Abstract
Background
Soil nematodes are one of the most important terrestrial faunal groups in Antarctica, as they are a major component of soil micro-food webs. Despite their crucial role in soil processes, knowledge of their species diversity and distribution is still incomplete. Taxonomic studies of Antarctic nematodes are fragmented, which prevents assessment of the degree of endemicity and distribution of the species, as well as other aspects of biogeography.
New information
The present study is focused on the nematode fauna of one of the three Antarctic sub-regions, the Maritime Antarctic and summarises all findings published up to April 2023. A species list that includes 44 species, belonging to 21 genera, 16 families and eight orders is provided. A review of the literature on terrestrial nematodes inhabiting the Maritime Antarctic showed that the sites are unevenly studied. Three islands (Signy, King George and Livingston Islands) revealed highest species richness, probably due to the highest rates of research effort. Most species and four genera (Antarctenchus, Pararhyssocolpus, Amblydorylaimus and Enchodeloides) are endemic, proving that nematode fauna of the Maritime Antarctic is autochthonous and unique. Several groups of islands/sites have been revealed, based on their nematode fauna. The study showed that species with a limited distribution prevailed, while only two species (Plectusantarcticus and Coomansusgerlachei) have been found in more than 50% of the sites. Based on the literature data, details on species localities, microhabitat distribution, plant associations and availability of DNA sequences are provided.
Keywords: endemics, distribution, DNA sequences, species
Introduction
Soil nematodes are one of the most important groups of the terrestrial fauna in Antarctica (Maslen and Convey 2006) as they are abundant, taxonomically and functionally diverse and occupy a central position in the soil micro-food webs (Adams et al. 2014) and may have an impact on nutrient cycling and carbon dioxide emission, when soils thaw for a longer period of the year under climate change (van den Hoogen et al. 2019). In the challenging environmental conditions of the Antarctic, their distribution is limited to ice-free areas, where they have evolved throughout millions of years of climatic fluctuations in refugia (Ebach et al. 2008, Convey et al. 2020, Stevens and Mackintosh 2023). The glaciations, long-term isolation, harsh climate and the patchy distribution of ice-free areas (present today where at least partially ice-free throughout repeated glacial maxima (Newman et al. 2009)) are the main factors affecting the Antarctic nematode fauna origin/genesis (i.e. the formation of fauna under the influence of multiple factors - historical, geographic and ecological) (Andrássy 1998, Convey and Peck 2019). In order to survive in the extreme environments, nematodes have developed exceptional cryptobiotic adaptations to manage freezing and desiccation stress (e.g. Pickup (1988), Wharton (1995), Treonis and Wall (2005), Kagoshima et al. (2019)).
Knowledge of the impact of climate change on nematode communities from extreme habitats and how they respond to these changes is insufficient (Freckman and Virginia 1997, Nielsen et al. 2011a). One of the main problems in predicting the effects of climate change in Antarctica is the limited knowledge on the diversity of terrestrial fauna, especially nematodes and the lack of comprehensive long-term studies (Gantait 2014). Data on species distribution and biogeography are not enough and the taxonomic information still remains confused or scarce (Andrássy 1998, Maslen and Convey 2006, Adams et al. 2014, Kagoshima et al. 2019). Nematodes possess high indicator potential for assessing various environmental changes in the soil environment because they are abundant, ubiquitous, utilise diverse trophic and live strategies and, thus, occupy key positions in soil micro-food webs (Neher 2001, Ferris et al. 2001, Neher 2010, Chauvin et al. 2020, Taylor et al. 2020, Ara Khanum et al. 2022, Du Preez et al. 2022). This highlights the need for research on the fauna of nematodes and their communities in extreme environments in view of the already occurring global change.
Antarctica represents three distinct climatic regions: the Sub-Antarctic, Maritime and Continental Antarctic (Holdgate 1977), with the Sub-Antarctic being the most favourable (mean air temperatures of most islands are low, but positive during the whole year), with the Continental Antarctic having the harshest conditions (the average monthly temperatures remain below freezing) (Convey 2017). The Continental Antarctic covers the territories of the continent, the Balleny Islands and the eastern side of the Antarctic Peninsula (Convey 2017). The Sub-Antarctic is the boundary zone that lies north of 56°S (Chown and Brooks 2019). The flora and fauna in this region are rather typical of temperate latitudes. In this paper, we focused on terrestrial nematodes from the Maritime Antarctic. This is a region with a strong influence of the Southern Ocean; it includes the western coast of the Antarctic Peninsula to ca. 72°S, the South Shetland, South Orkney and South Sandwich Islands and the isolated Bouvetøya and Peter I Øya (Convey 2006, Convey 2017). The Maritime Antarctic is characterised by more favourable conditions compared with the Continental Antarctic: mean air temperatures are positive for 1–4 months of the year (Convey 2017), the vegetation is predominantly cryptogamic (algae, mosses, liverworts, lichens); higher plants are represented by two species, Deschampsiaantarctica Desv. (Poaceae) and Colobanthusquitensis Bartl. (Caryophyllaceae) (Greene 1970, Longton 1979, Smith 1984).
Studies on nematodes in the Maritime Antarctic started at the beginning of the 20th century, with the contribution of the Romanian biologist Emil Racoviţă during the first scientific Antarctic expedition in winter (Andrássy 1998). This resulted in a description of the first Antarctic nematode species, Mononchusgerlachei (= Coomansusgerlachei (de Man 1904) Jairajpuri and Khan 1977), followed by three other species, Plectusantarcticus de Man 1904, P.belgicae de Man 1904 and Dorylaimus sp. Following these first data, targeted investigations on the terrestrial nematode fauna from this part of Antarctica started from the early 70s of the 20th century (Spaull 1972, Spaull 1973a, Spaull 1973b, Spaull 1973c, Loof 1975, Maslen 1979a, Maslen 1979b, Caldwell 1981, Maslen 1981, Spaull 1981, Pickup 1988, Shishida and Ohyama 1989, Tsalolikhin 1989, Pickup 1990, Janiec 1996, Peneva et al. 1996, Andrássy 1998, Peneva and Chipev 1999, Convey et al. 2000, Nedelchev and Peneva 2000, Peneva et al. 2002, Convey and Wynn-Williams 2002, Holovachov and Bostrom 2006, Maslen and Convey 2006, Nedelchev and Peneva 2007, Kito 2009, Peneva et al. 2009, Nielsen et al. 2011b, Velasco-Castrillón and Stevens 2014, Velasco-Castrillón et al. 2014a, Russell et al. 2014, Elshishka et al. 2015a, Elshishka et al. 2015b, Elshishka et al. 2017, Kagoshima et al. 2019, Newsham et al. 2020). Antarctic nematodes have been studied mainly in easy to access areas near to the research bases/stations; therefore, there are still many remote locations never sampled for nematodes which raise questions on how widespread the species are (Adams et al. 2014, Convey et al. 2020).
According to Andrássy (1998), numerous studies have reported species as new records with no morphological description making it impossible to confirm identifications, especially when the collected material is no longer available for subsequent examination. Further, this has an impact on the potential to assess fauna endemicity, which is critical for examining Antarctic biogeography within a global context (Andrássy 1998). There are numerous cases of misclassification and underestimation of the diversity for most microfaunal groups in Antarctica, likely due to poor taxonomic resolution caused by insufficient sampling and their difficult identification (Adams et al. 2006, Iakovenko et al. 2015, Carapelli et al. 2017, Short et al. 2022, Collins et al. 2023), as well as the low degree of the development and application of molecular taxonomy.
In recent years, molecular studies have become more important in these marginal habitats, as a powerful toolkit to complement the traditional taxonomy, species identification and descriptions and to assess biodiversity and biogeography (Courtright et al. 2000, Velasco-Castrillón et al. 2014b, Elshishka et al. 2015b, Elshishka et al. 2017, Czechowski et al. 2017, Velasco-Castrillón et al. 2018, Kagoshima et al. 2019).
The integrative approach (combining morphological and molecular data) is an effective way to understand the scale of endemism, evolution and distribution of the Antarctic nematode fauna. However, the main problem of not linking molecular data with morphology still remains for the vast majority of Antarctic nematode species.
The present paper aims to summarise all records of nematode species occurrence in the Maritime Antarctic between the years of 1904 and April 2023 as a basis for further studies and to present a snapshot of nematode species diversity in this part of the Antarctic.
Materials and methods
The nematode species list has been composed, based on literature data and refers to the Maritime Antarctic. This list includes all species recovered in the Maritime Antarctic, as well as the islands and sites from where each species was reported, along with data on microhabitats and plant associations, accession numbers of published sequences in GenBank also included, if available. The type of microhabitat is reported as in the original paper, the scientific names of the plants being adapted according to the current systematics (Ochyra 1998). Geographical coordinates are presented additionally for each site if missing in the original paper. For the literature search, online bibliography search engine Google Scholar and the academic databases Scopus, Web of Science and CABI were used with search keywords “terrestrial nematode species*” and “Maritime Antarctic*”. We focused on studies reporting nematode species (see Holovachov (2014a)) from the Maritime Antarctic and omitted those that provide data only at the generic or family level.
Several papers recording multiple unidentified taxa at generic level (Maslen and Convey 2006, Nielsen et al. 2011b) probably contain many undescribed nematode species from those regions suggesting that the nematode diversity there might be underestimated to a great extent. Overall, nematodes from 37 sites (34 islands and three localities on the Antarctic Peninsula) are included in the review. The taxonomic position of the Antarctic species was presented according to the current nematode nomenclature. Classification follows Andrássy (2005), Andrássy (2007) and Andrássy (2009); only for order Plectida classification follows Holovachov (2014b). The analyses are based on species presence/absence data and Wizard > Matrix display function in PRIMER v.7.0 software (Clarke and Gorley 2015). The Matrix display wizard performs a sequence of sample and species resemblance calculations and clustering and seriation steps resulting in a shade plot which visualises the species presence/absence data and sites similarity.
Checklists
Checklist of terrestrial nematodes from the Maritime Antarctic
Dorylaimida
DB2E6DDD-5A83-5FAD-AA05-491530706589
Nordiidae
02036AD4-B115-5ACB-9D3A-C365D4FFFBE3
Enchodeloides signyensis
(Loof, 1975) Elshishka, Lazarova, Radoslavov, Hristov, Peneva, 2017
F37166B3-37C5-55AA-A12F-64A130AA896F
Enchodelus signyensis Loof, 1975
Qudsianematidae
15105F37-9676-56CF-887B-328D15B98C4B
Eudorylaimus coniceps
Loof, 1975
181A5CE6-3861-500E-A087-F0FF38D50423
Eudorylaimus pseudocarteri
Loof, 1975
47CAF4A1-E28A-5E59-ACEB-31BCAA37A957
Eudorylaimus spaulli
Loof, 1975
5F4D2E55-53D8-538D-B503-A78CEEA105CE
Eudorylaimus verrucosus
Loof, 1975
3FB9DFB0-EEBC-545E-9436-944D9DBBEED3
Eudorylaimus cf. carteri
Andrassy, 1959 (Bastian, 1865)
13B2261E-A584-5999-BED0-190056CB4892
Pararhyssocolpidae
6C015E2C-BEF0-56CA-AF5D-8AE0D3404203
Pararhyssocolpus paradoxus
(Loof, 1975) Elshishka, Lazarova, Radoslavov, Hristov, Peneva, 2015
D9585A55-9B26-58B4-9516-DDC1BC9EB469
Eudorylaimus paradoxus Loof, 1975|Rhyssocolpus paradoxus (Loof, 1975) Andrássy, 1986
Dorylaimidae
671787FE-3946-5420-9619-FACA9D2FA213
Calcaridorylaimus signatus
(Loof, 1975) Andrássy, 1981
F5A9037F-584C-56F6-A44A-63714ACBC6D2
Mesodorylaimus signatus Loof, 1975
Mesodorylaimus antarcticus
Nedelchev and Peneva, 2000
EE444A29-4179-5887-B2A1-F41ADCCD81D6
Mesodorylaimus chipevi
Nedelchev and Peneva, 2000
9CD6CDD3-2EF8-5BAE-9D2A-2BCBC01ADE36
Mesodorylaimus imperator
Loof, 1975
9515D5C8-859D-5A66-A87E-9B42936469F3
Mesodorylaimus masleni
Nedelchev and Peneva, 2000
2A2130FE-3C7C-5B1D-9ADE-2B200FCE8D50
Aporcelaimidae
75CC4343-89B8-5900-94FC-936112411653
Amblydorylaimus isokaryon
(Loof, 1975) Andrássy, 1998
B044B3FC-B769-5B49-BA3E-B9BC91BE6581
Eudorylaimus isokaryon Loof, 1975
Aphelenchida
28744625-5C08-5291-98C0-3794406ED313
Aphelenchoididae
01714A28-F5E0-5D6B-B774-10899E47322D
Aphelenchoides haguei
Maslen, 1979
D4AC79D2-B450-5D13-9DCB-9E7FA2223288
Aphelenchoides vaughani
Maslen, 1979
A1301FE0-4F0B-5836-89C5-9246F2546124
Laimaphelenchus helicosoma
(Maslen, 1979) Peneva and Chipev, 1999
EFACFF62-976B-52D8-9526-5BA5A58C10E2
Aphelenchoides helicosoma Maslen, 1979
Alaimida
ADEC0201-921A-524A-B652-7B828938C15C
Amphidelidae
CBF633E8-7E83-5A54-A021-9C64861B55ED
Paramphidelus antarcticus
Tsalolikhin, 1989
D52EDC6A-5219-5C6E-A1F5-5F16A09027F6
Monhysterida
6748B1A9-7801-5159-8BB2-F159B2314D3E
Monhysteridae
7F94800F-CCCC-5B15-A41F-FBDDAE8C6CF3
Eumonhystera filiformis
(Bastian, 1865) Andrássy, 1981
87224B84-E25A-58B9-8B24-D9603402ED4E
Eumonhystera vulgaris
(de Man, 1880) Andrássy, 1981
33E17069-31EC-5865-A580-6DC42A82B918
Geomonhystera villosa
(Bütschli, 1873) Andrássy, 1981
6766505C-8E58-5AA4-BA98-A18390B2F1C0
Plectida
195B637E-A81D-5EC0-9837-4C8317666230
Plectidae
02148983-E217-5C83-8D3D-B5BA9EBF4DCD
Plectus antarcticus
de Man, 1904
E558E15A-D114-5D9C-BA37-37BC230AD109
Plectus cf. antarcticus
de Man, 1904
DF1FF3FE-15D0-5BDB-AFEC-B84A616E61AB
Plectus belgicae
de Man, 1904
49B0CF61-2DA8-5A0A-976F-96025CF7414C
Plectus cf. belgicae
de Man, 1904
A7BC5E1E-478B-5A29-8CE2-A308A5ABA7F0
Plectus insolens
Andrássy, 1998
ECAB5F55-AB36-5933-9BB3-4876420207EA
Plectus tolerans
Andrássy, 1998
06295DAB-BBB3-5019-A2F7-0C9B80497144
Plectus cf. tolerans
Andrássy, 1998
A9D39BED-6A38-5246-BBC1-7E7217EAE2F3
Plectus meridianus
Andrássy, 1998
BC659854-2A25-5484-9336-20F9490DBAC9
Plectus cf. meridianus
Andrássy, 1998
976487C9-C3EB-5CF8-B924-F0D5E86B7FB4
Plectus armatus
Bütschli, 1873
E97AA054-492C-5A21-B20B-198B5E58EB7B
Ceratoplectus armatus (Bütschli, 1873) Andrássy, 1984
Tylenchida
266FDC27-9624-596A-AD7A-24842C567356
Psilenchidae
A315F1BF-3304-582D-BB2C-CC87EF8EFC52
Antarctenchus hooperi
Spaull, 1972
8412463F-2F6C-5DB8-AF9A-7E4457E73FC2
Anguinidae
8DA64EBF-3D67-5C57-A8D9-9D104D5B8277
Ditylenchus parcevivens
Andrássy, 1998
E2215FCD-DAF8-5AB4-819E-30C2BA885C08
Rhabditida
51F7A518-984C-5256-93F6-F883548EFB18
Teratocephalidae
CDEA95B0-7F13-5883-9856-2C609C4E06F1
Teratocephalus tilbrooki
Maslen, 1979
3B584363-B128-5856-A2DB-DC6BDB952CA7
Teratocephalus pseudolirellus
Maslen, 1979
0BEC9F2C-E0DD-54EF-A34F-27716F289B59
Teratocephalus rugosus
Maslen, 1979
AD7D6030-A3BD-5870-A253-9F5E9C120EA1
Cephalobidae
82D975B4-05C7-5984-AD81-AAFC584A518B
Acrobeloides arctowskii
Holovachov and Boström, 2006
7C694E1F-086F-590F-9BBB-EF082420FBE7
Cervidellus cf. vexilliger
(de Man, 1880) Thorne, 1937
DE9D79DC-B582-5B55-B83A-1485B5BF627C
Rhabditidae
58F28B03-4A75-5365-B10C-05773A48BEB6
Cuticularia firmata
Andrássy, 1998
107613C0-FDBC-560E-9D60-60066C61F32B
Rhabditis krylovi
Tsalolikhin, 1989
D26C2B68-3D22-5133-A37C-DB9F260A2761
Rhabditis marina -group
410838AE-0F98-5B3C-AE3E-0CCB5F4D407A
Peloderidae
8C42EF21-19F7-5F43-A2E3-1A7D323A102B
Pelodera teres -group
F43B566C-DE04-54B9-881E-D65B10FA9A59
Pelodera strongyloides -group
C5A2BEFB-7A41-50C1-9050-C7E4AC3EA8F9
Pelodera parateres -group
D1D7791E-0B8D-5994-A75E-6FF0B8A08B8F
Mononchida
417FEFF6-89BD-5372-82DA-295F4AF64FC8
Mononchidae
CCA60FEF-1D5F-5DB5-A53E-25427B4590F4
Coomansus gerlachei
(de Man, 1904) Jairajpuri and Khan, 1977
4DDB6E9C-AEB1-54D8-95B5-28D16F93D36D
Mononchus gerlachei de Man, 1904|Clarkusgerlachei (de Man, 1904) Jairajpuri, 1970
Analysis
Results
To date, 44 species of terrestrial nematodes, belonging to 21 genera, 16 families and eight orders have been recorded in the Maritime Antarctic (Table 1, Fig. 1). Nematodes have been reported from 34 islands and three sites on the Antarctic Peninsula (Fig. 2). Several groups of islands/sites have been revealed, based on their nematode fauna. Those groups form a gradient from north (the group of Livingston, King George and Signy Islands) to south (the group of Adelaide, Charcot, Alexander, Leonie and Alamode Islands).
Table 1.
Distribution of terrestrial nematodes in the Maritime Antarctic.
* Taxonomic paper; ** Paper with molecular data; ***Paper with molecular and morphologica data
Terrestrial reference sites (SIRS) at Signy Island for long-term monitoring of the various biotic and abiotic components of Antarctic moss-peat communities (for full descriptions, see Tilbrook (1973)). These sites no longer exist and no studies have been done since the late 1980s.
SIRS 1 (Polytrichastrumalpinum (Hedwig), Chorisodontiumaciphyllum (Hook. f. & Wilson) Broth. (60°43.5'S, 45°35.6'W))
SIRS 2 (Sanioniauncinata (Hedw.), Warnstorfiasarmentosa (Wahlenb.), Warnstorfialaculosa (Müll. Hal.), Cephaloziellavarians (Gottsche) Steph. (60°43.7'S, 45°36'W))
1 Geographical coordinates according to original paper. 2 Geographical coordinates additionally added.
| Nematode species | Locality / Coordinates | Microhabitat and plant species | DNA / Accesion number in GenBank | Reference |
|
Enchodeloides
signyensis |
Signy Island (type locality) 1,260°43’S, 45°38’W |
Syntrichiafilaris (Müll. Hal.) (type habitat); D.antarctica; C.quitensis |
Loof (1975) * |
|
| Maslen (1979b) | ||||
| SIRS 1; SIRS 2 |
Caldwell (1981) | |||
| SIRS 1; SIRS 2 |
Maslen (1981) | |||
| Alamode Island 1,268°43'S, 67°32'W |
S.uncinata | Loof (1975)* | ||
| Maslen (1979b) | ||||
| Moss | Maslen and Convey (2006) | |||
| Alexander Island 271°0′0″S, 70°0′0″W |
Moss, lichen, soil, microbial mat | Maslen and Convey (2006) | ||
| Alexander Island171°52’40’’S, 68°15’57’’W | Convey and Wynn-Williams (2002) | |||
| Blaiklock Island 1,267°33’S, 67°00’W |
P.alpinum, Pohlianutans (Hedw.) | Loof (1975) * | ||
| Maslen (1979b) | ||||
| Coronation Island 1,260°38’S, 45°35’W |
D.antarctica | Loof (1975)* | ||
| Maslen (1979b) | ||||
| Charcot Island 1,269°45'S, 75°15'W |
Soil, moss clumps, algae, various lichens | Convey et al. (2000) | ||
| Moss, lichen, soil | Maslen and Convey (2006) | |||
| Dream Island 264°44′0″S, 64°14′0″W |
Moss mats with green algae | Shishida and Ohyama (1989)* | ||
| Elephant Island 1,261°10’S, 55°14’W |
D.antarctica; Polytrichum sp. |
Loof (1975)* | ||
| Maslen (1979b) | ||||
| Galindez Island 1,265°15'S, 64°15'W |
D.antarctica | Loof (1975)* | ||
| Maslen (1979b) | ||||
| King George Island 162°09‘32“S, 58°27‘58“W |
D.antarctica, C.quitensis, Sanionia sp., S.filaris, Syntrichiamagellanica (Mont.) | Russell et al. (2014) | ||
| King George Island 262°2′S, 58°21′W |
Moist brown soil without vegetation, surrounded by moss | 18S rDNA KY881720.1 28S rDNA KY881719.1 |
Elshishka et al. (2017)*** | |
| Livingston Island 262°36′S, 60°30′W |
D.antarctica; D.antarctica+S.uncinata; D.antarctica+S.uncinata+C.quitensis; P.alpinum; S.uncinata; Bryum sp.; Usnea sp.+P.alpinum; Cladonia sp.+S.uncinata+P.alpinum; Polytrichumjuniperinum Hedw.+S.uncinata; S.uncinata+Bartramiapatens Brid. |
Peneva et al. (2002)* | ||
| Elshishka et al. (2015a) | ||||
| Moss; Soil under moss crust; Soil |
Elshishka et al. (2017)* | |||
| Eudorylaimusconiceps | Signy Island (type locality) 1,260°43’S, 45°38’W |
S.filaris (type habitat); Andreaeagainii Card.; C.quitensis; W.laculosa and W.sarmentosa |
Loof (1975)* | |
| SIRS 1 | Maslen (1979b) | |||
| SIRS 2 | Maslen (1981) | |||
| Andreaea sp. | Pickup (1988) | |||
| Wharton and Block (1993) | ||||
| Alexander Island 271°0′0″S, 70°0′0″W |
Moss, lichen, soil, microbial mat |
Maslen and Convey (2006) | ||
| Ardley Island 162°12‘38“S, 58°56‘40“W |
Russell et al. (2014) | |||
| Charcot Island 1,269°45'S, 75°15'W |
Soil, moss clumps, algae, various lichens | Convey et al. (2000) | ||
| Moss, lichen, soil | Maslen and Convey (2006) | |||
| Coronation Island 1,260°38’S, 45°35’W |
D.antarctica | Loof (1975)* | ||
| Maslen (1979b) | ||||
| Elephant Island 1,261°10’S, 55°14’W |
D.antarctica | Loof (1975)* | ||
| Maslen (1979b) | ||||
| Emperor Island 1,267°52'S, 68°43'W |
S.uncinata and Bryumpseudotriquetrum (Hedw.) | Loof (1975)* | ||
| Maslen (1979b) | ||||
| Guebriant Island 267°48′S, 68°25′W |
Maslen (1979b) | |||
| Halfmoon Island 162°35‘45“S, 59°54‘06“W |
Russell et al. (2014) | |||
| King George Island 262°2′S, 58°21′W |
Mosses | Kito (2009) | ||
| King George Island 162°11‘48“S, 58°59‘28“W; 62°11‘50“S, 58°56‘33“W; 62°11‘53“S, 58°56‘47“W |
Russell et al. (2014) | |||
| Leonie Island 267°36′S, 68°21′W |
Mixture of soil, moss, lichen, liverworts, algae and cyanobacteria | 18S rDNA LC457670.1 LC457669.1 LC457668.1 LC457667.1 LC457666.1 LC457665.1 LC457664.1 LC457663.1 LC457662.1 LC457647.1 LC457646.1 LC457645.1 |
Kagoshima et al. (2019)** | |
| Livingston Island 262°36′S, 60°30′W |
Elshishka et al. (2015a) | |||
| E.pseudocarteri | Signy Island (type locality) 160°43’S, 45°38’W |
A.gainii (type habitat); W.laculosa and W.sarmentosa; D.antarctica |
Loof (1975)* | |
| Maslen (1979b) | ||||
| SIRS 1; SIRS 2 |
Maslen (1981) | |||
| Andreaea sp. | Pickup (1988) | |||
| Wharton and Block (1993) | ||||
| Adelaide Island 267°15′S, 68°30′W |
Moss, lichen | Maslen and Convey (2006) | ||
| Alexander Island 271°0′0″S, 70°0′0″W |
Moss, lichen, soil, microbial mat, freshwater | |||
| Ardley Island 162°12‘38“S, 58°56‘40“W |
Russell et al. (2014) | |||
| Charcot Island 1,269°45'S, 75°15'W |
Moss, lichen, soil | Maslen and Convey (2006) | ||
| Coronation Island 1,260°38’S, 45°35’W |
D.antarctica | Loof (1975)* | ||
| Maslen (1979b) | ||||
| Elephant Island 1,261°10’S, 55°14’W |
Polytrichum sp. | Loof (1975)* | ||
| Maslen (1979b) | ||||
| King George Island 262°2′S, 58°21′W |
Puddle | Tsalolikhin (1989)* | ||
| King George Island 162°09‘32“S, 58°27‘58“W; 62°11‘48“S, 58°59‘28“W; 62°11‘50“S, 58°56‘33“W |
Russell et al. (2014) | |||
| Livingston Island 162°39‘14“S, 60°36‘39“W |
||||
| E.spaulli | Alamode Island (type locality) 1,268°43'S, 67°32'W |
Soil around S.uncinata (type habitat) | Loof (1975)* | |
| Moss, lichen, soil, microbial mat | Maslen and Convey (2006) | |||
| Maslen (1979b) | ||||
| Adelaide Island 267°15′S, 68°30′W |
Moss, lichen | Maslen and Convey (2006) | ||
| Alexander Island 271°0′0″S, 70°0′0″W |
Moss, lichen, soil, microbial mat |
|||
| Anchorage Island 267°36′14.01″S, 68°12′32.78″W |
Moss, grass, lichen, soil, microbial mat, freshwater |
|||
| Blaiklock Island 1,267°33’S, 67°00’W |
P.alpinum, Pohlianutans | Loof (1975)* | ||
| Maslen (1979b) | ||||
| Charcot Island 1,269°45'S, 075°15'W |
Soil, moss clumps, algae, various lichens | Convey et al. (2000) | ||
| Moss, lichen, soil | Maslen and Convey (2006) | |||
| Coronation Island 1,260°38'S, 45°35'W |
D.antarctica | Loof (1975)* | ||
| Maslen (1979b) | ||||
| Deception Island 262°58′37″S, 60°39′0″W |
Maslen (1979b) | |||
| Elephant Island 1,261°10’S, 55°14’W |
D.antarctica; Polytrichum sp. |
Loof (1975)* | ||
| Maslen (1979b) | ||||
| Leonie Island 267°36′S, 68°21′W |
Moss, grass, lichen, soil, microbial mat, freshwater |
Maslen and Convey (2006) | ||
| Limpet Island 1,267°38'S, 68°19'W |
S.uncinata | Loof (1975)* | ||
| Maslen (1979b) | ||||
| Pourquoi pas Island 267°41S, 67°28′W |
Maslen (1979b) | |||
| Signy Island 1,260°43’S, 45°38’W |
S.filaris B . pseudotriquetrum; D.antarctica; W.laculosa and W.sarmentosa |
Loof (1975)* | ||
| Wharton and Block (1993) | ||||
| Andreaea sp. | Pickup (1988) | |||
| Maslen (1979b) | ||||
| SIRS 2 | Maslen (1981) | |||
| E.verrucosus | Elephant Island (type locality) 1,261°10’S, 55°14’W |
D.antarctica (type habitat) | Loof (1975)* | |
| Maslen (1979b) | ||||
| Adelaide Island 267°15′S, 68°30′W |
Moss, lichen, soil, microbial mat |
Maslen and Convey (2006) | ||
| Signy Island 160°43’S, 45°38’W |
Maslen (1979b) | |||
| SIRS 1; SIRS 2 |
Maslen (1981) | |||
| Wharton and Block (1993) | ||||
| Eudorylaimuscf.carteri | Livingston Island 262°36′S, 60°30′W |
Elshishka et al. (2015a) | ||
| Pararhyssocolpusparadoxus | Signy Island (type locality) 160°43’S, 45°38’W |
A.gainii (type habitat); S.filaris |
Loof (1975)* | |
| Maslen (1979b) | ||||
| SIRS 1; SIRS 2 |
Maslen (1981) | |||
| Adelaide Island 267°15′S, 68°30′W |
Moss, lichen, soil, microbial mat |
Maslen and Convey (2006) | ||
| Adelaide Island 167°34.429'S, 68°07.284'W |
C.varians and S. uncinata | Newsham et al. (2020) | ||
| Anchorage Island 267°36′14.01″S, 68°12′32.78″W |
Moss, grass, lichen, soil, microbial mat, freshwater |
Maslen and Convey (2006) | ||
| Blaiklock Island 1,267°33’S, 67°00’W |
Maslen (1979b) | |||
| Coronation Island 1,260°38'S, 45°35'W |
D.antarctica | Loof (1975)* | ||
| Maslen (1979b) | ||||
| Elephant Island 1,261°10’S, 55°14’W |
S.uncinata | Loof (1975)* | ||
| Maslen (1979b) | ||||
| Galindez Island 1, 265°15'S, 64°15'W |
D.antarctica | Loof (1975)* | ||
| Maslen (1979b) | ||||
| Guebriant Island 267°48′S, 68°25′W |
Maslen (1979b) | |||
| Intercurrence Island 1,263°55'S, 61°24'W |
Brachythecium sp. | Loof (1975)* | ||
| Maslen (1979b) | ||||
| King George Island 262°2′S, 58°21′W |
Kito (2009) | |||
| Soil | 18S rDNA KM092521.1 28S rDNA KM092522.1 |
Elshishka et al. (2015b)*** | ||
| King George Island 162°09‘32“S, 58°27‘58“W; 62°11‘48“S, 58°59‘28“W; 62°11‘50“S, 58°56‘33“W; 62°11‘53“S, 58°56‘47“W |
Russell et al. (2014) | |||
| Leonie Island 267°36′S, 68°21′W |
Moss, grass, lichen, soil, microbial mat, freshwater |
Maslen and Convey (2006) | ||
| Limpet Island 1,267°38'S, 68°19'W |
S.uncinata | Loof (1975)* | ||
| Maslen (1979b) | ||||
| Livingston Island 262°36′S, 60°30′W |
Elshishka et al. (2015a) | |||
|
Sanionia sp.; C.quitensis, D.antarctica, moss; D.antarctica, C.quitensis; D.antarctica, moss |
Elshishka et al. (2015b)* | |||
| Nelson Island 262°18′S, 59°3'W |
Moss | Elshishka et al. (2015b)* | ||
| Calcaridorylaimussignatus | Signy Island (type locality) 160°43’S, 45°38’W |
S.filaris (type habitat); B.pseudotriquetrum; C.quitensis; D.antarctica; Prasiolacrispa (Lightfoot) |
Loof (1975)* | |
| Maslen (1979b) | ||||
| SIRS1 | Caldwell (1981) | |||
| SIRS 1; SIRS 2 |
Maslen (1981) | |||
| Soil, moss, lichen, liverworts, algae and cyanobacteria | 18S rDNA LC457654.1 LC457653.1 LC457652.1 LC457651.1 LC457650.1 LC457649.1 LC457648.1 |
Kagoshima et al. (2019)** | ||
| Alamode Island 268°43'S, 67°32'W |
Moss, lichen, soil | Maslen and Convey (2006) | ||
| Alexander Island 271°0′0″S, 70°0′0″W |
Moss, lichen, soil, microbial mat |
|||
| Anchorage Island 267°36′14.01″S, 68°12′32.78″W |
Moss, grass, lichen, soil, microbial mat, freshwater |
|||
| Anvers Island, Biscoe Point 264°49′6.85″S, 63°46′32.29″W |
Soil around roots of D.antarctica | Shishida and Ohyama (1989)* | ||
| Coronation Island 1,260°38'S, 45°35'W |
D.antarctica | Loof (1975)* | ||
| Maslen (1979b) | ||||
| Dream Island 264°44′0″S, 64°14′0″W |
Moss mats with green algae | Shishida and Ohyama (1989)* | ||
| Galindez Island 1,265°15'S, 64°15'W |
D.antarctica | Loof (1975)* | ||
| Maslen (1979b) | ||||
| Leonie Island 267°36′S, 68°21′W |
Moss, grass, lichen, soil, microbial mat, freshwater |
Maslen and Convey (2006) | ||
| Mesodorylaimusantarcticus | Livingston Island (type locality) 162°39'22’’S, 60°21'13’’W |
Sanionia sp. (type habitat); D.antarctica; D.antarctica-Polytrichum sp.; A small moss tuft Sanionia sp.; A mix grass-moss spot D.antarctica+Sanionia sp. |
Nedelchev and Peneva (2000)* | |
| King George Island 162°09‘32“S, 58°27‘58“W; 62°11‘48“S, 58°59‘28“W; 62°11‘50“S, 58°56‘33“W |
Russell et al. (2014) | |||
| Neko Harbour, Antarctic Peninsula 164°50‘41“S, 62°31‘53“W |
||||
| M.chipevi | Livingston Island (type locality) 162°34'48’’S, 60°20'42’’W |
D.antarctica on the top of flat rock near sea (type habitat); Shallow soil with cover of green algae amongst grass on a rock; Small tuft of D.antarctica; Polytrichum sp.+S.uncinata+D.antarctica; A mix grass-moss spot D.antarctica+Sanionia sp.; A large pure grass spot D.antarctica |
Nedelchev and Peneva (2000)* | |
| Livingston Island 162°38'52’’S, 60°22'24’’W |
S.georgico-uncinata Müll. Hal. + D.antarctica | Nedelchev and Peneva (2007)* | ||
| King George Island 162°09‘32“S, 58°27‘58“W |
Russell et al. (2014) | |||
| M.imperator | Emperor Island (type locality) 1,267°52'S, 68°43'W |
S.uncinata and B.pseudotriquetrum (type habitat) | Loof (1975)* | |
| Maslen (1979b) | ||||
| Adelaide Island 267°15′S, 68°30′W |
Moss, lichen, soil | Maslen and Convey (2006) | ||
| Alamode Island 268°43'S, 67°32'W |
Moss | |||
| Alexander Island 271°0′0″S, 70°0′0″W |
Moss, grass, lichen, soil, microbial mat, freshwater |
|||
| Anchorage Island 267°36′14.01″S, 68°12′32.78″W |
Moss, grass, lichen, soil, microbial mat, freshwater |
Maslen and Convey (2006) | ||
| Charcot Island 1,269°45'S, 075°15'W |
Soil, moss clumps, algae, various lichens | Convey et al. (2000) | ||
| Cone Island 1,267°41'S, 69°10'W |
S.uncinata | Loof (1975)* | ||
| Maslen (1979b) | ||||
| Killinbeck Island 267°34′S, 68°5′W |
Moss, lichen, soil | Maslen and Convey (2006) | ||
| Leonie Island 267°36′S, 68°21′W |
Moss, grass, lichen, soil, microbial mat, freshwater |
|||
| M.masleni | Livingston Island (type locality) 162°39'46’’S, 60°23'29’’W |
A large area of D.antarctica (type habitat); A mix grass-moss spot D.antarctica + Sanionia sp. |
Nedelchev and Peneva (2000)* | |
| Amblydorylaimusisokaryon | Elephant Island (type locality) 1,261°10’S, 55°14’W |
D.antarctica (type habitat); Polytrichum sp. |
Loof (1975)* | |
| Maslen (1979b) | ||||
| Galindez Island 1,265°15'S, 64°15'W |
D.antarctica | Loof (1975)* | ||
| Maslen (1979b) | ||||
| Intercurrence Island 1,263°55'S, 61°24'W |
Brachythecium sp. | Loof (1975)* | ||
| Maslen (1979b) | ||||
| King George Island 262°2′S, 58°21′W |
Kito (2009) | |||
| Soil | Elshishka et al. (2015b)* | |||
| Livingston Island 262°36′S, 60°30′W |
Elshishka et al. (2015a) | |||
| Grass spot (D.antarctica); a moss- grass (D.antarctica-Polytrichum sp.) community; S.georgico-uncinata and D.antarctica; C.quitensis and D.antarctica, moss; D.antarctica and C.quitensis |
Elshishka et al. (2015b)* | |||
| Nelson Island 262°18′S, 59°3′W |
Moss | 18S rDNA KM092519.1 28S rDNA KM092520.1 |
Elshishka et al. (2015b)*** | |
| Aphelenchoideshaguei | Signy Island (type locality) 160°43’S, 45°38’W |
SIRS 1 (type habitat); SIRS 2 |
Maslen (1979a)* | |
| SIRS 1; SIRS 2 |
Maslen (1981) | |||
| Adelaide Island 267°15′S, 68°30′W |
Moss, lichen | Maslen and Convey (2006) | ||
| Adelaide Island 167°34.429'S, 68°07.284'W |
C.varians and S.uncinata | Newsham et al. (2020) | ||
| Alamode Island 268°43'S, 67°32'W |
Moss | Maslen and Convey (2006) | ||
| Alexander Island 271°0′0″S, 70°0′0″W |
Moss, lichen, soil, microbial mat | |||
| Anchorage Island 267°36′14.01″S, 68°12′32.78″W |
Moss, grass, lichen, soil, microbial mat, freshwater | |||
| Ardley Island 162°12‘38“S, 58°56‘40“W |
Russell et al. (2014) | |||
| Deception Island 162°58‘42“S, 60°33‘29“W |
||||
| King George Island 162°09‘32“S, 58°27‘58“W; 62°11‘48“S, 58°59‘28“W; 62°11‘50“S, 58°56‘33“W; 62°11‘53“S, 58°56‘47“W |
Russell et al. (2014) | |||
| Leonie Island 267°36′S, 68°21′W |
Moss, grass, lichen, soil, microbial mat, freshwater | Maslen and Convey (2006) | ||
| Livingston Island 162°39‘14“S, 60°36‘39“W |
Soil | Russell et al. (2014) | ||
| Livingston Island 262°36′S, 60°30′W |
Elshishka et al. (2015a) | |||
| A.vaughani | Signy Island (type locality) 160°43’S, 45°38’W |
SIRS 1; SIRS 2 |
Maslen (1979a)* | |
| SIRS 1; SIRS 2 |
Maslen (1981) | |||
| Alexander Island 271°0′0″S, 70°0′0″W |
Moss, lichen, soil, microbial mat, freshwater | Maslen and Convey (2006) | ||
| Livingston Island 262°36′S, 60°30′W |
Elshishka et al. (2015a) | |||
| Laimaphelenchushelicosoma | Signy Island (type locality) 160°43’S, 45°38’W |
SIRS 1 (type habitat); SIRS 2 |
Maslen (1979a)* | |
| SIRS 1 | Maslen (1981) | |||
| Adelaide Island 267°15′S, 68°30′W |
Moss, lichen | Maslen and Convey (2006) | ||
| Adelaide Island 167°34.429'S, 68°07.284'W |
C.varians and S.uncinata | Newsham et al. (2020) | ||
| Ardley Island 162°12‘38“S, 58°56‘40“W |
Russell et al. (2014) | |||
| King George Island 162°09‘32“S, 58°27‘58“W 62°11‘48“S, 58°59‘28“W 62°11‘50“S, 58°56‘33“W |
||||
| Russell et al. (2014) | ||||
| Livingston Island, 162°38'S, 60°20'W |
Primitive soil around roots of D.antarctica | Peneva and Chipev (1999)* | ||
| Paramphidelusantarcticus | King George Island (type locality) 262°2′S, 58°21′W |
Lichen (type habitat) | Tsalolikhin (1989)* | |
| Eumonhysterafiliformis | King George Island 162°09’S, 58°29'W |
Thaw ponds, with the bottom inhabited by W.sarmentosa | Janiec (1996) | |
| Livingston Island 262°36′S, 60°30′W |
Elshishka et al. (2015a) | |||
| E.vulgaris | King George Island 162°09’S, 58°29'W |
Puddle | Tsalolikhin (1989)* | |
| Moraine ponds, their shores are inhabited mainly by S.uncinata, W.sarmentosa and B.pseudotriquetrum; Moss banks of W.sarmentosa and W.laculosa; Thaw ponds, with the bottom inhabited by W.sarmentosa; Nearshore ponds, colonised by W.laculosa and W.sarmentosa |
Janiec (1996) | |||
| King George Island 162°09‘32“S, 58°27‘58“W; 62°11‘48“S, 58°59‘28“W; 62°11‘50“S, 58°56‘33“W; 62°11‘53“S, 58°56‘47“W |
Russell et al. (2014) | |||
| Livingston Island 262°36′S, 60°30′W |
Elshishka et al. (2015a) | |||
| Geomonhysteravillosa | Coronation Island 1,260°38'S, 45°35'W |
Maslen (1979b) | ||
| Ardley Island 162°12‘38“S, 58°56‘40“W |
Russell et al. (2014) | |||
| Elephant Islan 1,261°10’S, 55°14’W |
Maslen (1979b) | |||
| Galindez Island 1,265°15'S, 64°15'W |
||||
| Intercurrence Island 1,263°55'S, 61°24'W |
||||
| King George Island 162°09’S, 58°29'W |
W.sarmentosa and W.laculosa | Janiec (1996) | ||
| King George Island 162°11‘48“S, 58°59‘28“W 62°11‘50“S, 58°56‘33“W 62°11‘53“S, 58°56‘47“W |
Russell et al. (2014) | |||
| Livingston Island 162°39‘14“S, 60°36‘39“W |
||||
| Neko Harbour, Antarctic Peninsula 164°50‘41“S, 62°31‘53“W |
||||
| Signy Island 160°43’S, 45°38’W |
Maslen (1979b) | |||
| SIRS 1 | Caldwell (1981) | |||
| SIRS 1; SIRS 2 |
Maslen (1981) | |||
| Mixture of soil, moss, lichen, liverworts, algae and cyanobacteria | 18S rDNA LC457677.1 LC457676.1 LC457675.1 LC457674.1 LC457673.1 LC457672.1 LC457671.1 |
Kagoshima et al. (2019) | ||
| Plectusantarcticus | Danco Land coast, Beneden Head, Antarctic Peninsula (type locality) 264°46'S, 62°42'W |
Freshwater algae (type habitat) | de Man (1904)* | |
| Maslen (1979b) | ||||
| Moss from rock | Andrássy (1998)* | |||
| Adelaide Island 167°34′S, 68°07′W |
Moss, lichen, soil, microbial mat | Maslen and Convey (2006) | ||
| Cephaloziellavarians (Gottsche) | 18S rDNA LC457559.1 LC457558.1 LC457557.1 LC457556.1 LC457555.1 LC457554.1 |
Kagoshima et al. (2019)** | ||
| Adelaide Island 167°34.429'S, 68°07.284'W |
C.varians and S.uncinata | Newsham et al. (2020) | ||
| Alamode Island 268°43'S, 67°32'W |
Spaull (1973a) | |||
| Maslen (1979b) | ||||
| Moss | Maslen and Convey (2006) | |||
| Alexander Island, 171°52’40’’S, 68°15’57’’W |
Convey and Wynn-Williams (2002) | |||
| Alexander Island 271°0′0″S, 70°0′0″W |
Moss, lichen, soil, microbial mat | Maslen and Convey (2006) | ||
| Anchorage Island 267°36′14.01″S, 68°12′32.78″W |
Moss, grass, lichen, soil, microbial mat, freshwater | |||
| Avian Island 267°46"S, 68°54"W |
Spaull (1973a) | |||
| Maslen (1979b) | ||||
| Blaiklock Island 1,267°33’S, 67°00’W |
||||
| Charcot Island 1,269°45'S, 075°15'W |
Soil, moss clumps, algae, various lichens | Convey et al. (2000) | ||
| Moss, lichen, soil | Maslen and Convey (2006) | |||
| Cone Island 1,267°41'S, 69°10'W |
Maslen (1979b) | |||
| Coronation Island 1,260°38'S, 45°35'W |
||||
| Deception Island 262°58′37″S, 60°39′0″W |
Moss from basalt debris | Andrássy (1998)* | ||
| Maslen (1979b) | ||||
| Deception Island 162°58‘43“S, 60°33‘24“W |
Only erratic patches of mosses, lichens and algae | Russell et al. (2014) | ||
| Devil Island 163°47‘54“S, 57°17‘24“W |
Soil substrates of the very sandy with embedded gravel | |||
| Elephant Island 1,261°10’S, 55°14’W |
Spaull (1973a) | |||
| Maslen (1979b) | ||||
| S.uncinata | Andrássy (1998)* | |||
| Emperor Island 267°52'S, 68°43'W |
Spaull (1973a) | |||
| Maslen (1979b) | ||||
| Galindez Island 1,265°15'S, 64°15'W |
||||
| Guebriant Island 267°48′S, 68°25′W |
||||
| Intercurrence Island 1,263°55'S, 61°24'W |
Spaull (1973a) | |||
| Maslen (1979b) | ||||
| Killingbeck Island 267°34′S, 68°5′W |
Moss, lichen, soil | Maslen and Convey (2006) | ||
| King George Island 262°2′S, 58°21′W |
Soil around rhizosphere of grasses and under lichen | Tsalolikhin (1989)* | ||
| King George Island 162°09’S, 58°29'W |
Moraine ponds, their shores are inhabited mainly by S.uncinata, W.sarmentosa and B.pseudotriquetrum; Moss banks of W.sarmentosa and W.laculosa |
Janiec (1996) | ||
| King George Island 162°09‘32“S, 58°27‘58“W; 62°11‘48“S, 58°59‘28“W |
Mosses, lichens, D.antarctica | Russell et al. (2014) | ||
| Leonie Island 267°36′S, 68°21′W |
Moss, grass, lichen, soil, microbial mat, freshwater | Maslen and Convey (2006) | ||
| Limpet Island 1,267°38'S, 68°19'W |
Maslen (1979b) | |||
| Livingston Island 262°36′S, 60°30′W |
Elshishka et al. (2015a) | |||
| Pourqoui pas Island 267°41'S, 67°28′W |
Maslen (1979b) | |||
| Signy Island 160°43’S, 45°38’W |
Spaull (1973a) | |||
| Spaull (1973b) | ||||
| Spaull (1973c) | ||||
| Maslen (1979b) | ||||
| SIRS 1; SIRS 2 |
Maslen (1981) | |||
| SIRS1; SIRS2 |
Caldwell (1981) | |||
| Andreaea sp. | Pickup (1988) | |||
| Wharton and Block (1993) | ||||
|
Acrocladium sp.; D.antarctica |
Andrássy (1998)* | |||
| Plectuscf.antarcticus | Livingston Island 262°36′S, 60°30′W |
Elshishka et al. (2015a) | ||
| Signy Island 260°43’S, 45°36’W |
Mixture of soil, moss, lichen, liverworts, algae and cyanobacteria | 18S rDNA LC457687.1 LC457686.1 |
Kagoshima et al. (2019) | |
| P.belgicae | Cap Beneden, Danco Land, Antarctic Peninsula (type locality) 264°46''S, 62°42"W |
Algae fresh water (type habitat) | de Man (1904)* | |
| Adelaide Island 167°34′S, 68°07′W |
Moss, lichen, soil, microbial mat | Maslen and Convey (2006) | ||
| C.varians | 18S rDNA LC457565.1 LC457564.1 LC457563.1 LC457562.1 LC457561.1 LC457560.1 |
Kagoshima et al. (2019) | ||
| Adelaide Island 167°34.429'S, 68°07.284'W |
C.varians and S.uncinata | Newsham et al. (2020) | ||
| Anchorage Island 267°36′14.01″S, 68°12′32.78″W |
Moss, grass, lichen, soil, microbial mat, freshwater | Maslen and Convey (2006) | ||
| Charcot Island 1, 269°45'S, 075°15'W |
Moss, lichen, soil | |||
| Elephant Island 1, 261°10’S, 55°14’W |
P.juniperinum; S.uncinata |
Andrássy (1998)* | ||
| King George Island 262°2′S, 58°21′W |
B.pseudotriquetrum and Bartramiapatens | 18S rDNA LC457638.1 LC457637.1 LC457636.1 | Kagoshima et al. (2019) | |
| Livingston Island 262°36′S, 60°30′W |
Elshishka et al. (2015a) | |||
| Rhyolite Island, 269°40′S, 68°35′W |
Moss, grass | Maslen and Convey (2006) | ||
| Signy Island 260°43’S, 45°36’W |
Acrocladium sp.; Usnea sp.; SIRS 2 |
Andrássy (1998)* | ||
| Plectuscf.belgicae | Ardley Island 162°12‘38“S, 58°56‘40“W |
Russell et al. (2014) | ||
| Deception Island 162°58‘43“S, 60°33‘24“W; 62°58‘42“S, 60°33‘29“W |
||||
| King George Island 162°09‘32“S, 58°27‘58“W; 62°11‘48“S, 58°59‘28“W; 62°11‘50“S, 58°56‘33“W; 62°11‘53“S, 58°56‘47“W |
||||
| Livingston Island 162°39‘14“S, 60°36‘39“W |
||||
| Livingston Island 262°36′S, 60°30′W |
Elshishka et al. (2015a) | |||
| Petermann Island 165°10‘29“S, 64°08‘10“W |
Russell et al. (2014) | |||
| P.insolens | Signy Island (type locality) 260°43’S, 45°36’W |
Thin soil on rock covered with Acrocladium sp. (type habitat); roots of D.antarctica | Andrássy (1998)* | |
| Ardley Island 162°12‘38“S, 58°56‘40“W |
Soils, Sanionia sp., W.sarmentosa and Andreaearegularis Müll. Hal. | Russell et al. (2014) | ||
| Livingston Island 262°36′S, 60°30′W |
Elshishka et al. (2015a) | |||
| P.tolerans | Emperor Island (type locality) 267°52’S, 68°43’W |
S.uncinata (type habitat) | Andrássy (1998)* | |
| Alexander Island 271°0′0″S, 70°0′0″W |
Moss, lichen, soil, microbial mat, freshwater | Maslen and Convey (2006) | ||
| Anchorage Island 267°36′14.01″S, 68°12′32.78″W |
Moss, grass, lichen, soil, microbial mat, freshwater | |||
| Charcot Island 1,269°45'S, 075°15'W |
Moss, lichen, soil | |||
| King George Island 262°2′S, 58°21′W |
Kito (2009) | |||
| Leonie Island 267°36′S, 68°21′W |
Moss, grass, lichen, soil, microbial mat, freshwater | Maslen and Convey (2006) | ||
| Livingston Island 262°36′S, 60°30′W |
Elshishka et al. (2015a) | |||
| Rhyolite Island 269°40′S, 68°35′W |
Moss, grass | Maslen and Convey (2006) | ||
| Plectuscf.tolerans | Ardley Island 162°12‘38“S, 58°56‘40“W |
Russell et al. (2014) | ||
| King George Island 162°09‘32“S, 58°27‘58“W; 62°11‘48“S, 58°59‘28“W; 62°11‘50“S, 58°56‘33“W; 62°11‘53“S, 58°56‘47“W |
||||
| Livingston Island 162°39‘14“S, 60°36‘39“W |
||||
| P.meridianus | Terra Firma Island (type locality) 268°42′S, 67°32′W |
Lichen (type habitat) | Andrássy (1998)* | |
| Emperor Island 267°52’S, 68°43’W |
A carpet of S.uncinata | |||
| Livingston Island 262°36′S, 60°30′W |
Elshishka et al. (2015a) | |||
| Signy Island 260°43’S, 45°36’W |
Roots of D.antarctica | Andrássy (1998)* | ||
| Plectuscf.meridianus | Signy Island 260°43’S, 45°36’W |
Soil, moss, lichen, liverworts, algae and cyanobacteria | 18S rDNA LC457691.1 LC457690.1 LC457689.1 LC457688.1 |
Kagoshima et al. (2019) |
| P.armatus | Ardley Island 162°12‘38“S, 58°56‘40“W |
Russell et al. (2014) | ||
| Coronation Island 260°38'S, 45°35'W |
Maslen (1979b) | |||
| Elephant Island 261°10’S, 55°14’W |
||||
| Galindez Island 265°15'S, 64°15'W |
||||
| King George Island 162°11‘48“S, 58°59‘28“W |
Russell et al. (2014) | |||
| Livingston Island 262°36′S, 60°30′W |
Elshishka et al. (2015a) | |||
| Signy Island 160°43’S, 45°38’W |
D.antarctica; C.quitensis; mosses | Spaull (1973b) | ||
| Spaull (1973c) | ||||
| Maslen (1979b) | ||||
| Antarctenchushooperi | Signy Island (type locality) 1,260°43'S, 45°38'W |
A.gainii (type habitat); Brachythecium sp., Calliergon sp., S.filaris, Grimmiaantarctici Card., Ch.aciphyllum, P.juniperinum and D.antarctica |
Spaull (1972)* | |
|
Sanionia sp.+ Calliergon sp.+ Calliergidium sp.; Polytrichum sp.; Bryum sp. + Syntrichia sp. + Andreaea sp.; D.antarctica |
Spaull (1973a) | |||
| A.gainii, S.filaris, Calliergon-Calliergidium | Spaull (1973b) | |||
| Spaull (1973c) | ||||
| Maslen (1979b) | ||||
| SIRS 1; SIRS 2 |
Maslen (1981) | |||
| S.uncinata | Caldwell (1981) | |||
| Wharton and Block (1993) | ||||
| Coronation Island 1,260°38'S, 45°35'W |
D.antarctica | Spaull (1972)* | ||
| D.antarctica | Spaull (1973a) | |||
| Maslen (1979b) | ||||
| Ditylenchusparcevivens | Signy Island (type locality) 260°43’S, 45°38’W |
Fine silt (type habitat) | Andrássy (1998)* | |
| King George Island 162°09‘32“S, 58°27‘58“W; 62°11‘48“S, 58°59‘28“W; 62°11‘50“S, 58°56‘33“W; 62°11‘53“S, 58°56‘47“W |
Russell et al. (2014) | |||
| Livingston Island 262°36′S, 60°30′W |
Elshishka et al. (2015a) | |||
| Teratocephalustilbrooki | Signy Island (type locality) 1,260°43’S, 45°38’W |
SIRS 1 (type habitat); SIRS2 |
Maslen (1979a)* | |
| ‘Swamp’ moss carpets | Maslen (1979b) | |||
| SIRS 1; SIRS 2 |
Maslen (1981) | |||
| Andreaea sp. | Pickup (1988) | |||
| Wharton and Block (1993) | ||||
| Usnea sp. | Andrássy (1998)* | |||
| Adelaide Island 267°15′S, 68°30′W |
Moss, lichen | Maslen and Convey (2006) | ||
| Ardley Island 162°12‘38“S, 58°56‘40“W |
Russell et al. (2014) | |||
| Charcot Island 1, 269°45'S, 075°15'W |
Moss, lichen, soil | Maslen and Convey (2006) | ||
| Halfmoon Island 162°35‘45“S, 59°54‘06“W |
Soil | Russell et al. (2014) | ||
| King George Island 162°09‘32“S, 58°27‘58“W; 62°11‘48“S, 58°59‘28“W; 62°11‘50“S, 58°56‘33“W; 62°11‘53“S, 58°56‘47“W |
Soil | |||
| Leonie Island 267°36′S, 68°21′W |
Moss, grass, lichen, soil, microbial mat, freshwater | Maslen and Convey (2006) | ||
| T.pseudolirellus | Signy Island (type locality) 160°43’S, 45°38’W |
S.filaris (type habitat) | Maslen (1979a)* | |
| Alexander Island 271°0′0″S, 70°0′0″W |
Moss, lichen, soil, microbial mat, freshwater | Maslen and Convey (2006) | ||
| Charcot Island 269°45'S, 075°15'W |
Moss, lichen, soil | |||
| King George Island 262°2′S, 58°21′W |
Kito (2009) | |||
| Livingston Island 262°36′S, 60°30′W |
Elshishka et al. (2015a) | |||
| T.rugosus | Signy Island (type locality) 260°43’S, 45°38’W |
SIRS 1; SIRS 2 | Maslen (1979a)* | |
| Maslen (1979b) | ||||
| SIRS 1; SIRS 2 | Maslen (1981) | |||
| Ardley Island 162°12‘38“S, 58°56‘40“W |
Russell et al. (2014) | |||
| Deception Island 162°58‘42“S, 60°33‘29“W |
||||
| Devil Island 163°47‘54“S, 57°17‘24“W |
||||
| Halfmoon Island 162°35‘45“S, 59°54‘06“W |
||||
| King George Island 162°09‘32“S, 58°27‘58“W; 62°11‘48“S, 58°59‘28“W; 62°11‘53“S, 58°56‘47“W |
||||
| Livingston Island 262°36′S, 60°30′W |
Elshishka et al. (2015a) | |||
| Acrobeloidesarctowskii | King George Island (type locality) 158°29'30''W, 61°05'S |
Soil around roots of D.antarctica (type habitat) | Holovachov and Bostrom (2006)* | |
| King George Island 162°09‘32“S, 58°27‘58“W |
Russell et al. (2014) | |||
| Deception Island 162°58‘43“S, 60°33‘24“W; 62°58‘42“S, 60°33‘29“W |
Soil devoid of vegetation or with P.crispa | |||
| Cervidelluscf.vexilliger | King George Island 162°09‘32“S, 58°27‘58“W |
Russell et al. (2014) | ||
| Cuticulariafirmata | Signy Island (type locality) 260°43’S, 45°36’W |
Fine mud (type habitat); SIRS 2 |
Andrássy (1998)* | |
| Rhabditiskrylovi | King George Island (type locality) 262°2′S, 58°21′W |
Flowing lake (type habitat) | Tsalolikhin (1989)* | |
| Rhabditismarina-group | Deception Island 162°55‘43“S, 60°40‘48“W |
Russell et al. (2014) | ||
| Peloderateres group | Deception Island 162°58‘43“S, 60°33‘24“W; 62°58‘42“S, 60°33‘29“W |
Russell et al. (2014) | ||
| Livingston Island 162°39‘14“S, 60°36‘39“W |
||||
| Livingston Island 262°36′S, 60°30′W |
Elshishka et al. (2015a) | |||
| Neko Harbour, Antarctic Peninsula 164°50‘41“S, 62°31‘53“W |
Russell et al. (2014) | |||
| Paulet Island 163°34‘30“S, 55°46‘59“W |
Ornithogenic soils | |||
| Peloderastrongyloides group | Devil Island 163°47‘54“S, 57°17‘24“W |
Russell et al. (2014) | ||
| Livingston Island 262°36′S, 60°30′W |
Elshishka et al. (2015a) | |||
| Livingston Island 162°39‘14“S, 60°36‘39“W |
Russell et al. (2014) | |||
| Neko Harbour, Antarctic Peninsula 164°51‘45“S, 62°26‘47“W; 64°50‘41“S, 62°31‘53“W |
||||
| Peloderaparateres group | Deception Island 162°58‘42“S, 60°33‘29“W |
Russell et al. (2014) | ||
| King George Island 162°09‘32“S, 58°27‘58“W |
||||
| Coomansusgerlachei | Danco Land coast, Beneden Head Antarctic Peninsula (type locality) 264°46''S, 62°42"W |
Algae fresh water (type habitat) | de Man (1904)* | |
| Maslen (1979b) | ||||
| Andrássy (1998)* | ||||
| Caleta Cierva, Antarctic Peninsula 164°10’S, 60°57’W |
Soil around roots of D.antarctica | Chaves (1990)* | ||
| Antarctic Peninsula | Jiménez Guirado et al. (1998)* | |||
| Adelaide Island 167°34.429'S, 68°07.284'W |
C.varians and S. uncinata | Newsham et al. (2020) | ||
| Anchorage Island 267°36′14.01″S, 68°12′32.78″W |
Moss, grass, lichen, soil, microbial mat, freshwater |
Maslen and Convey (2006) | ||
| Anvers Island, Biscoe Point 264°49′6.85″S, 63°46′32.29″W |
Soil around roots of D.antarctica | Shishida and Ohyama (1989)* | ||
| Anvers Island, Old Palmer 264°45′48″S, 64°5′12″W |
Moss mats | |||
| Avian Island 267°46''S, 68°54"W |
Sanionia sp., Bryum sp. | Spaull (1973a) | ||
| Maslen (1979b) | ||||
| S.uncinata, B.pseudotriquetrum | Spaull (1981) | |||
| Coronation Island 1,260°38'S, 45°35'W |
D.antarctica | Spaull (1973a) | ||
| Maslen (1979b) | ||||
| Charcot Island 1,269°45'S, 75°15'W |
Soil, moss clumps, algae, various lichens | Convey et al. (2000) | ||
| Deception Island 262°58′37″S, 60°39′0″W |
Sanionia sp.; Polytrichum sp. |
Spaull (1973a) | ||
| Maslen (1979b) | ||||
| Melting snow moisten mosses in a shingle field | Andrássy (1998)* | |||
| Dream Island 264°44′0″S, 64°14′0″W |
Moss mats with green algae | Shishida and Ohyama (1989)* | ||
| Elephant Island 1,261°10’S, 55°14’W |
Brachythecium sp.; D.antarctica; Sanionia sp. |
Spaull (1973a) | ||
| Maslen (1979b) | ||||
| Soil bellow D.antarctica | Spaull (1981) | |||
| Galindez Island 1,265°15'S, 64°15'W |
Brachythecium sp.; D.antarctica; Bryum sp.; Sanionia sp. + Pohlia sp. |
Spaull (1973a) | ||
| Maslen (1979b) | ||||
| Brachytheciumaustrosalebrosum (C. Muell.) Par. | Spaull (1981) | |||
| Guebriant Island 267°48′S, 68°25′W |
Brachythecium sp., Bryum sp., Sanionia sp. |
Spaull (1973a) | ||
| Maslen (1979b) | ||||
| Intercurrence Island 1,263°55'S, 61°24'W |
Brachythecium sp., Bryum sp., Sanionia sp. | Spaull (1973a) | ||
| Maslen (1979b) | ||||
| Halfmoon Island 162°35‘45“S, 59°54‘06“W |
Russell et al. (2014) | |||
| King George Island 162°09’S, 58°29'W |
Moraine ponds, their shores are inhabited mainly by S.uncinata, W.sarmentosa and B.pseudotriquetrum | Janiec (1996) | ||
| Leonie Island 267°36′S, 68°21′W |
Moss, grass, lichen, soil, microbial mat, freshwater | Maslen and Convey (2006) | ||
| Mixture of soil, moss, lichen, liverworts, algae and cyanobacteria | 18S rDNA LC457644.1 LC457643.1 LC457642.1 LC457641.1 LC457640.1 LC457639.1 |
Kagoshima et al. (2019)** | ||
| Limpet Island 1,267°38'S, 68°19'W |
Brachythecium sp., Bryum sp., Sanionia sp. | Spaull (1973a) | ||
| Maslen (1979b) | ||||
| King George Island 262°2′S, 58°21′W |
Kito (2009) | |||
| King George Island 162°09‘32“S, 58°27‘58“W; 62°11‘48“S, 58°59‘28“W; 62°11‘50“S, 58°56‘33“W; 62°11‘53“S, 58°56‘47“W |
Russell et al. (2014) | |||
| Litchfield Island 264°46′S, 64°6′W |
Decaying moss mats with blue-green algae | Shishida and Ohyama (1989)* | ||
| Livingston Island 262°36′S, 60°30′W |
Soil under crisp of green algae; D.antarctica-S.uncinata; D.antarctica, P.alpinum |
Peneva et al. (1996)* | ||
| Livingston Island 162°39‘14“S, 60°36‘39“W |
Russell et al. (2014) | |||
| Livingston Island 262°36′S, 60°30′W |
Elshishka et al. (2015a) | |||
| Nelson Island 262°18′S, 59°3W |
Moss | 18S rDNA KM092523.1 28S rDNA KM092524.1 |
Elshishka et al. (2015b)** | |
| Signy Island 1,260°43’S, 45°38’W |
Sanionia sp.+ Calliergon sp.+ Calliergidium sp.; Polytrichum sp.; Bryum sp. + Syntrichia sp. +Andreaea sp.; D.antarctica | Spaull (1973a) | ||
| Spaull (1973b) | ||||
| Spaull (1973c) | ||||
| Maslen (1979b) | ||||
| S.uncinata and W.sarmentosa | Caldwell (1981) | |||
| SIRS 1; SIRS 2 |
Maslen (1981) | |||
| P.crispa from melt stream | Spaull (1981) | |||
| Sanionia sp. | Pickup (1988) | |||
|
P.crispa; S.uncinata |
Pickup (1990) | |||
| Calliergon sp. | Wharton and Block (1993) | |||
| Mixture of soil, lichen, liverworts, algae and cyanobacteria | 18S rDNA LC457661.1 LC457660.1 LC457659.1 LC457658.1 LC457657.1 LC457656.1 LC457655.1 |
Kagoshima et al. (2019)** |
Figure 1.
Terrestrial nematodes from the Maritime Antarctic - visual representation of the data matrix (shade plot): in the columns are the 37 sites and in the rows – 44 species. White and black spaces denote absence or presence of a particular species at a given site; sites and species are arranged according to the groups derived by the clustering analyses. Significant clusters were identified with SIMPROF test and visualised in red dashed lines and a range of coloured dots. Each colour represents a group of sites/islands with similar nematode fauna.
Figure 2.

Distribution of terrestrial nematodes in the Maritime Antarctic. In green are presented the sites with records of terrestrial nematodes.
The order Dorylaimida is the best represented order in this Antarctic Region with five families, six genera and 13 species. The order Mononchida is represented by only one family (one genus and species).
The families Aphelenchoididae, Cephalobidae, Monhysteridae, Plectidae, Qudsianematidae, Peloderidae and Rhabditidae have a cosmopolitan distribution and, in the Maritime Antarctic, they are represented by one to two genera and two to ten species. The family Plectidae is the most diverse (10 species). Seven families (Amphidelidae, Anguinidae, Aporcelaimidae, Mononchidae, Nordiidae, Pararhyssocolpidae and Psilenchidae) are represented by only one species each.
Almost all species and four genera (Antarctenchus, Pararhyssocolpus, Amblydorylaimus and Enchodeloides) are endemic. Four species generally known as cosmopolitan are reported in some ecological studies in the Maritime Antarctic: Eumonhysteravulgaris (de Man 1880) Andrássy (1981), E.filiformis (Bastian 1865) Andrássy (1981), Geomonhysteravillosa (Bütschli 1873) Andrássy (1981) and Plectusarmatus Bütschli 1873. Of these, a description and illustrations were provided only for E.vulgaris (Tsalolikhin 1989).
Most species (27) have limited distribution registered in up to five islands of the Maritime Antarctic. Cuticulariafirmata Andrássy 1998, Cervidelluscf.vexilliger, Rhabditiskrylovi Tsalolikhin 1989, a species of the Rhabditismarina-group, Mesodorylaimusmasleni Nedelchev and Peneva 2000, Eudorylaimuscf.carteri and Plectuscf.meridianus are recorded from one island only. Six species occurred in more than 30% of the sites (C.gerlachei, P.antarcticus, Pararhyssocolpusparadoxus (Loof 1975), Eudorylaimusspaulli Loof 1975, E.coniceps Loof 1975, Enchodeloidessignyensis (Loof 1975)) with C.gerlachei and P.antarcticus being the most widespread (reported from more than half of the sites) (Figs 3, 4, 5). There are no particular trends in the distribution of most common species (occurring in more than 22% of the sites, 1/4 of the species) related to longitude or latitude, only P.paradoxus and Mesodorylaimusimperator Loof 1975 have not been reported from the most southern sites, whereas G.villosa – from the most northern localities.
Figure 3.
Described species and their occurrences presented as percentages.
Figure 4.
Distribution of P.antarcticus, C.gerlachei and Pararhyssocolpusparadoxus in the Maritime Antarctic. In red are presented the sites with records of these species, in green are presented the sites with records of the other Antarctic terrestrial nematodes.
Figure 5.
Distribution of Eudorylaimusspaulli, E.coniceps and Enchodeloidessignyensis in the Maritime Antarctic. In red are presented the sites with records of these species, in green are presented the sites with records of the other Antarctic terrestrial nematodes.
In most of the literature sources, there are data on the microhabitats in which nematode species occurred. The nematodes have been recorded from various microhabitats: bare soil, microbial mats, moss, lichens and algae and soil around the two species of higher plants occurring in the Maritime Antarctic (Fig. 6).
Figure 6.
Some microhabitats in the Maritime Antarctic. A Colobanthusquitensis B Deschampsiaantarctica C Bare soil D Syntrichia sp. E Bryum sp. F D.antarctica, C.quitensis, mosses G Sanionia sp. H Polytrichum sp. J Usnea sp. Photographs by M. Elshishka (Livingston Island).
DNA data have been generated for 11 species, but sequences for only three of them (Amblydorylaimusisokaryon (Loof 1975), P.paradoxus and E.signyensis) are supported by full morphological descriptions as per the modern taxonomic standards (Elshishka et al. 2015b, Elshishka et al. 2017, Kagoshima et al. 2019).
The review of the literature related to terrestrial nematodes from the Maritime Antarctic showed that the different parts are unevenly studied and three islands, Livingston (31 species), King George (28 species) and Signy (25 species) exhibited the richest nematode fauna (Fig. 7). Signy Island is the best studied Antarctic island with 12 new species described. This is due to the intensive studies on the nematode fauna in the 1970s and 1980s undertaken by the British Antarctic Survey (Spaull 1973a, Spaull 1973b, Spaull 1973c, Loof 1975, Maslen 1979a, Maslen 1979b, Caldwell 1981, Maslen 1981, Pickup 1988, Pickup 1990 etc.).
Figure 7.
Bar chart visualising the described species (left axis) and literature sources (right axis) per each island/site.
Discussion
Our knowledge of the nematode species diversity in the Maritime Antarctic is still insufficient and fragmented. The different study efforts at the various sites do not allow gaining a clear picture of trends in the diversity and distribution of nematode species in the target Antarctic Region. Yet, the analysis provided on the basis of species presence/absence data revealed several groups of sites with similar nematode fauna forming a latitudinal gradient (Fig. 1). The high level of endemism at both the species and genus level is a characteristic feature of the nematode fauna of the region as was mentioned above. This high degree of endemism can be explained by the long-term isolation and the harsh conditions of the region (Convey et al. 2008, Nielsen et al. 2011a). It has been suggested that the Antarctic terrestrial fauna might have survived glaciation in ice-free areas and some species might be remnants of the fauna of the Gondwana super-continent (Andrássy 1998, Maslen and Convey 2006, Chown and Convey 2016, Convey et al. 2020).
The physical isolation and harsh environment of Antarctic terrestrial ecosystems is the major reason for the difficult colonisation by non-native biota (Convey and Peck 2019). In recent decades, human visits and activities in the Antarctic have provided ways (e.g. cargo, vehicles, scientific equipment, fresh food, clothing, people) to overcome these barriers (Lee and Chown 2009, Hughes et al. 2010, Chwedorzewska et al. 2013, Adams et al. 2014). So far, the probability that introduced invertebrates will become established and spread is considered to be quite low; most of them are not able to complete the life cycle and establish a stable population outside the station (Chwedorzewska et al. 2013). Although these organisms cannot survive outside at present, they are potential colonisers, which could be established in the future following the climate warming (Convey and Peck 2019). Тhe four cosmopolitan nematode taxa (E.vulgaris, E.filiformis, G.villosa and P.armatus) also reported from the Maritime Antarctic are considered to be of non-native origin by Andrássy (1998). Due to the absence or scarcity of data on the morphology of these species, at present, their origin cannot be confirmed. Future studies using an integrated taxonomic approach (i.e. simultaneous molecular and morphological characterisation) of materials obtained from pristine areas may help clarify their status. The gap in knowledge of nematode diversity, both in terms of taxonomy and distribution, is essential when assessing the introduction of non-native species. Nematode species richness in the Maritime Antarctic, which is underestimated (Nielsen et al. 2011b) may be compromised with increasing human impact in Antarctica.
The risk to Antarctic biodiversity is not limited to the transfer of alien species originating from other regions of Earth, but also concerns the transfer of native or endemic species from one part of Antarctica to another where they are not part of the indigenous biota (Convey 2008, Hughes et al. 2019, Hughes et al. 2020). This risk is greater because such species are likely to adapt well to the new location, unlike most non-native species that have been transferred to Antarctica from elsewhere (Convey 2015). The transfer of species across natural biogeographic boundaries can affect endemism in these areas. Antarctica is one of the few regions on the Planet where such boundaries still exist (Convey 2008). The nematode faunas of the Maritime and the Continental Antarctic are characterised by their uniqueness, as no overlap at the species level of the two local faunas exists (Andrássy 1998, Maslen and Convey 2006, Convey et al. 2020). This is indicative of an ancient geographical divide between these areas (Andrássy and Gibson 2007) and led Chown and Convey (2006) to define the Gressitt Line, which is located across the base of the Antarctic Peninsula.
So far, there is no evidence for the transfer and establishment of nematode species from the Continental to the Maritime Antarctic. Some nematological reports have included data on the presence of species that are emblematic of the Continental Antarctic (Plectusmurrayi Yeates 1970 and P.frigophilus Kirjanova 1958) in the Maritime part, without morphological data (see Velasco-Castrillón et al. (2014a)). In our study, these records are not included as they are most likely due to misidentification.
Regarding the biotope/microhabitat distribution of the species, the incomplete and insufficient data do not allow a definite conclusion, taking into account also the lack of research in the more inaccessible areas of the Antarctic Peninsula and the islands. Most likely the micro biotope distribution pattern is similar to that shown in the study of the nematode fauna of Cape Chelyuskin in the Arctic (Chernov et al. 1979), where species show very low biotopic associations and most of them inhabit all possible microhabitats (i.e. the majority of species are polytopic); this is also a characteristic feature of other groups of organisms in the polar regions (Chernov et al. 1979).
The major life strategy of organisms living in extreme environments is the development of tolerance and plasticity and not lack of competition and specialisation, which is typical of other biomes (Convey 1996, Chernov et al. 2011).
Comparing the two parts of the Antarctic shows that the nematode studies in the Maritime Antarctic are less represented, whereas investigations in the Continental Antarctic have been more intensive. However, the latter are primarily related to ecology (Adams et al. 2014, Velasco-Castrillón et al. 2014a, Velasco-Castrillón et al. 2018) and have identified to date 34 species of soil nematodes (Velasco-Castrillón et al. 2014a). The smaller number of species in the Continental Antarctic is associated with the harsher and more unfavourable environmental conditions. This zone includes ecosystems with the simplest terrestrial fauna on the Planet, where even nematodes are absent (Convey and McInnes 2005, Convey 2017).
The two opposite polar regions of the Earth are unevenly studied with respect to soil nematodes (Peneva et al. 2009, Holovachov 2014a). Despite the fewer taxonomic studies of terrestrial nematodes in the Arctic, 391 species have been recorded there (Holovachov 2014a). Key geographical and ecological features of both regions, such as geological history, climate, landscape, dispersal barriers and vegetation are responsible for the lower nematode diversity in the Antarctic than in the Arctic (Nielsen and Wall 2013).
Studies that include molecular data for the nematodes in the Maritime Antarctic are too rare to provide valuable information regarding nematode diversity, phylogenetics and endemism (Elshishka et al. 2015b, Elshishka et al. 2017, Kagoshima et al. 2019). The taxonomic position of only three Antarctic dorylaimid species, A.isokaryon, P.paradoxus and E.signyensis, was reconsidered on the basis of morphological and molecular characteristics of 18S rDNA (SSU rDNA) and the D2-D3 expansion fragments of 28S rDNA (LSU rDNA) (Elshishka et al. 2015b, Elshishka et al. 2017) and two new endemic genera were proposed (Pararhyssocolpus and Enchodeloides).
To advance the understanding of phylogeny and phylogeography of Antarctic nematodes, studies are required of other genes with higher evolutionary rates than 18S rDNA, such as 28S rDNA, the internal transcribed spacer (ITS in the ribosomal RNA locus) or the mitochondrial cytochrome c oxidase subunit I (COI). These genes should be included in future taxonomic analyses of Antarctic nematodes (Kagoshima et al. 2019).
The application of integrated taxonomy and DNA barcoding will substantially assist in nematode diversity studies, phylogenetics and especially the recognition of cryptic species. Further, comprehensive molecular studies will provide valuable information on the patterns of species distribution and for gaining additional knowledge on evolutionary processes and biogeography of Antarctic nematodes.
The scant studies of polar regions, in particular of the Maritime Antarctic, demand more intensive sampling and research, especially in the territories that have so far remained unexplored, in order to give a clearer and more adequate view of species diversity and trends in their microhabitat and geographical distribution. Therefore, further efforts aiming at targeted and systematic integrative studies are needed.
Supplementary Material
Acknowledgements
This study was funded by the Programme for career development of young scientists, Bulgarian Academy of Sciences, the project №80-10-214, National Centre for Polar Studies, Sofia University "St. Kliment Ohridski” and ANIDIV4, Bulgarian Academy of Sciences. The authors are thankful to Prof. Aneta Kostadinova from the Institute of Biodiversity and Ecosystem Research, Bulgarian Academy of Sciences, for the critical reading of the manuscript and helpful suggestions.
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