
Family Xenellidae De Coninck, 1965
Xennella filicaudata (Allgén, 1954)
Suborder Oncholaimina De Coninck, 1965
Family Oncholaimidae Filipjev, 1916
Adoncholaimus austrogeorgiae Allgén, 1959
478
Adoncholaimus crassicaudus Wieser, 1953
Adoncholaimus falklandiae Allgén, 1959
Adoncholaimus thalassophygas (de Man, 1876)
Curvolaimus decipiens Wieser, 1953
Metaparoncholaimus macrouraios Mawson, 1958
Metoncholaimoides squalus Wieser, 1953
Metoncholaimus antarcticus (Linstow, 1896)
Oncholaimellus carlbergi Allgén, 1947
Oncholaimus dujardinii de Man, 1876
Oncholaimus leptos Mawson, 1958
Oncholaimus longissimus Allgén, 1959
Oncholaimus notolangrunensis Allgén, 1959
Oncholaimus notoviridis Allgén, 1958
Oncholaimus notoxyuris Allgén, 1959
Oncholaimus paradujardini Allgén, 1959
Oncholaimus paraegypticus Mawson, 1956
Oncholaimus paralangrunensis (Allgén, 1947)
Oncholaimus paredron Mawson, 1958
Oncholaimus rotundicaudatus Allgén, 1959
Oncholaimus thysanouraios (Mawson, 1958)
Oncholaimus viridis (Bastian, 1865)
Pelagonema longicaudum (Allgén, 1953)
Pelagonema obtusicauda Filipjev, 1918
Pelagonema tenue (Kreis, 1928)
Pontonema cobbi Mawson, 1956
Pontonema leidyi Mawson, 1956
Pontonema propinquum (Allgén, 1930)
Pontonema serratodentatum Mawson, 1956
Viscosia antarctica Allgén, 1959
Viscosia brachydonta Allgén, 1959
Viscosia brevicaudata Mawson, 1958
Viscosia brevilaima Allgén, 1959
Viscosia carnleyensis (Ditlevsen, 1921)
Viscosia cryptodentata Allgén, 1959
Viscosia falklandiae Allgén, 1959
Viscosia glabra (Bastian, 1865) de Man, 1890
Viscosia graham Allgén, 1959
Viscosia langrunensis (de Man, 1890)
Viscosia parafalklandiae Allgén, 1959
Viscosia parapellucida (Cobb, 1898)
Viscosia propinqua Allgén, 1959
Viscosia similis Allgén, 1959
Viscosia subantarctica Allgén, 1959
Viscosia tenuilaima Allgén, 1959
Viscosia tenuissima Allgén, 1959
Viscosia viscosa (Bastian, 1865) de Man, 1890
Viscosia wieseri Mawson, 1958
Family Enchelidiidae Filipjev, 1918
Calyptronema axonolaimoides Allgén, 1959
Calyptronema mawsoni Mawson, 1958
Calyptronema retrocellatum (Wieser, 1953)
Catalaimus maxweberi (de Man, 1922)
Ditlevsenella tertia Wieser, 1953
Enchelidium filicolle Allgén, 1959
Eurystomina fenestella Wieser, 1953
Eurystomina filicaudatum (Allgén, 1959)
Eurystomina ornata (Eberth, 1863)
Eurystomina stenolaima (Ditlevsen, 1930)
Eurystomina tenuicaudata Allgén, 1932
Ledovitia fallae Mawson, 1958
Polygastrophora hexabulba (Filipjev, 1918)
Polygastrophora octobulba Micoletzky, 1930
Symplocostoma tenuicolle (Eberth, 1863)
Suborder Ironina Siddiqi, 1983
Family Ironidae de Man, 1876
Dolicholaimus marioni (de Man, 1888)
Syringolaimus striatocaudatus de Man, 1888
Thalassironus bipartitus Wieser, 1953
Family Leptosomatidae Filipjev, 1916
Deontostoma antarcticum (Linstow, 1892)
Deontostoma arcticum (Ssaweljev, 1912)
Deontostoma aucklandiae (Ditlevsen, 1921)
Deontostoma demani (Mawson, 1956)
Deontostoma timmerchioi Hope, 1974
Leptosomatides antarcticus Mawson, 1956
Leptosomatides conisetosus Schuurmans Stekhoven & Mawson, 1955
Leptosomatum arcticum Filipjev, 1916
Leptosomatum clavatum Platonova, 1958
Leptosomatum crassicutis Platonova, 1958
Leptosomatum gracile Bastian, 1865
Leptosomatum kerguelense Platonova, 1958
Leptosomatum sabangense Steiner, 1915
Paraleptosomatides elongatus Mawson, 1956
Paraleptosomatides spiralis Mawson, 1956
Platycomopsis dimorphica Mawson, 1956
Platycomopsis paracobbi Mawson, 1956
Pseudocella brachychaites Mawson, 1958
Pseudocella elegans (Ditlevsen, 1926)
Pseudocella panamaensis (Allgén, 1947)
Pseudocella polychaites (Mawson, 1958)
Pseudocella tabarini (Inglis, 1958)
Pseudocella trichodes (Leuckart, 1849)
Synonchus fasciculatus Cobb, 1893
Thoracostoma angustifissulatum Mawson, 1956
Thoracostoma anocellatum Schuurmans Stekhoven & Mawson, 1954
Thoracostoma arcticum Ssaveljev, 1912
Thoracostoma campbelli Ditlevsen, 1921
Thoracostoma chilense (Steiner, 1921)
Thoracostoma coronatum (Eberth, 1863) Marion, 1870
Thoracostoma falklandiae Allgén, 1959
Thoracostoma microfenestratum Allgén, 1959
Thoracostoma papillosum Ditlevsen, 1921
Thoracostoma parasetosum Mawson, 1958
Thoracostoma schizoepistylium Mawson, 1958
Thoracostoma setosum (Linstow, 1896)
Thoracostoma unifenestratum Allgén, 1959
Family Oxystominidae Chitwood, 1935
Halalaimus brachyaulax Mawson, 1958
Halalaimus ciliocaudatus Allgén, 1932
Halalaimus comatus Wieser, 1953
Halalaimus diacros Mawson, 1958
Halalaimus filicaudatus Allgén, 1959
Halalaimus fletcheri Mawson, 1958
Halalaimus gracilis de Man, 1888
Halalaimus longicollis Allgén, 1932
Halalaimus macquariensis Mawson, 1958
Halalaimus marri Mawson, 1958
Nemanema brachyure Allgén, 1959
Nemanema campbelli Allgén, 1932
Nemanema cylindraticaudatum de Man, 1922
Nemanema obtusicauda (Allgén, 1959)
Oxystomina antarctica Mawson, 1956
Oxystomina elongata Bütschli, 1874
Oxystomina filicaudata Allgén, 1959
Oxystomina greenpatchi Allgén, 1959
Oxystomina mirabilis Allgén, 1959
Oxystomina oxycaudata (Ditlevsen, 1926)
Oxystomina pulchella Vitiello, 1970
Oxystomina tenuicollis Allgén, 1959
Oxystomina vespertilio Wieser, 1953
Thalassoalaimus spissus (Allgén, 1932)
Suborder Tripyloidina De Coninck, 1965
Family Tripyloididae Filipjev, 1918
Bathylaimus australis Cobb, 1894
Bathylaimus austrogeorgiae Allgén, 1959
Paratripyloides viviparus (Cobb, 1930) Wieser, 1956
Order Triplonchida Cobb, 1920
Suborder Tobrilina Tsalolikhin, 1976
Family Rhabdodemaniidae Filipjev, 1934
Rhabdodemania calycolaimus Schuurmans Stekhoven & Mawson, 1955
Rhabdodemania minor (Southern, 1914)
Family Pandolaimidae Belogurov, 1980
Pandolaimus latilaimus (Allgén, 1929)
Subclass DORYLAIMIA Inglis, 1983
Order Mononchida Jairajpuri, 1969
Suborder Mononchina Kirjanova & Krall, 1969
Family Mononchidae Chitwood, 1937
Mononchus gerlachei (de Man, 1904)
Class CHROMADOREA
Subclass CHROMADORIA
Order Chromadorida Chitwood, 1933
Suborder Chromadorina Filipjev, 1929
Family Chromadoridae Filipjev, 1917
Acantholaimus quintus Gerlach, Schrage & Riemann, 1979
Actinonema longicaudatum (Steiner, 1918)
Actinonema pachydermatum Cobb, 1920
Atrochromadora microlaima (de Man, 1889)
Atrochromadora parva (de Man, 1893) Wieser, 1954
Chromadora nudicapitata (Bastian, 1865)
Chromadorella cobbiana Johnston, 1938
Chromadorella filiformis (Bastian, 1865)
Chromadorita brachypharynx (Allgén, 1932)
Chromadorita ceratoserolis Lorenzen, 1986
Chromadorita gracilis (Filipjev, 1922)
Chromadorita minor (Allgén, 1927) Wieser, 1954
Chromadorita mucrodonta (Steiner, 1916)
Chromadorita pharetra Ott, 1972
Dichromadora antarctica (Cobb, 1914) Timm, 1978
Dichromadora dissipata Wieser, 1954
Dichromadora parva Vermeeren, Vanreusel & Vanhove, 2004
Dichromadora polaris Vermeeren, Vanreusel & Vanhove, 2004
Dichromadora polarsternis Vermeeren, Vanreusel & Vanhove, 2004
Dichromadora quadripapillata Muthumbi & Vincx, 1998
Dichromadora southernis Vermeeren, Vanreusel & Vanhove, 2004
Dichromadora weddellensis Vermeeren, Vanreusel & Vanhove, 2004
Euchromadora amokurae (Ditlevsen, 1921)
Euchromadora denticulata (sp incertae sedis) Cobb, 1914
Euchromadora meridiana (sp incertae sedis) Cobb, 1914
Euchromadora vulgaris (Bastian, 1865)
Graphonema amokurae (Ditlevsen, 1921) Inglis, 1971
Neochromadora aberrans Cobb, 1930
Neochromadora complexa Gerlach, 1953
Neochromadora craspedota (Steiner, 1916)
Neochromadora edentata (Cobb, 1914)
Neochromadora notocraspedota Allgén, 1958
Neochromadora poecilosoma (de Man, 1893) Micoletzky, 1926
Prochromadorella antarctica (Cobb, 1914)
Prochromadorella conicaudata (Allgén, 1927) Wieser, 1954
Prochromadorella paramucrodonta (Allgén, 1929) Wieser, 1951
Spiliphera dolichura de Man, 1893
Spiliphera edentata (Cobb, 1914)
Spiliphera gracilicauda (de Man, 1893) Wieser, 1954
Spilophorella campbelli Allgén, 1928
Spilophorella paradoxa (de Man, 1888)
Family Neotonchidae Wieser & Hopper, 1966
Neotonchus chamberlaini Wieser & Hopper, 1966
Family Cyatholaimidae Filipjev, 1918
Cyatholaimus gracilis (Eberth, 1863)
Longicyatholaimus longicaudatus (de Man, 1876)
Marylynnia macrodentata (Wieser, 1959)
Marylynnia quadriseta Hopper, 1972
Metacyatholaimus spatiosus Wieser, 1954
Paracanthonchus arcospiculum Allgén, 1959
Paracanthonchus axonolaimoides Allgén, 1960
Paracanthonchus caecus (Bastian, 1865)
Paracanthonchus elongatus (de Man, 1906)
Paracanthonchus falklandiae Allgén, 1959
Paracanthonchus macrospiralis Allgén, 1959
Paracanthonchus paralongus Allgén, 1959
Paracanthonchus paramacrodon (Allgén, 1951)
Paracanthonchus spectabilis Allgén, 1931
Paracanthonchus stateni Allgén, 1930
Pomponema multipapillatum Filipjev, 1922
Praeacanthonchus kreisi (Allgén, 1929)
Praeacanthonchus punctatus (Bastian, 1865)
Family Selachnematidae Cobb, 1915
Cheironchus conicaudatus Allgén, 1959
Halichoanolaimus chordiurus Gerlach, 1955
Halichoanolaimus dolichurus Ssaweljev, 1912
Halichoanolaimus minor Ssaweljev, 1912
Halichoanolaimus ovalis (Ditlevsen, 1921)
Halichoanolaimus robustus (Bastian, 1865)
Order Desmodorida De Coninck, 1965
Suborder Desmodorina De Coninck, 1965
Family Desmodoridae Filipjev, 1922
Acanthopharynx merostomacha (Steiner, 1921)
Chromaspirina crinita Gerlach, 1952
Croconema stateni Allgén, 1928
Desmodora campbelli (Allgén, 1932)
Desmodora microchaeta (Allgén, 1929)
Desmodora minuta Wieser, 1954
Desmodora scaldensis de Man, 1889
Desmodorella abyssorum (Allgén, 1929)
Desmodorella aff. balteata Verschelde, Gourbault & Vincx, 1998
Desmodorella tenuispiculum (Allgén, 1928)
Laxus septentrionalis Cobb, 1914
Molgolaimus allgeni Allgén, 1935
Molgolaimus australis Fonseca, Vanreusel & Decraemer, 2006
Molgolaimus carpediem Fonseca, Vanreusel & Decraemer, 2006
Molgolaimus drakus Fonseca, Vanreusel & Decraemer, 2006
Molgolaimus exceptionregulum Fonseca, Vanreusel & Decraemer, 2006
Molgolaimus falliturvisus Fonseca, Vanreusel & Decraemer, 2006
Molgolaimus galluccii Fonseca, Vanreusel & Decraemer, 2006
Molgolaimus gigaslongicus Fonseca, Vanreusel & Decraemer, 2006
Molgolaimus gigasproximus Fonseca, Vanreusel & Decraemer, 2006
Molgolaimus liberalis Fonseca, Vanreusel & Decraemer, 2006
Molgolaimus macilenti Fonseca, Vanreusel & Decraemer, 2006
Molgolaimus mareprofundus Fonseca, Vanreusel & Decraemer, 2006
Molgolaimus nettoensis Fonseca, Vanreusel & Decraemer, 2006
Molgolaimus sabakii Muthumbi & Vincx, 1996
Molgolaimus sapiens Fonseca, Vanreusel & Decraemer, 2006
Molgolaimus tenuispiculum Ditlevsen, 1921
Molgolaimus unicus Fonseca, Vanreusel & Decraemer, 2006
Molgolaimus walbethi Fonseca, Vanreusel & Decraemer, 2006
Molgolaimus xuxunaraensis Fonseca, Vanreusel & Decraemer, 2006
Onyx ferox (Ditlevsen, 1921) Gerlach, 1951
Paradesmodora campbelli (Allgén, 1932) Gerlach, 1963
Pseudometachromadora longilaima (Schuurmans Stekhoven, 1950)
Pseudonchus symmetricus De Coninck, 1942
Spirinia gnaigeri Ott, 1977
Spirinia parasitifera (Bastian, 1865) Gerlach, 1963
Spirinia septentrionalis (Cobb, 1914) Gerlach, 1963
Spirinia tenuicauda (Allgén, 1959) Gerlach, 1963
Family Epsilonematidae Steiner, 1927
Archepsilonema celidotum Steiner, 1931
Bathyepsilonema brachycephalum Steiner, 1931
Bathyepsilonema drygalskii Steiner, 1931
Epsilonema cyrtum Steiner, 1931
Epsilonema docidocricum (Steiner, 1931)
Glochinema trispinatum Raes, Vanreusel & Decraemer, 2003
Family Draconematidae Filipjev, 1918
Cygnonema steineri Allen & Noffsinger, 1978
Draconactus suillus (Allgén, 1932) Allen & Noffsinger, 1978
Draconema antarcticum (Allen & Noffsinger, 1978)
Draconema cephalatum (Cobb, 1913)
Paradraconema antarcticum Allen & Noffsinger, 1978
Prochaetosoma campbelli (Allgén, 1932)
Prochaetosoma longicapitata Allgén, 1932
Family Microlaimidae Micoletzky, 1922
Bolbolaimus dentatus (Allgén, 1935)
Microlaimus dimorphus Chitwood, 1937
Microlaimus falklandiae Allgén, 1959
Microlaimus honestus (de Man, 1922)
Microlaimus kaurii Wieser, 1954
Microlaimus latilaimus Allgén, 1959
Microlaimus papilliferus Allgén, 1959
Microlaimus pinguis Wieser, 1954
Microlaimus sensus Wieser, 1954
Microlaimus texianus Chitwood, 1951
Family Monoposthiidae Filipjev, 1934
Monoposthia costata (Bastian, 1865)
Monoposthia desmodoroides Allgén, 1959
Monoposthia falklandiae Allgén, 1959
Monoposthia grahami Allgén, 1959
Monoposthia mirabilis Schulz, 1932
Monoposthia paramediterranea (Allgén, 1959)
Nudora campbelli (Schulz, 1935) Wieser, 1954
Order Desmoscolecida Filipjev, 1929
Family Desmoscolecidae Shipley, 1896
Antarcticonema comicapitatum Timm, 1978
Desmoscolex amaurus Lorenzen, 1972
Desmoscolex antarcticos Timm, 1970
Desmoscolex articulatus Timm, 1978
Desmoscolex campbelli Allgén, 1946
Desmoscolex cristatus (Allgén, 1932)
Desmoscolex frigidus Timm, 1978
Desmoscolex gerlachi Timm, 1970
Desmoscolex labiosus Lorenzen, 1969
Desmoscolex max Timm, 1970
Desmoscolex parafalklandiae Allgén, 1955
Desmoscolex spinosus Decraemer, 1976
Greeffiella antarctica Timm, 1978
Quadricoma avicapitata Timm, 1978
Quadricomoides magna (Timm, 1970)
Tricoma antarctica Timm, 1970
Tricoma curvicauda (Timm, 1978)
Tricoma maxima (Schepotieff, 1907)
Tricoma nematoides (Greeff, 1869)
Tricoma pontica (Filipjev, 1922)
Tricoma septentrionalis Timm, 1978
Usarpnema auriculatum Timm, 1978
Terschellingia longicaudata de Man, 1907
Terschellingia longispiculata Wieser & Hopper, 1967
Order Araeolaimida De Coninck & Schuurmans Stekhoven, 1933
Family Axonolaimidae Filipjev, 1918
Axonolaimus antarcticus Cobb, 1930
Axonolaimus austrogeorgiae Allgén, 1959
Axonolaimus spinosus (Bütschli, 1874)
Axonolaimus tenuicaudatus Allgén, 1959
Odontophora angustilaimoides Chitwood, 1951
Odontophora longisetosa (Allgén, 1928)
Odontophora peritricha Wieser, 1956
Odontophora polaris (Cobb, 1914)
Parodontophora pacifica (Allgén, 1947)
Parodontophora quadristicha (Schuurmans Stekhoven, 1950)
Family Comesomatidae Filipjev, 1918
Cervonema chilensis Chen & Vincx, 2000
Cervonema hermani Chen & Vincx, 2000
Cervonema papillatum Jensen, 1988
Cervonema shiae Chen & Vincx, 2000
Cervonema tenuicauda Schuurmans Stekhoven, 1950
Comesoma hermani Chen & Vincx, 1998
Comesoma tenuispiculum (Ditlevsen, 1921)
Dorylaimopsis magellanense Chen & Vincx, 1998
Dorylaimopsis punctatus Ditlevsen, 1918
Hopperia beaglense Chen & Vincx, 1998
Hopperia dorylaimopsoides Allgén, 1959
Laimella annae Chen & Vincx, 2000
Laimella filipjevi Jensen, 1979
Laimella longicauda Cobb, 1920
Order Monhysterida Filipjev, 1929
Laimella sandrae Chen & Vincx, 2000
Suborder Monhysterina De Coninck & Schuurmans Stekhoven, Laimella subterminata Chen & Vincx, 2000
1933
Metacomesoma cyatholaimoides Wieser, 1954
Family Monhysteridae de Man, 1876
Sabatieria celtica Southern, 1914
Halomonhystera disjuncta Bastian, 1865
Sabatieria coomansi Chen, 1999
Halomonhystera uniformis Cobb, 1914
Sabatieria curvispiculum (Allgén, 1959)
Longitubopharynx obtusicaudatus Allgén, 1959
Sabatieria falcifera Wieser, 1954
Monhystera macquariensis Allgén, 1929
Sabatieria furcillata Wieser, 1954
Thalassomonhystera parva (Bastian, 1865)
Sabatieria granifer Wieser, 1954
Sabatieria intermissa Wieser, 1954
Family Sphaerolaimidae Filipjev, 1918
Sabatieria kelletti Platt, 1983
Sphaerolaimus arcospiculum Allgén, 1959
Sabatieria lawsi Platt, 1983
Sphaerolaimus campbelli Allgén, 1927
Sabatieria mortenseni (Ditlevsen, 1921)
Sphaerolaimus gracilis de Man, 1876
Sabatieria ornata (Ditlevsen, 1918)
Sphaerolaimus hirsutus Bastian, 1865
Sabatieria parabyssalis Wieser, 1954
Sphaerolaimus pacificus Allgén, 1947
Sabatieria praedatrix de Man, 1907
Family Xyalidae Chitwood, 1951
Setosabatieria hilarula (de Man, 1922)
Austronema (Dub) spirurum Cobb, 1914
Vasostoma spiratum (Jensen, 1979)
Cobbia dentata Gerlach, 1953
Family Coninckiidae Lorenzen, 1981
Cobbia mawsoni Cobb, 1930
Coninckia macrospirifera Zhang, 1983
Daptonema acanthospiculum (Allgén, 1959)
Daptonema alternus (Wieser, 1956)
Family Diplopeltidae Filipjev, 1918
Daptonema dentatus (Wieser, 1956)
Araeolaimus australis Allgén, 1959
Daptonema filispiculum (Allgén, 1932)
Araeolaimus conicaudatus Allgén, 1959
Daptonema fistulatus (Wieser & Hopper, 1967)
Araeolaimus dubiosus Allgén, 1959
Daptonema normandicus (de Man, 1890)
Araeolaimus elegans (de Man, 1888)
Daptonema resimus (Wieser, 1959)
Araeolaimus obtusicaudatus Allgén, 1959
Daptonema septentrionalis (Cobb, 1914)
Araeolaimus ovalis Wieser, 1956
Daptonema tortus (Wieser & Hopper, 1967)
Araeolaimus paracylindricauda Allgén, 1959
Elzalia tenuis Allgén, 1959
Araeolaimus paradubiosus Allgén, 1959
Filipjeva crucis Blome & Schräge, 1985
Araeolaimus tenuicauda Allgén, 1959
Linhystera longa Pastor de Ward, 1985
Campylaimus inaequalis Cobb, 1920
Linhystera problematica Juario, 1974
Diplopeltis cirrhatus (Eberth, 1863)
Manganonema antarctica Fonseca, Vanreusel & Decraemer, Diplopeltula bulbosa Vitiello, 1972
2006
Diplopeltula cylindricauda (Allgén, 1932)
Paramonhystera biforma Wieser, 1956
Diplopeltula incisa (Southern, 1914)
Paramonhystera geraerti Chen & Vincx, 2000
Southerniella nojii Jensen, 1991.
Paramonhystera megacephala Wieser, 1956
Southerniella simplex (Allgén, 1932)
Paramonhystera proteus Wieser, 1956
Order Plectida Malakhov, 1982
Rhynchonema megamphida Boucher, 1974
Family Leptolaimidae Örley, 1880
Steineria pilosa Cobb, 1914
Camacolaimus ampullocaudatus Allgén, 1959
Steineridora loricata (Steiner, 1916)
Camacolaimus austrogeorgiae Allgén, 1959
Theristus acer Bastian, 1865
Camacolaimus cylindricauda Allgén, 1959
Theristus conicaudatus Allgén, 1959
Camacolaimus falklandiae Allgén, 1959
Theristus filicaudatus Allgén, 1959
Camacolaimus guillei de Bovée, 1977
Theristus horridus Steiner, 1916
Camacolaimus longicauda de Man, 1922
Theristus normandicus (de Man, 1890)
Camacolaimus macrocellatus Allgén, 1959
Theristus oistospiculum (Allgén, 1930)
Camacolaimus paratardus Allgén, 1959
Theristus paravelox Allgén, 1934
Camacolaimus spissus Allgén, 1959
Theristus pellucidus Allgén, 1939
Camacolaimus tardus de Man, 1889
Theristus problematicus (Allgén, 1928)
Camacolaimus zostericola (Filipjev, 1918)
Theristus velox (Bastian, 1865)
Ionema cobbi (Steiner, 1916)
Suborder Linhomoeina Andrássy, 1974
Leptolaimus antarcticus (Cobb, 1914)
Family Siphonolaimidae Filipjev, 1918
Leptoplectonema fuegoense Coomans & Raski, 1991
Siphonolaimus falklandiae Allgén, 1959
Necolaimus austrogeorgiae Allgén, 1959
Siphonolaimus smetti Chen & Vincx, 2000
Notocamacolaimus australis Allgén, 1959
Family Linhomoeidae Filipjev, 1922
Anticyathus septentrionalis (Cobb, 1914)
Desmolaimus conicaudatus Allgén, 1959
Desmolaimus macrocirculus Allgén, 1959
Desmolaimus propinquus Allgén, 1959
Disconema falklandiae Allgén, 1959
Linhomoeus elongatus Bastian, 1865
Metalinhomoeus biformis Juario, 1974
Metalinhomoeus filiformis (de Man, 1907)
Metalinhomoeus leptosoma Allgén, 1959
Metalinhomoeus longiseta Kreis, 1929
Metalinhomoeus retrosetosus Wieser, 1956
Metalinhomoeus setosus Chitwood, 1951
Metalinhomoeus tristis (Allgén, 1933)
Notosouthernia obtusicauda Allgén, 1959
Paralinhomoeus lepturus de Man, 1907
Paralinhomoeus macquariensis Allgén, 1929
Paralinhomoeus meridionalis Cobb, 1930
Paralinhomoeus tenuicaudatus (Bütschli, 1874)
Terschellingia claviger Wieser, 1956
Terschellingia communis de Man, 1888
Family Aegialoalaimidae Lorenzen, 1981
Aegialoalaimus conicaudatus Allgén, 1959
Aegialoalaimus elegans de Man, 1907
Aegialoalaimus paratenuicaudatus Allgén, 1959
Aegialoalaimus tenuicaudatus Allgén, 1932
Family Plectidae Örley, 1880
Plectus falklandiae Allgén, 1959
Plectus gisleni Allgén, 1951
Plectus grahami Allgén, 1951
Order Rhabditida Chitwood, 1933
Suborder Tylenchina Thorne, 1949
Family Cephalobidae Filipjev, 1934
Cephalobus incisocaudatus Allgén, 1951
Suborder Rhabditina Chitwood, 1933
Family Rhabditidae Örley, 1880
Rhabditis marina Bastian, 1865
Biogeographic Atlas of the Southern Ocean
479
THE BIOGEOGRAPHIC ATLAS OF THE SOUTHERN OCEAN
Scope
Biogeographic information is of fundamental importance for discovering marine biodiversity hotspots, detecting and understanding impacts of environmental changes, predicting future
distributions, monitoring biodiversity, or supporting conservation and sustainable management strategies.
The recent extensive exploration and assessment of biodiversity by the Census of Antarctic Marine Life (CAML), and the intense compilation and validation efforts of Southern Ocean
biogeographic data by the SCAR Marine Biodiversity Information Network (SCAR-MarBIN / OBIS) provided a unique opportunity to assess and synthesise the current knowledge on Southern
Ocean biogeography.
The scope of the Biogeographic Atlas of the Southern Ocean is to present a concise synopsis of the present state of knowledge of the distributional patterns of the major benthic and pelagic
taxa and of the key communities, in the light of biotic and abiotic factors operating within an evolutionary framework. Each chapter has been written by the most pertinent experts in their
field, relying on vastly improved occurrence datasets from recent decades, as well as on new insights provided by molecular and phylogeographic approaches, and new methods of analysis,
visualisation, modelling and prediction of biogeographic distributions.
A dynamic online version of the Biogeographic Atlas will be hosted on www.biodiversity.aq.
The Census of Antarctic Marine Life (CAML)
CAML (www.caml.aq) was a 5-year project that aimed at assessing the nature, distribution and abundance of all living organisms of the Southern Ocean. In this time of environmental change,
CAML provided a comprehensive baseline information on the Antarctic marine biodiversity as a sound benchmark against which future change can reliably be assessed. CAML was initiated
in 2005 as the regional Antarctic project of the worldwide programme Census of Marine Life (2000-2010) and was the most important biology project of the International Polar Year 2007-2009.
The SCAR Marine Biodiversity Information Network (SCAR-MarBIN)
In close connection with CAML, SCAR-MarBIN (www.scarmarbin.be, integrated into www.biodiversity.aq) compiled and managed the historic, current and new information (i.a. generated
by CAML) on Antarctic marine biodiversity by establishing and supporting a distributed system of interoperable databases, forming the Antarctic regional node of the Ocean Biogeographic
Information System (OBIS, www.iobis.org), under the aegis of SCAR (Scientific Committee on Antarctic Research, www.scar.org). SCAR-MarBIN established a comprehensive register of
Antarctic marine species and, with biodiversity.aq provided free access to more than 2.9 million Antarctic georeferenced biodiversity data, which allowed more than 60 million downloads.
The Editorial Team
Claude DE BROYER is a marine biologist at the Royal Belgian Institute of Natural
Sciences in Brussels. His research interests cover structural and ecofunctional
biodiversity and biogeography of crustaceans, and polar and deep sea benthic
ecology. Active promoter of CAML and ANDEEP, he is the initiator of the SCAR
Marine Biodiversity Information Network (SCAR-MarBIN). He took part to 19 polar
expeditions.
Philippe KOUBBI is professor at the University Pierre et Marie Curie (Paris,
France) and a specialist in Antarctic fish ecology and biogeography. He is the
Principal Investigator of projects supported by IPEV, the French Polar Institute.
As a French representative to the CCAMLR Scientific Committee, his main input
is on the proposal of Marine Protected Areas. His other field of research is on the
ecoregionalisation of the high seas.
Huw GRIFFITHS is a marine Biogeographer at the British Antarctic Survey. He
created and manages SOMBASE, the Southern Ocean Mollusc Database. His
interests include large-scale biogeographic and ecological patterns in space and
time. His focus has been on molluscs, bryozoans, sponges and pycnogonids as
model groups to investigate trends at high southern latitudes.
Ben RAYMOND is a computational ecologist and exploratory data analyst,
working across a variety of Southern Ocean, Antarctic, and wider research
projects. His areas of interest include ecosystem modelling, regionalisation
and marine protected area selection, risk assessment, animal tracking, seabird
ecology, complex systems, and remote sensed data analyses.
Cédric d’UDEKEM d’ACOZ is a research scientist at the Royal Belgian Institute
of Natural Sciences, Brussels. His main research interests are systematics of
amphipod crustaceans, especially of polar species and taxonomy of decapod
crustaceans. He took part to 2 scientific expeditions to Antarctica on board of the
Polarstern and to several sampling campaigns in Norway and Svalbard.
Anton VAN DE PUTTE works at the Royal Belgian Institute for Natural Sciences
(Brussels, Belgium). He is an expert in the ecology and evolution of Antarctic
fish and is currently the Science Officer for the Antarctic Biodiveristy Portal www.
biodiversity.aq. This portal provides free and open access to Antarctic Marine and
terrestrial biodiversity of the Antarctic and the Southern Ocean.
Bruno DANIS is an Associate Professor at the Université Libre de Bruxelles, where
his research focuses on polar biodiversity. Former coordinator of the scarmarbin.
be and antabif.be projects, he is a leading member of several international
committees, such as OBIS or the SCAR Expert Group on Antarctic Biodiversity
Informatics. He has published papers in various fields, including ecotoxicology,
physiology, biodiversity informatics, polar biodiversity or information science.
Bruno DAVID is CNRS director of research at the laboratory BIOGÉOSCIENCES,
University of Burgundy. His works focus on evolution of living forms, with and
more specifically on sea urchins. He authored a book and edited an extensive
database on Antarctic echinoids. He is currently President of the scientific council
of the Muséum National d’Histoire Naturelle (Paris), and Deputy Director at the
CNRS Institute for Ecology and Environment.
Susie GRANT is a marine biogeographer at the British Antarctic Survey. Her work
is focused on the design and implementation of marine protected areas, particularly
through the use of biogeographic information in systematic conservation planning.
Julian GUTT is a marine ecologist at the Alfred Wegener Institute Helmholtz
Centre for Polar and Marine Research, Bremerhaven, and professor at the
Oldenburg University, Germany. He participated in 13 scientific expeditions to
the Antarctic and was twice chief scientist on board Polarstern. He is member
of the SCAR committees ACCE and AnT-ERA (as chief officer). Main focii of his
work are: biodiversity, ecosystem functioning and services, response of marine
systems to climate change, non-invasive technologies, and outreach.
Christoph HELD is a Senior Research Scientist at the Alfred Wegener Institute
Helmholtz Centre for Polar and Marine Research, Bremerhaven. He is a specialist
in molecular systematics and phylogeography of Antarctic crustaceans, especially
isopods.
Graham HOSIE is Principal Research Scientist in zooplankton ecology at the
Australian Antarctic Division. He founded the SCAR Southern Ocean Continuous
Plankton Recorder Survey and is the Chief Officer of the SCAR Life Sciences
Standing Scientific Group. His research interests include the ecology and
biogeography of plankton species and communities, notably their response to
environmental changes. He has participated in 17 marine science voyages to
Antarctica.
Falk HUETTMANN is a ‘digital naturalist’ he works on three poles ( Arctic, Antarctic
and Hindu-Kush Himalaya) and elsewhere (marine, terrestrial and atmosphere).
He is based with the university of Alaska-Fairbank (UAF) and focuses primarily
on effective conservation questions engaging predictions and open access data.
Alexandra POST is a marine geoscientist, with expertise in benthic habitat
mapping, sedimentology and geomorphic characterisation of the seafloor.
She has worked at Geoscience Australia since 2002, with a primary focus on
understanding seafloor processes and habitats on the East Antarctic margin.
Most recently she has led work to understand the biophysical environment
beneath the Amery Ice Shelf, and to characterise the habitats on the George V
Shelf and slope following the successful CAML voyages in that region.
Yan ROPERT COUDERT spent 10 years at the Japanese National Institute of
Polar Research, where he graduated as a Doctor in Polar Sciences in 2001. Since
2007, he is a permanent researcher at the CNRS in France and the director of a
polar research programme (since 2011) that examines the ecological response
of Adélie penguins to environmental changes. He is also the secretary of the
Expert Group on Birds and Marine Mammals and of the Life Science Group of the
Scientific Committee on Antarctic Research.
AnT-ERA