Mar Biodiv
DOI 10.1007/s12526-014-0226-3
DIVERSITY OF MARINE MEIOFAUNA ON THE COAST OF BRAZIL
The state of the art of Xyalidae (Nematoda, Monhysterida)
with reference to the Brazilian records
Virág Venekey & Paula F. Gheller & Tatiana F. Maria & Marco C. Brustolin &
Noelia Kandratavicius & Danilo C. Vieira & Simone Brito & Guilherme S. Souza &
Gustavo Fonseca
Received: 28 August 2013 / Revised: 11 March 2014 / Accepted: 1 April 2014
# Senckenberg Gesellschaft für Naturforschung and Springer-Verlag Berlin Heidelberg 2014
Abstract In the current study we provide a reviewed list of
valid genera and species of Xyalidae, a widespread family of
mostly marine free-living nematodes. Comments are added
about the historic background and taxonomic situation of the
family, all valid genera and, when necessary, diagnostic characters are given. Additionally, information about distribution and
geographical location of species recorded along the Brazilian
coast is provided. Our review recognized 46 valid genera, 450
valid species and 73 descriptions without enough morphological
information for identification (species inquerendae). Nearly
80 % of the species inquerendae belong to Daptonema and
Theristus. To avoid homonymies, two Daptonema species were
renamed, Daptonema biwaensis (Tsalolikhin, 2002) new name
(former Mongolotheristus timoshkini) and Daptonema
vietnamensis (Gagarin and Thu, 2008) new name (former
D. curvatum sensu Gagarin and Thu, 2008). Cenolaimus
sapeloensis is transferred to Xyala sapeloensis comb. nov.
Along the Brazilian coast 28 genera and 41 species have been
recorded. The species Elzalia floresi, Metadesmolaimus tersus,
Paramonohystera stricta, Pseudosteineria scopae,
Rhynchonema cemae, R. veronicae, Steineria ericia, S.
marcorum, S. pavo, S. tripartita, Theristus acribus, T. flevensis,
V. Venekey (*)
Instituto de Ciências Biológicas, Universidade Federal do Pará
(UFPA), Av. Augusto Corrêa, 01, Guamá, CEP: 66075-110 Belém,
PA, Brazil
e-mail: venekey@ufpa.br
D. C. Vieira : S. Brito
Centro de Biologia Marinha da Universidade de São Paulo
(CEBIMAR-USP), Rod. Manuel Hyppólito do Rego. Km 131,5,
P. F. Gheller
Instituto Oceanográfico, Universidade de São Paulo (USP), Praça do
Oceanográfico, 191, Cidade Universitária, CEP: 05508-120 São
Paulo, SP, Brazil
e-mail: paulafgheller@gmail.com
CEP: 11600-000 São Sebastião, SP, Brazil
T. F. Maria
Departamento de Ecologia e Recursos Marinhos, Universidade
Federal do Estado do Rio de Janeiro (UNIRIO), Av. Pasteur, 458,
Urca, CEP:22290-240 Rio de Janeiro, RJ, Brazil
e-mail: tatiana_fabricio@yahoo.com.br
S. Brito
e-mail: simonebrito134@gmail.com
M. C. Brustolin
Centro de Estud os do Mar, Universidade Federal do Paraná (UFPR),
Avenida Beira Mar, s/n., Pontal do Sul, CEP 83255-976PO Box 61,
Pontal do Paraná, PR, Brazil
e-mail: marcobrustolin@gmail.com
N. Kandratavicius
Laboratorio de Oceanografía y Ecología Marina, Facultad de
Ciencias, Universidad de La Republica, Igua 4225, CP:
11400 Montevideo, Uruguay
e-mail: nkandra19@gmail.com
D. C. Vieira
e-mail: vieiradc@yahoo.com.br
G. S. Souza
Centro de Ciências Biológicas, Departamento de Zoologia,
Universidade Federal de Pernambuco (UFPE), Av. Prof. Moraes
Rego s/n, Cidade Universitária, CEP: 50670-920 Recife, PE, Brazil
e-mail: guilherme_georgi@hotmail.com
Present Address:
G. Fonseca
Instituto do Mar, Universidade Federal de São Paulo, Av. Alm.
Saldanha da Gama, 89, Ponta da Praia, CEP: 11030-400 Santos,
SP, Brazil
e-mail:
gfonseca@unifesp.br
gfonseca@unifesp.br
Mar Biodiv
T. macroflevensis, T. pertenuis, T. stranus, Trichotheristus
heterus, T. setosus and Zygonemella striata have the Brazilian
coast as the type locality. Among all species, three occurred
across three geographic regions, while the large majority was
restricted to one. Xyalidae is typically encountered in oceanic
sandy beaches, with only the species belonging to Daptonema,
Theristus, Trichotheristus and Zygonemella being recorded in
estuarine sediments. This observation suggests that the colonization of inland waters occurred multiple times along the evolutionary history.
Keywords Marine nematodes . Brazilian coast . Xyalidae
Introduction
Among the metazoans, nematodes are the most abundant organisms in all kind of benthic habitats (four of every five
multicellular animals on the planet are nematodes) so they can
hardly be ignored (Bongers and Ferris 1999). Hugot et al.
(2001) considered 26,646 known species of nematodes and
estimated the number of existing ones in 1 million. Although
in the last decades, many studies have focused on the systematics within the phylum (Lorenzen 1994; De Ley and Blaxter
2004), less attention has been given to the family level (but see
for instance Platonova and Mokievsky 1994 for Ironidae;
Decraemer et al. 1997 for Draconematidae; Fonseca and
Decraemer 2008 for Monhysteridae). In this study, we review
the Xyalidae, a family of mostly marine free-living nematodes.
Species descriptions of Xyalidae, as well as of most marine
nematodes, were largely made in the past century (e.g., Allgén
1929; Cobb 1920; Gerlach 1957a; Lorenzen 1977). Many of these
descriptions were based on few specimens (one to three individuals) or even in juveniles or females only, with relatively few
characters of taxonomic value. Such poor taxonomical accounts
made in the past are today impediments for the taxonomical
progress within the taxon. At the moment, we do not know which
are the valid species and genera belonging to Xyalidae. For instance, some genera such as Theristus Bastian, 1865 and
Daptonema Cobb, 1920, have each more than 100 nominal species
and a list of synonyms as large as the list of valid species (Gerlach
and Riemann 1973). Despite of this impediment, new genera and
species are still being described within the family (e.g., Armenteros
et al. 2009, 2010; Huang and Zhang 2010; Huang and Xu 2013).
Some authors already attempted to review taxonomically the
Xyalidae seeking for morphological relationships within the
group. Nicholas and Trueman (2002) recognized 33 genera in
their cladistic analysis whereas Fonseca and Bezerra (2013)
accepted 44 genera and also mentioned the number of valid
species of each genus. The total number of valid species also
varies enormously within Xyalidae. Since the checklist from
Gerlach and Riemann (1973), later updated by Lorenzen (1977),
no effort has been done to assemble the number of valid species
within this family. Many publications about nematodes (e.g.,
Castro et al. 2006; Botelho et al. 2007; Neres et al. 2010;
Fonseca and Bezerra 2013) have used large databases as
NeMys (Deprez et al. 2005), WoRMS (Appeltans et al. 2012)
or The Biology Catalog (available in https://insects.tamu.edu/
research/collection/hallan), which provide the numbers of
genera and species, but these compilations have not a
consensus in numbers of taxa. For instance, when looking for
the number of valid genera in Xyalidae, we found 41 in NeMys,
47 in WoRMS and 42 in The Biology Catalog.
In the current study, we provide a reviewed list of valid
genera and species of Xyalidae. Comments are added about
the situation of each genus, and, when necessary, diagnostic
characters are given. Additionally, we provide information
about distribution and geographical location of species recorded along the Brazilian coast.
Methods
The works of Gerlach and Riemann (1973) and Lorenzen
(1977) were used as the starting point of our review. All
taxonomical studies published thereafter were considered in
the present review. The new records were then double checked
with the information available on the three main virtual checklists: NeMys, WoRMS and The Biology Catalog databases.
The most relevant references and comments concerning
changes in the taxonomical position of species are given in the
section of each genus. A valid species list is presented for each
valid genus in which the type species is underlined and for each
species the sampling location of the original description is given
in parentheses. Species inquirendae, nomen nudum and incertae
sedis are cited in separate lists following the valid species list, and,
when necessary, names were adapted to attend to the Latin rules.
Occurrence of Xyalidae nematodes along the Brazilian coast is
provided. This information was retrieved from Venekey et al.
(2010) and other articles, as well as from PhD, M.Sc and
undergraduation theses. The genera and species names are currently
reported taking into account synonyms and changes in nomenclature. All species records were used to elaborate a map of occurrence
in Brazil with indication of author, year, locality and habitat.
Results and discussion
Historic background and general comments about Xyalidae
The Xyalidae Chitwood, 1951 occurs in nearly all types of marine
environments and frequently among the most abundant taxa. In
early classifications genera and species of Xyalidae were positioned in Monhysteridae de Man, 1876, but later Lorenzen (1978)
split up this family into Monhysteridae and Xyalidae. Lorenzen
(1994) established the holophyly of Xyalidae by the following
Mar Biodiv
holapomorphy: within species the anterior gonad is constantly to
the left of the intestine and the posterior gonad constantly to
the right of it. Additional features are: (1) cuticle always striated, (2) 6+4 sensillas almost always situated at the same level
and sometimes with additional cephalic setae and (3) stoma
usually funnel-shaped. Lorenzen (1978) and Lorenzen (1994)
recognized 34 valid genera within the family: Ammotheristus,
Amphimonhystera, Amphimonhystrella, Cenolaimus, Cobbia,
Dactylaimus, Daptonema, Echinotheristus, Elzalia, Filipjeva,
Gnomoxyala, Gonionchus, Hofmaenneria, Linhystera,
M e g a l a m p h i s , M e t a d e s m o l a i m u s , O m i c ro n e m a ,
Paramonohystera, Promonhystera, Pseudosteineria,
Retrotheristus, Rhynchonema, Scaptrella, Sphaerotheristus,
Spiramphinema, Steineria, Stylotheristus, Theristus,
Trichotheristus, Valvaelaimus, Wieserius, Xenolaimus, Xyala
and Zygonemella. Lorenzen (1994) considered Austronema,
Buccolaimus and Pulchranemella as dubious genera, following
Hope and Murphy (1972).
The description of Prorhynchonema Gourbault, 1982 and
Mongolotheristus Tsalolikhin, 1985 elevated the number of genera to 36. The genus Trichotheristus was later considered invalid
by Tchesunov (1990a). The description of Marisalbinema
Tchesunov, 1990 and Parelzalia Tchesunov, 1990 increased the
number of genera in Xyalidae to 37. Bussau (1993) described in
his PhD thesis three new genera: Capsula, Enchonema and
Manganonema. Fonseca et al. (2006) redescribed
Manganonema but Capsula and Enchonema remained in the
grey literature. Nevertheless, most nematologists recognize the
new taxa published by Bussau (1993) as valid given the quality
of his descriptions and the widespread divulgence. Andrássy
(1993) raised Mesotheristus, a subgenus of Theristus, to genus
level. Another three genera, Corononema Nicholas and Stewart,
1995; Gulanema Nicholas and Stewart, 1995 and Robustnema
Nicholas, 1996 were added. Later, Nicholas and Trueman (2002)
in a cladistics analysis of Xyalidae, did not consider Cenolaimus,
Dactylaimus, Hofmaenneria and Wieserius (incorrectly referred
as Wieseria by them) as valid because of the lack of adequate
descriptions and also questioned the validity of Spiramphinema
and Trichotheristus. They also omitted Mesotheristus and
Mongolotheristus without explanation. Meanwhile,
Sacrimarinema Shoshin, 2001; Arabanema Turpeenniemi,
Nasira and Maqbool, 2001; Dactylaimoides Blome, 2002,
Paragonionchus Blome, 2002 and Pseudechinotheristus
Blome, 2002 have been added to the family Xyalidae. Blome
(2002) considered Cenolaimus Cobb, 1933 invalid because of
insufficient description of the type species C. supersentiens
Cobb, 1933 and suggested that C. sapeloensis Nichols, 1979
belongs to Xyala or Gonionchus. The last additions to Xyalidae
were: Paramphimonhystrella Huang and Zhang, 2006;
Cienfuegia Armenteros, Vincx and Decraemer, 2009 and
Guitartia Armenteros, Vincx and Decraemer, 2010. Recently
Coomans and Eyualem-Abebe (2006) synonymized
Mesotheristus and Mongolotheristus with Daptonema and
Huang and Zhang (2006) recognized again Trichotheristus as a
valid genus.
The last revision of Xyalidae was published by Fonseca
and Bezerra (2013) which recognized 44 valid genera. These
authors considered again Hofmaenneria and Spiramphinema
as valid and synonymized Arabanema with Theristus. These
authors did not include Capsula and Guitartia but we considered them valid because both have well detailed and wellillustrated descriptions. Another genus has a situation, which
also should be commented on, is Megalamphis (Timm, 1961)
De Coninck, 1965. This genus was originally described with
the name Macramphis by Timm (1961) in the family
Sphaerolaimidae and renamed later for Megalamphis to avoid
homonymy with another already existing genus of terrestrial
nematode described previously by Altherr (1950). Since then
the genus is considered as a taxon of Sphaerolaimidae by
some authors (Timm 1961; Andrássy 1976) and of Xyalidae
by others (Lorenzen 1978; Nicholas and Trueman 2002). In
the present work we agree with the original description by
Timm (1961) and the last revision of Xyalidae (Fonseca and
Bezerra 2013) that considered it as a genus of
Sphaerolaimidae. Therefore, Megalamphis does not appear
in the list of valid genera and species and today the family
Xyalidae is characterized by 46 valid genera. The list of
genera and species is given in the Appendix.
Comments about valid genera of Xyalidae
Genus Ammotheristus Lorenzen, 1977
The genus Ammotheristus was established when in a revision of the Xyalidae family, based on a critical analysis of 56
species from North Sea and Baltic, Lorenzen (1977) found
that Amphimonhystera helgolandica Riemann, 1967 and
Amphimonhystera subtilis Lorenzen, 1972 differ from the
type species A. anechma in their colorless cuticle, conical tail
without terminal setae and in having a glandular posterior
widening of the pharynx (Lorenzen 1977). As a consequence,
Lorenzen (1977) established the genus Ammotheristus accommodating both A. helgolandicus and A. subtilis.
Number of valid species: 2.
Genus Amphimonhystera Allgén, 1929
The genus Amphimonhystera was established by Allgén
(1929) to accommodate the monhysterid nematode species
Monohystera anechma, described previously by Southern
(1914) in a survey of marine fauna of Ireland. Later another
new species, A. paranechma Allgén, 1935, was added.
Riemann (1967) reviewed the genus Amphimonhystera and
added a third species A. helgolandica. Lorenzen (1972) described a new species of Amphimonhystera, A. subtilis, however subsequently he concluded that A. subtilis and
A. helgolandica differ from the type species A. anechma
(Southern, 1914) Lorenzen, 1977 and established the genus
Ammotheristus Lorenzen, 1977. Posteriorly Fadeeva (1984),
Mar Biodiv
Bussau (1993) and Guo and Warwick (2001) described three
more species: Amphimonhystera galea, A. bella and
A. circula, respectively. More recently, Tchesunov and
Mokievsky (2005) described another three species: A.
marisalbi, A. molloyensis and A. pallida, and they also added
a redescription of A. galea. These authors also emended the
generic diagnosis and provided an identification key.
Number of valid species: 7.
Genus Amphimonhystrella Timm, 1961
The genus Amphimonhystrella was established when, in
the revision of free-living marine nematodes of the Bay of
Bengal, Timm (1961) described the new species
Amphimonhystrella megastoma Timm, 1961. This genus
was erected based on the large ovoid double-walled amphidial
fovea and on the deep and conical stoma with sclerotized
walls surrounded by the pharyngeal tissue. Later A. unita
Lorenzen, 1977 and A. bullacauda Tchesunov and Miljutina,
2005 were added. Tchesunov and Miljutina (2005) also
emended the generic diagnosis and provided an identification
key. More recently, Gagarin and Thanh (2009) described
A. parva.
Number of valid species: 4.
Genus Capsula Bussau, 1993
The monospecific genus Capsula was described by Bussau
(1993). In the last revision of Xyalidae made by Fonseca and
Bezerra (2013) this genus was omitted without explanation.
Nevertheless, we considered it a valid genus given its detailed
description and illustration. Within Xyalidae, Capsula is distinguished from the remaining taxa by showing a bipartite
head capsule strongly sclerotized. The wide buccal cavity
resembles those from Xyala, Gonionchus and Omicronema.
Additionally the three caudal glands open through a single
duct, and females may show a spermatheca.
Number of valid species: 1.
Genus Cienfuegia Armenteros, Vincx and Decraemer,
2009
The monospecific genus Cienfuegia was recently described
from the Caribbean Sea. The species has a characteristic
bipartite buccal cavity similar to the observed in
Diplolaimella Allgén, 1929 and Diplolaimelloides Meyl,
1954 from the family Monhysteridae. However Cienfuegia
has the anterior gonad constantly placed to the left of the
intestine. According to the position of the gonad this genus
has been positioned within Xyalidae.
Number of valid species: 1.
Genus Cobbia de Man, 1907
The genus Cobbia was erected by De Man (1907) with the
type species Cobbia trefusiaeformis, latter redescribed by
Lorenzen (1977). This genus differs from the other genera in
the family Xyalidae by a buccal cavity with three teeth and
setiform labial sensilla. Huang and Zhang (2010) comment
that the species in this genus can be distinguished from each
other by the combination of the size of the dorsal tooth, the
diameter and distance from the anterior end of the amphid, the
shape of spicules and gubernaculum and the length of the
labial and cephalic setae. Lo Russo and Pastor de Ward (2012)
described the most recent species, Cobbia macrodentata, and
also provided an identification key for all valid species.
Number of valid species: 8.
Genus Corononema Nicholas and Stewart, 1995
The genus Corononema was described based on two species, C. parvum and C. thai, from Australia and Thailand,
respectively. Corononema has a unique head which appears to
be crowned by the incised lips, set off from the cylindrical
buccal region by a deep groove. C. thai differs from
C. parvum by possessing a shorter and broader head.
Number of valid species: 2.
Genus Dactylaimoides Blome, 2002
The monospecific genus Dactylaimoides is known only
from the sandy beaches of Eastern Australia. Diagnose as in
Blome (2002) and Fonseca and Bezerra (2013).
Number of valid species: 1
Genus Daptonema Cobb, 1920
The genus Daptonema was described by Cobb (1920) with
type species Daptonema fissidens. Wieser (1956) consider this
genus a subgenus of Theristus Bastian, 1865. Lorenzen (1977)
agreed with Cobb (1920) and raised Daptonema to genus status
again and considered four other subgenera of Theristus
(Cylindrotheristus De Coninck, 1965; Mesotheristus
Chitwood and Murphy, 1964; Pseudotheristus Chitwood and
Murphy, 1964 and Spirotheristus Timm, 1961) and the genus
Tubolaimus Allgén, 1929 as synonyms of Daptonema. The
differences between Daptonema and Theristus are the shape
of the tail and the presence of terminal setae at the tail tip:
conico-cylindrical with terminal setae in the former, conical
without terminal setae in the later. Daptonema is currently the
genus with the highest number of valid and species inquerendae
within Xyalidae.
In the present study two species within genus Daptonema
were renamed. D. curvatum sensu Gagarin and Thu, 2008 was
renamed for D. vietnamensis because there was already a
species with this name within the genus (D. curvatum
Gerlach, 1956 described originally for Kiel Bay as Theristus
curvatus). Furthermore, as Mongolotheristus was synonymized with Daptonema by Coomans and Eyualem-Abebe
(2006), the new name Daptonema biwaensis was given for
Mongolotheristus timoshkini Tsalolikhin, 2002 to avoid homonymy with the previously existing Daptonema timoshkini
Gagarin, 2001.
Number of valid species: 116.
Genus Echinotheristus Thun and Riemann, 1967
The genus Echinotheristus and the two known species
were described for the sublittoral coarse sands of the North
Sea. They are distinguished from the other genera by having a
bladder-like precloacal supplements and the cuticle covered
by transverse rows of small spines. Lorenzen (1977) added
Mar Biodiv
some detailed comments on both species observing additional
specimens from the same area.
Number of valid species: 2.
Genus Elzalia Gerlach, 1957
The genus Elzalia was described by Gerlach (1957b) from
the São Paulo region (southern coast of Brazil) with the type
species E. floresi. Later Timm (1961) described the genus
Megalolaimus which was synonymized with Elzalia by
Hope and Murphy (1972). Vitiello (1971) described a new
species: E. mediterranea from the Mediterranean. Hope and
Murphy (1972) synonymized Megalolaimus Timm, 1961
with Elzalia, transferring M. speculifer to Elzalia. Timm’s
species description was based on a single female. Gerlach
and Riemann (1973) transferred Filipjeviella tenuis Allgén,
1959, to Elzalia which was also described from a single
female. Castillo-Fernandez and Lambshead (1990) revised
the genus Elzalia and described three new species from the
oil producing zone of Campeche Sound in the Gulf of Mexico:
E. polli, E. federici and E. kimae. In that study only species
with males were considered as valid, therefore, E. speculifer
(Timm, 1961) Hope and Murphy, 1972 and E. tenuis (Allgén,
1959) Gerlach and Riemann, 1973 were not included.
Recently, E. gerlachi and E. striatitenuis, were described from
the Yellow Sea, China, by Zhang and Zhang (2006) and
E. bipectinella and E. tubercullata from Thailand by Hope
and Aryuthaka (2009). Particularly the work of Zhang and
Zhang (2006) provides a pictorial and tabular key for identification of the valid species.
Number of valid species: 10.
Genus Enchonema Bussau, 1993
The monospecific genus Enchonema was described by
Bussau (1993) for the Eastern Pacific Basin of Peru (Central
Pacific Ocean). The only species Enchonema umbrosum was
found around 4,000 m depth in fine sediments, and males,
females and juveniles are described. Diagnose of the genus is
given in Fonseca and Bezerra (2013) (incorrectly spelled as
Echonema).
Number of valid species: 1
Genus Filipjeva Ditlevsen, 1928
The genus Filipjeva (in some works appearing as
Filipjevia) was described by Ditlevsen (1928) from East
Greenland based on F. artica. The emendation of F. artica
by Allgén (1929) was considered invalid by Chitwood (1960).
The genus Filipjeva was considered by Kreis (1934) and De
Coninck (1965) as belonging to the family Oncholaimidae but
Gerlach and Riemann (1973) assumed closer relationship with
Paramonohystera subgenus Leptogastrella Cobb, 1920.
F. meridionalis was descripted by Kreis (1932) from
Indonesia based only in two males and Schuurmans
Stekhoven (1950) described F. mediterranea with a single
juvenile from the Mediterranean. Vitiello (1970) described
F. parameridionalis from the Mediterranean and also revised
the genus recognizing all four species known until then as
valid. Posteriorly, F. crucis was described from Antarctica by
Blome and Schrage (1985). Tchesunov (1988) described
F. filipjevi and F. teratospicula from the Kandalaksh Bay of
the White Sea, off the Karelian coast (Russia). Here we
consider F. mediterranea as species inquirenda because of
its incomplete description.
Number of valid species: 6.
Genus Gnomoxyala Lorenzen, 1977
The monospecific genus Gnomoxyala was described by
Lorenzen (1977) in the revision of the family to accommodate
a new species from Baltic Sea. This species has a finely
striated cuticle, amphidial fovea absent and a cylindrical buccal cavity.
Number of valid species: 1.
Genus Gonionchus Cobb, 1920
The genus Gonionchus was described from a tide pool of
New Hamsphire, USA, and it is very similar to Xyala, but
differs from it by the presence of hyaline lips extended anteriorly. The species belonging to this genus was divided into
two groups according to Vincx (1986): (1) without longitudinal ridges along cuticle (G. africanus, G. australis, G.
cumbraensis, G. inaequalis, G. intermedius, G. paravillosus,
G. villosus) and (2) longitudinal ornamentations present
(G. alastairi, G. arabica, G. heipi, G. longicaudatus, G.
sensibilis). The species G. latentis was omitted in Vincx
(1986) without explanation and G. ecuadorensis is considered
nomen nudum since it is described in a PhD thesis (Procel
2007). Procel (2007) provides an identification key for all the
valid species within this genus.
Number of valid species: 13.
Genus Guitartia Armenteros, Vincx and Decraemer,
2010
Described from the Carribean Sea, the monospecific
Guitartia is the newest genus within Xyalidae. It is characterized by three long tooth-like structures in the stegostom,
second and third circle of anterior sensilla separated and
posterior genital branch of the female restricted to a long
post-vulvar sac. In the last revision of Xyalidae made by
Fonseca and Bezerra (2013) this genus was omitted without
explanation but, as it has a detailed diagnosis, we recognize it
as valid.
Number of valid species: 1.
Genus Gullanema Nicholas and Stewart, 1995
The monospecific genus Gullanema was described from
Australia and it can be differentiated from all other genera
within the family by having a very characteristic head that
narrows sharply to a cylindrical “turret-like” region surrounding the buccal cavity.
Number of valid species: 1.
Genus Hofmaenneria Gerlach and Meyl, 1957
The genus Hofmaenneria was erected to accommodate two
species whose systematic affiliation had not been clarified
sufficiently: Cylindrolaimus brachystoma Hofmmaner in
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Hofmanner and Menzel, 1914 and C. niddensis Skwarra,
1921. Schneider (1940) was the first to propose the genus
Hofmaenneria to include C. brachystoma and C. niddensis
but did not give a generic description. Later, Gerlach and Meyl
(1957) established the genus Hofmaenneria with
H. brachystoma (syn. C. brachystoma) as the type species
and also transferred C. niddensis to it. Later, Mulvey (1969)
described a new soil inhabiting species: H. hazanensis. The
systematic position of the genus has been matter of controversy. The genus has already been assigned to Sphaerolaimidae
(Andrássy 1981), to Xyalidae (Gerlach and Riemann 1973;
Lorenzen 1994) and also incerta sedis (Coomans and
Eyualem-Abebe 2006). In the present work we follow the
most recent review of Xyalidae (Fonseca and Bezerra 2013),
which recognized Hofmaenneria as a valid genus of Xyalidae.
At the moment nine species are recognized as valid. The
descriptions of H. hazeensis and H. longicuadata were based
only in females and, therefore, were presently considered
species inquirenda.
Number of valid species: 9.
Genus Linhystera Juario, 1974
The genus Linhystera was described by Juario (1974) from
the sublittoral zone of the German Bight (North Sea) with
doubts about its position within Xyalidae. Lorenzen (1977)
redescribed the genus with additional observations on the
specimens of L. problematica from Chile and determined the
correct position of the cephalic setae (6+4) and testis (anterior
placed to the left and posterior placed to the right side of the
intestine) confirming the systematic position of the genus.
Pastor de Ward (1985) described the second species L. longa
from sublittoral areas of Argentina. This species differs from
L. problematica by having a reduced posterior testis, a crown
of cervical setae and longer spicules.
Number of valid species: 2.
Genus Manganonema Bussau, 1993
The genus Manganonema and the type species
M. microcephalum were described by Bussau (1993) for
samples from 4,000 m depth from the Eastern Pacific Basin
of Peru. Fonseca et al. (2006) redescribed the genus adding the
description of five new species from different deep-sea locations (North and South-western Atlantic Ocean, Western
Indian Ocean, Weddell Sea and South Sandwich Trench in
the Southern Ocean). In this same study, another four species
were typed but no names were given (Fonseca et al. 2006).
Number of valid species: 6.
Genus Marisalbinema Tchesunov, 1990
The monospecific genus Marisalbinema was described for
the White Sea. Diagnose as in Fonseca and Bezerra (2013).
Number of valid species: 1.
Genus Metadesmolaimus Schuurmans-Stekhoven, 1935
The genus Metadesmolaimus was established with
M. labiosetosus based on a single juvenile specimen from
the Belgian coast (North Sea). Later two new species were
described, M. coronatus Schuurmans-Stekhoven, 1950 and
M. aversivulva Gerlach, 1953, but posteriorly Wieser (1956)
synonymized Metadesmolaimus with Theristus. Lorenzen
(1972) rehabilitated and redescribed Metadesmolaimus with
the description of four new species (M. aduncus,
M. heteroclitus, M. pandus and M. varians) and new combination of other three species: Paramonohystera canicula
Wieser and Hopper, 1967, Theristus hamatus Gerlach, 1956
and Theristus tarsus Gerlach, 1954. Lorenzen (1977) added a
species with another new combination: Theristus gelana
Warwick and Platt, 1973. Platt (1983) described M. gaelicus,
and provided a table comparing the main diagnostic features
of the valid species. Tchesunov (1990a) described
M. psammophilus and Tchesunov (1990b) described
M. similis, both species from the White Sea.
Number of valid species: 13.
Genus Omicronema Cobb, 1920
The genus Omicronema was established with the new
species O. litorium Cobb, 1920. Later three new species were
added, O. nidrosiensis Allgén, 1933 and Omicronema
truncatum Schuurmans-Stekhoven, 1950, which are considered species inquirenda and O. clavulatum Gerlach, 1957.
Vincx and Furstenberg (1988) considered Omicronema a junior synonym of Xyala. Stewart and Nicholas (1994) considered the relatively large amphids, without sharply defined
rims, and cryptospiral appearence; and the cuticular ornamentations of simple bars without crests as sufficient features to
separate Omicronema from Xyala and described three new
species: O. australis, O. coronalata and O. nana. More recently O. orientalis has been described (Gagarin and Thanh
2009).
Number of valid species: 6.
Genus Paragonionchus Blome, 2002
The monospecific genus Paragonionchus is known only
from the sandy beaches of Eastern Australia, and belongs
together with Dactylaimoides, Gonionchus and Xyala to the
group of Xyalidae genera with strongly annulated cuticle
forming longitudinal crests or peculiar shaped projections.
The unique character of the genus within the Xyalidae is the
deeply incised lips with cuticularised structures (archs, clasps,
strips). Diagnose as in Blome (2002) and Fonseca and Bezerra
(2013).
Number of valid species: 1
Genus Paramonohystera Steiner, 1916
The genus Paramononhystera (in some works spelled as
Paramonhystera) was initially proposed by Steiner (1916) as
a subgenus of Monhystera to accomodate the species
Monhystera (Paramonohystera) megacephala, which has
been described based on a single female. Paramonohystera
was raised to genus level by Filipjev (1918) and later Wieser
(1954) redescribed P. megacephala using many specimens
from Chile. Wieser (1956) considered Paramononhystera
Steiner, 1916 and Leptogastrella Cobb, 1920 synonyms,
Mar Biodiv
suggesting that the latter could be regarded as a subgenus of
the former. Gerlach and Riemann (1973) considered
Paramonhystera as used by Filipjev (1918) and many subsequent authors as an invalid emendation. Lorenzen (1977)
transferred P. breviseta Juario, 1974 to Retrotheristus.
Lorenzen (1994) considered Paramononhystera as the valid
name with two subgenera: Paramonohystera Steiner, 1916
and Leptogastrella Cobb, 1920, which is the classification
adopted here. Chen and Vincx (2000) described P. geraerti,
redescribed P. biforma Wieser, 1956 and P. megacephala, and
proposed an identification key for nine species. These authors
considered P. micramphis Schuurmans-Stekhoven, 1950 as
species inquirenda, included P. breviseta Juario, 1974 which
was transferred to Retrotheristus by Lorenzen (1977) and did
not included in their key P. pilosa Boucher, 1971,
P. parabutschlii (Timm, 1961) Pastor de Ward, 1985,
P. zizichi Pastor de Ward, 1985 and the species from
Leptogastrella.
Number of valid species: 18.
Genus Paramphimonhystrella Huang and Zhang, 2006
The genus Paramphimonhystrella was described based on
three new species from the Yellow Sea. It can be differentiated
from all other genera within the family by the buccal cavity,
which is conical elongated instead of typically conical.
P. elegans is the type species, and it is characterized by
amphids with ovoid fovea and spicules with a small hook at
the distal end. The other two species P. minor and P. sinica can
be distinguished from P. elegans by possessing spicules with a
sharp tip and spicules with a blunt tip, respectively.
Number of valid species: 3.
Genus Parelzalia Tchesunov, 1990
The genus Parelzalia was described by Tchesunov (1990b)
from the White Sea with the only species P. obscuramphis.
Diagnose of the genus is given in Fonseca and Bezerra (2013).
Number of valid species: 1.
Genus Promonhystera Wieser, 1956
The genus Promonhystera with its two known species was
described from Chile. A third species was described from The
North Sea for the genus, Promonhystera albigens Riemann,
1966, but it was later transferred to Daptonema.
Number of valid species: 2.
Genus Prorhynchonema Gourbault, 1982
The genus Prorhynchonema resembles Rhynchonema, but
differs from it by having a shorter buccal cavity. The type
species P. warwicki was described with reduced gonads and a
peculiar reproduction system formed by spermatophores. A
second species was posteriorly described: P. gourbaultae
Nicholas and Stewart, 1995.
Number of valid species: 2.
Genus Pseudechinotheristus Blome, 2002
The monospecific genus Pseudechinotheristus was described from specimens found in the intertidal medium to fine
sands of Australia. This genus is similar to Echinotheristus,
another genus of Xyalidae, but lacks the precloacal supplements and the transverse row of small spines on the cuticular
rings. Diagnose as in Blome (2002) and Fonseca and Bezerra
(2013).
Number of valid species: 1.
Genus Pseudosteineria Wieser, 1956
The genus Pseudosteineria was described based on two
species, P. anteferens and P. anticipans, and initially considered as a subgenus of Theristus, but later De Coninck (1965)
raised it to genus level. P. coronata Gerlach, 1955 and
P. inaequaspiculata Platonova, 1971 were described as
Theristus, but transferred to Pseudosteineria by Wieser
(1959) and Gerlach and Riemann (1973), respectively.
P. horrida Steiner, 1916 was initially described as
Monhystera and transferred to Pseudosteineria by Wieser
(1956). Three other species (P. scopae, P. pavo and
P. pulchra) were originally described as Steineria and transferred to Pseudosteineria by Fadeeva (1986), who also described P. sagittispiculata. Tchesunov (2000) described
P. ventropapilata. The differential diagnostic character of this
genus is the distribution of the additional subcephalic setae in
eight groups at the level of the amphideal fovea, which is
located posteriorly to the cephalic capsule. Recently Huang
and Li (2010) described two new species, P. sinica and
P. zhangi, and provided also an identification key.
Number of valid species: 12.
Genus Retrotheristus Lorenzen, 1977
The monospecific genus Retrotheristus was established
when Lorenzen (1977) transferred Paramonhystera breviseta
Juario, 1974 as a new combination based on the observation
of additional specimens of the North Sea. Diagnose as in
Fonseca and Bezerra (2013).
Number of valid species: 1.
Genus Rhynchonema Cobb, 1920
The genus Rhynchonema resembles Prorhynchonema, another genus also in Xyalidae. Main features that differentiate
Rhynchonema from Prorynchonema are the much attenuated
head, long and tubular buccal cavity, and the cuticle, which is
strongly annulated. The species Rhynchonema lyngei (Allgén,
1940) Gerlach, 1953 was originally described in Leptolaimus
and R. ronaldi is considered nomen nudum since it is described in a PhD thesis (Procel 2007). Diagnose of the genus
as in Nicholas and Stewart (1995) and Fonseca and Bezerra
(2013).
Number of valid species: 31.
Genus Robustnema Nicholas, 1996
The monospecific genus Robustnema was described as
common species for the mangrove mudflats in Australia.
The arrangement of the cephalic setae (six labial papillae,
six outer labial, and six cephalic setae in one circle), small
and unarmoures buccal cavity and low lips profile are a unique
combination in Xyalidae.
Number of valid species: 1.
Mar Biodiv
Genus Sacrimarinema Shoshin, 2001
The genus Sacrimarinema with three species was described from the largest freshwater lake in the world, Baikal
Lake. This genus resembles members of Theristus subgenus
Penzancia. Diagnose of the genus is given in Coomans and
Eyualem-Abebe (2006) and Fonseca and Bezerra (2013).
Number of valid species: 3.
Genus Scaptrella Cobb, 1917
The genus Scaptrella was described from California, USA,
by Cobb (1917). Species in this genus have jointed mandibles,
circular amphid, a single ovary, and spicules with only faint
accessory pieces. Later, two news species were described:
S. brevicaudata Gerlach, 1953 and S. tenuicaudata Gerlach,
1956, from Italy and Germany, respectively. Description of
S. tenuicaudata was based in a single female and, therefore, is
considered species inquirenda.
Number of valid species: 2.
Genus Sphaerotheristus Timm, 1968
The genus was erected by Timm (1968) from material
sampled in the Gulf of Thailand. In this study he describes
four new species, namely: S. bengalensis, S. pseudodentatus,
S. sonadiae and S. supoti and erects Cobbia macrostoma
Timm, 1963 as the type species. More recently, four new
species from Vietnam were added to the genus: S. parvus
Gagarin and Thanh, 2006; S. nothum Gagarin and Thanh,
2008; S. supplementatus Gagarin and Thanh, 2008 and
S. validum Gagarin and Thanh, 2008.
Number of valid species: 9.
Genus Spiramphinema Wieser, 1956
The genus Spiramphinema was described by Wieser
(1956) based on two species from Chile: S. convolutum and
S. longiseta. The main feature was the spiral amphid. Wieser
(1956) also transferred Monhystera microcephalon
Stekhoven, 1942 to Spiramphinema, but given the poor description this species should be considered as species
inquirenda. Nicholas and Trueman (2002) did not consider
Spiramphinema valid, suggesting a mistaken interpretation of
the amphid structure by Wieser (1956). Fonseca and Bezerra
(2013) considered valid and provided a detailed diagnoses of
this genus.
Number of valid species: 2.
Genus Steineria Mickoletzky, 1921
The genus Steineria was firstly established as a subgenus
of Monhystera with only three species (S. pilosa,
S. polychaeta and S. setossisima) and posteriorly raised to
genus by Stekhoven and De Conninck (1933). For some
years, Steineria setosissima was considered the type species
as designed by Stekhoven and De Conninck (1933). Later,
Mawson (1957) disagreed with the designation of the typespecies based on the fact that S. setosissima was not one of the
included species brought to the subgenus when this was first
established, then a new type species was designated
(S. polychaeta). Wieser and Hopper (1967) agreed with
Mawson (1957) and reinforced that the designated type species agreed with International Code of Zoological
Nomenclature. This genus differs from the other of family
Xyalidae by the additional subcephalic setae located at the
level of cephalic setae and arranged in 8-radiate symmetry.
Number of valid species: 22.
Genus Stylotheristus Lorenzen, 1977
The monospecific genus Stylotheristus was established
when Lorenzen (1977) transferred Paramonhystera mutila
Lorenzen, 1973 as a new combination based on the observation of additional specimens of the North Sea. Diagnose as in
Fonseca and Bezerra (2013).
Number of valid species: 1.
Genus Theristus Bastian, 1865
The genus Theristus is the oldest genus within Xyalidae
and since its description passed for some internal changes.
During the last century many taxa were placed as subgenera of
Theristus but Gerlach and Riemann (1973) recognized only
three subgenera of Theristus (Daptonema, Penzancia and
Theristus) and later Lorenzen (1994) accepted only two
(Penzancia and Theristus). Nowadays Theristus differs from
Daptonema only by the form of the tail and the terminal setae
at the tail tip: conical without terminal setae in the former,
conico-cylindrical with terminal setae in the later. The taxon is
currently the second genus with the highest number of valid
species within Xyalidae. At present, the total amount of species inquirenda is 27.
Number of valid species: 92.
Genus Trichotheristus Wieser, 1956
The genus Trichotheristus was initially erected as a subgenus of Theristus by Wieser (1956) to accommodate Steineria
mirabilis Stekhoven and De Coninck, 1933 and
S. paramirabilis Gerlach, 1955. Chitwood and Murphy
(1964) raised the taxon to genus level however; Wieser and
Hopper (1967) kept considering it as a subgenus, including
some species previously described in Mesotheristus (another
subgenus of Theristus): T. circumscriptus, T. erectus, T. laxus,
T. longisetosus, T. sanctimarteni, T. setifer and T. setosus.
Furthermore, Wieser and Hopper (1967) considered
Theristus heterus Gerlach, 1957 as species of the subgenus
Trichotheristus and also described three new species:
T. floridanus, T. erectus and T. galeatus. Other species was
described by Warwick (1970) to subgenus Trichotheristus:
T. p s a m m o i d e s . Tc h e s u n o v ( 1 9 9 0 a ) c o n s i d e r e d
Mesotheristus as a valid genus and Trichotheristus as an
invalid genus redistributing the species T. floridanus,
T. heterus, T. mirabilis and T. psammoides within
Daptonema; T. galeatus within Metadesmolaimus and
T. paramirabilis within Pseudosteineria. Huang and Zhang
(2006) described T. articulatus, recognizing Trichotheristus as
valid and provided an identification key. In the identification
key from Huang and Zhang (2006), T. psammoides and
T. paramirabilis were omitted. Since Trichotheristus is
Mar Biodiv
currently considered as a valid taxon, these two species previously transferred by Tchesunov (1990a) to Daptonema and
Pseudosteineria, respectively, are considered again as valid
species of the genus Trichotheristus.
Number of valid species: 14.
Genus Valvaelaimus Lorenzen, 1977
The genus Valvaelaimus was erected for Xyalidae in the
revision of the family made by Lorenzen (1977) to accommodate two species from Theristus, which have three teeth-like
structures in the buccal cavity.
Number of valid species: 2.
Genus Xenolaimus Cobb, 1920
The genus Xenolaimus was originally described with one
species, Xenolaimus striatus, from the Gulf Coast of
California and posteriorly redescribed by Wieser and Hopper
(1967). The other species belonging to this genus;
X. pauroamphus, was described by Nichols (1979). These
two species differ from each other by the size of the amphidial
fovea and absence of the gubernaculum in the latter.
Number of valid species: 2.
Genus Xyala Cobb, 1920
The genus Xyala was described by Cobb (1920).
Posteriorly, Vincx (1986) pointed for the existence of a large
similarity between Xyala and Omicronema and proposed that
Omicronema is a junior synonym for Xyala. Stewart and
Nicholas (1994) attested that there are morphological differences between Xyala and Omicronema. Species within Xyala
share the common characteristics of having a cuticule with
longitudinal crests and hyaline lips without protrusion,
amphidial fovea small and circular. The species Cenolaimus
sapeloensis Nichols, 1979 is transferred to this genus as Xyala
sapeloensis comb. nov., as suggested by Blome (2002). This
species was described with male specimens but differs from
the description of Cenolaimus, which is considered an invalid
genus.
Number of valid species: 13.
Genus Zygonemella Cobb, 1920
The monospecific genus Zygonemella was described by
Cobb (1920) from Costa Rica with Z. striata and later the
same species was recorded by Gerlach (1957a) in magrove
samples from Cananeia, Brazil. Recently Cunha et al. (2013)
re-examined the situation of this genus with morphological
and molecular studies.
Number of valid species: 1.
2001; Medeiros 1997; Venekey 2007; Maria et al. 2008;
Vicente 2008; Neres et al. 2010; Ataide 2012; Brustolin
et al. 2013; Cunha et al. 2013).
Along the Brazilian coastline 28 genera and 43 species of
Xyalidae were registered. From the valid genera of Xyalidae
Capsula, Cienfuegia, Corononema, Dactylaimoides,
Enchonema, Filipjeva, Guitartia, Gullanema, Manganonema,
Marisalbinema, Paragonionchus, Paramphimonhystrella,
Parelzalia, Pseudechinotheristus, Robustnema, Sacrimarinema,
Sphaerotheristus and Spiramphinema were not recorded in
Brazil. Among the species records, Elzalia floresi,
Metadesmolaimus tersus Paramonhystera stricta,
Pseudosteineria scopae, Rhynchonema cemae, R. olindensis,
Steineria ericia, S. marcorum, S. pavo, S. tripartita, Theristus
acribus, T. flevensis, T. macroflevensis, T. pertenuis, T.
stranus, Trichotheristus heterus, T. setosus and Zygonemella
striata have the Brazilian coast as type locality. Given the
paucity of data along the Brazilian coast, the distributional
range of species can be barely inferred. Although there are
three species registered at three different regions, supporting
previous observation of widespread distribution of marine
nematodes (e.g., Derycke et al. 2008; Bik et al. 2009), the
large majority (32 species) were recorded at one region. This
would support high levels of endemism along the Brazilian
coast. However, this conclusion cannot be made for two
main reasons: first, because our coast has been unevenly
covered with most of the records concentrated around Rio
de Janeiro and São Paulo (Fig. 1 and Table 1); and, second,
because for most of the restricted species, the type locality is
outside Brazil (see Appendix). An interesting aspect observed
along the Brazilian coast is that Xyalidae species occurred
mainly at oceanic sandy beaches rather than in estuarine
areas. Species belonging to Daptonema, Theristus,
Trichotheristus and Zygonemella were the only ones encountered in estuarine sediments (Table 1). Interestingly is that
these four genera are not necessarily monophyletic
(Nicholas and Trueman 2002; Fonseca and Bezerra
2013), suggesting that the colonization of inland waters
Records for the Brazilian coast
The first records of Xyalidae along Brazilian coastline were
done by Sebastian Gerlach (Gerlach 1954, 1956a, b, 1957a,
b). He sampled several sandy beaches, estuaries and lagoons
along the Brazilian coast (Pernambuco, Alagoas, João Pessoa,
Espírito Santo, Rio de Janeiro and São Paulo states). Since his
major contribution, nine punctual studies were done (Bezerra
Fig. 1 Map of Brazil showing location of records of Xyalidae species
Mar Biodiv
Table 1 Records of Xyalidae species along the Brazilian coast
Species
ID
PR
SP
RJ
Cobbia cf. dentata
Daptonema biggi
Daptonema curvatum
Daptonema furcatum
Daptonema normandicum
Daptonema oxycerca
Daptonema procerum
Daptonema resimum
Daptonema tenuispiculum
Daptonema trichinum
Daptonema vicinum
Elzalia floresi
Metadesmolaimus cf. labiosetosus
Metadesmolaimus coronatus
Metadesmolaimus tersus
Omicronema clavulatum
Omicronema litorium
5a
10
3
10
10, 12
9
10
12
3
3
10
5a
5a
5a
2, 5a, 5b, 5c
5a
5b, 6, 12
Paramonohystera stricta
Pseudosteineria scopae
Rhynchonema cemae
Rhynchonema cinctum
Rhynchonema veronicae
Scaptrella cf. brevicaudata
Scaptrella cincta
Steineria ericia
Steineria marcorum
Steineria pavo
Steineria tripartita
Theristus acribus
Theristus aff. rhynchonemoides
2, 5a
3, 8
15
5a
15
5a
3
2, 5a
2, 3, 5a, 5b
3, 5a
5a
2,5a
7
SB
Theristus flevensis
Theristus heterospiculoides
Theristus macroflevensis
Theristus metaflevensis
Theristus pertenuis
Theristus stranus
4, 5a, 12
5a
1,4
4, 5a
4, 5a, 12
5a
EST/SB
SB
EST
EST
EST/SB
EST/SB
SB
Trichotheristus heterus
Trichotheristus mirabilis
Trichotheristus psammoides
Trichotheristus setifer
Trichotheristus setosus
Xyala striata
Zygonemella striata
5a
SB
5a
SB
10
EST
10
EST
4, 5a, 10, 13 EST EST
5a
SB
4, 11, 12, 14
EST
ES
PE
PA
SB
EST
SB
EST
EST
SB
EST
EST
SB
SB
SB
EST
SB
SB
SB
SB
SB
SB
SB
SB
SB
SB
SB
SB
SB
SB
Gerlach 1957b
Vicente 2008
Gerlach 1956b
Vicente 2008
Vicente 2008; Ataide 2012
Maria et al. 2008
Vicente 2008
Ataide 2012
Gerlach 1956b
Gerlach 1956b
Vicente 2008
Gerlach 1957b
Gerlach 1957b
Gerlach 1957b
Gerlach 1957b
Gerlach 1957b
Gerlach 1957b; Medeiros 1997; Ataide 2012
Gerlach 1957b
Gerlach 1956b; Venekey 2007
Bezerra and Vincx 2014
Gerlach 1957b
Bezerra and Vincx 2014
Gerlach 1957b
Gerlach 1956b
Gerlach 1957b
Gerlach 1957b
Gerlach 1957b
Gerlach 1957b
Gerlach 1957b
Bezerra 2001
SB
SB
SB
SB
SB
SB
SB
References
SB
SB
SB
SB
SB
Gerlach 1957a, b; Ataide 2012
Gerlach 1957b
Gerlach 1957a
Gerlach 1957a
Gerlach 1957a
Gerlach 1957b
Vicente 2008
Gerlach 1957b
Vicente 2008
Gerlach 1957a
Gerlach 1957a, b; Vicente 2008; Brustolin et al. 2013
Gerlach 1957b
EST EST Gerlach 1957a; Neres et al. 2010; Cunha et al. 2013;
Venekey pers. com.
Habitats of occurrence were classified as sandy beaches (SB) or estuary (EST). PR Paraná, SP São Paulo, RJ Rio de Janeiro, ES Espirito Santo; PE
Pernambuco; PA Pará. ID location in the map (Fig. 1)
occurred multiple times along the evolutionary history
(Holterman et al. 2008). The same cannot be concluded
for the conquest of the terrestrial sediments. In terrestrial habitats only species belonging to Daptonema and
Mar Biodiv
Theristus have been reported (Coomans and EyualemAbebe 2006) and the taxonomical status of these genera
is questionable (Nicholas and Trueman 2002).
Acknowledgments The authors acknowledge FAPESP-2011/21289-3
(São Paulo Research Foundation) by financially supporting the “Workshop Taxonomy and Diversity of Marine Meiofauna–Brazil”. The authors
also thank Prof. Dr. Vladimir Gagarin for providing taxonomical literature and helping with the Russian articles, and Dr. Tania Nara Campinas
Bezerra for helping with the genus Rhynchonema. The editor and anonymous referees are acknowledged for their suggestions. Gustavo Fonseca
has been supported by FAPESP 2009/14019-0.
Appendix
Checklist of the valid genera and species belonging to the
family Xyalidae.
Family Xyalidae Chitwood, 1951
Genus Ammotheristus Lorenzen, 1977
VALID SPECIES LIST
1. Ammotheristus helgolandicus (Riemann, 1967)
Lorenzen, 1977 (Helgoland, North Sea)
2. Ammotheristus subtilis (Lorenzen, 1972) Lorenzen, 1977
(Helgoland, North Sea)
Genus Amphimonhystera Allgén, 1929
VALID SPECIES LIST
1. Amphimonhystera anechma (Southern, 1914) Lorenzen,
1977 (Helgoland, North Sea)
2. Amphimonhystera circula Guo and Warwick, 2001
(Bohai Sea, China)
3. Amphimonhystera galea Fadeeva, 1984 (White Sea,
Russia)
4. Amphimonhystera marisalbi Tchesunov and Mokievsky,
2005 (White Sea, Russia)
5. Amphimonhystera molloyensis Tchesunov and
Mokievsky, 2005 (Molloy Deep, Arctic Ocean)
6. Amphimonhystera pallida Tchesunov and Mokievsky,
2005 (Molloy Deep, Arctic Ocean)
7. Amphimonhystera bella Bussau, 1993 (Central Pacific
Ocean, Peru)
SPECIES INQUIRENDA LIST
1. Amphimonhystera paranechma Allgén, 1935 (Öresund)
Genus Amphimonhystrella Timm, 1961
VALID SPECIES LIST
1. Amphimonhystrella bullacauda Tchesunov and Miljutina,
2005 (Molloy Deep, Arctic Ocean)
2. Amphimonhystrella megastoma Timm, 1961 (Bay of
Bengal, Indian Ocean)
3. Amphimonhystrella parva Gagarin and Thanh, 2009
(Mekong River Delta, Vietnam)
4. Amphimonhystrella unita Lorenzen, 1977 (Skagen,
Denmark)
Genus Capsula Bussau, 1993
VALID SPECIES LIST
1. Capsula galeata Bussau, 1993 (Central Pacific Ocean,
Peru)
Genus Cienfuegia Armenteros, Vincx and Decraemer,
2009
VALID SPECIES LIST
1. Cienfuegia cachoi Armenteros, Vincx and Decraemer,
2009 (Cuba)
Genus Cobbia de Man, 1907
VALID SPECIES LIST
1. Cobbia caledonia Warwick and Platt, 1973 (Scotland)
2. Cobbia dentata Gerlach, 1953 (Italy)
3. Cobbia macrodentata Lo Russo and Pastor de Ward,
2012 (Argentina)
4. Cobbia sinica Huang and Zhang, 2010 (Yellow Sea,
China)
5. Cobbia trefusiaeformis (De Man, 1907) Lorenzen, 1977
(North Sea, Baltic)
6. Cobbia triodonta Filipjev, 1918 (Russia)
7. Cobbia truncata Wieser, 1959 (Washington, USA)
8. Cobbia urinator Wieser, 1959 (Washington, USA)
SPECIES INQUIRENDA LIST
1. Cobbia mawsoni Cobb, 1930 (Antarctica)
2. Cobbia scutata Wieser, 1956 (Chile)
3. Cobbia simplex (Allgén, 1929) Wieser, 1956 (Skagerrak,
Sweden)
NOMEN NUDUM LIST
1. Cobbia crenata Wieser, 1960 (Buzzards Bay, USA)
Genus Corononema Nicholas and Stewart, 1995
VALID SPECIES LIST
1. Corononema parvum Nicholas and Stewart, 1995
(Australia)
2. Corononema thai Nicholas and Stewart, 1995
(Thailand)
Mar Biodiv
Genus Dactylaimoides Blome, 2002
VALID SPECIES LIST
1. Dactylaimoides coronifer Blome, 2002 (Queensland,
Australia)
Genus Daptonema Cobb, 1920
VALID SPECIES LIST
1 . D a p t on e m a ac r i l a b i a t u m ( D e C o n i n c k a n d
Schuurmans-Stekhoven, 1933) Lorenzen, 1977 (North
Sea)
2. Daptonema adiecta (Schulz, 1932) Wieser, 1956 (Kiel
Bay)
3. Daptonema aegypticum (Gerlach, 1964) Lorenzen, 1977
(Red Sea)
4. Daptonema albigensis (Riemann, 1966) Hopper, 1968
(Prince Edward Island, Canada)
5. Daptonema altaicum Tsalolikhin, 1985 (Mongolia)
6. Daptonema alternum (Wieser, 1956) Lorenzen, 1977
(Chile)
7. Daptonema angulatum (Schuurmans-Stekhoven, 1950)
Wieser, 1956 (Mediterranean)
8. Daptonema aquaedulcis (Gagarin, 1987) Gagarin, 1993
(Russia)
9. Daptonema articulatum Lorenzen, 1977 (Helgoland,
North Sea)
10. Daptonema biggi (Gerlach, 1965) Lorenzen, 1977
(Svalbard)
11. Daptonema biwaensis (Tsalolikhin, 2002) (Biwa lake,
Japan) New name
12. Daptonema brevisetosum Thanh and Gagarin, 2009 (Me
Kong river, Vietnam)
13. Daptonema buetschlii Bresslau and Stekhoven in
Stekhoven, 1935 (North Sea)
14. Daptonema buetschlioides (Chitwood, 1951) Lorenzen,
1977 (Texas, USA)
15. Daptonema calcaneum (Schuurmans-Stekhoven, 1950)
Wieser, 1956 (Mediterranean)
16. Daptonema calceolatum (De Coninck and SchuurmansStekhoven, 1933) Lorenzen, 1977 (North Sea)
17. Daptonema circulum (Vitiello, 1971) Lorenzen, 1977
(Mediterranean)
18. Daptonema clavicaudatum (Gerlach, 1953) Lorenzen,
1981 (Chile)
19. Daptonema concordiense Pastor de Ward, 1985 (River
Deseado, Argentina)
20. Daptonema conicum (Filipjev, 1922) Lorenzen, 1977
(Azow Sea)
21. Daptonema crassissima Ditlevsen, 1911 (Denmark)
22. Daptonema curticauda (Tchesunov, 1980) Tchesunov,
1990 (Caspian Sea)
23. Daptonema curvatum Gerlach, 1956 (Kiel Bay)
24. Daptonema curvispicula Tchesunov and Miljutin, 2006
(Molloy Deep, Arctic Ocean)
25. Daptonema curvispiculum (Gerlach, 1953) Wieser, 1959
(Italia, Mediterranean)
26. Daptonema dentatum (Wieser, 1956) Lorenzen, 1977
(Chile)
27. Daptonema dihystera Gagarin and Thanh, 2005 (Nhue
River and West lake, Vietnam)
28. Daptonema divertens Boucher and Helléouet, 1977
(Pierre Noire, West Channel)
29. Daptonema dolichurus Nguyen, Thanh and Gagarin,
2004 (Can Gio mangrove, Vietnam)
30. Daptonema donsi (Allgén, 1948) Lorenzen, 1977
(Norway)
31. Daptonema dubium (Butschli, 1873) Lorenzen, 1977
(Germany)
32. Daptonema durum Gagarin and Nguyen, 2008 (Red
river delta Vietnam)
33. Daptonema ecphygmaticum (Wieser, 1959) Lorenzen,
1977 (Washington, USA)
34. Daptonema elaboratum (Chitwood, 1951) Lorenzen,
1977 (Texas, USA)
35. Daptonema elegans (Kreis, 1929) Lorenzen, 1977
(English Channel)
36. Daptonema elongatum Gagarin and Nguyen, 2008 (Red
river delta, Vietnam)
37. Daptonema eximium Gagarin and Lemzina, 1981 (Issikkull lake, Kyrgyzstan)
38. Daptonema exutum (Wieser, 1956) Lorenzen, 1977
(Chile)
39. Daptonema fallax (Lorenzen, 1972) Lorenzen, 1977
(Helgoland, North Sea)
40. Daptonema fissidens (Cobb, 1920) Lorenzen, 1977
(India)
41. Daptonema fistulatum (Wieser and Hopper, 1967)
Lorenzen, 1977 (Florida, USA)
42. Daptonema flagellicauda (Lorenzen, 1973) Lorenzen,
1977 (Helgoland, North Sea)
43. Daptonema foetidum Gagarin and Thanh, 2010 (China
Sea, South coast Vietnam)
44. Daptonema fortis Gagarin, 1993 (Lake Taimyr, Siberia,
Russia)
45. Daptonema furcatum (Juario, 1974) Lorenzen, 1977
(Helgoland, North Sea)
46. Daptonema gritsenkovi (Gagarin and Lemzina, 1981)
(Issik-kul lake, Kyrgyzstan)
47. Daptonema groenlandium (Ditlevsen, 1928) Lorenzen,
1977 (Greenland)
48. Daptonema gyrophurum (Wieser, 1956) Lorenzen, 1977
(Chile)
49. Daptonema hirsutum (Vitiello, 1967) Lorenzen, 1977
(English Channel)
50. Daptonema hirtum Gerlach, 1951 (North Sea)
Mar Biodiv
51. Daptonema hyalocella Aryuthaka and Kito, 2012
(Phuket Province, Thailand)
52. Daptonema iners Nguyen, Thanh and Gagarin, 2004
(Can Gio mangrove, Vietnam)
53. Daptonema intermedium Tchesunov, 1980 (Caspian
Sea)
54. Daptonema invagiferoum (Platt, 1973) Lorenzen, 1977
(Strangford lough, North Ireland)
55. Daptonema inversum Alekseev, 1984 (Kamchatka,
Russia)
56. Daptonema issykkulensis Gagarin and Lemzina, 1980
(Issyk-kul lake, Kyrgyzstan)
57. Daptonema karabugaensis Tchesunov, 1980 (Caspian
Sea)
58. Daptonema kornoeense (Allgén, 1929) Lorezen, 1977
(Skagerrak)
59. Daptonema leptogastrelloides Riemann, 1979 (Weser
estuary, Germany)
60. Daptonema limnobia Wu and Liang, 2000 (China)
61. Daptonema longiapophysis Huang and Zhang, 2010
(Yellow Sea, China)
62. Daptonema longissimecaudatum (Kreis, 1935)
Lorenzen, 1977 (Mediterranean)
63. Daptonema lopezi Pastor de Ward, 1985 (Deseado
River, Argentina)
64. Daptonema macrostoma Huang and Xu, 2013 (Yellow
Sea, China)
65. Daptonema maeoticum (Filipjev, 1922) Lorenzen, 1977
(Azow Sea)
66. Daptonema marylandicum (Timm, 1952) Wieser, 1959
(Maryland, USA)
67. Daptonema miamiense (Hopper, 1969) Lorenzen, 1977
(Florida, USA)
68. Daptonema microspiculum (Gerlach, 1953) Lorenzen
1977 (Italia, Mediterranean)
69. Daptonema modestum Tchesunov, 1990 (White Sea)
70. Daptonema nannospiculum Tchesunov, 1980 (Caspian
Sea)
71. Daptonema nanum (Lorenzen, 1972) Lorenzen, 1977
(Helgoland, North Sea)
72. Daptonema normandicum (De Man, 1890) Lorenzen,
1977 (English channel)
73. Daptonema obesum Gagarin, 2001 (Biwa lake,
Japan)
74. Daptonema osadchikhae (Tchesunov, 1980) Tchesunov,
1990 (Caspian Sea)
75. Daptonema ostentator Wieser and Hopper, 1967
(Florida, USA)
76. Daptonema oxycerca (De Man, 1888) Lorenzen, 1977
(North Sea)
77. Daptonema oxyuroides (Stekhoven, 1931) Gagarin and
Lemzina, 1981 (North Sea)
78. Daptonema papillatus Murphy, 1965 (Chile)
79. Daptonema paracircumscriptum Fadeeva and
Belogurov, 1987 (Sea of Japan)
80. Daptonema paradonsi (Allgén, 1949) Wieser, 1956
(Norway)
81. Daptonema paraelaboratum (Timm, 1952) Wieser,
1959 (Maryland, USA)
82. Daptonema paratortum (Vitiello, 1971) Lorenzen, 1977
(Mediterranean)
83. Daptonema planiere (Vitiello, 1971) Lorenzen, 1977
(Mediterranean)
84. Daptonema platonovae (Galtsova, 1976) Tschesunov,
1990 (White Sea)
85. Daptonema procerum (Gerlach, 1951) Lorenzen, 1977
(North Sea)
86. Daptonema prominens (Vitiello, 1971) Lorenzen, 1977
(Mediterranean)
87. Daptonema proprium (Lorenzen, 1972) Lorenzen, 1977
(Helgoland, North Sea)
88. Daptonema pseudotortum (Vitiello, 1971) Lorenzen,
1977 (Mediterranean)
89. Daptonema rectangulatum Pastor de Ward, 1985
(Deseado River, Argentina)
90. Daptonema resimum (Wieser, 1959) Lorenzen, 1977
(Washington, USA)
91. Daptonema riemanni Platt, 1973 (Strangford lough,
North Ireland)
92. Daptonema rigidum Gagarin, Thanh and Nguyen, 2005
(Chu river, Vietnam)
93. Daptonema robustum Tchesunov, 1980 (Caspian Sea)
94. Daptonema romanelloi Pastor de Ward, 1985
(Deseado river, Argentina)
95. Daptonema rusticum (Kreis, 1929) Lorenzen, 1977
(English Channel)
96. Daptonema salvum Gagarin, Thanh and Nguyen, 2005
(Chu river, Vietnam)
97. Daptonema setihyalocella Aryuthaka and Kito, 2012
(Phuket Province, Thailand)
98. Daptonema sibiricum Gagarin, 2000 (Russian Arctic)
99. Daptonema spirum (Gerlach, 1959) Lorenzen, 1977
(North Sea)
100. Daptonema stylosum (Lorenzen, 1973) Lorenzen, 1977
(Helgoland, North Sea)
101. Daptonema svalbardense (Gerlach, 1965) Lorenzen,
1977 (Svalbard)
102. Daptonema tenuispiculum (Ditlevsen, 1918) Lorenzen,
1977 (Danish Belt Sea)
103. Daptonema timmi (Timm, 1961) Gagarin and Nguyen,
2008 (Bay of Bengal)
104. Daptonema timoshkini Gagarin, 2001 (Biwa lake, Japan)
105. Daptonema tortum (Wieser and Hopper, 1967)
Lorenzen, 1977 (Florida, USA)
106. Daptonema tortuosum (Timm, 1961) Lorenzen, 1977
(Bay of Bengal)
Mar Biodiv
107. Daptonema trabeculosum (Schneider, 1906) Lorenzen,
1977 (Baltic sea)
108. Daptonema trecuspidatum (Wieser, 1959) Lorenzen,
1977 (Washington, USA)
109. Daptonema trichinum Gerlach, 1956 (Kiel Bay)
110. Daptonema uncinatus Wieser, 1959 (Washington,
USA)
111. Daptonema variasetosum Pavljuk, 1984 (West-Pacific)
112. Daptonema vicinum (Riemann, 1966) Lorenzen, 1977
(North Sea)
113. Daptonema vietnamensis (Gagarin and Thu, 2008)
(Red River delta, Vietnam) New name
114. Daptonema voskresenskii Tchesunov, 1990 (White
Sea)
115. Daptonema williamsi Vincx and Coomans, 1983
(Solomon Islands)
116. Daptonema xyaliforme (Wieser and Hopper, 1967)
Lorenzen, 1977 (Florida, USA)
SPECIES INQUIRENDA LIST
1. Daptonema arcospiculum (Allgén, 1947) Wieser, 1956
(Gulf of Panama)
2. Daptonema arcticum (Steiner, 1916) Filipjev, 1922
(Australia)
3. Daptonema australis Allgén, 1951 (Australia)
4. Daptonema bathylaimum Allgén, 1959 (Graham land,
Antarctica)
5. Daptonema buelkiensis (Schulz, 1932) Wieser, 1956
(Kiel Bay)
6. Daptonema cuspidospiculum (Allgén, 1932) Wieser,
1956 (Campbell Island)
7. Daptonema filicaudatum Allgén, 1959 (South Georgia)
8. Daptonema filispiculum Allgén, 1932 (Campbell Island)
9. Daptonema fimbriatum (Cobb, 1920) Hopper, 1969
(New Hampshire, USA)
10. Daptonema frigidum (Cobb, 1914) Wieser, 1956
(Antarctica)
11. Daptonema gracillimecaudatum (Allgén, 1946) Wieser,
1956 (Norway)
12. Daptonema grahami (Allgén, 1969) Lorenzen, 1977
(Graham land, Antarctica)
13. Daptonema lata (Cobb, 1894) Lorenzen, 1977
(Australia)
14. Daptonema longicaudatum (Filipjev, 1922) Lorenzen,
1977 (Black Sea)
15. Daptonema macrocirculum Allgén, 1959 (Graham land,
Antarctica)
16. Daptonema metasetosum (Allgén, 1929) (Skagerrak)
17. Daptonema naviculivorum (Cobb, 1930) Wieser, 1956
(Antarctica)
18. Daptonema norvegicum (Allgén, 1946) Wieser, 1956
(Norway)
19. Daptonema notoistospiculoides Allgén, 1959 (South
Georgia, Antarctica)
20. Daptonema notosetosum Allgén, 1959 (South Georgia,
Antarctica)
21. Daptonema paratenuispiculum (Allgén, 1942) Wieser,
1956 (Mediterranean)
22. Daptonema paraistospiculoides Allgén, 1959 (Falkland
Islands and South Georgia, Antarctica)
23. Daptonema polaris (Cobb, 1914) Wieser, 1956
(Antarctica)
24. Daptonema septentrionalis Cobb, 1914 (Antarctica)
25. Daptonema simplex Allgén, 1959 (South Georgia,
Antarctica)
26. Daptonema sinuosus Wieser, 1959 (Washington, USA)
27. Daptonema sphaerolaimoides (Schulz, 1932) Lorenzen,
1977 (Kiel Bay)
28. Daptonema sphaerolaimus (Allgén, 1935) Wieser, 1956
(Öresund)
29. Daptonema tenuicaudatum (Allgén, 1929) Lorezen,
1977 (Skagerrak)
30. Daptonema trichospiculum (Allgén, 1933) Wieser, 1956
(Norway)
NOMEN NUDUM LIST
1. Daptonema deconincki Sharma, 1985 (Netherlands)
Genus Echinotheristus Thun and Riemann, 1967
VALID SPECIES LIST
3. Echinotheristus cimbricus Thun and Riemann, 1967
(North Sea)
4. Echinotheristus teutonicus Thun and Riemann, 1967
(North Sea)
Genus Elzalia Gerlach, 1957
VALID SPECIES LIST
1. Elzalia federici Castillo-Fernández and Lambshead,
1990 (Gulf of Mexico)
2. Elzalia floresi Gerlach, 1957 (Ilha Bela, Brazil)
3. Elzalia gerlachi Zhang and Zhang, 2006 (Yellow sea,
China)
4. Elzalia kimae Castillo-Fernández and Lambshead, 1990
(Gulf of Mexico)
5. Elzalia mediterranea Vitiello, 1971 (Gulf of Lyon,
Mediterranean)
6. Elzalia poli Castillo-Fernández and Lambshead, 1990
(Gulf of Mexico)
7. Elzalia striatitenuis Zhang and Zhang, 2006 (Yellow
Sea, China)
8. Elzalia tenuis Allgén, 1959 (Graham Land, Antarctic
Peninsula)
Mar Biodiv
9. Elzalia bipectinella Hope and Aryuthaka, 2009 (Khung
Kraben Bay, East Thailand)
10. Elzalia tuberculata Hope and Aryuthaka, 2009 (Khung
Kraben Bay, East Thailand)
SPECIES INQUIRENDA LIST
1. Elzalia speculifer Timm, 1961 (Bay of Bengal)
Genus Enchonema Bussau, 1993
VALID SPECIES LIST
1. Enchonema umbrosum Bussau, 1993 (Central Pacific
Ocean, Peru)
Genus Filipjeva Ditlevsen, 1928
VALID SPECIES LIST
1. Filipjeva arctica Ditlevsen, 1928 (South of the Lille
Pendulum, East Greenland)
2. Filipjeva crucis Blome and Schräge, 1985 (King George
Island, Antarctic)
3. Filipjeva filipjevi Tchesunov, 1988 (Kandalaksh Bay,
White Sea, Karelian coast, Russia)
4. Filipjeva meridionalis Kreis, 1932 (Sunda Strasse Island,
Indonesia)
5. Filipjeva parameridionalis Vitiello, 1970 (Leon Gulf,
Mediterranean)
6. F i l i p j e v a t e r a t o s p i c u l a Tc h e s u n o v, 1 9 8 8
(Kandalaksh Bay, White Sea, Karelian coast,
Russia)
SPECIES INQUIRENDA LIST
1. Filipjeva mediterranea Stekhoven, 1950 (Bay of
Villefranche, Mediterranean)
Genus Gnomoxyala Lorenzen, 1977
VALID SPECIES LIST
1. Gnomoxyala spina Lorenzen, 1977 (Baltic Sea)
Genus Gonionchus Cobb, 1920
VALID SPECIES LIST
1. Gonionchus africanus Vincxand Furstenberg, 1988
(Sundays River, South Africa)
2. Gonionchus alastairi Stewart and Nicholas, 1994 (New
South Wales, Australia)
3. Gonionchus arabica NasiraandTurpeenniemi, 2003
(Arabian Sea, Pakistan)
4. Gonionchus australis Stewart and Nicholas, 1994 (New
South Wales, Australia)
5. Gonionchus cumbraensis Benwell, 1981 (Firth of
Clyde, Scotland)
6. Gonionchus heipi Vincx, 1986 (North Sea, Belgium and
Netherlands)
7. Gonionchus inaequalis Warwick and Platt, 1973
(Scotland)
8. Gonionchus intermedius Jensen, 1986 (East Flower
Garden, Gulf of Mexico)
9. Gonionchus latentis Fadeeva, 1984 (Melkovodnaya
Bay, Sea of Japan)
10. Gonionchus longicaudatus (Ward, 1972) Lorenzen,
1977 (Liverpool Bay, England)
11. Gonionchus paravilosus Blome, 1982 (Sylt Island,
North Sea)
12. Gonionchus sensibilis Lorenzen, 1977 (Helgoland,
Germany Island)
13. Gonionchus villosus Cobb, 1920 (New Hamsphire,
USA)
NOMEN NUDUM LIST
1. Gonionchus ecuadorensis Procel, 2007 (San Pedro
Makgarato, Ecuador)
Genus Guitartia Armenteros, Vincx and Decraemer,
2010
VALID SPECIES LIST
1. Guitartia tridentata Armenteros, Vincx and Decraemer,
2010 (Caribbean Sea)
Genus Gullanema Nicholas and Stewart, 1995
VALID SPECIES LIST
1. Gullanema fragilis Nicholas and Stewart, 1995 (New
South Wales, Australia)
Genus Hofmaenneria Gerlach and Meyl, 1957
VALID SPECIES LIST
1. H o f m a e n n e r i a b r a c h y s t o m a ( H o f m a n n e r i n
Hofmanner and Menzel, 1914) Gerlach and Meyl,
1957 (Yugoslavia)
2. Hofmaenneria elongata Gagarin, 1987 (Parabel River,
Russia)
3. Hofmaenneria gratiosa Alekseev, 1983 (Khanka Lake,
Russia)
4. Hofmaenneria keoladeoensis Khan, Hussain, Sultana and
Tahseen, 2005 (Rajasthan, India)
5. Hofmaenneria longispiculata Gagarin and Naumova,
2010 (Lake Baikal, Russia)
6. Hofmaenneria niddensis (Skwarra, 1921) Schneider,
1940 (Germany)
Mar Biodiv
7. Hofmaenneria obesa Gagarin and Naumova, 2010 (Lake
Baikal, Russia)
8. Hofmaenneria optata Alekseev, 1983 (Khanka Lake,
Russia)
9. Hofmaenneria sitnikovae Gagarin and Naumova, 2010
(Lake Baikal, Russia)
SPECIES INQUIRENDA LIST
1. Hofmaenneria hazanensis Mulvey, 1969 (Canadian
Arctic)
2. Hofmaenneria longicaudata Gagarin, 2000 (Borok,
Russia)
Genus Linhystera Juario, 1974
VALID SPECIES LIST
1. Linhystera longa Pastor de Ward, 1985 (Santa Cruz,
Argentina)
2. Linhystera problematica Juario, 1974 (German Bight)
3. Metadesmolaimus caniculus (Wieser and Hopper, 1967)
Lorenzen, 1972 (Florida, USA)
4. Metadesmolaimus coronatus Schuurmans-Stekhoven,
1950 (Mediterranean Sea)
5. Metadesmolaimus gaelicus Platt, 1983 (Ireland, UK)
6. Metadesmolaimus gelana (Warwick and Platt, 1973)
Lorenzen, 1977 (UK)
7. Metadesmolaimus hamatus (Gerlach, 1956) Lorenzen,
1972 (North Sea)
8. Metadesmolaimus heteroclitus Lorenzen, 1972
(Helgoland, North Sea)
9. Metadesmolaimus pandus Lorenzen, 1972 (North Sea)
10. Metadesmolaimus psammophilus Tchesunov, 1990
(White Sea)
11. Metadesmolaimus similis Tchesunov, 1990 (White Sea)
12. Metadesmolaimus tersus (Gerlach, 1956) Lorenzen,
1972 (São Sebastião, Brazil)
13. Metadesmolaimus varians Lorenzen, 1972 (Helgoland,
North Sea)
SPECIES INQUIRENDA LIST
Genus Manganonema Bussau, 1993
VALID SPECIES LIST
1. Manganonema antarctica Fonseca, Decraemer and
Vanreusel, 2006 (Southern Ocean)
2. Manganonema bussauensis Fonseca, Decraemer and
Vanreusel, 2006 (North Atlantic)
3. Manganonema media Fonseca, Decraemer and
Vanreusel, 2006 (SW Atlantic Ocean, Brazil; NE
Atlantic, Goban Spur)
4. Manganonema microcephalum Bussau, 1993 (Central
Pacific Ocean, Peru)
5. Manganonema pitilica Fonseca, Decraemer and
Vanreusel, 2006 (SW Atlantic Ocean, Brazil)
6. Manganonema robustus Fonseca, Decraemer and
Vanreusel, 2006 (SW Atlantic Ocean, Brazil)
Genus Marisalbinema Tchesunov, 1990
VALID SPECIES LIST
1. Metadesmolaimus labiosetosus Schuurmans-Stekhoven,
1935 (North Sea)
Genus Omicronema Cobb, 1920
VALID SPECIES LIST
1. Omicronema australis Stewart and Nicholas, 1994
(Australia)
2. Omicronema clavulatum Gerlach, 1957 (Rio de Janeiro,
Brazil)
3. Omicronema coronalata Stewart and Nicholas, 1994
(Australia)
4. Omicronema litorium Cobb, 1920 (California, USA)
5. Omicronema nana Stewart and Nicholas, 1994
(Australia)
6. Omicronema orientalis Gagarin and Thanh, 2009
(Mekong River Delta, Vietnam)
SPECIES INQUIRENDA LIST
1. Marisalbinema galtsovae Tchesunov, 1990 (White
Sea)
Genus Metadesmolaimus Schuurmans-Stekhoven, 1935
VALID SPECIES LIST
1. Metadesmolaimus aduncus Lorenzen, 1972 (Helgoland,
North Sea)
2. Metadesmolaimus aversivulva Gerlach, 1953
(Mediterranean Sea)
1. Omicronema nidrosiens Allgén, 1933 (Norway)
2. Omicronema truncatum Schuurmans-Stekhoven, 1950
(Mediterranean Sea)
Genus Paragonionchus Blome, 2002
VALID SPECIES LIST
1. Paragonionchus sclerolabiatus Blome, 2002 (New South
Wales)
Mar Biodiv
Genus Paramonohystera Steiner, 1916
VALID SPECIES LIST
1. Paramonohystera biforma Wieser, 1956 (Chile)
2. Paramonohystera concinna Lorenzen, 1977 (North and
Baltic Seas)
3. Paramonohystera elliptica Filipjev, 1918 (Black Sea)
4. Paramonohystera eurycephalus Huang and Wu, 2011
(Yellow Sea, China)
5. Paramonohystera geraerti Chen andVincx, 2000 (Strait
of Magellan, Chile)
6. Paramonohystera halerba Fadeeva and Belogurov,
1987 (Sea of Japan)
7. Paramonohystera levicula (Lorenzen, 1972) Lorenzen,
1977 (Helgoland, North Sea)
8. Paramonohystera longicaudata Timm, 1963 (Arabian Sea)
9. Paramonohystera megacephala (Steiner, 1916) Filipjev,
1918 (Barents Sea)
10. Paramonohystera parabutschlii (Timm, 1961) Pastor de
Ward, 1985 (Mediterranean)
11. Paramonohystera paranormandica (Micoletzky, 1922)
(Mediterranean)
12. Paramonohystera pellucida Cobb, 1920 (Chile)
13. Paramonohystera pilosa Boucher, 1971 (France)
14. Paramonohystera proteus Wieser, 1956 (Chile)
15. Paramonohystera stricta (Gerlach, 1956) (Santos,
Brazil)
16. Paramonohystera tschilenkoi Platonova, 1971 (Sea of
Japan)
17. Paramonohystera zizichi Pastor de Ward, 1985
(Argentina)
18. Paramononhystera wieseri Ott, 1977 (Bermuda)
SPECIES INQUIRENDA
1. Paramonohystera micramphis Schuurmans-Stekhoven,
1950 (Mediterranean)
NOMEN NUDUM LIST
1. Paramonohystera mystacoderma Wieser, 1960
Genus Paramphimonhystrella Huang and Zhang, 2006
VALID SPECIES LIST
1. Paramphimonhystrella elegans Huang and Zhang, 2006
(Yellow Sea)
2. Paramphimonhystrella minor Huang and Zhang, 2006
(Yellow Sea)
3. Paramphimonhystrella sinica Huang and Zhang, 2006
(Yellow Sea)
Genus Parelzalia Tchesunov, 1990
VALID SPECIES LIST
1. Parelzalia obscuramphis Tchesunov, 1990 (White Sea)
Genus Promonhystera Wieser, 1956
VALID SPECIES LIST
1. Promonhystera faber Wieser, 1956 (Chile)
2. Promonhystera tricuspidata Wieser, 1956 (Chile)
Genus Prorhynchonema Gourbault, 1982
VALID SPECIES LIST
1. Prorhynchonema gourbaultae Nicholas and Stewart,
1995 (New South Wales, Australia)
2 . P ro rh y n c h o n e m a w a r w i c k i G o u r b a u l t , 1 9 8 2
(Guadeloupe, France)
Genus Pseudechinotheristus Blome, 2002
VALID SPECIES LIST
1. Pseudechinotheristus nudus Blome, 2002 (Queensland)
Genus Pseudosteineria Wieser, 1956
VALID SPECIES LIST
1. Pseudosteineria anteferens (Wieser, 1956) De Coninck,
1965 (Chile)
2. Pseudosteineria anticipans (Wieser, 1956) De Coninck,
1965 (Washington, USA)
3. Pseudosteineria coronata (Gerlach, 1955) Wieser, 1959
(San Salvador)
4. Pseudosteineria horrida (Steiner, 1916) Wieser, 1956
(Barents Sea)
5. Pseudosteineria inaequispiculata (Platonova, 1971)
Gerlach and Riemann, 1973 (Sea of Japan)
6. Pseudosteineria pavo (Gerlach, 1957) Fadeeva, 1986
(Ilha Bela, Brazil)
7. Pseudosteineria pulchra (Mawson, 1957) Techsunov,
2000 (Encouter Bay, Australia)
8. Pseudosteineria sagittispiculata Fadeeva, 1986 (Sea of
Japan)
9. Pseudosteineria scopae (Gerlach, 1956) Wieser, 1959
(Pernambuco, Brazil)
10. Pseudosteineria sinica Huang and Li, 2010 (Yellow Sea,
China)
11. Pseudosteineria zhangi Huang and Li, 2010 (Yellow
Sea, China)
12. Pseudosteineria ventropapilata Tchesunov, 2010 (White
Sea)
Mar Biodiv
Genus Retrotheristus Lorenzen, 1977
VALID SPECIES LIST
1. Retrotheristus breviseta (Juario, 1974) Lorenzen, 1977
(German Bight)
Genus Rhynchonema Cobb, 1920
VALID SPECIES LIST
1. Rhynchonema amakusanum Aryuthaka, 1989 (Oniike,
Japan)
2. Rhynchonema ambianorum Boucher, 1974 (Eastern
English Channel)
3. Rhynchonema brevituba Gerlach, 1953 (Mediterranean
Sea)
4. Rhynchonema cemae Bezerra and Vincx, 2014 (Olinda,
Brazil)
5. Rhynchonema ceramotos Boucher, 1974 (Eastern
English Channel)
6. Rhynchonema chiloense Lorenzen, 1975 (South Chile)
7. Rhynchonema cinctum Cobb, 1920 (Salavery, Peru)
8. Rhynchonema collare Nicholas and Stewart, 1995 (New
South Wales, Australia)
9. Rhynchonema deconincki Vitiello, 1967 (Roscoff,
France)
10. Rhynchonema díspar Gourbault, 1982 (Guadeloupe,
France)
11. Rhynchonema falciferum Boucher, 1974 (English
Channel)
12. Rhynchonema fossum Lorenzen, 1975 (Santa Marta,
Caribbean Columbia)
13. Rhynchonema gerlachi Vitiello, 1967 (Europa Island,
IndianOcean)
14. Rhynchonema hirsutum Hopper, 1961 (Gulf of Mexico)
15. Rhynchonema impar Lorenzen, 1975 (Santa Marta,
Caribbean Columbia)
16. Rhynchonema kikuchii Aryuthaka, 1989 (Oniike, Japan)
17. Rhynchonema longituba Gerlach, 1953 (Mediterranean
Sea)
18. Rhynchonema lyngei (Allgén, 1940) Gerlach, 1953
(Baltic Sea)
19. Rhynchonema megamphida Boucher, 1974 (Western
English Channel)
20. Rhynchonema moorea Boucher, 1974 (Society Islands,
Central Pacific)
21. Rhynchonema ornatum Lorenzen, 1975 (Salamanca
Island, Caribbean Columbia)
22. Rhynchonema quemer Boucher, 1974 (Eastern English
Channel)
23. Rhynchonema scutatum Lorenzen, 1972 (Helgoland,
North Sea)
24. Rhynchonema semiserratum Lorenzen, 1975 (South
Chile)
25. Rhynchonema separatum Lorenzen, 1975 (South Chile)
26. Rhynchonema sieverti Gourbault, 1982 (Guadeloupe,
France)
27. Rhynchonema subsetosum Murphy, 1964 (North Pacific
Coast)
28. Rhynchonema tomakinense Nicholas and Stewart, 1995
(New South Wales, Australia)
29. Rhynchonema tremendum Lorenzen, 1975 (South Chile)
30. Rhynchonema veronicae Bezerra and Vincx, 2014
(Olinda, Brazil)
31. Rhynchonema xiamenensis Huang and Liu, 2002
(Xiamen Island, China)
SPECIES INQUIRENDA
1. Rhynchonema wieseri Hopper, 1961 (Peru, Chile)
NOMEN NUDUM LIST
1. Rhynchonema ronaldi Procel, 2007 (Ecuador)
Genus Robustnema Nicholas, 1996
VALID SPECIES LIST
1. Robustnema fosteri Nicholas, 1996 (Australia)
Genus Sacrimarinema Shoshin, 2001
VALID SPECIES LIST
1. Sacrimarinema allae Shoshin, 2001 (Lake Baikal,
Russia)
2. Sacrimarinema ljajiae Shoshin, 2001 (Lake Baikal,
Russia)
3. Sacrimarinema tatushae Shoshin, 2001 (Lake Baikal,
Russia)
Genus Scaptrella Cobb, 1917
VALID SPECIES LIST
1. Scaptrella brevicaudata Gerlach, 1953 (Rimini, Adriatic
coast)
2. Scaptrella cincta Cobb, 1917 (San Francisco Bay,
California)
SPECIES INQUIRENDA LIST
1. Scaptrella tenuicaudata Gerlach, 1956 (Kiel Bay)
Genus Sphaerotheristus Timm, 1968
VALID SPECIES LIST
1. Sphaerotheristus bengalensis Timm, 1968 (Sunderbans,
East Pakistan)
Mar Biodiv
2. Sphaerotheristus macrostoma (Timm, 1963) Timm, 1968
(Bang Po, Gulf of Thailand)
3. Shaerotheristus nothum Gagarin and Thanh, 2008
(Camestuary, HaiPhong, Vietnam)
4. Sphaerotheristus parvus Gagarin and Thanh, 2006 (llai
Phong Prov, Vietnam)
5. Sphaerotheristus pseudodentatus Timm, 1968
(Sunderbans, East Pakistan)
6. Sphaerotheristus sonadiae Timm, 1968 (Bay of Bengal,
East Pakistan)
7. Sphaerotheristus supoti Timm, 1968 (Sunderbans, East
Pakistan)
8. Sphaerotheristus supplementatus Gagarin and Thanh,
2008 (Mekong River)
9. Sphaerotheristus validum Gagarin and Thanh, 2008
(Camestuary, HaiPhong, Vietnam)
Genus Spiramphinema Wieser, 1956
VALID SPECIES LIST
1. Spiramphinema convolutum Wieser, 1956 (Chile)
2. Spiramphinema longiseta Wieser, 1956 (Chile)
SPECIES INQUIRENDA LIST
1. Spiramphinema microcephalon (Stekhoven, 1942)
Wieser, 1956 (Mediterranean)
Genus Steineria Micoletzky, 1922
VALID SPECIES LIST
1. Steineria aegyptica Andrássy, 1959 (Red Sea)
2. Steineria ampullacea Wieser and Hopper, 1967 (Florida,
USA)
3. Steineria chiliensis Murphy, 1966 (Chile)
4. Steineria cobbi Wieser, 1956 (Chile)
5. Steineria copiosa Faadeva, 1991 (Sea of Japan)
6. Steineria ericia Gerlach, 1956 (Guarujá and São
Sebastião, Brazil)
7. Steineria gerlachi Wieser, 1956 (Washington, USA)
8. Steineria longicaudata Vitiello, 1971 (Gulf of Lyon,
Mediterranean)
9. Steineria marcorum Gerlach, 1956 (Guarujá, Brazil)
10. Steineria parapolychaeta Gerlach, 1953 (Chile)
11. Steineria pectinata Wieser, 1956 (Washington, USA)
12. Steineria phimifera Wieser, 1959 (Washington, USA)
13. Steineria pilosa (Cobb, 1914) Micoletzky, 1921 (Cape
Royds, Antarctica)
14. Steineria polychaeta (Steiner, 1915) Micoletzky, 1922
(Indonesia)
15. Steineria polychaetoides Gerlach, 1951 (North Sea)
16. Steineria pontica Gröza-Rojancovski, 1972 (Black Sea)
17. Steineria punctata Gerlach, 1955 (San Salvador)
18. Steineria setosissima (Cobb, 1894) Stekhoven and De
Connick, 1933 (Australia)
19. Steineria simplex Timm, 1963 (Arabian Sea)
20. Steineria sinica Huang and Wu, 2011 (Yellow Sea,
China)
21. Steineria sterreri Ott, 1977 (Bermuda)
22. Steineria tripartita Gerlach, 1957 (Ilha Bela, Brazil)
Genus Stylotheristus Lorenzen, 1977
VALID SPECIES LIST
1. Stylotheristus mutilus (Lorenzen, 1973) Lorenzen, 1977
(Helgoland, North Sea)
Genus Theristus Bastian, 1865
VALID SPECIES LIST
1. Theristus acer (Bastian, 1865) Bussau, 1990 (English
Channel)
2. Theristus acribus Gerlach, 1956 (São Paulo littoral,
Brazil)
3. Theristus acriformis De Man, 1922 (North Sea)
4. Theristus aculeatus Schulz, 1935 (Kiel Bay)
5. Theristus agilis (De Man, 1880) Filipjev, 1918
(Netherlands)
6. Theristus altenbachi (Jensen, 1991) (Norwegian Sea)
7. Theristus ambronensis Schulz, 1936 (North Sea)
8. Theristus anisocirculus Blome, 1982 (Sylt Island, North
Sea)
9. Theristus anisotrichus Lorenzen, 1972 (Helgoland,
North Sea)
10. Theristus anoxybioticus Jensen, 1995 (Hirsholm Islands,
Denmark)
11. Theristus athesinus Andrássy, 1962 (Italy)
12. Theristus balticus Lorenzen, 1973 (Kiel Bay)
13. Theristus bastiani Wieser in Gerlach and Riemann, 1973
(Kiel Bay)
14. Theristus biarcospiculoides Blome, 1982 (Sylt Island,
North Sea)
15. Theristus biarcospiculus Timm, 1952 (Maryland, USA)
16. Theristus bipunctatus Schneider, 1906 (Baltic Sea)
17. Theristus blandicor Rachor, 1971 (Wesser estuary,
Germany)
18. Theristus borosi Andrássy, 1958 (Egypt)
19. Theristus brevisetosus Alekseev in Alekseev, 1992
(Lake Khanka, Russia)
20. Theristus bujumbura Tsalolikhin in Tsalolikhin, 1989
(Lake Taganyika, Tanzania)
21. Theristus calx Wieser and Hopper, 1967 (Florida)
22. Theristus caudasaliens Adams and Tyler, 1980 (Cape
Elizabeth, USA)
23. Theristus complexus Jayasree and Warwick, 1977 (Firth
of Clyde, Scotland)
Mar Biodiv
24. Theristus conicaudatus Allgén, 1959 (South Georgia,
Antarctica)
25. Theristus copulatus Jensen, 1986 (Gulf of
Mexico)
26. Theristus darkovi Tsalolikhin, 1993 (Langano lake,
Ethiopia)
27. Theristus denticulatus Warwick, 1970 (English
Channel)
28. Theristus discolensis Bussau, 1993 (Central Pacific
Ocean, Peru)
29. Theristus diversispiculus Gerlach, 1953 (Chile)
30. Theristus ensifer Gerlach, 1951 (St. Peter Ording,
Germany, North Sea)
31. Theristus fimbriatoides (Chtiwood and Murphy, 1964)
Gerlach and Riemann, 1973 (Chile)
32. Theristus flevensis (Schuurmans-Stekhoven, 1935)
Nasira and Turpeenniemi, 2003 (North Sea)
33. Theristus franzbergeri Schiemer, 1984 (Danube river,
Austria)
34. Theristus gracillis (De Man, 1876) Filipjev, 1918 (Gulf
of Naples, Mediterranean)
35. Theristus heterospiculoides Gerlach, 1953 (Italia,
Mediterranean)
36. Theristus heterospiculus (Allgén, 1932) Gerlach, 1953
(Campbell Island)
37. Theristus inermis Gerlach, 1953 (Italia, Mediterranean)
38. Theristus interstitialis Warwick, 1970 (English Channel)
39. Theristus lineatus Gerlach, 1965 (Svalbard)
40. Theristus littoralis Filipjev, 1922 (Black Sea)
41. Theristus longicollis Blome, 1982 (Sylt Island, North
Sea)
42. Theristus longisetifer Kito and Aryuthaka, 1998
(Chanthabury, Gulf of Thailand)
43. Theristus longisetosus (Stekhoven and De Coninck,
1933) Lorenzen, 1977 (North Sea)
44. Theristus longispicula Platonova, 1971 (Sea of Japan)
45. Theristus longus Platt, 1973 (Strangford lough, North
Island)
46. Theristus lorenzeni Pastor de Ward, 1985 (Deseado
River, Argentina)
47. Theristus macer (Lorenzen, 1973) (North Sea)
48. Theristus macroflevensis Gerlach, 1954 (Rodrigo de
Freitas Lake, Brazil)
49. Theristus manicatus Wieser, 1956 (Chile)
50. Theristus marinae Tchesunov, 1981 (Caspian Sea)
51. Theristus megalaimoides Wieser, 1956 (Chile)
52. Theristus melnikovi Tchesunov, 1986 (Central Arctic
Basin)
53. Theristus metaflevensis Gerlach, 1955 (El Salvador)
54. Theristus meyli Riemann, 1966 (Elbe Estuary, North
Sea)
55. Theristus minimus Gagarin and Thanh, 2011 (Red River,
North Vietnam)
56. Theristus modicus Wieser, 1956 (Chile)
57. Theristus monstrosus Gerlach, 1954 (Mediterranean)
58. Theristus norwedicus (Allgén, 1933) Wieser, 1956
(Norway)
59.Theristus oistospiculum Allgén, 1932 (Campbell
Island)
60. Theristus orientalis Gagarin and Thanh, 2005 (Cam
Estuary, Hai Phong, Vietnam)
61. Theristus otoplanobius (Gerlach, 1951) Nasira and
Turpeenniemi, 2003 (Kiel Bay)
62. Theristus pakistanesis (Turpeenniemi, Nasira and
Maqbool, 2001) Fonseca and Bezerra, 2013 (Arabian
Sea, Pakistan)
63. Theristus pannonicus (Andrássy, 1985) Gagarin and
Holovachov, 2002 (Keszthely, Hungary)
64. Theristus parambronensis Timm, 1952 (Maryland,
USA)
65. Theristus paravelox Allgén, 1934 (Tarva, Norway)
66. Theristus pertenuis Bresslau and Stekhoven in
Stekhoven, 1935 (North Sea)
67. Theristus pictus Gerlach, 1951 (Sylt Island, North Sea)
68. Theristus polychaetophilus Hopper, 1966 (Florida,
USA)
79. Theristus pratti Murphy and Canaris, 1964 (Indian
Ocean, Kenya)
70. Theristus problematicus (Allgén, 1928) Wieser, 1956
(Campbell Island)
71. Theristus profundus Blome, 1982 (Sylt Island, North
Sea)
72. Theristus psammophilus Gagarin and Naumova, 2012
(Lake Baikal, Russia)
73. Theristus pyronasi Gerlach, 1965 (Svalbard)
74. Theristus rezaki Jensen, 1986 (Gulf of Mexico)
75. Theristus rhynchonemoides Hopper, 1961 (Gulf of
Mexico)
76. Theristus roscoffiensis Vitiello, 1967 (English Channel)
77. Theristus ruffoi (Andrássy, 1959) (Italia)
78. Theristus scanicus Allgén, 1949 (Sweden)
79. Theristus sonnae Bussau, 1993 (Central Pacific Ocean,
Peru)
80. Theristus stranus (Gerlach, 1957) (São Paulo coast,
Brazil)
81. Theristus strelzovi Galtsova, Platonova, Streltsova and
Petukhov, 1980 (Barents Sea)
82. Theristus subacer Pavljuk, 1984 (Sea of Japan)
83. Theristus subcurvatus Lorenzen, 1977 (Anholt Island,
Denmark)
84. Theristus terricola Andrássy, 1985 (Budapest, Hungary)
85. Theristus tessae Heyns and Coomans, 1989 (Namibia)
86. Theristus tsalolikhini Gagarin and Naumova, 2012
(Baikal lake, Russia)
87. Theristus turbidus Gagarin and Thanh, 2010 (China Sea,
South Coast Vietnam)
Mar Biodiv
88.
89.
90.
91.
Theristus velox Bastian, 1865 (English Channel)
Theristus vesentiniae Andrássy, 1962 (Italy)
Theristus wegelinae Andrássy, 1962 (Germany)
Theristus wimmeri (Wieser, 1959) (Washington,
USA)
SPECIES INQUIRENDA LIST
1. Theristus bathylaimoides Allgén, 1959 (Falkland
Islands)
2. Theristus bidontolaimus Allgén, 1959 (Graham Land,
Antarctica)
3. Theristus brachysetosus Allgén, 1959 (Graham Land,
Antarctica)
4. Theristus brevicollis (Cobb, 1894) Filipjev, 1922
(Australia)
5. Theristus chitinolaimus (Allgén, 1932) Wieser, 1956
(Campbell Island)
6. Theristus falklandiae Allgén, 1959 (Falkland
Islands)
7. Theristus helveticus (Steiner, 1914) Andrássy, 1981
(Switzerland)
8. Theristus kaszabi Andrássy, 1977 (Mongolia)
9. Theristus latissimus (Filipjev, 1922) Wieser, 1956
(Black Sea)
10. Theristus leptosoma Allgén, 1950 (Skagerrak)
11. Theristus lingi (Hoeppli and Chu, 1932) Andrássy, 1960
(Chile)
12. Theristus macropapillatus Allgén, 1959 (Falkland
Islands)
13. Theristus maspapillatus (Cobb, 1891) Filipjev, 1918
(Gulf of Aden)
14. Theristus megalaimus (Stewart, 1914) Wieser, 1956
(Bay of Bengal)
15. Theristus meridianus (Cobb, 1914) Filipjev, 1922
(Antarctica)
16. Theristus neglectus (Cobb, 1930) Wieser, 1956
(Antarctica)
17. Theristus nidrosiensis (Allgén, 1933) Wieser, 1956
(Norway)
18. Theristus obtusicephalus Stekhoven, 1950
(Mediterranean)
19. Theristus pacificus (Johnston, 1938) Filipjev, 1918
(Australia)
20. Theristus parasetosus Allgén, 1928 (Kristineberg Bay,
Sweden)
21. Theristus parasiticus (Penso, 1938) Andrássy, 1981
(Libya)
22. Theristus parvulus Timm, 1952 (Maryland, USA)
23. Theristus pellucidus Allgén, 1939 (Bud, Norway)
24. Theristus sabulicola (Filipjev, 1918) Wieser, 1956
(Black Sea)
25. Theristus strichotricha Stekhoven, 1950 (Mediterranean)
26. Theristus tenuicaudatus Allgén, 1951 (Tautra Island,
Norway)
27. Theristus strandi Allgén, 1934 (Öresund)
NOMEN NUDUM LIST
1. Theristus frictor Wieser and Kanwisher, 1961
2. Theristus multisetosus Wieser, 1960
3. Theristus camelopardis Wieser, 1959
SPECIES INCERTAE SEDIS LIST
1. Theristus quadripapillatus Decraemer and Coomans,
1978 (Lizard island, Australia)
Genus Trichotheristus Wieser, 1956
VALID SPECIES LIST
1. Trichotheristus articulatus Huang and Zhang, 2006
(Hong Kong, China)
2. Trichotheristus circumscriptus Wieser, 1959
(Washington, USA)
3. Trichotheristus erectus (Wieser and Hopper,
1967) Gerlach and Rieman, 1973 (Biscay Bay,
Spain)
4. Trichotheristus floridanus (Wieser and Hopper, 1967)
Gerlach and Rieman, 1973 (Biscayney Bay, Spain)
5. Trichotheristus galeatus (Wieser and Hopper, 1967)
Gerlach and Rieman, 1973 (Virginia, USA)
6. Trichotheristus heterus (Gerlach, 1957) Wieser and
Hopper, 1967 (Itanhaen, Brazil)
7. Trichotheristus laxus (Wieser, 1956) Wieser and
Hopper, 1967 (Chile)
8. Trichotheristus longisetosus (Stekhoven and De
Coninck, 1933) Wieser and Hopper (North Sea)
9. Trichotheristus mirabilis (Stekhoven and De Coninck,
1933) Wieser, 1956 (North Sea)
10. Trichotheristus paramirabilis (Gerlach, 1955) Wieser,
1959 (San Salvador, Bahamas)
11. Trichotheristus psammoides (Warwick, 1970) Gerlach
and Rieman, 1973 (North Atlantic)
12. Trichotheristus sanctimarteni (Timm, 1957) Wieser and
Hopper, 1967 (San Martin Island)
13. Trichotheristus setifer (Gerlach, 1952) Wieser and
Hopper, 1967 (Mediterranean)
14. Trichotheristus setosus (Büstchli, 1874) Wieser and
Hopper, 1967 (Kiel Bay)
Genus Valvaelaimus Lorenzen, 1977
VALID SPECIES LIST
1. Valvaelaimus euxinus (Filipjev, 1918) Lorenzen, 1977
(Black Sea)
Mar Biodiv
2. Valvaelaimus maior (Gerlach, 1956) Lorenzen, 1977
(Kiel Bay)
Genus Xenolaimus Cobb, 1920
VALID SPECIES LIST
1. Xenolaimus pauroamphus Nichols, 1979 (Georgia, USA)
2. Xenolaimus striatus Cobb, 1920 (Florida, USA)
Genus Xyala Cobb, 1920
VALID SPECIES LIST
1. Xyala aestuariensis Vincx and Furstenberg, 1988
(Sundays River, South Africa)
2. Xyala barbata Ward, 1972 (Irish Sea)
3. Xyala brevibucca Stewart and Nicholas, 1994 (New
South Wales, Australia)
4. Xyala finneyae King, Mundo-O campo and De Ley,
2010 (Santa Clara, Gulf of Mexico)
5. Xyala hamleyi Stewart and Nicholas, 1994 (Queesland
and Nouthern Territory, Australia)
6. Xyala imparis Boucher and Helleouet, 1977 (Pierre
Noire, West Channel)
7. Xyala macramphis Stewart and Nicholas, 1994 (New
South Wales, Australia)
8. Xyala oxybiotica Jensen, 1986 (East Flower Garden,
Gulf of Mexico)
9. Xyala psammonalis Vincx and Furstenberg, 1988
(Sundays River, South Africa)
10. Xyala riemanni Boucher and Helleouet, 1977 (Pierre
Noire, West Channel)
11. Xyala sapeloensis (Nichols, 1979) Blome, 2002
(Georgia State, USA) New combination
12. Xyala smo Warwick and Platt, 1973 (Scotland)
13. Xyala striata Cobb, 1920 (Massachussetts, USA)
SPECIES INQUIRENDA LIST
1. Xyala exigua Wieser, 1956 (Chile)
Genus Zygonemella Cobb, 1920
VALID SPECIES LIST
1. Zygonemella striata Cobb, 1920 (Costa Rica, Pacific
Coast)
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