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Tania Nara Bezerra, Nic Smol, Renata M. S. Alves, Ellen Pape. (2026). Review of Halalaimus (Nematoda, Enoplida, Oxystominidae) and description of two new species from the abyssal nodule-bearing Clarion-Clipperton Zone. Zookeys. 1284.187212.
566502
Tania Nara Bezerra, Nic Smol, Renata M. S. Alves, Ellen Pape
2026
Review of Halalaimus (Nematoda, Enoplida, Oxystominidae) and description of two new species from the abyssal nodule-bearing Clarion-Clipperton Zone
Zookeys
1284.187212.
Publication
Available for editors  PDF available [request]
This review examines the geographical distribution, morphology, and taxonomy of the genus Halalaimus. It consolidates existing descriptions of the currently 102 valid species, resolves nomenclatural inconsistencies, and introduces two new species: Halalaimus marcosi sp. nov. and Halalaimus pierri sp. nov. Halalaimus marcosi sp. nov. is characterised by its large body, papilliform inner labial sensilla, outer labial setae shorter than cephalic setae spaced one head diameter apart, and a conic-filiform tail with the posterior two-thirds filiform. Halalaimus pierri sp. nov. is distinguished by its very strongly tapered anterior end, indistinct inner labial sensilla, and outer labial setae longer than the cephalic setae. Long-standing challenges in species-level identification, mainly due to incomplete descriptions and confused nomenclature, are addressed through a detailed reassessment that corrects duplicated names, assigns replacement names to misattributed species, and incorporates previously overlooked taxa. Halalaimus borealis Gagarin, 2020 is considered a junior homonym and is renamed H. gagarini nom. nov. This study provides interactive global distribution maps for all Halalaimus species, a table summarising species ranges and habitat information, and a table listing 36 morphometric and morphological traits per species. Updated identification keys are presented, including a new key for deep-sea species, offering a practical tool for nematode taxonomists and identifiers worldwide.
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2026-07-03 22:41:52Z
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2026-07-20 08:49:38Z
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Halalaimus de Man, 1888 (additional source)
Halalaimus (Pachyodora) Wieser, 1953 accepted as Halalaimus de Man, 1888 (additional source)
Halalaimus absconditus Bussau, 1993 (additional source)
Halalaimus abyssus Bussau, 1993 (additional source)
Halalaimus aciculus Gagarin & Thanh, 2014 (additional source)
Halalaimus acuminatus (Cobb, 1933) Hope & Murphy, 1972 (additional source)
Halalaimus aedificandistudiosus Bussau, 1993 (additional source)
Halalaimus alatus Timm, 1952 (additional source)
Halalaimus algeriensis Coomans & Jacobs, 1983 (additional source)
Halalaimus americanus Keppner, 1992 (additional source)
Halalaimus amphidellus Vitiello, 1970 (additional source)
Halalaimus amphistrius Vitiello, 1970 (additional source)
Halalaimus anne Sergeeva, 1972 (additional source)
Halalaimus anulatus Wieser, 1960 (additional source)
Halalaimus arundinaceus Bussau, 1993 (additional source)
Halalaimus bayensis Keppner, 1992 (additional source)
Halalaimus borealis (Steiner, 1916) Lorenzen, 1981 (additional source)
Halalaimus borealis Gagarin, 2020 accepted as Halalaimus gagarini Bezerra, Smol, Alves & Pape, 2026 (additional source)
Halalaimus brachyaulax Mawson, 1958 (additional source)
Halalaimus brevispiculum Sergeeva, 1973 (additional source)
Halalaimus brimi Keppner, 1992 (additional source)
Halalaimus bulbocaudatus Keppner, 1992 (additional source)
Halalaimus capitulatus Boucher, 1977 (additional source)
Halalaimus caroliniensis Chitwood, 1936 (additional source)
Halalaimus ciliocaudatus Allgén, 1932 (additional source)
Halalaimus cirrhatus Gerlach, 1953 (additional source)
Halalaimus climactericus Wieser, 1953 (additional source)
Halalaimus comatus Wieser, 1953 (additional source)
Halalaimus cubanus Andrássy, 1973 (additional source)
Halalaimus curvicaudatus Juario, 1974 (additional source)
Halalaimus cuthemon Blood et al., 1992 (status source)
Halalaimus delamarei Vitiello, 1970 (additional source)
Halalaimus deseadensis Pastor de Ward, 1998 (additional source)
Halalaimus diacros Mawson, 1958 (additional source)
Halalaimus dimorphus Turpeenniemi, 1997 (additional source)
Halalaimus diplocephalus Filipjev, 1927 (additional source)
Halalaimus dolgovi Alekseev & Linnik, 1994 (additional source)
Halalaimus droebachiensis Allgén, 1931 (additional source)
Halalaimus durus Gagarin & Thanh, 2004 (additional source)
Halalaimus egregius Bussau, 1993 (additional source)
Halalaimus estuarii Sinha, Choudhury & Baqri, 1985 (additional source)
Halalaimus filicaudatus Allgén, 1959 (additional source)
Halalaimus filicollis Timm, 1961 (additional source)
Halalaimus filicorpus Vitiello, 1970 (additional source)
Halalaimus filiformis (Allgen, 1929) Schuurmans Stekhoven, 1935 (additional source)
Halalaimus filum Gerlach, 1962 (additional source)
Halalaimus fletcheri Mawson, 1958 (additional source)
Halalaimus florescens Gerlach, 1967 (additional source)
Halalaimus floridanus Keppner, 1992 (additional source)
Halalaimus gagarini Bezerra, Smol, Alves & Pape, 2026 (new combination reference)
Halalaimus gerlachi Keppner, 1992 (additional source)
Halalaimus gidanensis Nasira & Turpeenniemi, 2002 (additional source)
Halalaimus gracilis de Man, 1888 (additional source)
Halalaimus horridus Gerlach, 1956 (additional source)
Halalaimus isaitshikovi (Filipjev, 1927) Schuurmans Stekhoven, 1935 (additional source)
Halalaimus jaltensis Sergeeva, 1973 (additional source)
Halalaimus leptoderma Platonova, 1971 (additional source)
Halalaimus leptosoma (Southern, 1914) Schuurmans Stekhoven, 1935 (additional source)
Halalaimus lineatoides Timm, 1961 (additional source)
Halalaimus lineatus Timm, 1961 (additional source)
Halalaimus longamphidus Huang & Zhang, 2005 (additional source)
Halalaimus longicauda accepted as Halalaimus longicaudatus (Filipjev, 1927) Schneider, 1939 (additional source)
Halalaimus longicaudatus (Filipjev, 1927) Schneider, 1939 (additional source)
Halalaimus longicollis Allgén, 1932 (additional source)
Halalaimus longinquus Bussau, 1993 (additional source)
Halalaimus longipharynx Gagarin & Nguyen Vu Thanh, 2018 (additional source)
Halalaimus longisetosus Hopper, 1963 (additional source)
Halalaimus longistriatus Timm, 1961 (additional source)
Halalaimus lutarus Vitiello, 1970 (additional source)
Halalaimus luticolus Timm, 1961 (additional source)
Halalaimus macquariensis Mawson, 1958 (additional source)
Halalaimus marcosi Bezerra, Smol, Alves & Pape (original description)
Halalaimus marri Mawson, 1958 (additional source)
Halalaimus meyersi Wieser & Hopper, 1967 (additional source)
Halalaimus minimus Gagarin, 2016 (additional source)
Halalaimus minor Gagarin & Thanh, 2004 (additional source)
Halalaimus minusculus Tchesunov, 1978 (additional source)
Halalaimus monstrocaudatus Vitiello, 1970 (additional source)
Halalaimus nigrilapidarius Boucher, 1977 (additional source)
Halalaimus oblongus Bussau, 1993 (additional source)
Halalaimus orientalis Gagarin, 2016 (additional source)
Halalaimus pachyderma (Filipjev, 1927) Schuurmans Stekhoven, 1935 (additional source)
Halalaimus pachydermatus (Cobb, 1920) Schneider, 1939 (additional source)
Halalaimus pachyodoroides Vitiello, 1970 (additional source)
Halalaimus papillifer Gerlach, 1956 (additional source)
Halalaimus paracomatus Keppner, 1992 (additional source)
Halalaimus parafletcheri Keppner, 1992 (additional source)
Halalaimus parapapillifer Pastor de Ward, 1986 (additional source)
Halalaimus parasetosus Pastor de Ward, 1986 (additional source)
Halalaimus parvulus Gagarin & Thanh, 2018 (additional source)
Halalaimus parvus Chitwood, 1936 (additional source)
Halalaimus pierri Bezerra, Smol, Alves & Pape (original description)
Halalaimus ponticus Filipjev, 1922 (additional source)
Halalaimus praestans Bussau, 1993 (additional source)
Halalaimus rectispiculatus (Platonova, 1971) Gerlach & Riemann, 1974 (additional source)
Halalaimus relatus Gerlach, 1967 (additional source)
Halalaimus sagarensis Sinha, Choudhury & Baqri, 1986 (additional source)
Halalaimus sapeloi Wieser, 1964 (additional source)
Halalaimus sarsi Gerlach, 1967 (additional source)
Halalaimus scleratus Timm, 1952 (additional source)
Holotype UGMD UGMD_104484, geounit Clarion Clipperton Zone, identified as Halalaimus marcosi Bezerra, Smol, Alves & Pape
Holotype UGMD UGMD_104489, geounit Clarion Clipperton Zone, identified as Halalaimus pierri Bezerra, Smol, Alves & Pape
 New combination

"Gagarin (2020) described a new species under the name Halalaimus borealis Gagarin, 2020, apparently overlooking ... [details]

 Spelling

H. papillifer (a valid species) is written on page 8 and Table 3 of Bezerra et al (2026) instead of H. ... [details]

 Spelling

Misspelled in Bezerra et al (2026) page 8 and Table 3: written H. papillifer instead of H. parapapillifer.  [details]

 Status

"Gerlach and Riemann (1974) listed three species as nomina nuda: H. annulatus Wieser, 1960; H. sapeloi Wieser, 1964 ... [details]

 Status

"Gagarin (2020) described a new species under the name Halalaimus borealis Gagarin, 2020, apparently overlooking ... [details]

 Status

"Blood et al. (1992) listed three species H. sapeloi, listed by Wieser (1964) and regarded as nomen nudum by ... [details]

 Status

"Blood et al. (1992) listed three species H. sapeloi, listed by Wieser (1964) and regarded as nomen nudum by ... [details]

 Status

"Gerlach and Riemann (1974) listed three species as nomina nuda: H. annulatus Wieser, 1960; H. sapeloi Wieser, 1964 ... [details]

 Status

"Blood et al. (1992) listed three species H. sapeloi, listed by Wieser (1964) and regarded as nomen nudum by ... [details]

 Taxonomic remark

Male described by Gerlach, 1957 - Page 6 and 7. H. ciliocaudatus sensu Gerlach, 1957 a species ... [details]

 Taxonomic remark

"The following redescriptions: Allgén (1947) provides only a few measurements without information on the anterior ... [details]

 Taxonomic remark

"In the original description, H. isaitshikovi Filipjev, 1927 is unique in possessing a distinct protrusion of the ... [details]

 Taxonomic remark

"In the original description of H. longicaudatus Filipjev, 1927, the species is characterised by a small preanal ... [details]

 Taxonomic remark

"H. setosus Timm, 1961: Pastor de Ward (1998) described this species with four cephalic setae instead of six, ... [details]

 Taxonomic remark

Specimen is incomplete and the description insufficient.  [details]