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Zootaxa 2183: 1–99 (2009)
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Unresolved cases of type fixation, synonymy and homonymy in
harpacticoid copepod nomenclature (Crustacea: Copepoda)
RONY HUYS
Department of Zoology, Natural History Museum, Cromwell Road, London SW7 5BD, U.K.
E-mail: rjh@nhm.ac.uk
Magnolia Press
Auckland, New Zealand
Accepted by A. Minelli: 20 Jun. 2009; published: 6 Aug. 2009
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RONY HUYS
Unresolved cases of type fixation, synonymy and homonymy in harpacticoid copepod nomenclature
(Crustacea: Copepoda)
(Zootaxa 2183)
99 pp.; 30 cm.
6 Aug. 2009
ISBN 978-1-86977-399-1 (paperback)
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Table of contents
Abstract ................................................................................................................................................................................ 4
Introduction .......................................................................................................................................................................... 5
Cases of homonymy ............................................................................................................................................................. 6
Anoplosoma Sars, 1911c (Family Ameiridae) .............................................................................................................. 7
Poria Lang, 1965 (Family Canthocamptidae) .............................................................................................................. 7
Sewellia Lang, 1965 (Family Dactylopusiidae) ............................................................................................................ 8
Ichnusella Cottarelli, 1971 (Family Leptopontiidae).................................................................................................... 8
Dahlakia Por, 1986a (Family Canthocamptidae).......................................................................................................... 9
Unavailable genus-group names .......................................................................................................................................... 9
Rhizothrix Brady & Robertson, 1876 (Family Rhizotrichidae) .................................................................................. 10
Psammastacus Nicholls, 1935 (Family Leptastacidae) .............................................................................................. 10
Paranannopus Lang, 1936a (Family Pseudotachidiidae) ........................................................................................... 10
Paraidya Sewell, 1940 (Family Tisbidae) .................................................................................................................. 12
Alteuthellopsis Lang, 1944 (Family Peltidiidae)......................................................................................................... 12
Halectinosoma Lang, 1944 (Family Ectinosomatidae)............................................................................................... 12
Heterolaophonte Lang, 1944 (Family Laophontidae)................................................................................................. 14
Idyellopsis Lang, 1944 (Family Idyanthidae) ............................................................................................................. 15
Paralaophonte Lang, 1944 (Family Laophontidae) ................................................................................................... 15
Robertgurneya Lang, 1944 (Family Miraciidae) ........................................................................................................ 16
Rheocamptus Borutzky, 1948 (Family Canthocamptidae).......................................................................................... 16
Apodopsyllus Kunz, 1962 (Family Paramesochridae) ................................................................................................ 17
Kliopsyllus Kunz, 1962 (Family Paramesochridae).................................................................................................... 18
Scottopsyllus Kunz, 1962 (Family Paramesochridae)................................................................................................. 19
Cladorostrata Shen & Tai, 1963 (Family Miraciidae)................................................................................................ 20
Scottolana Por, 1967 (Family Canuellidae) ................................................................................................................ 20
Psyllocamptus (Langpsyllocamptus) Kunz, 1975b (Family Ameiridae) .................................................................... 22
Micropsammis Mielke, 1975 (Family Pseudotachidiidae).......................................................................................... 22
Barbaracletodes Becker, 1979 (Family Cletodidae incertae sedis) ........................................................................... 22
Ameiropsyllus Bodin, 1979 (Family Ameiridae) ........................................................................................................ 23
Chilaophonte Mielke, 1985 (Family Laophontidae)................................................................................................... 23
Psammonitocrella Rouch, 1992 (Family Ameiridae) ................................................................................................. 23
Tectacingulum Harris, 1994 (Family Porcellidiidae) .................................................................................................. 24
Stygonitocrella (Fiersiella) Suárez-Morales & Iliffe, 2005 (Family Ameiridae) ....................................................... 24
Pilocamptus Wells, 2007 (Family Canthocamptidae)................................................................................................. 25
Junior subjective synonyms ............................................................................................................................................... 26
Idomene Philippi, 1843 (Family Pseudotachidiidae) .................................................................................................. 26
Alteutha Baird, 1846b (Family Peltidiidae) ................................................................................................................ 27
Family Tisbidae Stebbing, 1910.................................................................................................................................. 28
Subfamily Cerviniopsinae Brotskaya, 1963................................................................................................................ 29
Namakosiramia Ho & Perkins, 1977 (Family Laophontidae) .................................................................................... 30
Family Huntemanniidae Por, 1986a............................................................................................................................ 30
Orphaned taxonomic groupings ......................................................................................................................................... 31
Amphiascus Sars, 1905a (Family Miraciidae) ............................................................................................................ 31
Teissierella Monard, 1935a (Family Miraciidae)........................................................................................................ 32
Hastigerella Nicholls, 1935 (Family Ectinosomatidae).............................................................................................. 33
Interleptomesochra Lang, 1965 (Family Ameiridae) ................................................................................................. 34
Addendum: Junior homonyms in Calanoida and Cyclopoida............................................................................................ 36
Parathalassius Dussart, 1986 ..................................................................................................................................... 36
Berea Yamaguti, 1963 ................................................................................................................................................. 36
Acknowledgements ............................................................................................................................................................ 36
References .......................................................................................................................................................................... 37
UNRESOLVED CASES IN HARPACTICOID COPEPOD NOMENCLATURE
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Abstract
Type fixation for each of the 601 valid genera (17 placed incertae sedis) and 13 genera of doubtful identity (genera
inquirenda) in the Harpacticoida (Crustacea, Copepoda) has been verified. Twenty-four genus-group names published
after 1930 lack the mandatory type fixation and are therefore unavailable. With the exception of Kliopsyllus Kunz, 1962
which is replaced by its senior synonym Emertonia Wilson, 1932, such names are made available here by either
attributing the original name to the first author(s) who explicitly fixed a type species (Psammastacus Nicholls, 1935;
Alteuthellopsis Lang, 1944; Idyellopsis Lang, 1944; Paralaophonte Lang, 1944; Robertgurneya Lang, 1944;
Cladorostrata Shen & Tai, 1963; Micropsammis Mielke, 1975; and the subgenera Rheocamptus Borutzky, 1948 and
Scottopsyllus Kunz, 1962;) or by adopting the name taking the present authorship and date (Paranannopus Lang, 1936a;
Paraidya Sewell, 1940; Apodopsyllus Kunz, 1962; Scottolana Por, 1967; Barbaracletodes Becker, 1979; Ameiropsyllus
Bodin, 1979; Chilaophonte Mielke, 1985; Psammonitocrella Rouch, 1992; Tectacingulum Harris, 1994; and the
subgenera Intermedopsyllus Kunz, 1962 (corrected spelling Intermediopsyllus) and Fiersiella Suárez Morales & Iliffe,
2005). In two cases a ruling by the International Commission on Zoological Nomenclature will be required to avoid
upsetting a long-accepted name in its accustomed meaning (Halectinosoma Lang, 1944; Heterolaophonte Lang, 1944).
The recently proposed generic name Pilocamptus Wells, 2007 does not satisfy the provisions of ICZN Art. 13.1 and is
here made available by explicit citation of a bibliographic reference that provides a diagnosis purported to differentiate
the taxon. Rhizothrix Brady & Robertson, 1876 is an unavailable name which was first made available by Sars (1909a).
The unavailable generic name Scottopsyllus Kunz, 1962 has no potentially valid synonym and is replaced by the next
oldest available name from among its subgenera, i.e. Wellsopsyllus Kunz, 1981 (ICZN Art. 23.3.5). The unavailable
subgeneric name Psyllocamptus (Langpsyllocamptus) Kunz, 1975b is not reinstated because it denotes a taxon that is
based exclusively on plesiomorphies.
New replacement names have been proposed for preoccupied generic names in the harpacticoid families
Canthocamptidae (Poria Lang, 1965; Dahlakia Por, 1986a), Dactylopusiidae (Sewellia Lang, 1965), and Leptopontiidae
(Ichnusella Cottarelli, 1971). The preoccupied generic name Anoplosoma Sars, 1911c (family Ameiridae) is replaced by
a previously proposed, but subsequently forgotten, replacement name, Anoplosomella Strand, 1929. Nomina nova are
also suggested for Parathalassius Dussart, 1986 (Calanoida: Centropagidae) and Berea Yamaguti, 1963 (Cyclopoida:
Chondracanthidae) which have entered into homonymy with previously established names.
The junior synonym Alteutha Baird, 1846b is considered valid, taking precedence as a nomen protectum over the
older names Sterope Goodsir, 1845 and Carillus Goodsir, 1845 (nomina oblita). Similar reversal of precedence applies to
the family-group names Peltidiidae Claus, 1860 and Tisbidae Stebbing, 1910 which are junior subjective synonyms of
Steropinae Dana, 1854 and Scutellidiinae Claus, 1889, respectively. Since the type of Idomene Philippi, 1843 is
identified as a member of the Clausidiidae (Cyclopoida), the generic name Xouthous Thomson, 1883 is reinstated to
accommodate all remaining species currently placed in Idomene. The forgotten copepod genus Microchelonia Brady,
1918 is placed in the family Laophontidae and considered a senior subjective synonym of Namakosiramia Ho & Perkins,
1977.
The family-group name Pontostratiotidae A. Scott, 1909 (type: Pontostratiotes Brady, 1883) is a senior subjective
synonym of Cerviniopseinae Brotskaya, 1963 (type: Cerviniopsis Sars, 1903) and the former is consequently reinstated
at the subfamilial level. The family-group name Huntemanniidae Por, 1986a (type: Huntemannia Poppe, 1884) is a
junior subjective synonym of Nannopinae Brady, 1880a (type: Nannopus Brady, 1880a) and the latter is reinstated as the
valid name at family rank and with the spelling corrected to Nannopodidae. The family-name Paranannopinae Por, 1986a
is a nomen nudum based on an unavailable generic name and is replaced by Danielsseniinae Huys & Gee in Huys et al.,
1996.
Four orphaned taxonomic groupings created by the removal of the type species – but not of the remaining species
included in a genus – require an existing (previously invalid) or new generic name. Amphiascus Sars, 1905a is a senior
objective synonym of Paramphiascopsis Lang, 1944 and must be restricted to the species currently included in the latter;
a new genus Sarsamphiascus (type: Dactylopus minutus Claus, 1863) is proposed to receive all remaining Amphiascus
species. The new generic names Monardius gen. nov. and Glabrotelson gen. nov. are proposed for the orphaned
taxonomic groupings resulting from the removal of the types of Teissierella Monard, 1935a to Robertsonia Brady, 1880a,
and of Hastigerella Nicholls, 1935 to Arenosetella Wilson, 1932, respectively. Leptomesochra Sars, 1911b is a senior
subjective synonym of Interleptomesochra Lang, 1965 and must be restricted to the latter’s taxonomic concept; the
previously unavailable generic name Leptameira Lang, 1936d is reinstated under the present authorship and date to
assemble all remaining Leptomesochra species.
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Type species are fixed for a number of genera that were proposed before 1931 without original type designation:
Marshia Herrick, 1895; Cerviniopsis Sars, 1903; Pseudobradya Sars, 1904a; Alteuthella A. Scott, 1909; Diosaccopsis
Brian, 1925a; and Nannopodella Monard, 1928.
Xouthous sarsi sp. nov. is proposed for Idomene forficata Philippi, 1843 sensu Sars (1906a). Tachidius longicornis
Olofsson, 1917 is reinstated and placed in the genus Geeopsis Huys in Huys et al., 1996 as G. longicornis (Olofsson,
1917) comb. nov. Micropsammis galapagoensis Mielke, 1997 is transferred to Telopsammis as T. galapagoensis (Mielke,
1997) comb. nov.
Key words: Harpacticoida, zoological nomenclature, type fixation, new replacement names, unavailable names, junior
synonyms, orphaned taxa, Bereacanthus nom. nov., Bereraia nom. nov., Dahlakocamptus nom. nov., Dussartopages
nom. nov., Hanikraia nom. nov., Sewelliapusia nom. nov., Glabrotelson gen. nov., Monardius gen. nov., Sarsamphiascus
gen. nov.
Introduction
With the recent publication of the comprehensive checklist of harpacticoids of the world (Wells 2007), the
classification of the group has – at least for the foreseeable future – acquired a level of stability it had not
witnessed for the last half-century. The ongoing search for natural order has now generated a testable
taxonomic framework of the Harpacticoida that can be challenged with alternative character sets such as
molecular sequence data. Wells (2007) recognized about 4,300 species placed in 589 genera and 56 families,
representing a five-fold increase in the number of species since the last comprehensive treatment by Lang
(1948). Given the vast number of synonyms, homonyms, new combinations and unavailable names that have
accumulated since the first harpacticoid copepod description by O.F. Müller (1776), it is to be expected that
not all nomenclatural problems have been resolved thus far. Wells’s (2007) thorough approach revealed
several unresolved cases of homonymy at the species level and new replacement names were proposed by the
author where required; the situation is, however, less satisfactory at genus level. Several harpacticoid genusgroup names are preoccupied by senior homonyms, and new replacement names have already been proposed
for some of them (Özdikmen & Pesce 2006; Özdikmen 2008, 2009). Dual designation of the same namebearing type or subsequent removal of the type species to another genus has resulted in a number of
unresolved cases of synonymy. One example of such “orphaned” taxonomic groupings that has recently come
to light is the genus Echinocamptus Chappuis, 1929a, the type species of which was also subsequently
designated as the type of the subgenus Bryocamptus (Limocamptus) Chappuis, 1929a (Wells 2007). However,
by far the largest source of nomenclatural confusion stems from the lack of attention to typification, which
provides stability and objective continuity in the application of names. It has recurrently been ignored that
genus-group names established after 1930 must be accompanied by the explicit fixation of a type species
(ICZN Art. 13.3) in order to make them available. Failure to comply with this requirement has introduced –
even until very recently – an unexpectedly high number of nomina nuda in the harpacticoid literature. Some
of these cases have been resolved (Galassi et al. 1999; Reid et al. 2003) but many unavailable genus-group
names still await type fixation.
The steady addition of new taxa shows that the dynamism of harpacticoid taxonomy is clearly set to
continue. Since Wells’s (2007) monumental checklist twenty-eight genera have already been added:
Ancorabolina George, 2006b; Apistophonte Gheerardyn & Fiers in Gheerardyn et al., 2006b; Carcinocaris
Cottarelli, Bruno & Berrera, 2006; Foweya Gee, 2006; Propephonte Gheerardyn & Fiers in Gheerardyn et al.,
2006b; Pseudechinopsyllus George, 2006a; Spiniferaphonte Gheerardyn & Fiers in Gheerardyn et al., 2007;
Aequinoctiella Cottarelli, Bruno & Berera, 2008; Arcticocarella Kornev & Chertoprud, 2008; Ciplakastacus
Sak, Karaytuğ & Huys, 2008; Mesopontia Sak, Huys & Karaytuğ, 2008; Onychopontia Sak, Huys &
Karaytuğ, 2008; Pesceus Özdikmen, 2008; Pordfus Özdikmen, 2008; Raowellsia Özdikmen, 2008;
Vermicaris Kornev & Chertoprud, 2008; Vibriopsyllus Kornev & Chertoprud, 2008; Acuticoxa Huys &
Kihara, 2009; Inermiphonte Huys & Lee, 2009; Marbefia Huys & Lee, 2009; Chaulionyx Kihara & Huys,
UNRESOLVED CASES IN HARPACTICOID COPEPOD NOMENCLATURE
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2009a; Paranaiara Kihara & Huys, 2009b; Dahmsopottekina Özdikmen, 2009; Muohuysia Özdikmen, 2009;
Monodicaris Schminke, 2009; Paranitocrella Tang & Knott, 2009; Nyxis Willen, 2009; Keraia Willen &
Dittmar, 2009 (Cottarelli et al. 2006, 2008; Gee 2006; George 2006a, 2006b; Gheerardyn et al. 2006b, 2007;
Huys & Kihara 2009; Huys & Lee 2009; Kihara & Huys 2009a, 2009b; Kornev & Chertoprud 2008;
Özdikmen 2008, 2009; Sak et al. 2008a, 2008b; Schminke 2009; Tang & Knott 2009; Willen 2009; Willen &
Dittmar 2009). In addition, the subgenus Parabradya Lang, 1944 was upgraded to generic status (Seifried et
al. 2007), the new replacement name Neomrazekiella Özdikmen & Pesce, 2006 was proposed for the
preoccupied subgeneric name Mrazekiella Brehm, 1949 (Özdikmen & Pesce 2006) and the genus Kinnecaris
Jakobi, 1972 was reinstated (Schminke 2008). Finally, Psammoleptastacus Pennak, 1942 (formerly a junior
subjective synonym of Arenopontia Kunz, 1937) and Neoleptastacus Nicholls, 1945b (formerly a subgenus of
Arenopontia) were reinstated as valid genera by Sak et al. (2008a). The proposal of the genus Eolaophonte by
Apostolov (1990) does not feature in any subsequent monograph or checklist and has escaped most workers’
attention till now (cf. Bodin 1997; Boxshall & Halsey 2004; Wells 2007). Both Phycolaophonte Pallares,
1975a and Eolaophonte Apostolov, 1990 were relegated to junior subjective synonyms of Coullia Hamond,
1973a by Huys (2009). Gheerardyn et al. (in press) relegated the recently proposed genus Archaeotisbe
Kornev & Chertoprud, 2008 to a junior subjective synonym of Atergopedia Martínez Arbizu & Moura, 1998.
In accordance with Wells’s (1967) earlier course of action (overruled by Kunz (1975a)) the genus Lineosoma
Wells, 1965b was synonymised with Noodtiella Wells, 1965b by Kihara and Huys (2009a). The genus
Cristacoxa Huys, 1990a was relegated to a junior subjective synonym of Noodtorthopsyllus Lang, 1965 by
Huys and Kihara (2009).
The purpose of this paper is to resolve, where possible and appropriate, outstanding nomenclatural issues
in harpacticoid taxonomy – at genus-group level and above. The problems addressed include cases of
homonymy, non-availability of names owing to lack of type fixation, and miscellaneous matters centered
mostly on subjective junior synonyms and “orphaned” taxonomic groupings. The solutions variously involve
proposal of new replacement names for homonyms, clarification of actual authorship of names made available
after their initial proposal, assumption of authorship of an unavailable name, and proposal of new names for
“orphaned” taxonomic assemblages. In three such cases a ruling by the International Commission on
Zoological Nomenclature will be required and proposals have been submitted to ensure that stability and
universality in the application of those names are not threatened (Huys 2008a, 2008b; Huys & Clark 2009).
The nomenclatural changes proposed in this paper are summarized in Table 1. The rationale behind these
changes is explained in detail below. Articles and Recommendations cited in the text refer to the fourth edition
of the International Code of Zoological Nomenclature (ICZN 1999), the provisions of which supersede those
of the previous editions of the Code with effect from 1 January 2000. The validity of type fixation for each of
the 601 genera of Harpacticoida currently in use was verified and the old and current binomen of the type
species as well as the nomenclatural act that established it are summarized in Table 4. Finally, the opportunity
was taken to point out two unresolved homonymy issues at genus-group level in other copepod orders.
Cases of homonymy
At least five harpacticoid generic names have entered into homonymy with previously established genusgroup names proposed for non-copepodan taxa. In each of these cases, both the prior nominal taxon name
(junior homonym) and its new replacement name proposed below have the same type species (ICZN Art.
67.8). The recently proposed family-group name Zosimidae Seifried, 2003 (type genus Zosime Boeck, 1873)
(cf. Seifried 2003: 100) is also a junior homonym of Zosiminae Alcock, 1898 (Decapoda, family Xanthidae;
type genus Zosimus A.-G. Desmarest, 1823) (cf. Alcock 1898: 77). Such a case resulting from similarity but
not identity of the names of the respective type genera must be referred to the Commission for a ruling to
remove homonymy (ICZN Art. 55.3.1). An application to preserve the family-group name Zosimidae
Seifried, 2003 by choosing a new grammatical stem from the name of the type genus Zosime (according to
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ICZN Recommendation 29A), taking the original authorship and date, has been submitted (Huys & Clark
2009).
Anoplosoma Sars, 1911c (Family Ameiridae)
Sars (1911c: 431) established the genus Anoplosoma Sars, 1911c for a new species A. sordidum Sars, 1911c
(type by monotypy) and placed it in the Ancorabolidae. Lang (1944, 1948) transferred the genus to the
Ameiridae (subfamily Stenocopiinae) while Por (1964a: 99) added a second species, A. stryx Por, 1964a. Sars
(1911c) himself used two different spellings for the generic name, Anoplosoma in the heading of the generic
diagnosis (p. 431) and the figure legend of Supplement Plate 49, and Anaplosoma in the combination with the
type species name in the text (p. 432). The latter spelling was adopted by some authors (e.g. Pesta 1927: 41)
but should be ranked as an incorrect original spelling (lapsus calami) since Sars (1911c: 432) clearly stated
“… it is from this character [the absence of any obvious armature on the body; Greek anoplos, meaning
unarmed] that the generic name here proposed has been derived…”.
Strand (1929: 9) pointed out that Anoplosoma Sars, 1911c was preoccupied by Anoplosoma Agassiz, 1846
and proposed the new replacement name Anoplosomella. This nomen novum has not been cited since and does
not feature in Boxshall and Halsey’s (2004) list of unused generic names. Agassiz (1846: 25, 1848: 70)
corrected the spelling of Anoplosomatum Grube, 1840 (type species: Anoplosomatum utriculus Grube, 1840)
to Anoplosoma. According to ICZN Arts 32.2 and 32.5, Grube’s (1840: 47) spelling of the name
Anoplosomatum is the correct original one and hence Agassiz’s subsequent spelling is an unjustified
emendation (Art. 33.2.3). Anoplosoma Agassiz, 1846 therefore becomes a junior objective synonym (as an
unnecessary replacement name) of Anoplosomatum Grube, 1840 and can enter into homonymy with other
genus-group names. There are only a few citations of Grube’s genus in the literature and all use the original
spelling without reference to Agassiz’s subsequent spelling (e.g. Siebold 1845: 321; Diesing 1859: 754, 766).
Initially thought to represent a “transition” from the Echinodermata to the worms, Baird (1868: 99)
subsequently listed Anoplosomatum under “Genera of Sipunculidae not sufficiently established, and the
position of which is doubtful” before Delage and Hérouard (1897: 24) eventually considered it a synonym of
Phascolosoma Leuckart, 1828 (Sipuncula: Phascolosomatidae).
Although the senior homonym has not been used as a valid name after 1899 (ICZN Art. 23.9.1.1), the
generic name Anoplosoma Sars, 1911c cannot be maintained as a nomen protectum on the basis of prevailing
usage since it has not been used in at least 25 works, published by at least 10 authors since 1960 (i.e. the
immediately preceding 50 years) (ICZN Art. 23.9.1.2). Strand’s (1929: 9) replacement of it by Anoplosomella
Strand, 1929 is confirmed, and the resulting new combinations must be cited as Anoplosomella sordida (Sars,
1911c) comb. nov. [type] and A. stryx (Por, 1964a) comb. nov. Note that the former is formally treated here as
a new combination since Strand (1929) had not mentioned the type species Anoplosoma sordidum when he
proposed the replacement name Anoplosomella.
Poria Lang, 1965 (Family Canthocamptidae)
Por (1964a: 115) synonymised the monotypic genera Hemimesochra Sars, 1920b (type species:
Hemimesochra clavularis Sars, 1920b) and Mesopsyllus Por, 1960b (type species: Mesopsyllus atargatis Por,
1960b) and added a third species, Hemimesochra derketo Por, 1964a. Lang (1965: 423) dismissed this course
of action by resurrecting Mesopsyllus for M. atargatis, restricting Hemimesochra to H. clavularis and fixing
H. derketo as the type and only species of a new genus Poria Lang, 1965. All three genera were removed from
the Cletodidae by Por (1986a) and placed in a new subfamily Hemimesochrinae in the Canthocamptidae.
Poria Lang, 1965 cannot be maintained as a valid name since it is a junior homonym of Poria Mulsant, 1850
(Coleoptera: Coccinellidae). Mulsant (1850: 885) included eight species in Poria but refrained from type
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fixation. Crotch (1874) subsequently listed Poria cyanea Mulsant, 1850 as the type species. The senior
homonym Poria Mulsant, 1850 is still a valid name that is widely used in the coccinellid literature (e.g.
Fürsch 1990; Lawrence & Newton 1995). Here the new replacement name Hanikraia nom. nov. (gender:
feminine), alluding to the location of the type locality (Israel; Eastern Mediterranean; off Rosh Hanikra) is
substituted for Poria Lang, 1965, containing Hanikraia derketo (Por, 1964a) comb. nov. as its type and only
valid species. Huys and Thistle (1989) suggested that the species identified as “Cletodidae sp. indet.” by Wells
(1965a) also belongs to Poria, but Bodin (1997) considered it to be an unidentifiable juvenile specimen. In
accordance with Wells’s (2007) checklist it is here regarded as species incertae sedis in Hanikraia pending the
re-examination of additional material.
Sewellia Lang, 1965 (Family Dactylopusiidae)
Sewell (1940: 227) described the new species Dactylopusia tropica Sewell, 1940 on the basis of an
unspecified number of females. Krishnaswamy (1957: 37) described the hitherto unknown male based on
non-type material. Hence, his statement that the “… male allotype will be lodged in the Zoological Survey…”
is obviously in violation with the Code (ICZN Art. 72.1.1). Sewell (1940: 229) believed the species formed a
“connecting link” between the genera Dactylopusia Norman, 1903a and Dactylopusioides Brian, 1928b. Lang
(1965: 168) proposed a new genus Sewellia Lang, 1965 to accommodate D. tropica as its type and only
species (by original designation), overlooking that Hora (1932: 315) had already used the same genus-group
name for a genus of river loaches (Cypriniformes, Balitoridae). Sewellia Lang, 1965 is therefore a junior
homonym of Sewellia Hora, 1932 (type: Balitora lineolata Valenciennes in Cuvier and Valenciennes, 1846,
by monotypy). A new replacement name, Sewelliapusia nom. nov. (named after R. B. Seymour Sewell;
gender: feminine), is proposed to resolve this case of homonymy and hence the type species should be cited as
Sewelliapusia tropica (Sewell, 1940) comb. nov. The genus has remained monotypic since its inception and is
currently placed in the family Dactylopusiidae (Willen 2000).
Ichnusella Cottarelli, 1971 (Family Leptopontiidae)
Cottarelli (1971: 58) established this genus for his new species Ichnusella eione Cottarelli, 1971 (type species
by original designation) and Psammopsyllus pasquinii Cottarelli, 1969, overlooking that the generic name
Ichnusella had already been introduced by Dieni and Massari (1966: 170) for a genus of fossil Foraminiferida.
Since the Code applies to Metazoa and also to protistan taxa when workers treat them as animals for the
purposes of nomenclature (ICZN Art. 1.1.1), Ichnusella Cottarelli, 1971 is a junior homonym of Ichnusella
Dieni & Massari, 1966 (type species by original designation: Ichnusella trocholinaeformis Dieni & Massari,
1966), both being derived from the ancient name of Sardinia (Greek Ίχνοΰσα). A new replacement name,
Bereraia nom. nov. (gender: feminine), is proposed, named after Dr Raffaella Berera (Università della Tuscia,
Viterbo, Italy), in recognition of her valuable contributions to the knowledge of the genus (Berera & Cottarelli
2003; Berera et al. 2001; Cottarelli et al. 1998). The genus currently includes the following species:
Psammopsyllus pasquinii Cottarelli, 1969 = Bereraia pasquinii (Cottarelli, 1969) comb. nov.; Ichnusella
eione Cottarelli, 1971 = B. eione (Cottarelli, 1971) comb. nov. [type]; Ichnusella longifurca Cottarelli, Bruno
& Berera, 1998 = B. longifurca (Cottarelli, Bruno & Berera, 1998) comb. nov.; Ichnusella tertia Cottarelli,
Bruno & Berera, 1998 = B. tertia (Cottarelli, Bruno & Berera, 1998) comb. nov.; Ichnusella improvisa Berera,
Cottarelli & Bruno, 2001 = B. improvisa (Berera, Cottarelli & Bruno, 2001) comb. nov.; and Ichnusella ionica
Berera & Cottarelli, 2003 = B. ionica (Berera & Cottarelli, 2003) comb. nov.
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Dahlakia Por, 1986a (Family Canthocamptidae)
Por (1968: 45) described a new species of Cletodidae, Cletocamptus xenuus Por, 1968, from the Dahlak
Archipelago but expressed reservations about its generic assignment. In a later paper (Por 1986a: 123), he
proposed a new genus Dahlakia Por, 1986a in the subfamily Hemimesochrinae (Canthocamptidae) and
designated C. xenuus as its type and only species. Dahlakia Por, 1986a is a junior homonym of Dahlakia
Biggs, 1971 (type species by original designation: Dahlakia leilae Biggs, 1971) (Gastropoda: Cerithiidae).
Houbrick (1978) considered Biggs’s (1971: 221) genus a junior subjective synonym of Bittium Leach in Gray,
1847 while some authors have used Dahlakia as a subgenus of Bittium (e.g. Bosch et al. 1995). In a later
paper Houbrick (1992: 270) proposed that the type species of Dahlakia is in fact a probable synonym of
Cerithium scabridum Philippi, 1848, and therefore excluded the genus from the Bittiinae (family Cerithiidae).
It is proposed here to remove Dahlakia Por, 1986a from its homonymy by replacing it with the new
replacement name Dahlakocamptus nom. nov. (gender: masculine), containing Dahlakocamptus xenuus (Por,
1968) comb. nov. as its type and sole species.
Unavailable genus-group names
In order to be available every new genus-group name published after 1930 must be accompanied by the
fixation of a type species in the original description. Such mandatory type fixation (ICZN Art. 13.3) has been
ignored for a number of harpacticoid genus-group names and its strict application does not appear to have had
much impact on copepod nomenclature until recently. Both Galassi et al. (1999) and Reid et al. (2003)
recognized that Petkovski’s (1976) generic names Nitocrellopsis Petkovski, 1976 and Stygonitocrella
Petkovski, 1976 had been established without type fixation and consequently made them available under their
own authorship by fixing Nitocrellopsis rouchi Galassi, De Laurentiis & Dole-Olivier, 1999 and Nitocrella
montana Noodt, 1965 as the respective types.
A taxon denoted by an unavailable name requires a replacement name which – in the absence of
potentially valid synonyms – can either be the old name or a newly proposed name (ICZN Art. 23.3.5). I have
elected to validate the old names instead of proposing new substitute names which are likely to be
destabilizing even if they are allowed by the Code. The adoption of a previously unavailable name as the valid
name of a taxon establishes it as a new name with its own authorship and date (ICZN Glossary: nomen
nudum). The Code (Arts 11.9.3 and 51.3) is not explicit about the nomenclatural implications of such course
of action but its logical interpretation entails that (1) since this does not involve homonymy between two
available names (in which case type fixation would be carried over to the new replacement name), the adopted
unavailable name is not a nomen novum but must be indicated by the term gen. nov. (thereby satisfying the
provisions of ICZN Art. 16.1 for names published after 1999); the term nom. nov. should only be used to
indicate a new replacement name (ICZN Recommendation 16A); (2) since species-group names must be
published in combination with a generic name, but the latter need not be valid or even available (ICZN Art.
11.9.3.1) the unavailability of genus-group names has no implications for its originally or subsequently
included species other than that they must be treated as new combinations; hence the authorship and date of
the author(s) making the name available must be cited in parentheses (ICZN Art. 51.3); and (3) since the name
was originally unavailable, type fixation is by original rather than subsequent designation.
In the introduction to his monograph, Lang (1948: 7) stated that whenever the type species of a genus was
not explicitly indicated by him, he considered the first species to have been described in that genus as the type
(“Wenn der Genotypus für eine Gattung nicht angegeben wird, betrachte ich die zuerst beschriebene Art als
solcher.”). Such a rule is not among the allowed criteria for ascertaining type fixation in the original
publication (ICZN Art. 68), and thus the status of being the “oldest species” assigned to one of Lang’s genera
does not in and of itself confer type status on that species. According to ICZN Art. 67.5 the term “designation”
in relation to fixation of a type species must be rigidly construed, which applies to Art. 69 for subsequent
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designation as well. Additional problems associated with Lang’s (1948) “rule” are that for several genera
listed by him (Aegisthus Giesbrecht, 1891; Alteuthella A. Scott, 1909; Ameiropsis Sars, 1907b; Cerviniopsis
Sars, 1903; Idyanthe Sars, 1909c; Idyella Sars, 1905b; Psammastacus Nicholls, 1935), more than one
originally included species shared the same publication date, and that these species also had the oldest names
among their congeners. In one case (Pseudobradya Sars, 1904a) the oldest nominal species listed by Lang
(1948: 237) was not even originally included (Bradya brevicornis T. Scott, 1894a).
Rhizothrix Brady & Robertson, 1876 (Family Rhizotrichidae)
The binomen Rhizothrix curvata was first mentioned as a nomen nudum in a species list by Brady and
Robertson (1876: 197). Brady (1880a: 98) abandoned the intended generic name and described the species as
Enhydrosoma curvatum, attributing authorship to “Brady & Robertson”. Sars (1909a: 302) erroneously
claimed that Brady and Robertson (1876) had established the genus Rhizothrix and unfortunately this error
has been perpetuated in the literature (e.g. Lang 1948; Bodin 1997; Boxshall & Halsey 2004; Wells 2007;
Kornev & Chertoprud 2008). In fact, by providing a detailed generic diagnosis and therefore satisfying the
provisions of ICZN Art. 12.1, Sars (1909a) himself had made the genus-group name available. Consequently
this author should be credited with the authorship as Huys et al. (1996: 40) had recognized already.
The type species of Rhizothrix Sars, 1909a (by monotypy) is Enhydrosoma curvatum Brady, 1880a =
Rhizothrix curvata (Brady, 1880a). It should be noted that the species cannot be attributed to “Brady &
Robertson, 1880” or “Brady & Robertson in Brady, 1880a” since an outside person (i.e. other than an author
of the work) can only be credited with authorship if he/she is alone responsible for the name and for satifying
the criteria of availability (ICZN Art. 50.1.1). The same rule applies to the type species of the genera
Jonesiella Brady, 1880a (Pseudotachidiidae), Normanella Brady, 1880a (Normanellidae), Robertsonia Brady,
1880a (Miraciidae) and Stylicletodes Lang, 1936b (Cletodidae), all of which have been erroneously attributed
to Brady & Robertson by Brady (1880a) (Table 4).
Boxshall and Halsey (2004: 378) remarked that the traditional spelling of the family name, Rhizothricidae
Por, 1986a, reflects an incorrect derivation that must be amended to Rhizotrichidae.
Psammastacus Nicholls, 1935 (Family Leptastacidae)
Nicholls (1935) proposed the genus Psammastacus Nicholls, 1935 for two new species, P. confluens Nicholls,
1935 and P. brevicaudatus Nicholls, 1935, but did not designate a type species, without which the generic
name remained unavailable as a nomen nudum. Lang’s (1948: 7) rule of designating the oldest species as type
(unless stated otherwise) does not satisfy the provisions of the Code (see above). The first authors to formally
fix a type species were Apostolov and Marinov (1988: 288) who designated P. confluens and consequently reestablished the generic name. They also met the provisions of ICZN Art. 13.1.1 by providing a generic
diagnosis (in Bulgarian) which serves to differentiate Psammastacus Apostolov & Marinov, 1988 from the
other leptastacid genera. Accordingly, the latter generic name should take their authorship and date. Huys
(1992: 121) showed that the original description of P. brevicaudatus was based on copepodids of P. confluens.
Given this synonymy, the genus Psammastacus Apostolov & Marinov, 1988 remains monotypic with
Psammastacus confluens (Nicholls, 1935) comb. nov. as the type.
Paranannopus Lang, 1936a (Family Pseudotachidiidae)
Lang (1936b: 476; published 21st September) is usually considered as the authority who introduced the
generic name Paranannopus, however the new name had already been proposed in an earlier paper (Lang
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1936a: 51; published 20th March) to accommodate Nannopus abyssi Sars, 1920c and Paranannopus sarsi
Lang, 1936a. In neither of Lang's (1936a, b) papers was a type species fixed, rendering the generic name
Paranannopus Lang, 1936a unavailable. The first author attempting at fixing a type was Lang (1948: 7, 1330)
himself by citing Nannopus abyssi as the earliest described species originally included in the genus, however
his habitual proclivities are insufficient to recognize his type fixation as valid (see above). Huys et al. (1996:
244) also attempted to reinstitute the genus Paranannopus by attributing authorship to “Huys & Gee, 1996” –
however, the completion of this paper which included a revision of the genus has not been published yet.
Although the name Paranannopus Huys & Gee in Huys et al., 1996 was accompanied by a diagnosis stating
in words characters that are purported to differentiate the taxon (and hence meeting the provisions of ICZN
Art. 13.1.1), it is to be considered a nomen nudum since no type was explicitly selected (Wells 2007: 85; ICZN
Art. 13.3). The generic name (gender: masculine) is here made available, taking the authorship and date of the
present paper, by fixing Nannopus abyssi as the type species by original designation and by making reference
to Huys et al.’s (1996: 244) generic diagnosis as the standard for differentiating the taxon from its confamilial
members. Being unavailable, both Paranannopus Lang, 1936a [nomen nudum] and Paranannopus Huys &
Gee in Huys et al., 1996 [nomen nudum], cannot enter into the synonymy of Paranannopus gen. nov., which
is here explicitly indicated as intentionally new (ICZN Art. 16.1). The genus currently includes 22 valid
species which must be cited as new combinations as follows: Nannopus abyssi Sars, 1920c = Paranannopus
abyssi (Sars, 1920c) comb. nov. [type]; Paranannopus sarsi Lang, 1936a = Paranannopus sarsi (Lang,
1936a) comb. nov.; P. echinipes Smirnov, 1946 = P. echinipes (Smirnov, 1946) comb. nov.; P. minutus
Smirnov, 1946 = P. minutus (Smirnov, 1946) comb. nov.; P. bahusiense [sic] Por, 1964b = P. bahusiensis (Por,
1964b) comb. nov.; P. philistinus Por, 1964c = P. philistinus (Por, 1964c) comb. nov.; P. caheti Soyer, 1965 =
P. caheti (Soyer, 1965) comb. nov.; P. triarticulatus Wells, 1965a = P. triarticulatus (Wells, 1965a) comb.
nov.; P. atlanticus Coull, 1973a = P. atlanticus (Coull, 1973a) comb. nov.; P. wellsi Soyer, 1976 = P. wellsi
(Soyer, 1976) comb. nov.; P. longithorax Becker, 1979 = P. longithorax (Becker, 1979) comb. nov.; P.
reductus Becker, 1979 = P. reductus (Becker, 1979) comb. nov.; P. truncatus Becker, 1979 = P. truncatus
(Becker, 1979) comb. nov.; P. plumosus Schriever, 1983 = P. plumosus (Schriever, 1983) comb. nov.; P.
denticulatus Schriever, 1985 = P. denticulatus (Schriever, 1985) comb. nov.; P. hicksi Schriever, 1985 = P.
hicksi (Schriever, 1985) comb. nov.; P. kunzi Schriever, 1985 = P. kunzi (Schriever, 1985) comb. nov.; P.
singulosetosus Schriever, 1985 = P. singulosetosus (Schriever, 1985) comb. nov.; P. trisetosus Schriever, 1985
= P. trisetosus (Schriever, 1985) comb. nov.; P. uniarticulatus Schriever, 1985) = P. uniarticulatus (Schriever,
1985) comb. nov.; P. variabilis Schriever, 1985 = P. variabilis (Schriever, 1985) comb. nov.; P. arndwilleni
Willen, 2005 = P. arndwilleni (Willen, 2005) comb. nov. The species described by Wells (1965a: 19) under the
name Paranannopus langi Wells, 1965a [= Paranannopus langi (Wells, 1965a) comb. nov.] is considered
species incertae sedis in this genus (cf. Wells 2007: 21).
Huys et al. (1996: 236), building on the non-availability of the generic name Paranannopus Lang, 1936a
[nomen nudum], introduced the new family name Danielsseniidae to replace Paranannopidae Por, 1986a
[nomen nudum], since family-group names taking their stem from an unavailable (and thus invalid) generic
name, also become unavailable (ICZN Arts 11.7.1.1 and 63–64). In addition, it was believed at the time that
continued use of the family name Paranannopidae would have been preferential for the sake of stability, but
would also have the less favourable implication of basing the family on a type genus that (1) is very advanced
and not really representative for the family, (2) is mostly known from females only, and (3) misleadingly
alludes to a relationship with the genus Nannopus Brady, 1880a, currently placed in the family
Huntemanniidae (= Nannopodidae, see below). Alternatively, the second option offered the possibility of
designating another genus that is both well known and representative of the family. The family name
Danielsseniidae Huys & Gee in Huys et al., 1996 was accompanied by a differential diagnosis (Huys et al.
1996: 236; ICZN Art. 13.1.1) and was based on the available and still valid generic name Danielssenia Boeck,
1873 (ICZN Art. 13.2). Although the family name was not explicitly indicated as intentionally new or
accompanied by a citation of the type genus (these provisions only apply to family-group names published
after 1999; ICZN Arts. 16.1–2), the family-group name Danielsseniidae is valid. The family Paranannopidae
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Por, 1986a [nomen nudum] was relegated to a subfamily of the Pseudotachidiidae Lang, 1936c by Willen
(1999, 2000); hence at this rank it should be cited as Danielsseniinae Huys & Gee in Huys et al. (1996). Note
also that the ending of the generic name Paranannopus is derived from the Grrek stem pous (πους), meaning
foot, and hence the correct spelling of the family name should have been Paranannopodidae.
Paraidya Sewell, 1940 (Family Tisbidae)
Sewell (1940: 163) established the genus Paraidya Sewell, 1940 in the Tisbidae for two new species,
Paraidya major Sewell, 1940 and P. minor Sewell, 1940. Since the publication of Paraidya was not
accompanied by the mandatory type fixation, the generic name does not satisfy the provisions of ICZN Art.
13.3 and must therefore be considered unavailable. Vervoort (1964: 103) listed the type species for each genus
then regarded valid in the Tisbidae. Unfortunately he did not do so for Paraidya since he claimed that the
taxon fell within the boundaries of the genus Tisbe Lilljeborg, 1853 as redefined by Lang (1948: 364). This
view gained support from Lang (1965: 145) who formally relegated Paraidya to a junior subjective synonym
of Tisbe. Humes and Ho (1969b: 128) disputed this course of action and reinstated the genus for the two
originally included species and a new species P. occulta Humes & Ho, 1969b but refrained from subsequent
type designation. Finally, Humes (1981a) redescribed both P. major and P. minor but did not make the genusgroup name available either. In the absence of any available synonyms that could potentially substitute for
Paraidya Sewell, 1940 [nomen nudum], the generic name is validated here by fixing Paraidya major Sewell,
1940 as the type species (ICZN Art. 13.3) and by making explicit reference to Humes and Ho’s (1969b: 128)
list of characters that differentiate Paraidya gen. nov. from the closely related genus Tisbe (ICZN Art. 13.1.2).
Paraidya gen. nov. includes the following new combinations: Paraidya major Sewell, 1940 = Paraidya major
(Sewell, 1940) comb. nov.; P. minor Sewell, 1940 = P. minor (Sewell, 1940) comb. nov.; and P. occulta
Humes & Ho, 1969b = P. occulta (Humes & Ho, 1969b) comb. nov.
Alteuthellopsis Lang, 1944 (Family Peltidiidae)
Lang (1944: 11) introduced the generic name Alteuthellopsis for the single species, Eupelte oblivia A. Scott,
1909 but did not present a diagnosis or a bibliographic reference to such a description, hence the name
remained unavailable. It was not until 1948 that Lang (p. 457) satisfied the provisions of ICZN Art. 13.1 by
providing a generic diagnosis for Alteuthellopsis and fixing Eupelte oblivia as the type species (by monotypy).
The generic name must therefore be attributed to Lang (1948). Humes (1981b: 227) added a second species,
Alteuthellopsis corallina Humes, 1981.
Halectinosoma Lang, 1944 (Family Ectinosomatidae)
Lang (1944: 6) divided the genus Ectinosoma Boeck, 1865 into two subgenera, Ectinosoma (type:
Ectinosoma melaniceps Boeck, 1865) and the newly proposed Halectinosoma for which he did not fix a type.
Although Lang (1944) designated a type for each of the two species-groups delimited within this latter
subgenus, viz., the sarsii-group (type: Ectinosoma sarsii Boeck, 1873) and the curticorne-group (type:
Ectinosoma curticorne Boeck, 1873), the subgenus Halectinosoma was denoted by an unavailable name and
remained so in Lang’s (1948: 194) monograph. Lang (1944) did not include any other species in these groups
but expanded them significantly in his 1948 monograph by adding twelve species to the sarsii-group and six
to the curticorne-group. To the former group of species he referred Ectinosoma sarsii Boeck, 1873; Tachidius
abrau Krićagin, 1878; Ectinosoma chrystalii T. Scott, 1894a; E. propinquum Scott & Scott, 1896; E.
herdmani Scott & Scott, 1896; E. armiferum Scott & Scott, 1896; E. finmarchicum T. Scott, 1903a; E.
neglectum Sars, 1904a; E. elongatum Sars, 1904a; E. brunnea Brady, 1907 (an incorrect original spelling of
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brunneum); E. proximum Sars, 1919; E. angulifrons Sars, 1919; and E. tenerum Sars, 1920a. As has been
pointed out by Karanovic and Pesce (2001), Vervoort (1962: 399) explicitly fixed Ectinosoma sarsii Boeck,
1873 as type species of the subgenus Halectinosoma. Lang (1965: 11), who upgraded Halectinosoma to
generic status, did not mention Vervoort’s (1962) designation. Halectinosoma is available from Vervoort
(1962), who cited (p. 255) the diagnosis for Halectinosoma in Lang (1944: 6) in this connection and
designated a type species, and it takes the authorship Halectinosoma Vervoort, 1962 (ICZN Arts 13.1.1 and
13.3).
There are, however, a number of unresolved issues associated with the name:
(1) Boeck’s (1873) original description of the type species Halectinosoma sarsii (Boeck, 1873) is poor and
lacks information on the structure of the maxilla and maxilliped, which are used to distinguish between the
very closely related genera Halectinosoma and Pseudobradya Sars, 1904a (Huys et al. 1996). According to
Clément and Moore (1995: 256) it is uncertain whether the species belongs to Halectinosoma and
consequently it should be placed as species incertae sedis in the genus. According to ICZN Art. 67.2.5, a
nominal species is deemed not to be originally included if it was cited as a species incertae sedis, however
since Halectinosoma sarsii was not cited with that status by Vervoort (1962), his type fixation cannot be
invalidated on that ground. According to Scott and Scott (1896) the new species described as Ectinosoma
spinipes by Brady (1880a: 9–10, Plate XXXVI, figs. 1–10) is a junior subjective synonym of Ectinosoma
sarsii but like many other earlier records of H. sarsii this identification is probably incorrect (Clément &
Moore 1995). Most workers have adopted Sars’s (1904a: 30, Plate XVI) redescription of Ectinosoma sarsii as
the standard of reference for correct identification but Clément and Moore (1995) showed there were major
discrepancies between Sars’s material and the original description given by Boeck (1873) and consequently
renamed it Halectinosoma pseudosarsi. There are no verifiable published records of Ectinosoma sarsii since
Boeck’s (1873) type material no longer exists.
(2) Since H. sarsii cannot be the objective standard of reference for the application of the name
Halectinosoma, another species, which can best serve stability and universality, should be fixed. Dussart and
Defaye (1988: 11) inadvertently cited Tachidius abrau Krićagin, 1878 as the type species but, being one of the
very few freshwater species of the genus, it is not representative. A ruling by the Commission will be required
to set aside the respective type designations by Vervoort (1962) and Dussart and Defaye (1988) and fix a new
type species for this genus.
(3) The genus Pararenosetella Lang, 1944 is a senior subjective synonym of Halectinosoma Vervoort, 1962.
Initially proposed for the type and only originally included species, Ectinosoma erythrops Brady, 1880a (Lang
1944: 6), a further four species were included in the genus by Lang (1948: 255): Ectinosoma gracile Scott &
Scott, 1896; E. longicorne Scott & Scott, 1896; E. tenuireme Scott & Scott, 1896; and E. leptoderma Klie,
1929. Chappuis (1954a: 38) described Pararenosetella meridionalis Chappuis, 1954a while Noodt (1955a:
87–89) added the new species Pararenosetella psammae Noodt, 1955a and transferred Ectinosoma oblongum
Kunz, 1949 to the genus Pararenosetella. In two subsequent papers Noodt (1958: 58; 1964: 131) raised the
number of species to ten by adding Pararenosetella litoralis Noodt, 1958 and P. ghardaqensis Noodt, 1964.
Lang (1965: 10, 547) dissolved the genus, claiming there were two distinct, not closely related lineages of
species in Pararenosetella, and reallocated the species to three previously described genera. Ectinosoma
erythrops, E. gracile, E. longicorne, E. tenuireme and E. oblongum were transferred to the genus
Halectinosoma, Pararenosetella litoralis and P. ghardaqensis to the genus Ectinosoma and Pararenosetella
meridionalis, P. psammae and Ectinosoma leptoderma to the genus Hastigerella Nicholls, 1935. The species
figured as Pararenosetella sp. (?) by Wells (1963: 12) was designated as the type and only species (under the
new binomen Lineosoma iscensis Wells, 1965b, an incorrect original spelling of iscense) of the genus
Lineosoma Wells, 1965b (Wells 1965b: 33). Despite Lang’s (1965) rejection of the genus, two more species
were added to Pararenosetella, however both P. monniotae Guille & Soyer, 1966 and P. clavata Rao &
Ganapati, 1969 were formally placed in the genus Hastigerella by Bodin (1967: 13) and Bodin (1976: 8),
respectively. Since the type, Ectinosoma erythrops, is currently included in Halectinosoma, Pararenosetella is
the only valid replacement name for the latter (ICZN Art. 23.3.5). Adopting this name would, however, cause
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instability as it would upset a long-accepted name in its accustomed meaning.
The nomenclatural problems outlined above are currently being presented to the Commission, asking to
use its Plenary Power (a) to give the generic name Halectinosoma Vervoort, 1962 precedence over the generic
name Pararenosetella Lang, 1944, whenever they are considered to be synonyms, and (b) to set aside all
previous fixations of type species for Halectinosoma and to designate Ectinosoma chrystalii Scott, 1894a as
the type species (Huys 2008a). Since prevailing usage of names is to be maintained until the ruling of the
Commission is published (ICZN Art. 28.1), Halectinosoma sarsii is still listed as the type species in Table 4.
Heterolaophonte Lang, 1944 (Family Laophontidae)
In his revision of the Laophontidae, Nicholls (1941b: 98) divided the type genus Laophonte Philippi, 1840
into five subgenera: Laophonte Philippi, 1840 (type species Laophonte cornuta Philippi, 1840 by monotypy),
Mesolaophonte Nicholls, 1941b (type species Laophonte littorale Scott & Scott, 1893b by original
designation; an incorrect original spelling of littoralis), Metalaophonte Nicholls, 1941b (type species
Laophonte depressa Scott, 1894b by original designation), Monolaophonte Nicholls, 1941b (type species
Laophonte curvata Douwe, 1929 by monotypy) and Neolaophonte Nicholls, 1941b (type species Laophonte
trilobata Willey, 1929 by original designation).
Lang (1944: 34) proposed the generic name Heterolaophonte but did not fix a type species. He divided the
genus into seven species-groups and designated a ‘Typus’ for each: (1) stroemii-group (type Cyclops stroemii
Baird, 1837), (2) minuta-group (type Laophonte minuta Boeck, 1873), (3) littoralis-group (type L. littoralis
Scott & Scott, 1893b), (4) quinquespinosa-group (type L. quinquespinosa Sewell, 1924), (5) discophoragroup (type L. discophora Willey, 1929), (6) campbelliensis-group (type L. campbelliensis Lang, 1934) and
(7) tenuispina-group (type L. tenuispina Lang, 1934). Wells et al. (1982: 178) proposed a new genus
Quinquelaophonte Wells, Hicks & Coull, 1982 (type by original designation Laophonte quinquespinosa
Sewell, 1924) for the species of the quinquespinosa-group. Nowadays, the other groups are no longer
recognized as taxonomically useful units (Wells 2007: 85).
Many of the names and nomenclatural acts proposed by Lang (1948) take precedence in the 1944
preamble to his monograph. Lang’s (1944) paper was not widely disseminated (but nevertheless satisfied the
criteria of publication) and most post-1948 authors have ignored it or were not aware of its existence (a
notable exception is Vervoort (1964) who was unfairly criticised by Lang (1965: 547) for his allegedly
“imperfect knowledge” of the literature!). Recent workers (Wells & Rao 1987; Huys & Willems 1989; Huys
1990a, 1992; Huys & Conroy-Dalton 1996; Bodin 1997; Seifried 2003; Wells 2007) have started crediting
Lang (1944) with the authorship of the respective names and acts but it has remained unnoticed that some
generic names, such as Heterolaophonte and Paralaophonte (see below), were not explicitly made available
by that work. As pointed out by Vervoort (1964: 333), Lang (1948: 1368) formally designated Cyclops
stroemii Baird, 1837 as the type species of Heterolaophonte and hence the authorship and date of the generic
name Heterolaophonte should be attributed to Lang (1948). Vervoort and Holthuis (1983: 56) subsequently
pointed out that Norman (1903a: 368) had already designated C. stroemii as the type species of Dactylopusia
Norman, 1903a (family Dactylopusiidae) (a new replacement name for Dactylopus Claus, 1863 non Gill,
1859). Since Norman (1903a), when designating C. stroemii as the type species of Dactylopusia, had intended
the dactylopusiid species identified by Claus (1863: 126, plate XVI, Figs 1–6) as such and not the real C.
stroemii, Vervoort and Holthuis (1983) asked the Commission to use its plenary power to set aside all
previous type fixations made for Dactylopusia Claus, 1863 and to designate Dactylopus tisboides Claus, 1863
as type species. The Commission voted in favour of Vervoort and Holthuis’s (1983) application and, as a
result, the generic name Heterolaophonte Lang, 1948 and the specific name of its type species, Cyclops
stroemii Baird, 1837, were placed on the Official Lists of Generic and Specific Names in Zoology,
respectively (Melville 1985).
Since Lang (1948) assigned both Laophonte littoralis Scott & Scott, 1893b (spelled incorrectly by Scott
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and Scott (1893b: 238) as littorale) (type of Mesolaophonte Nicholls, 1941b) and Laophonte curvata Douwe,
1929 (type of Monolaophonte Nicholls, 1941b) to the genus Heterolaophonte, the generic names
Mesolaophonte and Monolaophonte are currently senior subjective synonyms of Heterolaophonte. Nicholls’s
(1941b) subgeneric division was based solely on the endopodal armature of the female third pair of swimming
legs and his system has been criticised for its artificiality by Lang (1948: 1620) and Vervoort (1964: 314).
Both authors abolished Nicholls’s subgenera but they were not formally synonymised with or subsumed
within existing genera in prevailing usage. Since Mesolaophonte Nicholls, 1941b and Monolaophonte
Nicholls, 1941b are perfectly legitimate and available names, they cannot be ignored. The subgeneric name
Mesolaophonte has only been used twice as a valid name since Lang (1948). Krishnaswamy (1959: 29)
assigned his new species Laophonte pseudoculata Krishnaswamy, 1959 (incorrect original spelling pseudooculata) to Laophonte (Mesolaophonte) and Raibaut (1962) suggested a similar subgeneric assignment for
Laophonte commensalis Raibaut, 1962. The replacement of the well-known and universally accepted name
Heterolaophonte Lang, 1948 by one of its two underused senior subjective synonyms would result in many
new combinations and undue confusion in the nomenclature and taxonomy of the Laophontidae in general,
and of its most speciose genus in particular. Unfortunately, since the conditions of ICZN Art. 23.9.1 are not
met (the senior synonyms have been used as valid names after 1899), reversal of precedence is only possible
by a Commission’s ruling (ICZN Art. 23.9.3). To promote stability an application (Huys 2008b) is currently
being presented to the Commission, asking to use its Plenary Power to give the generic name Heterolaophonte
Lang, 1948 precedence over the names Mesolaophonte Nicholls, 1941b and Monolaophonte Nicholls, 1941b,
whenever it and either of the other two are considered to be synonyms. If in the light of future revisions,
Heterolaophonte is found not to be congeneric with Mesolaophonte and/or Monolaophonte both senior genusgroup names are still available to denote the two taxa as originally proposed by Nicholls (1941b).
Idyellopsis Lang, 1944 (Family Idyanthidae)
Lang (1944: 11) proposed the generic name Idyellopsis Lang, 1944 and fixed Idyellopsis typica Lang, 1944 as
the type without describing or figuring it until 1948. According to ICZN Art. 13.4, the combined description
or definition of a new nominal genus and a single included new nominal species, if marked by “gen. nov., sp.
nov.” or an equivalent expression, is deemed to confer availability on each name. However, no such
expression is apparent from Lang’s (1944) diagnosis (the genus is marked by “Gen. Idyellopsis, nov.” while
the species is not mentioned until the end of the generic diagnosis as “Typus I. typica n. sp.”). Since the name
I. typica is not accompanied by a description or definition that states in words characters that are purported to
differentiate the species (ICZN Art. 13.1.1), or by a bibliographic reference to such a published statement
(ICZN Art. 13.1.2), the name of the type species is effectively unavailable. Consequently, the generic name
Idyellopsis Lang, 1944 also becomes unavailable since it does not meet the provisions of ICZN Art. 13.3 for
genus-group names published after 1930 (type fixation is mandatory). The first author to make Idyellopsis
available was Lang (1948: 413) who provided a differential diagnosis for the genus and its type species (by
indication), Idyellopsis typica; hence the date and authorship of both should rest with Lang (1948). The genus
has remained monotypic since.
Paralaophonte Lang, 1944 (Family Laophontidae)
The genus-group name Paralaophonte was first published by Lang (1944: 36) but without any valid type
fixation. Lang (1944) divided the genus in four species-groups and designated a ‘type species’ for three of
them: (1) brevirostris-group (no type designated), (2) perplexa-group (type: Laophonte perplexa T. Scott,
1899a), (3) gracilipes-group (type: Laophonte gracilipes Brady, 1910), and (4) karmensis-group (type:
Laophonte karmensis Sars, 1911c). Vervoort (1964: 334) pointed out that Paralaophonte Lang, 1944 is an
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unavailable generic name which was subsequently validated by Lang (1948: 1386) by the indication of a type
species (Cleta brevirostris Claus, 1863) and providing an accompanying generic diagnosis. The authorship
and date of this genus should, therefore, be attributed as Paralaophonte Lang, 1948. For similar reasons Lee
and Huys (1999b: 315) remarked that Paronychocamptus Lang, 1944 is also a nomen nudum which was
subsequently made available by Lang (1948: 1380).
Robertgurneya Lang, 1944 (Family Miraciidae)
The genus Robertgurneya was first diagnosed by Lang (1944: 20) but without a rigidly construed fixation of a
type species. The author divided the genus in two species-groups and designated a type for each: similis-group
(type: Stenhelia simulans Norman & T. Scott, 1905) and spinulosus-group (type: Amphiascus spinulosus Sars,
1911a). Lang (1948: 697) suggested that both species groups may eventually be recognised as separate genera
and under those circumstances he would reserve the name Robertgurneya for the similis-group and propose a
new name Robertgurneyella for the spinulosa-group. Such conditional proposal cannot be considered a
rigidly construed type designation under the Code (ICZN Art. 67.5.3) and neither can his habitual designation
of the earliest described species (Stenhelia similis A. Scott, 1896) as the type (Lang 1948: 7). Under ICZN
Arts 13.1 and 13.3, the first authors to make the genus-group name Robertgurneya available were Apostolov
and Marinov (1988: 173) who cited “Robertgurneya similis A. Scott, 1896” [sic] as the type and provided a
differential diagnosis of the genus (in Bulgarian). The correct author attribution for Robertgurneya must
therefore be “Apostolov & Marinov, 1988” and the unavailable generic name Robertgurneya Lang, 1944
[nomen nudum] cannot enter into its synonymy. The genus includes the following species and subspecies:
Stenhelia similis A. Scott, 1896 = Robertgurneya similis (A. Scott, 1896); Robertgurneya similis
bulbamphiascoides Noodt, 1955b = R. similis bulbamphiascoides (Noodt, 1955b) comb. nov.; Stenhelia
eythraea A. Scott, 1902 = R. erythraea (A. Scott, 1902) comb. nov.; Stenhelia simulans Norman & T. Scott,
1905 = R. simulans (Norman & T. Scott, 1905) comb. nov.; Amphiascus spinulosus Sars, 1911a = R. spinulosa
(Sars, 1911a) comb. nov.; Amphiascus dictydiophorus Monard, 1924 = R. dictydiophora (Monard, 1924)
comb. nov.; Amphiascus rostratus Gurney, 1927 = R. rostrata (Gurney, 1927) comb. nov.; Amphiascus
dactylifer Wilson, 1932 = R. dactylifer (Wilson, 1932) comb. nov.; Amphiascus ilievecensis Monard, 1935a =
R. ilievecensis (Monard, 1935a) comb. nov.; Amphiascus falklandiensis Lang, 1936c = R. falklandiensis
(Lang, 1936c); Amphiascus ecaudatus Monard, 1936 = R. ecaudata (Monard, 1936); Robertgurneya remanei
Klie, 1950 = R. remanei (Klie, 1950) comb. nov.; Robertgurneya oligochaeta Noodt, 1955b = R. oligochaeta
(Noodt, 1955b) comb. nov.; Amphiascoides? arabicus Noodt, 1964 = R. arabica (Noodt, 1964) comb. nov.;
Robertgurneya diversa Lang, 1965 = R. diversa (Lang, 1965) comb. nov.; Robertgurneya hopkinsi Lang,
1965 = R. hopkinsi (Lang, 1965) comb. nov.; Robertgurneya smithi Hamond, 1973b = R. smithi (Hamond,
1973b) comb. nov.; Robertgurneya soyeri Apostolov, 1974 = R. soyeri (Apostolov, 1974) comb. nov.;
Robertgurneya brevipes Wells & Rao, 1987 = R. brevipes (Wells & Rao, 1987) comb. nov.
Rheocamptus Borutzky, 1948 (Family Canthocamptidae)
Dussart and Defaye (1990: 148) recognized four subgenera in the genus Bryocamptus Chappuis, 1929a:
Bryocamptus Chappuis, 1929a (type: Canthocamptus minutus Claus, 1863), Arcticocamptus Chappuis, 1929a
(type: Cyclopsine alpestris Vogt, 1845), Limocamptus Chappuis, 1929a (type: see below for valid type
fixation), and Rheocamptus Borutzky, 1948 (type: see below for valid type fixation). Wells (2007: 92) adopted
this subdivision with the exception of the latter subgenus. The species previously placed in Bryocamptus
(Rheocamptus) were allocated to the nominotypical subgenus Bryocamptus but this course of action was not
accompanied by a supporting argument. Until the validity of Rheocamptus has been properly assessed it is
preferable to maintain it as a valid subgenus.
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Borutzky (1948: 1669) proposed the subgenus Rheocamptus in an identification key to the Bryocamptus
species of Lake Baikal (Russia) and listed two species included in it: B. (R.) baicalensis Borutzky, 1931d and
B. (R.) rylovi Borutzky, 1931d. Neither was formally fixed as the type species. The genus-group name was
made available by Borutzky (1952: 190) who explicitly cited Canthocamptus zschokkei Schmeil, 1893 [=
Bryocamptus zschokkei (Schmeil, 1893)] as the type species (ICZN Art. 13.3) and provided a short diagnosis
by which the subgenus can be differentiated from the other subgenera in Bryocamptus (ICZN Art. 13.1.1).
Although a nominal species is only eligible to be fixed as the type species of a nominal genus or subgenus if it
is an originally included nominal species (ICZN Art. 67.2), this rule is not applicable here because
Rheocamptus Borutzky, 1948 [nomen nudum] is a permanently unavailable name. The authorship and date of
the genus-group name Rheocamptus should rest with Borutzky (1952) who fixed the type by original
designation.
Apodopsyllus Kunz, 1962 (Family Paramesochridae)
Kunz (1962: 254) proposed the generic name Apodopsyllus for Apodopsyllus africanus Kunz, 1962 and four
species previously placed in the genus Leptopsyllus T. Scott, 1894b: L. littoralis Nicholls, 1939b; L. spinipes
Nicholls, 1939b; L. arenicola Chappuis, 1954c; and L. reductus Petkovski, 1955 [subsequently replaced by L.
perplexus Wells, 1963 (nomen novum)]. Kunz neglected to fix a type species and, despite several reviews and
substantial additions subsequent to his work (e.g. Lang 1965; Cottarelli 1973; Coull & Hogue 1978; Gómez
2002a), the generic name Apodopsyllus Kunz, 1962 [nomen nudum] has remained unavailable until now.
Apodopsyllus gen. nov., here explicitly indicated as an intentionally new name (ICZN Art. 16.1), denotes a
taxon that can be differentiated from other genera in the Paramesochridae by the combination of characters
outlined in Kunz’s (1962: 254) generic diagnosis. Apodopsyllus panamensis Mielke, 1984a is here fixed as the
type species by original designation. Since the provisions of both ICZN Arts 13.1.2 and 13.3 are now
satisified the generic name Apodopsyllus is here made available with the present authorship and date. The
genus has seen the addition of 19 species and one subspecies since its initial proposal as a nomen nudum by
Kunz (1962). The respective new combinations are as follows: Leptopsyllus littoralis Nicholls, 1939b =
Apodopsyllus littoralis (Nicholls, 1939b) comb. nov.; Leptopsyllus spinipes Nicholls, 1939b = A. spinipes
(Nicholls, 1939b) comb. nov.; Leptopsyllus madrasensis Krishnaswamy, 1951 = A. madrasensis
(Krishnaswamy, 1951) comb. nov.; Leptopsyllus arenicola Chappuis, 1954c = A. arenicola (Chappuis, 1954c)
comb. nov.; Leptopsyllus adaptatus Krishnaswamy, 1957 = A. adaptatus (Krishnaswamy, 1957) comb. nov.;
Leptopsyllus depressus Krishnaswamy, 1957 = A. depressus (Krishnaswamy, 1957) comb. nov.; Apodopsyllus
africanus Kunz, 1962 = A. africanus (Kunz, 1962) comb. nov.; Apodopsyllus perplexus Wells, 1963 = A.
perplexus (Wells, 1963) comb. nov.; Apodopsyllus schulzi Noodt, 1964 = A. schulzi (Noodt, 1964) comb. nov.;
Apodopsyllus vermiculiformis Lang, 1965 = A. vermiculiformis (Lang, 1965) comb. nov.; Apodopsyllus
lynceorum Cottarelli, 1973 = A. lynceorum (Cottarelli, 1973) comb. nov.; Apodopsyllus camptus Wells, 1971
= A. camptus (Wells, 1971) comb. nov.; Apodopsyllus africanus listensis Mielke, 1975 = A. africanus listensis
(Mielke, 1975) comb. nov.; Apodopsyllus bermudensis Coull & Hogue, 1978 = A. bermudensis (Coull &
Hogue, 1978) comb. nov.; Apodopsyllus unguiformis Coull & Hogue, 1978 = A. unguiformis (Coull & Hogue,
1978) comb. nov.; Apodopsyllus aberrans Mielke, 1984a = A. aberrans (Mielke, 1984a) comb. nov.;
Apodopsyllus arcuatus Mielke, 1984b = A. arcuatus (Mielke, 1984b) comb. nov.; Apodopsyllus panamensis
Mielke, 1984a = A. panamensis (Mielke, 1984a) comb. nov. [type]; Apodopsyllus biarticulatus Cottarelli &
Altamura, 1986 = A. biarticulatus (Cottarelli & Altamura, 1986) comb. nov.; Apodopsyllus chilensis Mielke,
1987 = A. chilensis (Mielke, 1987) comb. nov.; Apodopsyllus cubensis Mielke, 1988 = A. cubensis (Mielke,
1988) comb. nov.; Apodopsyllus melitae Kunz, 1992 = A. melitae (Kunz, 1992) comb. nov.; Apodopsyllus
alejandrovillalobosi Gómez, 2002a = A. alejandrovillalobosi (Gómez, 2002a) comb. nov.; Apodopsyllus
pseudocubensis Gómez, 2002a = A. pseudocubensis (Gómez, 2002a) comb. nov.; and Apodopsyllus
samuelgomezi Gómez, 2002a = A. samuelgomezi (Gómez, 2002a) comb. nov.
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Kliopsyllus Kunz, 1962 (Family Paramesochridae)
As part of his revision of the family Paramesochridae Kunz (1962: 251) proposed the generic name
Kliopsyllus for four species that had previously been included in other paramesochrid genera: Leptopsyllus
coelebs Monard, 1928; Paramesochra holsatica Klie, 1929 (including the subspecies P. holsatica varians
Kunz, 1951); Leptopsyllus constrictus Nicholls, 1935 (including the subspecies Paramesochra constricta
orotavae Noodt, 1958); and Paramesochra major Nicholls, 1939b. Unfortunately Kunz (1962) failed to fix
the type, rendering the genus-group name Kliopsyllus unavailable. The genus has witnessed the addition of 31
species and 5 subspecies (Wells 2007; Kornev & Chertoprud 2008), was extensively reviewed by Kunz
(1981), and was regarded as a senior subjective synonym of Krishnapsyllus Kunz, 1974 by Wells et al. (1975)
(even though – as a nomen nudum – it could not have entered the synonymy of other generic names). The first
authorities to make the generic name Kliopsyllus available are Apostolov and Marinov (1988: 248–249) who
designated Paramesochra holsatica Klie, 1929 as the type species by original designation and provided an
accompanying differential diagnosis (in Bulgarian). Since the provisions of ICZN Arts 13.1.1 and 13.3 are
met, the generic name should be cited as Kliopsyllus Apostolov & Marinov, 1988. However, as an available
name the latter now formally becomes the junior subjective synonym of Krishnapsyllus Kunz, 1974 (type
species by original designation: Krishnapsyllus furcovaricatus Kunz, 1974). None of these names can be
adopted as the valid name for the genus they denote since both are also junior subjective synonyms of
Emertonia Wilson, 1932 (type species by original designation: Emertonia gracilis Wilson, 1932).
Nicholls (1935: 385) listed Emertonia as a synonym of Leptopsyllus T. Scott, 1894b, while Kunz’s (1938:
228) acceptance of the former genus was conditional, depending upon the accuracy of Wilson’s (1932: 256)
description of E. gracilis. In a personal communication to A. G. Nicholls (August 1937) C. B. Wilson wrote
“Let me suggest that there are really three valid genera as follows, Leptopsyllus with the single species
typicus, Paramesochra with the species dubia, intermedia, herdmani, robertsoni, (and) Emertonia with the
species gracilis, minor, constrictus [sic], holsatica, acutata” (Nicholls 1945c: 100). Based on the different
endopodal segmentation of the swimming legs, Nicholls (1945c: 103) accepted the genus as valid and
included two more species, Leptopsyllus coelebs Monard, 1935a and Paramesochra pygmaea Nicholls,
1939b. His statement that “… E. pygmaea may be regarded as the genotype” has no nomenclatural validity
even if Wilson (1932) had failed to fix the type species (ICZN Art. 67.5). Re-examination of Wilson’s type
material in the National Museum of Natural History, Washington, D.C., led Lang (1948: 879) to rank
Emertonia as a genus incertae sedis in the family Paramesochridae. Conversely, Krishnaswamy (1957: 115)
followed Nicholls’s (1945c) judgement and added three new species to the genus: E. capensis Krishnaswamy,
1957, E. minuta Krishnaswamy, 1957 and E. pseudogracilis Krishnaswamy, 1957. Both Lang (1965: 377) and
Kunz (1981: 11–13) rejected this generic assignment and removed all three species to Kliopsyllus.
Examination by the present author of the remaining syntypes of E. gracilis deposited in the Natural History
Museum, London confirmed that this species conforms entirely with Kunz’s (1962: 251; 1981: 11) and
Apostolov and Marinov’s (1988: 249) diagnoses of Kliopsyllus. Consequently, the senior synonym Emertonia
is reinstated here as the valid name for the genus it denotes. Its constituent species and subspecies are as
follows: Paramesochra holsatica Klie, 1929 = Emertonia holsatica (Klie, 1929) comb. nov.; Emertonia
gracilis Wilson, 1932 [type]; Leptopsyllus coelebs Monard, 1935a = E. coelebs (Monard, 1935a) comb. nov.;
Leptopsyllus constrictus Nicholls, 1935 = E. constricta (Nicholls, 1935) comb. nov.; Paramesochra major
Nicholls, 1939b = E. major (Nicholls, 1939b) comb. nov.; Paramesochra pygmaea Nicholls, 1939b = E.
pygmaea (Nicholls, 1939b) comb. nov.; Paramesochra holsatica varians Kunz, 1951 = E. holsatica varians
(Kunz, 1951) comb. nov.; Paramesochra longisetosa Krishnaswamy, 1951 = E. longisetosa (Krishnaswamy,
1951); Emertonia capensis Krishnaswamy, 1957; Emertonia minuta Krishnaswamy, 1957; Emertonia
pseudogracilis Krishnaswamy, 1957; Paramesochra enalius Krishnaswamy, 1957 = E. enalia
(Krishnaswamy, 1957) comb. nov.; Paramesochra constricta orotavae Noodt, 1958 = E. constricta orotavae
(Noodt, 1958) comb. nov.; Paramesochra pontica Şerban, 1959 = E. pontica (Şerban, 1959) comb. nov.;
Paramesochra perharidiensis Wells, 1963 = E. perharidiensis (Wells, 1963) comb. nov.; Paramesochra
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psammobionta Noodt, 1964 = E. psammobionta (Noodt, 1964) comb. nov.; Paramesochra psammophila
Noodt, 1964 = E. psammophila (Noodt, 1964) comb. nov.; Kliopsyllus idiotes Wells, 1967 = E. idiotes (Wells,
1967) comb. nov.; Kliopsyllus holsaticus longicaudatus Galhano, 1970 = E. holsatica longicaudata (Galhano,
1970) comb. nov.; Krishnapsyllus furcavaricatus Kunz, 1974 = E. furcavaricata Kunz, 1974; Kliopsyllus
paraholsaticus Mielke, 1975 = E. paraholsatica (Mielke, 1975) comb. nov.; Kliopsyllus longifurcatus
Scheibel, 1975 = E. longifurcata (Scheibel, 1975) comb. nov.; Kliopsyllus spiniger Wells, Kunz & Rao, 1975
= E. spiniger (Wells, Kunz & Rao, 1975) comb. nov.; Kliopsyllus masryi Bodin, 1979 = E. masryi (Bodin,
1979) comb. nov.; Kliopsyllus californicus Kunz, 1981 = E. californica (Kunz, 1981) comb. nov.; Kliopsyllus
debilis Kunz, 1981 = E. debilis (Kunz, 1981) comb. nov.; Kliopsyllus insularis Kunz, 1981 = E. insularis
(Kunz, 1981) comb. nov.; Kliopsyllus spiniger ornatus Kunz, 1981 = E. spiniger ornata (Kunz, 1981) comb.
nov.; Kliopsyllus atlanticus Kunz, 1983 = E. atlantica (Kunz, 1983) comb. nov.; Kliopsyllus miguelensis
Kunz, 1983 = E. miguelensis (Kunz, 1983) comb. nov.; Kliopsyllus constrictus pacificus Mielke, 1984a = E.
constricta pacifica (Mielke, 1984a) comb. nov.; Kliopsyllus holsaticus listensis Mielke, 1984b = E. holsatica
listensis (Mielke, 1984b) comb. nov.; Kliopsyllus panamensis Mielke, 1984a = E. panamensis (Mielke,
1984a) comb. nov.; Kliopsyllus regulexstans Mielke, 1984b = E. regulexstans (Mielke, 1984b) comb. nov.;
Kliopsyllus similis Mielke, 1984b = E. similis (Mielke, 1984b) comb. nov.; Kliopsyllus unguiseta Mielke,
1984b = E. unguiseta (Mielke, 1984b) comb. nov.; Kliopsyllus acutifurcatus Mielke, 1985 = E. acutifurcata
(Mielke, 1985) comb. nov.; Kliopsyllus chilensis Mielke, 1985 = E. chilensis (Mielke, 1985) comb. nov.;
Kliopsyllus constrictus egypticus Mitwally & Montagna, 2001 = E. constricta egyptica (Mitwally &
Montagna, 2001) comb. nov.; Kliopsyllus andeep Veit-Köhler, 2004 = E. andeep (Veit-Köhler, 2004) comb.
nov.; Kliopsyllus diva Veit-Köhler, 2005 = E. diva (Veit-Köhler, 2005) comb. nov.; Kliopsyllus brevicaudatus
Kornev & Chertoprud, 2008 = E. brevicaudata (Kornev & Chertoprud, 2008) comb. nov.; Kliopsyllus minor
Vasconcelos, Veit-Köhler, Drewes & Parreira dos Santos = E. minor (Vasconcelos, Veit-Köhler, Drewes &
Parreira dos Santos) comb. nov.; Kliopsyllus schminkei Veit-Köhler & Drewes, 2009 = E. schminkei (VeitKöhler & Drewes, 2009) comb. nov.
Scottopsyllus Kunz, 1962 (Family Paramesochridae)
Kunz (1962: 253) proposed the genus Scottopsyllus to accommodate three species, which originally had been
placed in Leptopsyllus T. Scott, 1894b and were subsequently attributed to the robertsoni-group of the genus
Paramesochra T. Scott, 1892 (Lang 1948: 872): Leptopsyllus minor Scott & Scott, 1895a; L. robertsoni Scott
& Scott, 1895a, and L. herdmani Thompson & Scott, 1900. Since there is no trace of a type fixation in the
original publication the genus-group name Scottopsyllus as proposed by Kunz (1962) remains unavailable
(nomen nudum). Lang’s (1944: 26) designation of L. robertsoni as type of the robertsoni-group of
Paramesochra (strictly speaking the equivalent of Scottopsyllus in Kunz’s (1962) sense) has no nomenclatural
significance. The first valid type fixation is that by Apostolov and Marinov (1988: 251) who made the genus
available by providing a generic diagnosis (in Bulgarian) that states in words characters that are purported to
differentiate the taxon (ICZN Art. 13.1.1) and by fixing Leptopsyllus minor Scott & Scott, 1895a as the type
species by original designation (ICZN Art. 13.3). Consequently the genus-group name should be cited as
Scottopsyllus Apostolov & Marinov, 1988.
Kunz (1981) divided Scottopsyllus in three subgenera: Scottopsyllus, Intermedopsyllus Kunz, 1962, and
Wellsopsyllus Kunz, 1981. Apostolov and Marinov’s (1988) type fixation for Scottopsyllus also applies to its
nominotypical subgenus Scottopsyllus Apostolov & Marinov, 1988 (ICZN Art. 67.1.1). Kunz (1981: 7) fixed
Paramesochra gigas Wells, 1965a as the type of the subgenus Wellsopsyllus by original designation.
Intermedopsyllus was originally proposed as a new genus (Kunz 1962: 252) for Leptopsyllus intermedius
Scott & Scott, 1895b and Paramesochra minuta Nicholls, 1939b but since no mention of type fixation was
made in either of Kunz’s (1962, 1981) papers, the genus-group name has remained unavailable. The
subgeneric name Intermediopsyllus subgen. nov. (corrected spelling!) is formally made available herein as an
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intentionally new name (ICZN Art. 16.1) by fixing Leptopsyllus intermedius as the type (ICZN Art. 13.3) and
by making bibliographic reference to Kunz’s (1981: 14) diagnosis which differentiates the taxon from the
other subgenera, Wellsopsyllus Kunz, 1981 and Scottopsyllus Apostolov & Marinov, 1988 (ICZN Art. 13.1.2).
The species described by Kunz (1992: 86) under the name Scottopsyllus (? Intermedopsyllus) smirnovi is
considered species incertae sedis in this subgenus (cf. Wells 2007: 75).
The Principle of Priority requires that if a name in use for a taxon is found to be unavailable it must be
replaced by the next oldest available name from among its synonyms, including the names of the contained
taxa of the same group (ICZN Art. 23.3.5). Since Scottopsyllus Kunz, 1962 was found to be unavailable and
has no potentially valid synonym (i.e. that contains the type species of the former genus), the name of the
genus and nominotypical subgenus becomes Wellsopsyllus Kunz, 1981. The revised species composition of
the genus Wellsopsyllus is summarized in Table 2.
Cladorostrata Shen & Tai, 1963 (Family Miraciidae)
Shen and Tai (1963: 419) established the genus Cladorostrata for two new species, C. brevipoda Shen & Tai,
1963 and C. longipoda Shen & Tai, 1963, but neither was fixed as the type species in the original publication
and hence the name Cladorostrata Shen & Tai, 1963 is to be treated as a nomen nudum. The generic name
remained unavailable until Tai and Song (1979: 185–186) (possibly inadvertently) fixed C. brevipoda Shen &
Tai, 1963 as the type species by original designation in combination with providing a generic diagnosis (in
Chinese), thereby satisfying the provisions of ICZN Arts 13.1.1 and 13.3. This original designation was also
adopted by Shen (1984: 210). The generic name Cladorostrata should therefore be attributed to Tai and Song
(1979) and the names of the original authors should be enclosed in parentheses in the changed combinations
as Cladorostrata brevipoda (Shen & Tai, 1963) and Cladorostrata longipoda (Shen & Tai, 1963).
Scottolana Por, 1967 (Family Canuellidae)
Por (1967: 105) proposed the genus Scottolana to accommodate three species that had originally been placed
in the genus Sunaristes Hesse, 1867: Sunaristes inopinata Thompson & Scott, 1903; S. longipes Thompson &
Scott, 1903; and S. bulbosus Por, 1964a. He also considered Canuella scotti Sewell, 1940 and Sunaristes
curticaudata Thompson & Scott, 1903 as potential candidate members of the genus. Subsequently, Coull
(1972: 209) added Canuella canadensis Willey, 1923 and Wells (1976: 18) assigned Canuella bulbifera
Chislenko, 1971 to Scottolana without making a firm recommendation for this placement. In a later review of
the Canuellidae, Por (1984: 14) restricted the genus – which he puzzlingly cited as a ‘new combination’ – to
just two species, Scottolana longipes (Thompson & Scott, 1903) and S. uxoris Por, 1983a, and maintained
Canuella scotti as a candidate member for inclusion. He provisionally reassigned Scottolana bulbosa (Por,
1964a) to Sunaristes, proposed a new genus Coullana Por, 1984 for S. canadensis (Willey, 1923), and
regarded the status of both S. curticaudata (Thompson & Scott, 1903) and S. inopinata (Thompson & Scott,
1903) as too problematic for further consideration. The genus currently includes 14 species (Mu & Huys
2004; Wells 2007).
Por (1967) did not designate a type species for the genus Scottolana, nor has any subsequent author done
so. Apart from the fact that the lack of a formal type fixation made his genus-group name unavailable, it is
also clear that his diagnosis of the genus did not satisfy the provisions of ICZN Art. 13.1.1. Por’s (1967: 105)
definition of the genus (“… these species in which the first leg-bearing segment can be fused to the
cephalothorax can be best characterized by the reduction of the armature of P IV while the other legs have still
the usual armature of Canuella”) is very uninformative, to the extent that no positive assignment of a species
to this genus can be made, and no characters are given that can unequivocally differentiate the genus from
other taxa in the Canuellidae. Mu and Huys (2004: 2) mentioned that an application (Case 3218) had been
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submitted to the International Commission on Zoological Nomenclature, proposing the conservation of the
generic name Scottolana Por, 1967 and the fixation of Sunaristes bulbosus Por, 1964a as type species.
However, since the latter was selected on the erroneous ground that it had to be one of the originally included
nominal species to be eligible for type fixation (ICZN Art. 69.2 is obviously not applicable to unavailable
names; see also Art. 67.2.1), the application was subsequently withdrawn.
Given the artificial species composition of the genus Scottolana, an objective standard of reference for the
application of the name is highly desirable, yet difficult to define (Mu & Huys 2004). In the interest of
nomenclatural stability the generic name Scottolana (gender: feminine) is re-established here as intentionally
new (ICZN Art. 16.1), taking the authorship and date of the present paper. Scottolana geei Mu & Huys, 2004
[= Scottolana geei (Mu & Huys, 2004) comb. nov.] is formally fixed as the type species (ICZN Art. 13.3),
being representative of a core group of species referred to as the longipes-group (Mu & Huys 2004: 33). This
group includes S. longipes (Thompson & Scott, 1903), S. longipes (Thompson & Scott, 1903) sensu Por
(1964a), S. longipes (Thompson & Scott, 1903) sensu Wells (1967), S. dissimilis Fiers, 1982, S. uxoris Por,
1983a and S. longipes (Thompson Scott, 1903) sensu Wells and Rao (1987), and is characterized by at leat six
apomorphies (Mu & Huys 2004: 33). In order to satisfy the provisions of ICZN Art. 13.1 a generic diagnosis
is given below:
Canuellidae. Leg 1-bearing somite not fused to cephalosome. Rostrum elongate. Urosome without distinct
spinule rows; hyaline frills plain. Genital double-somite of female with subcuticular ribs laterally. Female
genital field large; copulatory pores posteriorly displaced and covered by flaps with free lateral margins;
gonopores covered by opercula with one seta. Anal somite with weakly developed operculum. Caudal rami
usually sexually dimorphic.
Antennule haplocer in male. Antennary exopod 8- or 9-segmented, as long as endopod, setae plumose or
pinnate; endopod distinctly or indistinctly 3-segmented. Mandibular exopod 3-segmented or (indistinctly) 4segmented. Maxillule with 2 setae on coxal epipodite.
Legs 1–4 with 3-segmented rami. Coxae of legs 1–3 (and sometimes leg 4) with inner seta or spine. Leg 2
enp-1 with apophysis fitting into groove with bordering spinule rows on enp-2. Leg 4 endopod longer than
exopod. Sometimes with sexual dimorphism on leg 3 enp-3 or leg 4 exopod. Setal formulae of swimming legs
as follows:
Exopod
Endopod
Leg 1
0.1.313
1.1.222
Leg 2
0.1.322
1.1.221
Leg 3
0.1.122
1.1.121
Leg 4
0.1.121
1.0.121
Leg 5 with 4 setae in both sexes. Male leg 6 large and complex; each produced into large spinous process
bearing short naked seta on ventral surface; additional element arising from inner proximal part of P6, fused at
base, very long. First abdominal somite of male with midventral spinulose ridge.
The genus Scottolana includes the following new combinations: Sunaristes inopinata Thompson & Scott,
1903 = Scottolana inopinata (Thompson & Scott, 1903) comb. nov.; Sunaristes longipes Thompson & Scott,
1903 = Scottolana longipes (Thompson & Scott, 1903) comb. nov.; Canuella scotti Sewell, 1940 = Scottolana
scotti (Sewell, 1940) comb. nov.; Sunaristes bulbosus Por, 1964a = Scottolana bulbosa (Por, 1964a) comb.
nov.; Canuella brevifurca Wells, 1967 = Scottolana brevifurca (Wells, 1967) comb. nov.; Canuella bulbifera
Chislenko, 1971 = Scottolana bulbifera (Chislenko, 1971) comb. nov.; Scottolana dissimilis Fiers, 1982 = S.
dissimilis (Fiers, 1982) comb. nov.; Scottolana glabra Fiers, 1982 = S. glabra (Fiers, 1982) comb. nov.;
Scottolana uxoris Por, 1983a = S. uxoris (Por, 1983a) comb. nov.; Scottolana antillensis Fiers, 1984 = S.
antillensis (Fiers, 1984) comb. nov.; Scottolana oleosa Wells & Rao, 1987 = S. oleosa (Wells & Rao, 1987)
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comb. nov.; Scottolana rostrata Wells & Rao, 1987 = S. rostrata (Wells & Rao, 1987) comb. nov.; Scottolana
tumidiseta Wells & Rao, 1987 = S. tumidiseta (Wells & Rao, 1987) comb. nov.; and Scottolana geei Mu &
Huys, 2004 = S. geei (Mu & Huys, 2004) comb. nov. [type].
Psyllocamptus (Langpsyllocamptus) Kunz, 1975b (Family Ameiridae)
T. Scott (1899b: 254) proposed the genus Psyllocamptus for Psyllocamptus fairliensis T. Scott, 1899b which,
according to Lang (1948: 826), is a junior subjective synonym of Mesochra propinqua T. Scott, 1896 =
Psyllocamptus propinquus (T. Scott, 1896) (type by monotypy). Subsequent to the addition of five more
species, Kunz (1975b: 188) divided the genus in the nominotypical subgenus Psyllocamptus and a new
subgenus Langpsyllocamptus, to which he assigned P. triarticulatus Lang, 1965, P. quinquespinosus Coull,
1970 and two new species, P. longisetosus and P. quadrospinosus. The genus-group name Langpsyllocamptus
Kunz, 1975b cannot be used as an available subgeneric name since no type was designated by Kunz (1975b).
Neither Ceccherelli (1988) who provided a key to species, nor Gómez (2002b) who made a significant
contribution to the systematics of the genus Psyllocamptus, made the subgenus Langpsyllocamptus available.
Instead of re-introducing the latter name by satisfying the provisions of ICZN Arts 13.1, 13.3 and 16.1 it is
opted here for abolishing Kunz’s subgeneric division since it was based on a single character, i.e. the
endopodal segmentation of leg 1. The subgenus Langpsyllocamptus is characterized solely by the
plesiomorphic 3-segmented state of this character and consequently is likely to be a paraphyletic taxon.
Gómez (2002b) recently added credence to this supposition by describing two new species, P. bermudae
Gómez, 2002b and P. totoramensis Gómez, 2002b, which seem to link the two subgenera. Note that Bodin
(1997) erroneously listed P. fuegiensis Pallares, 1982a under the subgenus Langpsyllocamptus; perhaps for
this reason, it was also left out in Gómez’s (2002b: 86) recent key to species of the subgenus Psyllocamptus.
Micropsammis Mielke, 1975 (Family Pseudotachidiidae)
Mielke (1975: 40) established this genus for two new species, Micropsammis noodti Mielke, 1975 and M.
secunda Mielke, 1975 but neglected to fix a type species, rendering the generic name Micropsammis
unavailable. Gee and Huys (1991: 1154) proposed a new genus Telopsammis for M. secunda and inadvertently
made the generic name Micropsammis available by designating M. noodti as the type species (p. 1146) and by
providing an amended generic diagnosis (p. 1145) (ICZN Arts 13.1.1 and 13.3). Huys et al. (1996), Willen
(2000) and Boxshall and Halsey (2004) continued to credit Mielke (1975) with the authorship but based on
the type fixation the genus should be cited as Micropsammis Gee & Huys, 1991 with the new combination,
Micropsammis noodti (Mielke, 1975), as the type and only originally included species. Mielke (1997) argued
against the recognition of the genus Telopsammis but his opinion appears to have gathered no support (Willen
2000; Boxshall & Halsey 2004; Wells 2007; Kornev & Chertoprud 2008). Five out of seven synapomorphies
shared by Telopsammis and Leptotachidia Becker, 1974 [characters (2)–(5) and (7) identified by Gee and
Huys (1991: 1153)] are also displayed by Micropsammis galapagoensis Mielke, 1997, and this suggests it
cannot be maintained in Micropsammis. It is here tentatively removed to Telopsammis as T. galapagoensis
(Mielke, 1997) comb. nov. A third species, Telopsammis pelobionta, was recently described by Kornev and
Chertoprud (2008: 183).
Barbaracletodes Becker, 1979 (Family Cletodidae incertae sedis)
Becker (1979: 10) established Barbaracletodes in the Cletodidae to accommodate two new species, B.
barbara Becker, 1979 and B. carola Becker, 1979, but did not designate a type species. The genus is here
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expressly made available as Barbaracletodes gen. nov. (ICZN Art. 16.1) by fixing B. barbara as the type,
consequently taking the authorship and publication date of the present paper, and by explicit reference to
Becker’s (1979: 10) original generic diagnosis (ICZN Arts 13.1.2 and 13.3) which serves to distinguish it
from other harpacticoid genera. Although B. barbara is named in honour of a woman – Barbara BeckerSchüßler – the name of this species cannot be amended from barbara to barbarae (ICZN Art. 31.1.2—if a
noun in the genitive case) since the Code (Art. 31.1) allows nouns in apposition as well as genitive
constructions for specific names based on personal names. Barbaracletodes barbara and B. carola are clearly
examples of the former. Although Recommendation 31A advises against using personal names in apposition,
once such a name has been proposed it cannot be changed.
Por (1986a) considered Barbaracletodes a member of his Cletodidae sensu stricto. In their key to families
Huys et al. (1996: 92) placed the genus in the Canthocamptidae, an assignment that was also adopted by
Boxshall and Halsey (2004). Gee (1998) excluded Barbaracletodes from the Cletodidae but did not place it
elsewhere. Since none of these authors gave a formal justification for their courses of action, Wells (2007)
considered Barbaracletodes a genus incertae sedis in the Cletodidae.
Ameiropsyllus Bodin, 1979 (Family Ameiridae)
Bodin (1979: 334) established the new genus Ameiropsyllus to accommodate Ameiropsis ariana Monard,
1928 and a new species, Ameiropsyllus monardi Bodin, 1979, but unfortunately neglected to fix a type
species, thus rendering the genus unavailable. Both included species are known from the female only but
since Bodin’s (1979) description is superior over Monard’s (1928), Ameiropsyllus monardi is here fixed as the
type of Ameiropsyllus gen. nov., which is expressly proposed as new under its original name (ICZN Art. 16.1),
taking authorship and date of the present paper. The taxon can be differentiated from other genera in the
Ameiridae by the characters outlined in Bodin’s (1979: 334) diagnosis, hence satisfying the provisions of
ICZN Art. 13.1.2. The two species included are to be cited as new combinations, Ameiropsyllus ariana
(Monard, 1928) comb. nov. and A. monardi (Bodin, 1979) comb. nov. In accordance with Wells (2007) [but
see Lang (1965: 336) for a dissenting view], the variety described under the name Ameira ariana var.
alexandrina Steuer, 1943 should remain subspecies incertae sedis in the genus Ameiropsyllus.
Chilaophonte Mielke, 1985 (Family Laophontidae)
This genus was proposed by Mielke (1985: 240) for two new species, Chilaophonte concepcionensis Mielke,
1985 and C. maiquillahuensis Mielke, 1985. Regrettably, the mandatory requirement of type fixation was not
met by the author, which renders the genus-group name unavailable. It is here made available under its
original name as Chilaophonte gen. nov. (ICZN Art. 16.1) with C. maiquillahuensis Mielke, 1985 as the
newly fixed type species (ICZN Art. 13.3). The taxon can be differentiated from other genera in the family
Laophontidae by explicit reference to Mielke’s (1985: 240) original generic diagnosis (ICZN Art. 13.1.2).
Both included species should be cited as new combinations: Chilaophonte concepcionensis (Mielke, 1985)
comb. nov. and C. maiquillahuensis (Mielke, 1985) comb. nov.
Psammonitocrella Rouch, 1992 (Family Ameiridae)
Rouch (1992: 149) proposed Psammonitocrella for two new species, P. boultoni Rouch, 1992 and P.
longifurcata Rouch, 1992. Although its taxonomic position has been the subject of debate (Martínez Arbizu &
Moura 1994; Lee & Huys 2002; Wells 2007) the genus has until now been left without the mandatory type
fixation and the generic name Psammonitocrella Rouch, 1992 is thus unavailable. In accordance with the
provisions of ICZN for new names published after 1999 (Arts 13.1.2, 13.3 and 16.1) it is here made available
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as Psammonitocrella gen. nov. by fixing Psammonitocrella boultoni Rouch, 1992 as the type species and by
explicit reference to Rouch’s (1992: 149) original designation which serves to differentiate the taxon from
other ameirid genera. Consequently, the generic name takes the authorship and date of the present paper and
both species should be cited as new combinations: Psammonitocrella boultoni (Rouch, 1992) comb. nov. and
P. longifurcata (Rouch, 1992) comb. nov.
Tectacingulum Harris, 1994 (Family Porcellidiidae)
Harris (1994: 304) proposed the genus Tectacingulum to accommodate two new species T. nigrum Harris,
1994 and T. tumidum Harris, 1994. Huys et al. (1996) did not accept any of the new genera established by
Harris (1994) and Harris and Robertson (1994) in the absence of a full phylogenetic analysis of the highly
speciose genus Porcellidium Claus, 1860. Walker-Smith (2001) and Wells (2007) both reinstated
Tectacingulum as a valid genus. Unfortunately, Harris (1994) did not fix a type species, and this oversight
renders the genus-group name unavailable. Tectacingulum gen. nov. is here expressly made available as a new
generic name (ICZN Art. 16.1) by fixing T. tumidum Harris, 1994 as the type species and by explicit reference
to Harris’s (1994: 304) diagnosis which suffices to differentiate the taxon from other genera in the
Porcellidiidae (ICZN Arts 13.1.2 and 13.3). The genus takes the authorship and date of the present paper and
includes the following new combinations: Tectacingulum tumidum (Harris, 1994) comb. nov. and T. nigrum
(Harris, 1994) comb. nov.
It should be noted that both Murramia Harris, 1994 – established for the new species M. bicincta Harris,
1994 and M. magna Harris, 1994 – and Acutiramus Harris & Robertson, 1994 – established for Porcellidium
acuticaudatum Thompson & Scott, 1903, P. brevicaudatum Thompson & Scott, 1903, P. ovatum Haller, 1879
sensu Geddes (1968b) (incertae sedis: cf. Wells 2007: 102), Acutiramus quinquelineatus Harris & Robertson,
1994 and A. rufolineatus Harris & Robertson, 1994 – are also unavailable for the same reason. Since these
genera are now regarded as junior subjective synonyms of Porcellidium (Walker-Smith 2001) they are not
dealt with here any further. Moreover, in his study of the chelicerae of the pterygotid eurypterids, Ruedemann
(1935) established the subgenus Acutiramus (upgraded to generic level by Størmer (1974)). According to
Tollerton (1997) the correct citation of the type species is Pterygotus buffaloensis Pohlman, 1881, by original
designation = Pterygotus cummingsi Grote & Pitt, 1875. The eurypterid generic name Acutiramus
Ruedemann, 1935 would have clear priority over the copepod generic name Acutiramus if a future worker
adopts the latter name and – intentionally or accidentally – makes it available under a different authorship and
date.
Stygonitocrella (Fiersiella) Suárez-Morales & Iliffe, 2005 (Family Ameiridae)
Reid et al. (2003) pointed out that the genus-group name Stygonitocrella Petkovski, 1976 was unavailable
since no type had been designated. In the interest of preserving prevailing usage they re-established the genus
by fixing Nitocrella montana Noodt, 1965 as the type species and by presenting an updated generic diagnosis
(ICZN Arts 13.1.1, 13.3 and 16.1). Suárez-Morales and Iliffe (2005) continued to attribute authorship to
Petkovski (1976), and proposed a subdivision of the genus in two new subgenera, Eustygonitocrella SuárezMorales & Iliffe, 2005 and Fiersiella Suárez-Morales & Iliffe, 2005. As pointed out by Wells (2007: 89),
since the subgenus Eustygonitocrella contains the type species of the genus, Stygonitocrella montana (Noodt,
1965), it must be replaced by the nominotypical subgenus and carry the name, author and date of the genus
(ICZN Art. 44.1), i.e. Stygonitocrella (Stygonitocrella) Reid, Hunt & Stanley, 2003. Moreover, the subgeneric
name Eustygonitocrella does satisfy the provisions of ICZN Art. 13.1.1 (it is accompanied by a differential
diagnosis) but not those of Art. 13.3 (type fixation was not rigidly construed) and hence is permanently
unavailable. Suárez-Morales and Iliffe’s (2005: 217) statement in the etymology section that “… the subgenus
was named using the Greek prefix ‘‘eu’’ meaning ‘‘the original’’, to denote the inclusion, in this group, of the
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type species of the genus Stygonitocrella (S. montana), designated by Reid et al. (2003)” does not qualify as a
type fixation (ICZN Art. 67.5).
The status of the second subgenus, Fiersiella, is also problematic. Suárez-Morales and Iliffe (2005: 224)
designated Nitocrella dubia Chappuis, 1937 as “… the representative species of the subgenus…”. However,
type designation must be rigidly construed (ICZN Art. 67.5) preferably by using the term “type species” or a
strictly equivalent term in another language (espèce-type, Typusart, specie tipo, etc.) (ICZN Recommendation
67A) to avoid ambiguity. According to these provisions Suárez-Morales and Iliffe’s (2005) designation cannot
be considered valid and renders the genus-group name Fiersiella unavailable. It is here re-established as
Fiersiella subgen. nov. by the explicit fixation of Stygonitocrella sequoyahi Reid, Hunt & Stanley, 2003 as the
type species, taking the authorship and date of the present paper. Stygonitocrella sequoyahi is by far the most
accurately described species in the subgenus and a much better candidate for the name-bearing type than
Nitocrella dubia (the oldest species in the subgenus). The subgenus Fiersiella can be differentiated by the
characters listed in the original diagnosis given by Suárez-Morales and Iliffe (2005: 223) (ICZN Art. 13.1.1).
According to these authors the subgenus includes Nitocrella dubia Chappuis, 1937 = Stygonitocrella
(Fiersiella) dubia (Chappuis, 1937), N. karamani Petkovski, 1959 = S. (F.) karamani (Petkovski, 1959), N.
ljovuschkini Borutzky, 1967 = S. (F.) ljovuschkini (Borutzky, 1967), N. colchica Borutzky & MikhailovaNeikova, 1970 = S. (F.) colchica (Borutzky & Mikhailova-Neikova, 1970), Stygonitocrella petkovskii Pesce,
1985a = S. (F.) petkovskii (Pesce, 1985a) and S. (F.) sequoyahi Reid, Hunt & Stanley, 2003. Wells (2007:
37–38) inadvertently assigned these species to the subgenus Stygonitocrella and vice versa.
Pilocamptus Wells, 2007
Chappuis (1929a: 43–45) fixed the types for Bryocamptus Chappuis, 1929a (i.e. Canthocamptus minutus
Claus, 1863) and Echinocamptus Chappuis, 1929a (i.e. Canthocamptus echinatus Mrázek, 1893) (and thus for
their respective nominotypical subgenera, ICZN Art. 67.1.1) but not for the new subgenus Arcticocamptus
Chappuis, 1929a in the former or the new subgenus Limocamptus Chappuis, 1929a in the latter. Borutzky
[1952: 212; 1964 (translation): 193] subsequently fixed Cyclopsine alpestris Vogt, 1845 as the type of
Arcticocamptus.
Chappuis (1929a: 45) included the following species in the subgenus Echinocamptus (Echinocamptus):
Canthocamptus echinatus Mrázek, 1893; C. pilosus Douwe, 1910; C. schroederi Douwe, 1915; C.
georgevitchi Chappuis, 1924b; and C. kamerunensis Kiefer, 1928. In the subgenus Echinocamptus
(Limocamptus) he included Canthocamptus echinatus var. luenensis Schmeil, 1894; C. hiemalis Pearse, 1905;
C. hoferi Douwe, 1908; C. praegeri Scourfield, 1912; C. dacicus Chappuis, 1924a; C. calvus Brehm, 1927;
and C. horai Chappuis, 1928a. Lang (1944: 28) re-assigned the subgenus Limocamptus to Bryocamptus and
explicitly designated Canthocamptus echinatus Mrázek, 1893 (the type species of Echinocamptus) as the type
of Bryocamptus (Limocamptus) while maintaining Echinocamptus as a valid genus (note that this taxon
became hereby an “orphaned” grouping). This type fixation is invalid since C. echinatus was not originally
included in the subgenus and consequently is not eligible to be fixed as the type species (ICZN Art. 67.2). The
next author to fix a type species for the genus-group name Limocamptus is Borutzky [1952: 234; 1964
(translation): 214] who maintained Chappuis’s (1929a) subgeneric division of Echinocamptus and validly
designated Canthocamptus echinatus var. luenensis Schmeil, 1894 as the type of Echinocamptus
(Limocamptus). However, Lang (1948: 1102) claimed that Echinocamptus (L.) luenensis (Schmeil, 1894) was
nothing more than a variety of Echinocamptus (E.) echinatus and that both forms certainly did not belong to
separate subgenera. Except for Borutzky (1952, 1964) this synonymy was widely adopted in the freshwater
copepod literature (e.g. Dussart & Defaye 1990) and effectively rendered both genus-group names
Echinocamptus and Limocamptus objective synonyms of one another (ICZN Art. 61.3.2). Since they were
established simultaneously, but proposed at different ranks, in the genus group, the name proposed at higher
rank (Echinocamptus) takes precedence (ICZN Art. 24.1).
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Since Lang (1944, 1948) had removed Canthocamptus echinatus (the type of Echinocamptus) and all
species placed in Echinocamptus (Limocamptus) to a taxon of subgeneric rank in Bryocamptus, the valid
name of this subgenus is Bryocamptus (Echinocamptus) and not Bryocamptus (Limocamptus) as generally
accepted (e.g. Dussart & Defaye 1990; Wells 2007). The subgenus includes the following species:
Bryocamptus (Echinocamptus) echinatus (Mrázek, 1893); B. (E.) hiemalis (Pearse, 1905) [and its subspecies
hiemalis hiemalis (Pearse, 1905); h. verestschagini (Borutzky, 1931d); h. brevifurcatus Coker, 1934; h.
yunnanensis (Borutzky, 1952); h. elongatus (Shen & Tai, 1964a); and h. yetii Löffler, 1968]; B. (E.) hoferi
(Douwe, 1908); B. (E.) praegeri (Scourfield, 1912); B. (E.) dacicus (Chappuis, 1924a); B. (E.) douwei
(Willey, 1925a); B. (E.) nivalis (Willey, 1925b); B. (E.) calvus (Brehm, 1927); B. (E.) horai (Chappuis,
1928a); B. (E.) morrisoni (Chappuis, 1929c) [and its subspecies morrisoni morrisoni (Chappuis, 1929c) and
m. elegans (Chappuis, 1929c)]; B. (E.) smirnovi (Borutzky, 1931d); B. (E.) parvus (Borutzky, 1931d); B. (E.)
viduus Kiefer, 1952; B. (E.) hostensis (Borutzky, 1972); B. (E.) pacificus Ishida, 1992; and B. (E.) lacustris
Wells, 2007.
As pointed out only recently by Wells (2007: 93), Lang’s (1944, 1948) removal of the type of
Echinocamptus to the genus Bryocamptus left the remaining Echinocamptus species assembled in an
orphaned taxon which requires a new generic name. Wells (2007: 93) formally proposed the name
“Pilocamptus n. gen.” and fixed Canthocamptus pilosus Douwe, 1910 as the type species. However,
according to ICZN Art. 13.1 this name is unavailable since its proposal was not accompanied by a description
or definition that states in words characters that are purported to differentiate the taxon, or a bibliographic
reference to such a published statement. The generic name Pilocamptus gen. nov. is here re-instated as
intentionally new, taking the authorship and date of the present paper. The taxon it denotes can be
differentiated from other genera in the family Canthocamptidae by the diagnosis given by Lang (1948: 1113)
for the genus Echinocamptus. The genus includes the following species: Canthocamptus pilosus Douwe, 1910
= Pilocamptus pilosus (Douwe, 1910) [type species by original designation]; C. schröderi Douwe, 1915 = P.
schroederi (Douwe, 1915) comb. nov.; C. Georgevitchi Chappuis, 1924b = P. georgevitchi (Chappuis, 1924b)
comb. nov.; C. kamerunensis kamerunensis Kiefer, 1928 = P. kamerunensis kamerunensis (Kiefer, 1928)
comb. nov.; Echinocamptus (Echinocamptus) kamerunensis villosus Chappuis, 1932 = P. kamerunensis
villosus (Chappuis, 1933a) comb. nov.; E. (E.) africanus Chappuis, 1933a = P. africanus (Chappuis, 1933a)
comb. nov.; E. (E.) Alluaudi Chappuis, 1933a = P. alluaudi (Chappuis, 1933a) comb. nov.; E. (E.) trichotus
Chappuis, 1933a = P. trichotus (Chappuis, 1933a) comb. nov.; Echinocamptus Jeanneli Chappuis, 1936 = P.
jeanneli (Chappuis, 1936) comb. nov.; E. monticola Chappuis, 1936 = P. monticola (Chappuis, 1936) comb.
nov.; E. verrucosus Chappuis, 1936 = P. verrucosus (Chappuis, 1936) comb. nov., E. (s. str.) vulgaris
Chappuis, 1936 = P. vulgaris (Chappuis, 1936) comb. nov.; E. Pauliani Chappuis, 1956 = P. pauliani
(Chappuis, 1956) comb. nov.; E. monodi Dussart, 1974 = P. monodi (Dussart, 1974) comb. nov.; and E.
hypophyllus Defaye & Heymer, 1996 = P. hypophyllus (Defaye & Heymer, 1996) comb. nov.
Junior subjective synonyms
Idomene Philippi, 1843 (Family Pseudotachidiidae)
Philippi (1843: 64) proposed this genus for a new species Idomene forficata based on a single individual. The
original description is fragmentary and insufficient for unequivocal identification, hence most authors have
invariably referred to Sars’s (1906a: 134) detailed redescription and used his diagnosis of the genus as the
major standard of reference for adding subsequent species, either newly described or reassigned from other
genera regarded as junior synonyms of Idomene. This shift in attention had the unfortunate consequence that
noone either scrutinized the real identity of Philippi’s (1843) type species or confirmed its conspecificity with
what Sars (1906a) had identified as I. forficata. In spite of their poor quality, Philippi’s (1843) illustrations
provide sufficiently compelling evidence that I. forficata is not a harpacticoid but rather a poecilostome
cyclopoid. The 7-segmented antennule with four setae on the basal segment, the 3-segmented antennary
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endopod, the mandibular gnathobase with distinct blades (his Fig. 4f, labelled “ein Kaufuss?”), and the
laterally located, uniramous fifth legs (with 2 outer margin spines, 1 apical seta and 1 inner spine) not only
unequivocally exclude I. forficata from the oligoarthran Harpacticoida but also identify it as a member of the
family Clausidiidae in the Cyclopoida. The armature pattern and shape of the maxilliped is another line of
evidence supporting such assignment. It is conceivable that Idomene Philippi, 1843 is the senior synonym of
either Hemicyclops Boeck, 1873 or Hersiliodes Canu, 1888. It is here considered a genus incertae sedis in the
Clausidiidae. Since Philippi’s type material no longer exists a neotype can unfortunaly not be designated.
Three available harpacticoid genus-group names have entered into the synonymy of Idomene (Lang 1934:
28; 1936c: 14): Xouthous Thomson, 1883 (type: Xouthous novaezealandiae Thomson, 1883), Megarthrum
Norman & Scott, 1906 (type: Dactylopusia purpurocincta Norman & Scott, 1905) and Idomenella T. Scott,
1906a (type: Dactylopus coronatus T. Scott, 1894b). Xouthous, being the oldest available name, is here
proposed as the valid name for Idomene Philippi, 1843 sensu Sars (1906a).
In addition to the type, the following species should be referred to the genus Xouthous: Dactylopus
coronatus T. Scott, 1894b = Xouthous coronatus (T. Scott, 1894b) comb. nov.; Dactylopus pectinatus Scott &
Scott, 1898 = Xouthous pectinatus (Scott & Scott, 1898) comb. nov.; Dactylopus antarcticus Giesbrecht, 1902
= Xouthous antarcticus (Giesbrecht, 1902) comb. nov.; Dactylophusia laticaudata Thompson & Scott, 1903 =
Xouthous laticaudatus (Thompson & Scott, 1903) comb. nov.; Dactylopusia purpurocincta Norman & Scott,
1905 = Xouthous purpurocinctus (Norman & Scott, 1905) comb. nov.; Idomene pusilla Brady, 1910 =
Xouthous pusillus (Brady, 1910) comb. nov.; Megarthrum simulans Brady, 1910 = Xouthous simulans (Brady,
1910) comb. nov.; Idomene borealis Sars, 1911a = Xouthous borealis (Sars, 1911a) comb. nov.; Dactylopusia
ferrieri T. Scott, 1912 = Xouthous ferrieri (T. Scott, 1912) comb. nov.; Idomene intermedia Lang, 1934 =
Xouthous intermedius (Lang, 1934) comb. nov.; Xouthous maldiviæ Sewell, 1940 = Xouthous maldiviae
Sewell, 1940; Idomene scotti Lang, 1948 = Xouthous scotti (Lang, 1948) comb. nov.; Idomene parasimulans
Médioni & Soyer, 1968 = Xouthous parasimulans (Médioni & Soyer, 1968) comb. nov.; and Idomene
cookensi Pallares, 1975b = Xouthous cookensis (Pallares, 1975b) comb. nov. [incorrect original spelling
corrected by Wells (2007: 85)]. Since Philippi’s (1843) text and figures do not bear any relationship to
Idomene forficata sensu Sars (1906a) a new name is required for the latter for which I propose, in the absence
of any available junior synonyms, Xouthous sarsi sp. nov. The new species can be differentiated from its
congeners by the specific characters given in Sars’s (1906a: 134–135, Plate LXXXII) description (ICZN Art.
13.1.2).
Following Wells (2007) the genus should also include Idomene australis Brady, 1910 and Idomene
antarctica (Giesbrecht, 1902) sensu Dahms and Schminke (1992) (both being species inquirendae) as well as
Dactylophusia aemula Thompson & Scott, 1903 and Idomene kabylica Monard, 1936 (both being species
incertae sedis).
Alteutha Baird, 1846b (Family Peltidiidae)
Goodsir (1845) established the genus Sterope Goodsir, 1845 for three new species, S. ovalis, S. armatus and S.
interruptus, and created another genus, Carrillus Goodsir, 1845, for the type and only species C. oblongus
Goodsir, 1845. Boeck (1865: 272) synonymized Carillus [subsequent emendation of Carrillus Goodsir, 1845
by Vosmaer (1885: 218)] with Alteutha Baird, 1846b (type species: Cyclops depressus Baird, 1837). Similarly,
Claus (1889: 6) regarded Sterope also as a synonym of Alteutha. Both A. oblonga (Goodsir, 1845) and A.
interrupta (Goodsir, 1845) are now considered valid species while S. ovalis and S. armatus are treated as
species incertae sedis in Alteutha (Wells 2007: 77). However, both Sterope and Carillus are senior subjective
synonyms of Alteutha. Although the publication date is usually quoted as 1845 (Lang 1948; Wells 2007), in
reality Baird’s (1846b: 155) diagnosis was not published until June 1846 while Goodsir’s genera were
published in November 1845 and hence, in accordance with the Principle of Priority, take precedence over
Alteutha. In the interest of nomenclatural stability, prevailing usage can be maintained since both senior
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synonyms have not been used as valid names since 1899 (ICZN Art. 23.9.1.1) and Alteutha has been used as a
valid name in at least 25 works, published by at least 10 authors in the last 50 years (ICZN Art. 23.9.1.2) [e.g.
Por 1960a; Wiborg 1964; Lang 1965; Pallares 1968a; Fives 1969; Apostolov 1973; Gharagozlou-Van
Ginneken 1976; Wells 1976, 2007; Hicks 1977, 1982, 1986a; Coffin 1981; McClelland 1982; Apostolov &
Marinov 1988; Dahms 1992, 1993a; Song & Chang 1995; Huys et al. 1996; Bodin 1997; Mitwally &
Montagna 2001; Apostolov & Pandourski 2002; Duggan et al. 2006; Kane 2007; Michels 2007; Veit-Köhler
& Fuentes 2007a, 2007b]. In accordance with ICZN Art. 23.9.2 the junior synonym Alteutha Baird, 1846b is
considered the valid name (nomen protectum) which has precedence over the older names Sterope Goodsir,
1845 and Carillus Goodsir, 1845 (nomina oblita). Since this is a case of subjective synonymy, whenever
Sterope and/or Carillus are not regarded as synonyms of Alteutha, the older synonyms may be used as valid
names (ICZN Art. 23.9.2).
It should also be noted that Dana (1854: 1179) proposed the subfamily Steropinae for the genera Zaus
Goodsir, 1845 and Sterope (type genus) (he is less explicit about the inclusion of Porcellidium Claus, 1860).
This family-group name is a senior subjective synonym of the name Peltidiidae (type genus Peltidium
Philippi, 1839) which is generally attributed to Sars (1904b) (e.g. Lang 1948; Seifried 2003; Boxshall &
Halsey 2004). Wells (2007: 101) recently pointed out that the authorship and date of the name lies with Claus
(1860: 5) who first published it in vernacular format as “Familie der Peltidien” and subsequently latinized it –
as Peltididae or Peltidinae – in his later work (Claus 1862: 89; 1863: 92, 137; 1891: 429). However, in none of
these contributions did Claus make the name Peltidiidae available since a family-group name when first
published must be formed from the stem of an available generic name then used as valid in the new familygroup taxon; the use of the stem alone in forming the name is only accepted as evidence that the author used
the generic name as valid in the new family-group taxon if there is no evidence to the contrary (ICZN Art.
11.7.1.1). Claus (1860, 1862, 1863, 1891) did not include Peltidium as a valid genus in his family-group name
but regarded it a possible synonym of Oniscidium Claus, 1860 (one of the five genera included, the others
being Alteutha, Porcellidium, Zaus and Eupelte Claus, 1860). The first author to make the name Peltidiidae
available is Boeck (1873: 58) who proposed the subfamily Peltidinae in the family Harpactidae,
accommodating the genera Peltidium (including its junior synonym Oniscidium), Scutellidium Claus, 1866
and Aspediscus [sic] Norman, 1869.
Since the name Steropinae has not been used as a valid name after 1899 and the conditions of ICZN Art.
23.9.1.2 are met (Peltidiidae is cited in e.g. Lang 1965; Geddes 1968b; Wells 1970, 1976, 2007; Hicks 1977,
1982, 1986a; Humes 1981b, 1984; Apostolov & Marinov 1988; Dahms 1992, 1993a; Huys et al. 1996; Bodin
1997; Mitwally & Montagna 2001; Seifried 2003, 2004; Boxshall & Halsey 2004; Varela 2005; SuárezMorales et al. 2006; Veit-Köhler & Fuentes 2007a–b; Gheerardyn et al. 2008; Ivanenko et al. 2008), the
junior synonym Peltidiidae is considered the valid name (nomen protectum) which has precedence over the
older name Steropinae (nomen oblitum).
Family Tisbidae Stebbing, 1910
Philippi (1843: 58) proposed the genus Idya Philippi, 1843 (type species by monotypy: Idya barbigera
Philippi, 1843) while Lilljeborg (1853: 191) created the generic name Tisbe for Cyclops furcatus Baird, 1837
(type species by monotypy). Boeck (1865: 257) considered both names synonyms of each other. Sars (1905b:
89) suspected that I. barbigera was conspecific with C. furcatus, a notion that was confirmed by Lang (1948:
369) who considered the former a junior subjective synonym of the latter, effectively rendering Idya and Tisbe
objective synonyms (ICZN Art. 61.3.3). Sars (1909c: 21), suggested a slight alteration of the name Idya to
Idyæa (an incorrect original spelling of Idyaea) to avoid the homonymy with Idya Fréminville, 1809
(Ctenophora), however, Norman and Scott (1906: 183) had already reinstated the junior synonym Tisbe.
The family-group name Idyinae, proposed by Brady (1880a: 170) and upgraded to family level by Sars
(1904b: 78), is invalid because it was based on the generic name Idya Philippi, 1843, which is a junior
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homonym of the ctenophoran genus Idya Fréminville, 1809 (ICZN Art. 39). Stebbing (1910: 544), referring to
the family name Idyidae, stated “The necessity for a change in the family name will be seen from the
references for the leading genus” and proposed the family-group name Tisbidae (type genus Tisbe Lilljeborg,
1853) as an alternative. Although Stebbing (1910) did not give a description or definition, the name
“Tisbidae” satisfies all criteria required for availability (ICZN Art. 12.2.4) since it was published before 1931
and based on the then (and still) valid generic name Tisbe.
Claus (1862: 89; 1863: 137) proposed the family name “Peltididae” to accommodate the genera Alteutha,
Eupelte, Oniscidium, Porcellidium and Zaus and subsequently (Claus 1866: 20) added the new genus
Scutellidium Claus, 1866 (type by monotypy: S. tisboides Claus, 1866) to it. Although Claus did not make the
name “Peltididae” available in any of these publications (see above under Alteutha) his subsequent
subdivision of the family deserves attention. Claus (1889: 3) divided the “Peltididae” into two subordinate
taxa of subfamilial rank: in the Peltidinae he included Alteutha, Eupelte and Oniscidium, while Porcellidium,
Scutellidium and Zaus were placed in the Scutellidinae (incorrect original spelling of Scutellidiinae). The
latter subfamily which takes the authorship and date of Claus (1889) is an available name since it is based on
the then (and still) valid generic name Scutellidium (ICZN Art. 12.2.4). In addition, Scutellidiinae Claus, 1889
is a senior subjective synonym of Tisbidae Stebbing, 1910. However, the senior synonym has not been used as
a valid name after 1899 and the junior synonym has been widely used in the last 50 years by a large number of
authors (e.g. Avdeev 1983, 1986; Becker & Schriever 1979; Bodin 1976, 1997; Boxshall 1979; Bresciani
1970; Bresciani & Lützen 1994; Coull 1973b; Dahms 1993b; Dahms & Dieckmann 1987; Gee & Fleeger
1986; Gómez et al. 2004; Hicks 1971; Humes & Ho 1969b; Humes & Voight 1997; Huys 1996b; Lang 1965;
Lee & Huys 1999c; López-González et al. 2000; Puello Cruz et al. 2004; Vervoort 1964; Volkmann-Rocco
1971; Wells 1976, 2007). Since the provisions of ICZN Art. 23.9 are met, the name Tisbidae Stebbing, 1910
must be considered the valid name (nomen protectum) which has precedence over the older name
Scutellidiinae Claus, 1889 (nomen oblitum). Being a case of subjective synonymy, whenever the latter is not
regarded as a synonym of the former, Scutellidiinae may be used as valid (ICZN Art. 23.9.2).
Based solely on the respective publication dates, Sars’s (1910) family-group name “Idyaeidae” (taking the
stem of Idyaea Sars, 1909) mentioned in the heading of Supplement Plate 10 (Idyaea tenella) (Natural History
Museum copy date stamped 20 July 1910) is potentially a senior objective synonym of the family name
Tisbidae Stebbing, 1910 (published on 15 December 1910). Since the provisions of ICZN Art. 23.9 are met
the same reversal of precedence applies as for the name Scutellidiinae (see above); the name Tisbidae
Stebbing, 1910 must be considered the valid name (nomen protectum) which has precedence over the older
name Idyaeidae Sars, 1910 (nomen oblitum).
Subfamily Cerviniopsinae Brotskaya, 1963
Brotskaya (1963: 793) proposed the new subfamily Cerviniopsinae in the Cerviniidae (now Aegisthidae),
accommodating the genera Herdmania Thompson, 1893 [a senior objective synonym of Hemicervinia Lang,
1935 according to Por (1964a)]; Pontostratiotes Brady, 1883; Cerviniopsis Sars, 1903 (type); Pseudocervinia
Brotskaya, 1963 [a junior subjective synonym of Cervinia Norman in Brady, 1878 according to Seifried
(2003)]; and Herdmaniopsis Brotskaya, 1963. It should be noted that the stem of –opsis in Cerviniopsis is
–opse-, thus the correct name should have been Cerviniopseinae and Cerviniopsinae is to be considered an
incorrect original spelling. According to Seifried and Schminke (2003: 32) members of the Aegisthinae are
probably nested within the Cerviniopseinae, rendering the latter subfamily paraphyletic. However, pending a
phylogenetic analysis at species level they maintained the subfamilial subdivision of the Aegisthidae. Wells
(2007: 17) likewise continued to recognize the Cerviniopseinae as a valid subfamily, accommodating
Cerviniopsis, Pontostratiotes, Hemicervinia, Herdmaniopsis, Ameliotes Por, 1969, Stratiopontotes Por, 1969
and Tonpostratiotes Itô, 1982.
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A. Scott (1909: 232) proposed the family-group name Pontostratiotidae and stated that “… the type of this
family is Pontostratiotes abyssicola Brady, 1883”. This species was fixed as the type of Pontostratiotes by
monotypy (Brady 1883: 105). Since objectivity provided by typification is continuous through the hierarchy
of names, extending in ascending order from the species group to the family group (ICZN Art. 61.1.2), the
family-group name Pontostrationinae A. Scott, 1909 (type genus Pontostratiotes) takes priority over
Cerviniopseinae Brotskaya, 1963.
Namakosiramia Ho & Perkins, 1977 (Family Laophontidae)
Ho and Perkins (1977: 368) proposed the genus Namakosiramia Ho & Perkins, 1977 for a new species N.
californiensis Ho & Perkins, 1977 (type by original designation) and established the family
Namakosiramiidae in the order Siphonostomatoida to accommodate it. Huys (1988a: 1520, 1522) placed the
family in the order Harpacticoida and relegated it to a junior subjective synonym of the family Laophontidae
T. Scott, 1905a. Kim (1991: 429) added a second species, N. koreensis Kim, 1991, to the genus.
Brady (1918: 34), in a brief supplementary note to his report on the Copepoda collected during the
Australasian Antarctic Expedition 1911–1914, proposed the genus Microchelonia for a single species, M.
glacialis Brady, 1918 (type by monotypy). The genus was not mentioned in Lang’s (1948) monograph and
does not feature in the Nomenclator Zoologicus (Neave 2005). It was listed as a genus inquirendum by
Boxshall and Halsey (2004: 844) but without ordinal or familial assignment. Although Brady (1918: 34)
stated that M. glacialis may “…form the type of an entirely new division of the Copepoda…”, the illustration
of the first leg (labelled as the “anterior antenna”; Plate 15, Fig. 4) indicates that Microchelonia is a member
of the harpacticoid family Laophontidae. The general dorso-ventrally depressed habitus, the shape of the P1
endopod (and claw), the powerful P2 (shown in the habitus drawings; Plate 15, Figs 1–2), the reduced P2–P3
(labelled as “posterior antenna”; Plate 15, Fig. 5) and P5 (labelled as “maxilliped”; Plate 15, Fig. 6), and the
short caudal rami bearing one long principal seta unequivocally identify M. glacialis as a member of the genus
Namakosiramia which should consequently sink as a junior objective synonym of Microchelonia Brady,
1918. The genus contains three species: M. glacialis, M. californiensis (Ho & Perkins, 1977) comb. nov. and
M. koreensis (Kim, 1991) comb. nov.
Family Huntemanniidae Por, 1986a
Por’s (1986a) review of the Cletodidae, including the proposal of four new families and the reassignment of
some species to the Canthocamptidae, has not gained universal acceptance and needs further testing (Wells
2007). In the meantime, his family-group names cannot be ignored as they were accompanied by diagnoses
and are to be used as valid names according to ICZN Art. 13.2 until future revisionary work proves otherwise.
As part of his review, Por (1986a: 421) established the family Huntemanniidae for the genera Nannopus
Brady, 1880a, Huntemannia Poppe, 1884 [type], Pontopolites T. Scott, 1894b, Metahuntemannia Smirnov,
1946, Beckeria Por, 1986b and possibly Pseudocletodes Scott & Scott, 1893b. Except for Lang (1948: 1248),
it has escaped most authors’ attention that Brady (1880a: 100) had already established a new subfamily
Nannopinae (incorrect original spelling Nannopinæ) within the Harpacticidae for Nannopus Brady, 1880a
(type genus) and Platychelipus Brady, 1880a (now placed in the Laophontidae). According to the Principle of
Coordination applied to family-group names (ICZN Art. 36.1) Brady (1880a) is deemed also to have
simultaneously established the coordinate family name Nannopidae (here corrected to Nannopodidae since
the second part of the generic name Nannopus is based on the Greek stem πους, meaning foot) which takes the
same authorship and date. Since the family-group name Huntemanniidae was used by Por (1986a) to include
the generic name Nannopus, it must sink as a junior synonym of Nannopodinae Brady, 1880a. The Principle
of Priority requires that the latter be reinstated as the valid name with change of rank to family (ICZN Art.
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23.1). The family name Nannopodidae is equivalent to Por’s (1986a) Huntemanniidae (evidently only if
Nannopus and Huntemannia are considered to be confamilial) with the exception that (1) Dahms and Pottek
(1992) have relegated Beckeria to a junior subjective syonym of Metahuntemannia; (2) three genera have
been added since: Rosacletodes Wells, 1985 [= Echinocletodes Pallares, 1982a nec Lang (1936b); cf. Huys et
al. (1996: 75)], Laophontisochra George, 2002 and Acuticoxa (cf. Huys & Kihara 2009), and (3) both
Metahuntemannia and Dahmsopottekina Özdikmen, 2009 [= Talpina Dahms & Pottek, 1992 nec Hagenow
(1840); see above] were assigned to the subfamily Hemimesochrinae in the Canthocamptidae (Huys & Kihara
2009).
Orphaned taxonomic groupings
The application of each genus-group name is determined by reference to the type species of the nominal taxon
that it denotes (ICZN Art. 42.3). Removal of the type species – but not of the remaining species included in a
genus – creates orphaned taxonomic groupings which require an existing (previously invalid) or new
substitute generic name. In addition to the Echinocamptus case explained above (see under Pilocamptus)
another four cases of orphaned taxa have remained unnoticed in harpacticoid systematics.
Amphiascus Sars, 1905a (Family Miraciidae)
Sars (1905a: 380) proposed the genus Amphiascus Sars, 1905a to include Dactylopus longirostris Claus,
1863, D. minutus Claus, 1863, D. debilis Giesbrecht, 1881 and a new species Amphiascus pacificus Sars,
1905a. Nicholls (1941b: 69) transferred D. minutus to Amphiascopsis Gurney, 1927 and D. debilis to
Amphiascoides Nicholls, 1941b, and he designated D. longirostris as the type species of Amphiascus (on the
ground of page precedence in Sars’s (1905a) publication). Lang (1944: 19), who was unaware of Nicholls’s
(1941b) subsequent designation, unfortunately also fixed D. longirostris as type of Paramphiascopsis Lang,
1944. The latter therefore becomes a junior objective synonym of Amphiascus Sars, 1905 (ICZN Arts 61.3.3
and 67.11). Other workers have overlooked Nicholls’s type fixation and incorrectly treated D. minutus as the
type species (e.g. Marinov & Apostolov 1988: 153). Accepting Nicholls’s (1941b) subsequent designation
and Lang’s (1944, 1948) revision of Amphiascus would imply that only the species currently assigned to
Paramphiascopsis should be included in Amphiascus. A new generic name is therefore required to receive all
“orphaned” species that are currently included in Amphiascus sensu Lang (1944, 1948).
The only genus-group name that has been cited in the literature as a junior subjective synonym of, and
potential substitute name for, Amphiascus sensu Lang (1944, 1948) is Mesamphiascus Nicholls, 1941b (Lang
1965: 253). This genus is probably one of the most unnatural ever established in the Miraciidae, to the extent
that Nicholls (1941b: 79) himself admitted having difficulties in selecting a type species. His statement that
“… perhaps parvus Sars (1906, p. 162, pl. ciii) is suitable, occupying a more or less central position in the
genus, and having a fairly wide distribution” does not qualify for an explicit designation that is rigidly
construed (ICZN Art. 67.5; see also Vervoort 1964: 191). Consequently, the genus-group name
Mesamphiascus is unavailable and therefore cannot enter into the synonymy of Amphiascus. A new genus,
Sarsamphiascus gen. nov., is proposed here to accommodate all species previously placed in Amphiascus by
Lang (1948) and subsequent authors. Dactylopus minutus Claus, 1863 is formally fixed as the type species.
The revised species compositions of Amphiascus and Sarsamphiascus gen. nov., including all new
combinations, are given in Table 3. In order to satisfy the provisions of ICZN Art. 13.1 the new genus must be
accompanied by (a) a description or definition that states in words characters that are purported to differentiate
it, or (b) a bibliographic reference to such a published statement. Reference is made here to Lang’s (1948:
644) generic diagnosis of Amphiascus which is equivalent to the diagnosis of Sarsamphiascus gen. nov.
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Teissierella Monard, 1935a (Family Miraciidae)
The genus Teissierella was proposed by Monard (1935a: 24) to accommodate a new species T. celtica
Monard, 1935a, which he regarded as “transitionary” between Robertsonia Brady, 1880a and Amphiascus
Sars, 1905a. He also believed Stenhelia knoxi Thompson & Scott, 1903 and, with reservations, Amphiascus
bulbifer Sars, 1911a (now in Haloschizopera Lang, 1944), Stenhelia irrasa A. Scott, 1902 and Dactylopus
propinquus T. Scott, 1894a (both now in Robertsonia) could be referred to this genus. Monard’s (1935a)
expression “n. g. n. sp.” applied to Teissierella celtica effectively designates this species as the name-bearing
type of the genus (ICZN Art. 68.2.1). In the same year Monard (1935b: 26) added the new species, T.
salammboi and described a second taxon under the name “T. irrasa (A. Scott) var.”. In a footnote (p. 27) he
stated that Teissierella (with two egg-sacs) could turn out to be a synonym of Robertsonia if Sars’s (1909b:
335) assertion of the presence of only one egg-sac in Robertsonia tenuis Brady, 1880a were proved wrong.
Sewell (1940: 286) discussed the relationships between both genera and described a new species Teissierella
adduensis Sewell, 1940.
Lang (1944: 14) listed Teissierella as a junior synonym of Robertsonia and in his monograph (Lang 1948:
638) considered both T. celtica and T. salammboi as species incertae sedis in Robertsonia, the former as being
possibly conspecific with Robertsonia knoxi (Thompson & Scott, 1903), the latter as a potential member of
the genus Diosaccopsis Brian, 1925a. Monard’s (1935b) Teissierella irrasa var. was regarded as conspecific
with Robertsonia angolensis (Monard, 1934). Krishnaswamy (1957: 51) assigned T. adduensis to
Robertsonia, albeit without any factual justification, but this course of action has now been validated by Wells
and Rao (1987: 99).
Following Roe’s (1958: 248) redescription, the type species Teissierella celtica is now formally
considered a valid species of Robertsonia (Hamond 1973c: 433; Wells 2007: 25). Despite Teissierella thus
being a junior subjective synonym of the latter, two new species have subsequently been added: Bodin (1964:
157) described T. massiliensis Bodin, 1964 while Apostolov (1968: 400) added T. pontica Apostolov, 1968.
The latter was subsequently considered a junior subjective synonym of Robertsonia knoxi (Thompson &
Scott, 1903) by Apostolov and Marinov (1988: 151).
The latest state of affairs was summarized by Wells (2007: 25) who listed Teissierella as a genus incertae
sedis in the subfamily Diosaccinae (Miraciidae), containing T. salammboi, T. massiliensis and T. pontica.
Since the type species is not included (see above) a new generic name is required for this orphaned grouping.
Teissierella salammboi Monard, 1935b is here fixed as the type of the new genus Monardius gen. nov. (in
memory of Dr André Monard; gender: masculine) which contains the following new combinations:
Monardius salammboi (Monard, 1935b) comb. nov. [type] and M. massiliensis (Bodin, 1964) comb. nov.
Apostolov’s (1968: 400–401) description of T. pontica shows that this species cannot be placed in Monardius
(e.g. antennary exopod 2-segmented; leg 1 exp-3 with 5 elements and endopod 3-segmented; endopodal
armature of legs 2–4; form of setae on leg 5 etc.); in addition, Wells (2007: 550) pointed out that Apostolov
(1968: Tafel 3) showed two inner setae on the middle exopodal segment of legs 2–4 which is, if proven
correct, a character not found in any other copepod (Huys and Boxshall 1991: 23, 28). Apostolov and
Marinov’s (1988: 151) generic assignment is adopted here with the caveat that T. pontica is not necessarily a
synonym of R. knoxi; pending re-examination of the type material it is best regarded a species incertae sedis
in Robertsonia. In order to satisfy the provisions of ICZN Art. 13.1 a generic diagnosis is given below
summarizing the characters that are purported to differentiate Monardius from other genera in the Miraciidae.
Miraciidae. Rostrum elongate, extending beyond second antennulary segment. Antennule 8-segmented in
female, with numerous pinnate setae. Antenna with abexopodal seta on allobasis; exopod 3-segmented with
setal formula [1-1-3]. Mandibular palp with 1-segmented rami and 3 setae on basis. Caudal rami squarish or
wider than long, with well developed setae IV and V.
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Leg 1 with prehensile 2-segmented endopod; enp-1 longer than enp-2, about as long as exopod. Leg 2
endopod modified in ♂; 2-segmented. Setal formulae of swimming legs as follows:
Exopod
Endopod
Leg 1
0.1.022
1.1.120
Leg 2
1.1.223
1.2.121 (♀) or 1.321 (♂)
Leg 3
1.1.223
1.1.221
Leg 4
1.1.223
1.1.121
Leg 5 with distinct exopod and baseoendopod; endopodal lobe with 5 (♀) or 2 (♂) elements; exopod with 5–6
(♀) or 6 (♂) elements; elements typically short, stubby and hirsute.
Hastigerella Nicholls, 1935 (Family Ectinosomatidae)
Nicholls (1935: 381) established the genus Hastigerella for a new species Hastigerella palpilabra Nicholls,
1935 (type species by monotypy) collected in Kames Bay, Millport (Scotland), and considered it most closely
related to the genus Arenosetella Wilson, 1932. The principal diagnostic difference between the genera
Arenosetella and Hastigerella is the presence of anal hooks or claws in the latter. McLachlan and Moore
(1978: 198) relegated H. palpilabra to a junior synonym of Ectinosoma tenuissima Klie, 1929 (incorrect
original spelling of tenuissimum) and – based on their assertion that Nicholls (1935) had overlooked the anal
claws – transferred this species to Arenosetella. They retained Hastigerella as a valid generic name and
designated Ectinosoma leptoderma Klie, 1929 as the new type species. This course of action is in violation
with the Code since, once fixed, name-bearing types are stable and provide objective continuity in the
application of names (ICZN Art. 61.1.3).
It is, however, not clear from McLachlan and Moore’s (1978) publication whether they had also reexamined Nicholls’s type specimens of H. palpilabra to substantiate their claim of the presence of anal claws
in this species. Since Nicholls’s (1935) descriptions are excellent by contemporary standards it would be
surprising if he indeed had overlooked such conspicuous stuctures. If on the other hand his observation proves
to be correct, there would be no grounds for removing H. palpilabra from Hastigerella and thus no need to
resolve a non-existent issue – simply a need to reverse an error. The only material of H. palpilabra from
Kames Bay that has ever been deposited in the Natural History Museum in London (NHM reg. nos
1947.10.14.28–30) consists of a jar containing five vials with a considerable number of specimens and a label
stating that the material was transferred to ethanol by C.G.M., suggesting Dr Colin Moore had examined it
(Nicholls’s material was originally preserved in formalin). The vials each contained a label mentioning the
name “longisetosus” but the jar also contained an additional label in A.G. Nicholls’s handwriting, stating
“longisetosus = MS name for Hastigerella palpilabra”. There is no doubt that this material represents the type
series (syntypes) designated by Nicholls and was subsequently re-examined by McLachlan and Moore (1978).
Re-examination confirmed the latter authors’ observation; however, the anal claws are less well developed
and curved than in e.g. Arenosetella germanica Kunz, 1937, and are also aligned to the margins of the anal
opening which is probably the reason why Nicholls (1935) had overlooked them.
Adopting McLachlan and Moore’s (1978) synonymy of H. palpilabra renders Hastigerella a junior
subjective synonym of Arenosetella and therefore an invalid name. The orphaned taxonomic grouping
equivalent to Hastigerella sensu McLachlan and Moore (1978) needs a new generic name for which I propose
Glabrotelson gen. nov. (Latin glaber = bald, alluding to the absence of anal hooks which differentiates the
genus from Arenosetella; gender: neuter). Hastigerella mehuinensis Mielke, 1986 is here formally fixed as the
type of the new genus-group name. The generic diagnosis of Glabrotelson gen. nov. coincides with that given
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by Huys et al. (1996: 188) for the genus Hastigerella (according to ICZN Art. 13.1.2 such bibliographic
reference is sufficient to satisfy the criteria of availability).
The genus Glabrotelson contains the following species and subspecies: Ectinosoma leptoderma Klie,
1929 = G. leptoderma (Klie, 1929) comb. nov.; Pararenosetella meridionalis Chappuis, 1954a = G.
meridionale (Chappuis, 1954a) comb. nov.; Pararenosetella psammae Noodt, 1955a = G. psammae (Noodt,
1955a) comb. nov.; Hastigerella abbotti Lang, 1965 = G. abbotti (Lang, 1965) comb. nov.; Pararenosetella
monniotae Guille & Soyer, 1966 = G. monniotae (Guille & Soyer, 1966) comb. nov.; Pararenosetella clavata
Rao & Ganapati, 1969 = G. clavatum (Rao & Ganapati, 1969) comb. nov.; Arenosetella noodti Rao &
Ganapati, 1969 = G. noodti (Rao & Ganapati, 1969) comb. nov.; Arenosetella setosus [sic] Rao & Ganapati,
1969 = G. setosum (Rao & Ganapati, 1969) comb. nov.; Hastigerella bengalensis Rao, 1972 = G. bengalense
(Rao, 1972) comb. nov.; Hastigerella bodini Apostolov, 1974 = G. bodini (Apostolov, 1974) comb. nov.;
Hastigerella bozici Soyer, 1974 = G. bozici (Soyer, 1974) comb. nov.; Hastigerella chappuisi Soyer, 1974 = G.
chappuisi (Soyer, 1974) comb. nov.; Hastigerella scheibeli Mielke, 1975 = G. scheibeli (Mielke, 1975) comb.
nov.; Hastigerella soyeri Bodin, 1976 = G. soyeri (Bodin, 1976) comb. nov.; Hastigerella abbotti
santacruzensis Mielke, 1979 = G. abbotti santacruzense (Mielke, 1979) comb. nov.; Hastigerella mehuinensis
Mielke, 1986 = G. mehuinense (Mielke, 1986) comb. nov. [type]; Hastigerella antarctica Dahms & Schminke,
1992 = G. antarcticum (Dahms & Schminke, 1992) comb. nov.
According to Wells’s (2007) checklist Arenosetella unisetosa Wells, 1965a (= G. unisetosum (Wells,
1965a) comb. nov.) and Hastigerella sp. sensu Mielke (1986) are to be considered species inquirendae in
Glabrotelson.
Interleptomesochra Lang, 1965 (Family Ameiridae)
Sars (1911b: 418) proposed the genus Leptomesochra Sars, 1911b to accommodate Mesochra macintoshi
Scott & Scott, 1895b (incorrect original spelling “MacIntoshi”), Normanella attenuata A. Scott, 1896, and
two new species Leptomesochra tenuicornis Sars, 1911b and L. confluens Sars, 1911c, but neglected type
fixation. According to Lang (1936d: 134) L. confluens was sufficiently different from its congeners to exclude
it from the genus Leptomesochra, an opinion that had already been voiced earlier by Gurney (1932: 50).
Consequently, Lang (1936d: 135) proposed the genus Leptameira (type species by monotypy: Leptomesochra
confluens) which remained monotypic until Nicholls (1939a: 282) added Leptameira attenuata Nicholls,
1939a. Lang’s (1936d: 134–135) superficial statement that “…Leptomesochra confluens Sars unterscheidet
sich von den übrigen Arten im Bau des 1. Maxillipeden und des 5. Beinpaares und muß in eine besondere
Gattung … gestellt werden” does not satisfy the provisions of ICZN Art. 13.1.1. since no specific diagnostic
characters were mentioned; consequently, it does not make the name Leptameira available. Conversely,
Nicholls (1939a: 282) explicitly mentioned the fused rami in leg 5 as a differentiating character for
Leptameira, however, since he refrained from type fixation, his paper also failed to make the generic name
available. After having consulted Klie’s (1929: 359) previously unnoticed description of the male of
Leptomesochra macintoshi (Scott & Scott, 1895b), Lang (1944: 24; 1948: 785) himself relegated Leptameira
to a junior subjective synonym of Leptomesochra. He also added Leptomesochra infima Monard, 1928 to the
latter genus but overlooked Nicholls’s (1939a) description of Leptameira attenuata. Lang’s (1948: 7, 839)
inexplicit designation of the oldest originally included species, Mesochra macintoshi, as the type of
Leptomesochra is invalid for reasons outlined above. The first author to have validly fixed the type is Vervoort
(1964: 270) who cited Normanella attenuata A. Scott, 1896 as the type species. Vervoort (1964: 271) also
included L. nasuta Sewell, 1940, L. africana Kunz, 1951, L. eulitoralis Noodt, 1952, L. elongata Božić, 1955,
L. pygmaea Vervoort, 1964 and L. nichollsi Noodt, 1955a in the genus, the latter being a new substitute name
for Leptomesochra attenuata (Nicholls, 1939a) (a secondary subjective homonym of L. attenuata (A. Scott,
1896) (cf. Noodt 1955a: 96).
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Lang (1965: 338) revised the genus Leptomesochra and subdivided it into three genera based on the
endopodal segmentation of legs 3–4. In the genus Praeleptomesochra Lang, 1965 he grouped Leptomesochra
africana (type by original designation) and Praeleptomesochra similis Lang, 1965, both of which display 3segmented endopods in legs 3–4. He also referred Leptomesochra pygmaea to this genus in a postscript to his
work (Lang 1965: 548). In the genus Leptomesochra he included species which exhibit 2-segmented
endopods (Mesochra macintoshi, Leptomesochra nasuta, L. confluens, L. infima and Leptameira attenuata).
Finally, in the genus Interleptomesochra Lang, 1965 he included species with a 2-segmented leg 3 endopod
and a 3-segmented leg 4 endopod (Normanella attenuata, Leptomesochra tenuicornis, L. elongata, L.
eulitoralis and Interleptomesochra reducta Lang, 1965) and fixed L. tenuicornis as the type species by
original designation. However, the inclusion of N. attenuata in this genus effectively renders the generic name
Interleptomesochra Lang, 1965 a junior subjective synonym of Leptomesochra. Hence, according to current
ameirid systematics the generic concept of Leptomesochra Sars, 1911b in reality coincides with Lang’s (1965:
375) diagnosis of Interleptomesochra. In addition to the five species referred to this genus by Lang (1965) and
which are listed above, Leptomesochra also includes two other species that were originally assigned to
Interleptomesochra, i.e I. noodti Galhano, 1968 = Leptomesochra noodti (Galhano, 1968) comb. nov. and I.
boguensis Lindgren, 1975 = L. boguensis (Lindgren, 1975) comb. nov. As a result of this nomenclatural
change an alternative generic name is required to accommodate the species of the orphaned taxon
“Leptomesochra sensu Lang (1965)”, i.e. Mesochra macintoshi, Leptomesochra nasuta, L. confluens, L.
infima and Leptameira attenuata. Rather than introducing an entirely new name I have elected to make
Leptameira Lang, 1936d available, taking the date and authorship of the present paper. It can be differentiated
from the other genera in the Ameiridae by the diagnosis given below.
Ameiridae. Body slender. Rostrum typically small (except L. nasuta). Antennule 8- or 9-segmented in female,
without pinnate setae; haplocer in male, with 3 segments distal to geniculation. Antenna with discrete basis or
allobasis; exopod 1-segmented, with 2–3 setae. Mandibular palp without defined exopod; basis with 1–3
elements. Caudal rami short, at most 1.5 times as long as wide.
Leg 1 with prehensile 3- (L. infima, L. hirsuta) or 2-segmented endopod; enp-1 much longer than
remaining segments and longer than exopod; exp-2 with or without outer spine, without inner seta; exp-3 with
4–5 elements. Legs 2–4 with 3-segmented exopods and 2-segmented endopods; no sexual dimorphism. Setal
formulae of swimming legs as follows:
Exopod
Endopod
Leg 2
0.1.(0–1)2(2–3)
(0–1).(1–2)2(0–1)
Leg 3
0.(0–1).(0–1)2(2–3)
(0–1).(1–2)2(0–1)
Leg 4
0.1.(1–2)2(2–3)
(0–1).12(0–1)
Leg 5 exopod and baseoendopod fused (L. confluens, L. attenuata, L. nasuta and L. eremeensis) or distinct
(other species) in ♀, always fused in ♂ forming bilobate plate; endopodal lobe with 3–4 (♀) or 3 (♂)
elements; exopod(al lobe) with 5–6 (♀) or 3 or 5 (♂) elements. Leg 6 with 3 setae in ♂.
Leptameira gen. nov. (type species: Leptomesochra confluens Sars, 1911c) includes the following new
combinations: Mesochra macintoshi Scott & Scott, 1895b = Leptameira macintoshi (Scott & Scott, 1895b)
comb. nov., Leptomesochra confluens Sars, 1911c = Leptameira confluens (Sars, 1911c) comb. nov.,
Leptomesochra infima Monard, 1928 = Leptameira infima (Monard, 1928) comb. nov., Leptameira attenuata
Nicholls, 1939a = Leptameira attenuata (Nicholls, 1939a) comb. nov.; Leptomesochra nasuta Sewell, 1940 =
Leptameira nasuta (Sewell, 1940) comb. nov., Leptomesochra hirsuta Wiborg, 1964 = Leptameira hirsuta
(Wiborg, 1964) comb. nov., Leptomesochra theodoridis Soyer, 1966 = Leptameira theodoridis (Soyer, 1966)
comb. nov., Leptomesochra eremeensis Kornev & Chertoprud, 2008 = Leptameira eremeensis (Kornev &
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Chertoprud, 2008) comb. nov., and Leptomesochra kunzi Kornev & Chertoprud, 2008 = Leptameira kunzi
(Kornev & Chertoprud, 2008) comb. nov.
Noodt (1955a: 96) proposed the new replacement name Leptomesochra nichollsi for Leptomesochra
attenuata (Nicholls, 1939a) (originally combined with the generic name Leptameira Lang, 1936d) which is a
junior secondary homonym of Leptomeschra attenuata (A. Scott, 1896) (originally combined with the generic
name Normanella). Although Nicholls’s (1939a) species name was originally combined with an unavailable
generic name, this does not affect its availability (ICZN Art. 11.9.3.1). Based on the argument that both
species were no longer regarded congeneric, Lang (1965: 338) reinstated Nicholls’s (1939a) specific name
and considered L. nichollsi a junior subjective synonym. Since the substitute name Leptomesochra nichollsi is
no longer in use (Bodin 1997; Wells 2007) it is indeed to be rejected according to ICZN Art. 59.3 and hence
Leptameira attenuata Nicholls, 1939a is to be considered the valid name for the species it denotes.
Addendum: Junior homonyms in Calanoida and Cyclopoida
Parathalassius Dussart, 1986
Özdikmen (2008: 266) noted that Parathalassius Dussart, 1986 (type species by monotypy: P. fagesi Dussart,
1986) (Calanoida: Centropagidae) is a junior homonym of Parathalassius Mik, 1891 (type species P. blasigii
Mik, 1891) (Diptera: Empididae) and proposed the new replacement name “Dusartius”. Since this name was
clearly intended as a patronym in honour of the late Prof. Bernard H. Dussart (“Etymology: from B.H. Dusart
who is the current author of pre-existing genus Parathalassius”), Özdikmen’s (2008) spelling is to be
considered a lapsus calami which has no separate availability and cannot be used as a substitute name (ICZN
Art. 32.4). The correction of incorrect original spellings resulting from inadvertent errors is mandatory and
hence Dusartius has to be emended to Dussartius in accordance with ICZN Art. 32.5.1, retaining the
authorship and date of the original spelling. Unfortunately Dussartius Özdikmen, 2008 is a junior homonym
of the monotypic Dussartius Kiefer, 1978 (type species: Eudiaptomus baeticus Dussart, 1967; see Kiefer
(1978: 156)) (Calanoida: Diaptomidae) and hence requires a new replacement name for which I propose
Dussartopages nom. nov. (gender: masculine). Dussartopages fagesi (Dussart, 1986) comb. nov. is the type
and sole species of the genus.
Berea Yamaguti, 1963
In a posthumous publication, Péringuey (1926: 626) proposed the genus-group name Berea in the Carabidae
(Coleoptera) for Argutoridius trivialis Boheman, 1858 and a new species Berea caffer (no type designated).
Yamaguti (1963: 280) established the genus Berea for Triphyllocanthus ancoralis Bere, 1936 (type by
monotypy) in the family Chondracanthidae (Cyclopoida). Since Berea Yamaguti, 1963 is a junior homonym
of Berea Péringuey, 1926, it cannot be used as a valid name. It is here replaced by the new substitute name
Bereacanthus nom. nov. (named after the late Dr Ruby Bere; gender: masculine). The genus contains
Bereacanthus ancoralis (Bere, 1936) comb. nov. [type], Acanthochondria tenuis Pearse, 1952 = Bereacanthus
tenuis (Pearse, 1952) comb. nov., Pseudochondracanthus nellcauseyae Causey, 1955 = B. nellcauseyae
(Causey, 1955) comb. nov. and Berea clava Ho & Sey, 1997 = B. clava (Ho & Sey, 1997) comb. nov.
Acknowledgements
Natalia J. Vandenberg (Systematic Entomology Laboratory, U.S. Department of Agriculture, Washington,
D.C.), David Reid and David McL Roberts (Department of Zoology, NHM, London) are acknowledged for
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providing relevant literature references. Miguel A. Alonso-Zarazaga (Museo Nacional de Ciencias Naturales
Madrid, ICZN Commissioner), Mark J. Grygier (Lake Biwa Museum, ICZN Commissioner), Peter Ng Kee
Lin (National University of Singapore, ICZN Commissioner) and David Notton (ICZN Secretariat) are
gratefully acknowledged for advice and constructive criticism, Prof. John B. J. Wells (University of
Wellington) for his constant encouragement, and Prof. Alessandro Minelli (University of Padova) for efficient
editorial handling.
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Prior nominal taxon
Proposed nominal taxon
Type
Cerviniopseinae Brotskaya, 1963
Huntemanniidae Por, 1986a
Paranannopodinae Por, 1986a
Pontostratiotinae A. Scott, 1909
Nannopodidae Brady, 1880a
Danielsseniinae Huys & Gee in Huys et al., 1996
Pontostratiotes Brady, 1883
Nannopus Brady, 1880a
Danielssenia Boeck, 1873
Idomene Philippi, 1843 a
Rhizothrix Brady & Robertson, 1876
Amphiascus Sars, 1905a b
Anoplosoma Sars, 1911c
Leptomesochra Sars, 1911b c
Teissierella Monard, 1935a d
Hastigerella Nicholls, 1935 e
Psammastacus Nicholls, 1935
Xouthous Thomson, 1883
Rhizothrix Sars, 1909a
Sarsamphiascus gen. nov.
Anoplosomella Strand, 1929
Leptameira gen. nov.**
Monardius gen. nov.
Glabrotelson gen. nov.
Psammastacus Apostolov & Marinov, 1988
Paranannopus Lang, 1936a
Paraidya Sewell, 1940
Alteuthellopsis Lang, 1944
Halectinosoma Lang, 1944
Idyellopsis Lang, 1944
Paralaophonte Lang, 1944
Robertgurneya Lang, 1944
Rheocamptus Borutzky, 1948*
Apodopsyllus Kunz, 1962
Paranannopus gen. nov.**
Paraidya gen. nov.**
Alteuthellopsis Lang, 1948
Halectinosoma Vervoort, 1962
Idyellopsis Lang, 1948
Paralaophonte Lang, 1948
Robertgurneya Apostolov & Marinov, 1988
Rheocamptus Borutzky, 1952
Apodopsyllus gen. nov.**
Xouthous novaezealandiae Thomson, 1883
Enhydrosoma curvatum Brady, 1880a = Rhizothrix curvata (Brady, 1880a)
Dactylopus minutus Claus, 1863 = Sarsamphiascus minutus (Claus, 1863) comb. nov.
Anoplosoma sordidum Sars, 1911c = Anoplosomella sordida (Sars, 1911c) comb. nov.
Leptomesochra confluens Sars, 1911c = Leptameira confluens (Sars, 1911c)
Teissierella salammboi Monard, 1935b = Monardius salammboi (Monard, 1935b) comb. nov.
Hastigerella mehuinensis Mielke, 1986 = Glabrotelson mehuinense (Mielke, 1986) comb. nov.
Psammastacus confluens Nicholls, 1935 = Psammastacus confluens (Nicholls, 1935) comb. nov. sensu
novo
Nannopus abyssi Sars, 1920c = Parannopus abyssi (Sars, 1920c) comb. nov.
Paraidya major Sewell, 1940 = Paraidya major (Sewell, 1940) comb. nov. sensu novo
Eupelte oblivia A. Scott, 1909 = Alteuthellopsis oblivia (A. Scott, 1909)
Ectinosoma sarsii Boeck, 1873 = Halectinosoma sarsii (Boeck, 1873)
Idyellopsis typica Lang, 1948
Cleta brevirostris Claus, 1863 = Paralaophonte brevirostris (Claus, 1863)
Stenhelia similis A. Scott, 1896 = Robertgurneya similis (A. Scott, 1896)
Canthocamptus Zschokkei Schmeil, 1893 = Bryocamptus zschokkei (Schmeil, 1893)
Apodopsyllus panamensis Mielke, 1984a = Apodopsyllus panamensis (Mielke, 1984a) comb. nov. sensu
novo
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UNRESOLVED CASES IN HARPACTICOID COPEPOD NOMENCLATURE
TABLE 1. Summary of nomenclatural changes proposed in this paper (* denotes subgeneric names; ** denotes previously used, unavailable names made available in this paper
by the present author). New replacement names are indicated by the suffix “nom. nov.” only when they substitute for a junior homonym; in all other cases (synonyms, previously
unavailable names) the substitute name is followed by the suffix “gen. nov.”; the suffix “sensu novo” is applied when a new combination results in an identical binomen but with
different authorship and date.
a
sensu Sars (1906a), Lang (1948) and subsequent authors; b sensu Lang (1944, 1948) and subsequent authors; c sensu Lang (1965); d sensu Bodin (1997) and Wells (2007); e
sensu McLachlan & Moore (1978).
67
Proposed nominal taxon
Type
Intermedopsyllus Kunz, 1962*
Kliopsyllus Kunz, 1962
Scottopsyllus Kunz, 1962
Scottopsyllus Kunz, 1962*
Cladorostrata Shen & Tai, 1963
Interleptomesochra Lang, 1965
Poria Lang, 1965
Sewellia Lang, 1965
Scottolana Por, 1967
Ichnusella Cottarelli, 1971
Langpsyllocamptus Kunz, 1975b*
Micropsammis Mielke, 1975
Namakosiramia Ho & Perkins, 1977
Barbaracletodes Becker, 1979
Ameiropsyllus Bodin, 1979
Chilaophonte Mielke, 1985
Intermediopsyllus subgen. nov.**
Emertonia Wilson, 1932
Wellsopsyllus Kunz, 1981
Scottopsyllus Apostolov & Marinov, 1988
Cladorostrata Tai & Song, 1979
Leptomesochra Sars, 1911b
Hanikraia nom. nov.
Sewelliapusia nom. nov.
Scottolana gen. nov.**
Bereraia nom. nov.
Psyllocamptus T. Scott, 1899b*
Micropsammis Gee & Huys, 1991
Microchelonia Brady, 1918
Barbaracletodes gen. nov.**
Ameiropsyllus gen. nov.**
Chilaophonte gen. nov.**
Dahlakia Por, 1986a
Psammonitocrella Rouch, 1992
Dahlakocamptus nom. nov.
Psammonitocrella gen. nov.**
Tectacingulum Harris, 1994
Fiersiella Suárez-Morales &
Iliffe, 2005*
Pilocamptus Wells, 2007
Tectacingulum gen. nov.**
Fiersiella subgen. nov.**
Leptopsyllus intermedius Scott & Scott, 1895b = Wellsopsyllus intermedius (Scott & Scott, 1895b)
Emertonia gracilis Wilson, 1932
Paramesochra gigas Wells, 1965a = Wellsopsyllus gigas (Wells, 1965a)
Leptopsyllus minor Scott & Scott, 1895a = Scottopsyllus minor (Scott & Scott, 1895a)
Cladorostrata brevipoda Shen & Tai, 1963 = Cladorostrata brevipoda (Shen & Tai, 1963)
Normanella attenuata A. Scott, 1896 = Leptomesochra attenuata (A. Scott, 1896)
Hemimesochra derketo Por, 1964a = Hanikraia derketo (Por, 1964a) comb. nov.
Dactylopusia tropica Sewell, 1940 = Sewelliapusia tropica (Sewell, 1940) comb. nov.
Scottolana geei Mu & Huys, 2004 = Scottolana geei (Mu & Huys, 2004) comb. nov. sensu novo
Ichnusella eione Cottarelli, 1971 = Bereraia eione (Cottarelli, 1971) comb. nov.
Mesochra propinqua T. Scott, 1896 = Psyllocamptus propinquus (T. Scott, 1896)
Micropsammis noodti (Mielke, 1975)
Microchelonia glacialis Brady, 1918
Barbaracletodes barbara Becker, 1979 = Barbaracletodes barbara (Becker, 1979) comb. nov. sensu novo
Ameiropsyllus monardi Bodin, 1979 = Ameiropsyllus monardi (Bodin, 1979) comb. nov. sensu novo
Chilaophonte maiquillahuensis Mielke, 1985 = Chilaophonte maiquillahuensis (Mielke, 1985) comb. nov.
sensu novo
Cletocamptus xenuus Por, 1968 = Dahlakocamptus xenuus (Por, 1968) comb. nov.
Psammonitocrella boultoni Rouch, 1992 = Psammonitocrella boultoni (Rouch, 1992) comb. nov.
sensu novo
Tectacingulum tumidum Harris, 1994 = Tectacingulum tumidum (Harris, 1994) comb. nov. sensu novo
Stygonitocrella sequoyahi Reid, Hunt & Stanley, 2003
Pilocamptus gen. nov.**
Canthocamptus pilosus Douwe, 1910 = Pilocamptus pilosus (Douwe, 1910) comb. nov.
Berea Yamaguti, 1963
Dussartius Özdikmen, 2008
Bereacanthus nom. nov.
Dussartopages nom. nov.
Triphyllocanthus ancoralis Bere, 1936 = Bereacanthus ancoralis (Bere, 1936) comb. nov.
Parathalassius fagesi Dussart, 1986 = Dussartopages fagesi (Dussart, 1986) comb. nov.
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68 · Zootaxa 2183 © 2009 Magnolia Press
Prior nominal taxon
HUYS
Prior nominal taxon
Proposed nominal taxon
SUBGENUS WELLSOPSYLLUS KUNZ, 1981
Paramesochra gigas Wells, 1965a [type]
Kliopsyllus runtzi Soyer, 1975a
Wellsopsyllus gigas (Wells, 1965a) comb. nov.
Wellsopsyllus runtzi (Soyer, 1975a) comb. nov.
Kliopsyllus abyssalis Becker, 1979
Wellsopsyllus abyssalis (Becker, 1979) comb. nov.
SUBGENUS SCOTTOPSYLLUS APOSTOLOV & MARINOV, 1988
Leptopsyllus minor Scott & Scott, 1895a [type]
Wellsopsyllus minor (Scott & Scott, 1895a) comb. nov.
Leptopsyllus robertsoni Scott & Scott, 1895a
Leptopsyllus herdmani Thompson & Scott, 1900
Wellsopsyllus robertsoni (Scott & Scott, 1895a) comb. nov.
Wellsopsyllus herdmani (Thompson & Scott, 1900) comb. nov.
Scottopsyllus pararobertsoni Lang, 1965
Scottopsyllus langi Mielke, 1984b
Wellsopsyllus pararobertsoni (Lang, 1965) comb. nov.
Wellsopsyllus langi (Mielke, 1984b) comb. nov.
Scottopsyllus langi langi Mielke, 1984b
Wellsopsyllus langi langi (Mielke, 1984b) comb. nov.
Zootaxa 2183 © 2009 Magnolia Press ·
Scottopsyllus langi continentalis Kunz, 1992
Scottopsyllus praecipuus Veit-Köhler, 2000
Wellsopsyllus langi continentalis (Kunz, 1992) comb. nov.
Wellsopsyllus praecipuus (Veit-Köhler, 2000) comb. nov.
Scottopsyllus depressus Kornev & Chertoprud, 2008
Wellsopsyllus depressus (Kornev & Chertoprud, 2008) comb. nov.
+ species inquirenda: Scottopsyllus minor (Scott & Scott, 1895a) sensu Kunz (1981)
SUBGENUS INTERMEDIOPSYLLUS HUYS, 2009 MIHI
Leptopsyllus intermedius Scott & Scott, 1895b [type]
Paramesochra minuta Nicholls, 1939b
Wellsopsyllus intermedius (Scott & Scott, 1895b) comb. nov.
Wellsopsyllus minutus (Nicholls, 1939b) comb. nov.
+ species incertae sedis: Scottopsyllus (? Intermedopsyllus) smirnovi Kunz, 1992
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UNRESOLVED CASES IN HARPACTICOID COPEPOD NOMENCLATURE
TABLE 2. Species allocated to the subgenera of Wellsopsyllus Kunz, 1981. Inclusion of species inquirendae and species incertae sedis
follows Wells (2007).
69
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TABLE 3. Species allocated to Amphiascus Sars, 1905a and Sarsamphiascus gen. nov. Inclusion of species
inquirendae and species incertae sedis follows Wells (2007).
Prior nominal taxon
Proposed nominal taxon
AMPHIASCUS SARS, 1905A
Dactylopus longirostris Claus, 1863 [type]
Amphiascus Giesbrechti Sars, 1906b
Amphiascus pallidus Sars, 1906b
Paramphiascopsis ekmani Lang, 1965
Paramphiascopsis soyeri Lang, 1965
Paramphiascopsis paromolae Soyer, 1973
Paramphiascopsis triarticulatus Moore, 1976
Paramphiascopsis waihonu Hicks, 1986b
Amphiascus longirostris (Claus, 1863) comb. nov.
Amphiascus giesbrechti Sars, 1906b
Amphiascus pallidus Sars, 1906b
Amphiascus ekmani (Lang, 1965) comb. nov.
Amphiascus soyeri (Lang, 1965) comb. nov.
Amphiascopsis paromolae (Soyer, 1973) comb. nov.
Amphiascopsis triarticulatus (Moore, 1976) comb. nov.
Amphiascopsis waihonu (Hicks, 1986b) comb. nov.
SARSAMPHIASCUS GEN. NOV.
minutus-group
Dactylopus minutus Claus, 1863 [type]
Sarsamphiascus minutus (Claus, 1863) comb. nov.
Dactylopus tenuiremis Brady, 1880a
Sarsamphiascus tenuiremis (Brady, 1880a) comb. nov.
Amphiascus brevis Sars, 1909c
Sarsamphiascus brevis (Sars, 1909c) comb. nov.
Amphiascus congener Sars, 1909c
Sarsamphiascus congener (Sars, 1909c) comb. nov.
Amphiascus caudaespinosus Brian, 1927a
Sarsamphiascus caudaespinosus (Brian, 1927a) comb. nov.
Amphiascus hirtus Gurney, 1927
Sarsamphiascus hirtus (Gurney, 1927) comb. nov.
Amphiascus ultimus Monard, 1928
Sarsamphiascus ultimus (Monard, 1928) comb. nov.
Amphiascus gracilis Lang, 1936c
Sarsamphiascus gracilis (Lang, 1936c) comb. nov.
Amphiascus demersus Nicholls, 1939a
Sarsamphiascus demersus (Nicholls, 1939a) comb. nov.
Amphiascus graciloides Klie, 1950
Sarsamphiascus graciloides Klie, 1950) comb. nov.
Amphiascus paracaudaespinosus Roe, 1958
Sarsamphiascus paracaudaespinosus (Roe, 1958) comb. nov.
Amphiascus longarticulatus Marcus, 1974
Sarsamphiascus longarticulatus (Marcus, 1974) comb. nov.
Amphiascus discrepans Mielke, 1989
Sarsamphiascus discrepans (Mielke, 1989) comb. nov.
+ species inquirendae:
Amphiascus caudaespinosus Brian, 1927a sensu Pesce (1985b)
varians-group
Stenhelia varians Norman & T. Scott, 1905
Amphiascus propinqvus Sars, 1906b
Amphiascus tenellus Sars, 1906c
Amphiascus polaris Sars, 1909c
Amphiascus angustipes Gurney, 1927
Amphiascus gauthieri Monard, 1936
70 · Zootaxa 2183 © 2009 Magnolia Press
Sarsamphiascus varians (Norman & T. Scott, 1905) comb. nov.
Sarsamphiascus propinquus (Sars, 1906b) comb. nov.
Sarsamphiascus tenellus (Sars, 1906c) comb. nov.
Sarsamphiascus polaris (Sars, 1909c) comb. nov.
Sarsamphiascus angustipes (Gurney, 1927) comb. nov.
Sarsamphiascus gauthieri (Monard, 1936) comb. nov.
HUYS
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Prior nominal taxon
Proposed nominal taxon
Amphiascus dentiformis Coull, 1971b
Sarsamphiascus dentiformis (Coull, 1971b) comb. nov.
Amphiascus lobatus Hicks, 1971
Sarsamphiascus lobatus (Hicks, 1971) comb. nov.
Amphiascus elongatus Itô, 1972
Sarsamphiascus elongatus (Itô, 1972) comb. nov.
Amphiascus profundus Becker & Schriever, 1979
Sarsamphiascus profundus (Becker & Schriever, 1979) comb. nov.
Amphiascus tainui Hicks, 1989
Sarsamphiascus tainui (Hicks, 1989) comb. nov.
+ species inquirendae:
Amphiascus tenellus Sars, 1906c sensu Noodt (1964)
Amphiascus angustipes Gurney, 1927 sensu Bodin (1964)
+ species incertae sedis: Stenhelia perplexa Thompson & Scott, 1903
Mesamphiascus ampullifer Humes, 1953
pacificus-group
Amphiascus pacificus Sars, 1905a
Sarsamphiascus pacificus (Sars, 1905a) comb. nov.
Amphiascus parvus Sars, 1906b
Sarsamphiascus parvus (Sars, 1906b) comb. nov.
Amphiascus sinuatus Sars, 1906c
Sarsamphiascus sinuatus (Sars, 1906c) comb. nov.
Amphiascus undosus Lang, 1965
Sarsamphiascus undosus (Lang, 1965) comb. nov.
Amphiascus kawamurai Ueda & Nagai, 2005
Sarsamphiascus kawamurai (Ueda & Nagai, 2005) comb. nov.
+ species inquirendae:
Amphiascus humphriesi Roe, 1960
Amphiascus parvus Sars, 1906b sensu Rouch (1962)
Amphiascus parvus Sars, 1906b sensu Noodt (1964)
amblyops-group
Amphiascus amblyops Sars, 1911a
Sarsamphiascus amblyops (Sars, 1911a) comb. nov.
Sarsamphiascus species inquirendae
Amphiascus monodi Monard, 1952
Amphiascus sp. sensu Pesta (1959)
Amphiascus sp. I sensu Griga (1961)
Amphiascus sp. II sensu Griga (1961)
Sarsamphiascus species incertae sedis
Robertgurneya intermedia Božić, 1955
UNRESOLVED CASES IN HARPACTICOID COPEPOD NOMENCLATURE
Zootaxa 2183 © 2009 Magnolia Press ·
71
Genus-group name (subgenera indented)
Type species (original binomen)
Type species (valid binomen)
Abnitocrella Karanovic, 2006
Abscondicola Fiers, 1990
Acrenhydrosoma Lang, 1944
Actopsyllus Wells, 1967
Aculeopsis Huys & Willems, 1989
Adenopleurella Huys, 1990c
Acuticoxa Huys & Kihara, 2009
Aegisthus Giesbrecht, 1891
Aequinoctiella Cottarelli, Bruno & Berera, 2008
Afrocamptus Chappuis, 1932b
Afrolaophonte Chappuis, 1960
Afroleptastacus Huys, 1992
Afrosenia Huys & Gee, 1996a
Aigondiceps Fiers, 1995
Algensiella Cottarelli & Baldari, 1987a
Allophyllognathopus Kiefer, 1967
Alteutha Baird, 1846b
Alteuthella A. Scott, 1909
Alteuthellopsis Lang, 1948
Alteuthoides Hicks, 1986a
Ambunguipes Huys, 1990b
Ameira Boeck, 1865
Ameiropsis Sars, 1907b
Ameiropsyllus Huys, 2009 mihi
Ameliotes Por, 1969
Amenophia Boeck, 1865
Amerolaophontina Fiers, 1991b
Amonardia Lang, 1944
Abnitocrella halsei Karanovic, 2006
Abscondicola humesi Fiers, 1990
Cletodes perplexa T. Scott, 1899b
Acrenhydrosoma perplexum (T. Scott, 1899b)
Actopsyllus longipes Wells, 1967
Aculeopsis longisetosa Huys & Willems, 1989
Adenopleurella brevipes Huys, 1990c
Acuticoxa ubatubae Huys & Kihara, 2009
Aegisthus mucronatus Giesbrecht, 1891
Aequinoctiella cavalettii Cottarelli, Bruno & Berera, 2008
Eucamptus uncinatus Chappuis, 1932a
Afrocamptus uncinatus (Chappuis, 1932a)
Afrolaophonte monodi Chappuis, 1960
Afroleptastacus clandestinus Huys, 1992
Danielssenia spinipes Wells, 1967
Afrosenia spinipes (Wells, 1967)
Diagoniceps bocki Lang, 1948
Aigondiceps bocki (Lang, 1948)
Algensiella boitanii Cottarelli & Baldari, 1987a
Allophyllognathopus brasiliensis Kiefer, 1967
Cyclops depressus Baird, 1837
Alteutha depressa (Baird, 1837)
Alteuthella pellucida A. Scott, 1909
Eupelte oblivia A. Scott, 1909
Alteuthellopsis oblivia (A. Scott, 1909)
Alteuthoides kootare Hicks, 1986a
Thalestris rufocincta Norman in Brady, 1880a
Ambunguipes rufocincta (Norman in Brady, 1880a)
Ameira longipes Boeck, 1865
Ameiropsis brevicornis Sars, 1907b
Ameiropsyllus monardi Bodin, 1979
Ameiropsyllus monardi (Bodin, 1979)
Ameliotes malagassicus Por, 1969
Amenophia peltata Boeck, 1865
Laophontina reducta Coull & Zo, 1980
Amerolaophontina reducta (Coull & Zo, 1980)
Dactylopus similis Claus, 1866
Amonardia similis (Claus, 1866)
Type fixation
original designation
original designation
original designation
original designation
original designation
original designation
original designation
subsequent designation
original designation
monotypy
monotypy
original designation
original designation
original designation
monotypy
monotypy
monotypy
subsequent designation
monotypy
original designation
original designation
subsequent designation
subsequent designation
original designation
original designation
monotypy
original designation
original designation
Notes
(1, 2)
(3)
(4)
(1, 5)
(4)
(6, 24)
(7)
(1, 8)
(4)
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TABLE 4. Harpacticoid genus-group names with original and valid (only listed if different in spelling or combination from original) binominal names of type species
and mode of type fixation: original designation = with explicit fixation in original publication (ICZN Art. 68.2); indication = original designation without explicit
fixation but originally included species with species-group name typicus or –a or –um becomes the type (ICZN Art. 68.2.2); monotypy = genus-group name established
for a single species which was not explicitly designated as the type (ICZN Art. 68.3); subsequent designation = by the first author(s) who subsequently validly
designate(s) one of the originally included species as the type species; subsequent monotypy = if only one species was first subsequently included in a genus established
without included species; designation by ICZN = by use of the plenary power of the International Commission on Zoological Nomenclature (ICZN Art. 81). Genera of
uncertain taxonomic position are classified as genera incertae sedis; those of doubtful identity requiring further investigation as genera inquirenda. *: The use of the
genitive ending -i in a subsequent spelling of a species-group name that is a genitive based upon a personal name in which the correct original spelling ends with -ii, is
here treated as an incorrect subsequent spelling, even if the change in spelling is deliberate (ICZN Art. 33.4). Despite its potentially negative conotation (Evenhuis 2008)
I have used the term “mihi” to denote names made available in this paper by the present author; this was considered particularly desirable in the case of taxa previously
denoted by unavailable names which are made available and validated herein.
HUYS
Zootaxa 2183 © 2009 Magnolia Press ·
Type species (original binomen)
Type species (valid binomen)
Amphiascoides Nicholls, 1941b
Amphiascopsis Gurney, 1927
Amphiascus Sars, 1905a
Amphibiperita Fiers & Rutledge, 1990
Anapophysia Huys & Gee 1996b
Ancorabolus Norman, 1903b
Ancorabolina George, 2006b
Andromastax Conroy-Dalton & Huys, 1999
Anisostenhelia Mu & Huys, 2002
Anoplosomella Strand, 1929
Antarcticobradya Huys, 1987
Antarctobiotus Chappuis, 1930
Antiboreodiosaccus Lang, 1944
Antillesia Humes, 1958
Antrocamptus Chappuis, 1957
Apistophonte Gheerardyn & Fiers in Gheerardyn et al., 2006b
Apodonsiella Hicks, 1988b
Apodopsyllus Huys, 2009 mihi
Applanola Huys & Lee, 2000
Aquilastacus Huys & Conroy-Dalton, 2005
Arawella Cottarelli & Baldari, 1987b
Arbutifera Huys & Kunz, 1988
Archesola Huys & Lee, 2000
Archilaophonte Willen, 1995
Archileptastacus Huys, 1992
Archinitocrella Karanovic, 2006
Archisenia Huys & Gee, 1993
Archizausodes Bouck, Thistle & Huys, 1999
Arcticocarella Kornev & Chertoprud, 2008
Arenocaris Nicholls, 1935
Arenolaophonte Lang, 1965
Arenopontia Kunz, 1937
Arenosetella Wilson, 1932
Arenotopa Chappuis & Rouch, 1960
Argestes Sars, 1910
Argestigens Willey, 1935
Arthropsyllus Sars, 1909b
Arthuricornua Conroy-Dalton, 2001
Asellopsis Brady & Robertson, 1873
Aspinothorax Moura & Martínez Arbizu, 2003
Atergopedia Martínez Arbizu & Moura, 1998
Dactylopus debilis Giesbrecht, 1881
Amphiascoides debilis (Giesbrecht, 1881)
Dactylopus cinctus Claus, 1866
Amphiascopsis cinctus (Claus, 1866)
Dactylopus longirostris Claus, 1863
Amphiascus longirostris (Claus, 1863)
Mesochra neotropica Jakobi, 1956
Amphibiperita neotropica (Jakobi, 1956)
Psammis borealis Klie, 1939
Anapophysia borealis (Klie, 1939)
Ancorabolus mirabilis Norman, 1903b
Ancorabolina chimaera George, 2006b
Andromastax muricatus Conroy-Dalton & Huys, 1999
Stenhelia (Stenhelia) asetosa Thistle & Coull, 1979 Anisostenhelia asetosa (Thistle & Coull, 1979)
Anoplosoma sordidum Sars, 1911c
Anoplosomella sordida (Sars, 1911c)
Parastenhelia (?) tenuis Brady, 1910
Antarcticobradya tenuis (Brady, 1910)
Attheyella (Canthocamptus) Koenigi Pesta, 1928 Antarctobiotus koenigi (Pesta, 1928)
Dactylopus crassus Giesbrecht, 1902
Antiboreodiosaccus crassus (Giesbrecht, 1902)
Antillesia cardisomae Humes, 1958
Antrocamptus Coiffaiti Chappuis, 1957
Antrocamptus coiffaiti Chappuis, 1957
Apistophonte wasiniensis Gheerardyn & Fiers in Gheerardyn et al., 2006b
Apodonsiella indica Hicks, 1988b
Apodopsyllus panamensis Mielke, 1984a
Apodopsyllus panamensis (Mielke, 1984a)
Laophonte hirsuta Thompson & Scott, 1903
Applanola hirsuta (Thompson & Scott, 1903)
Aquilastacus serratus Huys & Conroy-Dalton, 2005
Arawella alexandri Cottarelli & Baldari, 1987b
Delamarella phyllosetosa Kunz, 1984
Arbutifera phyllosetosa (Kunz, 1984)
Laophonte typhlops Sars, 1908a
Archesola typhlops (Sars, 1908a)
Archilaophonte maxima Willen, 1995
Leptastacus aberrans dichatoensis Mielke, 1985
Archileptastacus dichatoensis (Mielke, 1985)
Archinitocrella newmanensis Karanovic, 2006
Danielssenia sibirica Sars, 1898
Archisenia sibirica (Sars, 1898)
Zausodes biarticulatus Itô, 1979a
Archizausodes biarticulatus (Itô, 1979a)
Arcticocarella reducta Kornev & Chertoprud, 2008
Arenocaris bifida Nicholls, 1935
Arenolaophonte stygia Lang, 1965
Arenopontia subterranea Kunz, 1937
Arenosetella spinicauda Wilson, 1932
Arenotopa ghanai Chappuis & Rouch, 1960
Argestes mollis Sars, 1910
Argestigens uniremis Willey, 1935
Arthropsyllus serratus Sars, 1909b
Arthuricornua anendopodia Conroy-Dalton, 2001
Asellopsis hispida Brady & Robertson, 1873
Asellopsis hispidus Brady & Robertson, 1873
Aspinothorax insolentis Moura & Martínez Arbizu, 2003
Atergopedia vetusta Martínez Arbizu & Moura, 1998
Type fixation
Notes
original designation
subsequent designation
subsequent designation
original designation
original designation
monotypy
original designation
original designation
original designation
monotypy
original designation
monotypy
original designation
original designation
monotypy
original designation
monotypy
original designation
original designation
original designation
monotypy
original designation
original designation
monotypy
original designation
original designation
original designation
original designation
original designation
monotypy
monotypy
monotypy
original designation
monotypy
monotypy
monotypy
monotypy
original designation
monotypy
original designation
monotypy
(9)
(10)
(4)
(11)
(4)
(12)
(12)
(4)
(13)
(14)
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UNRESOLVED CASES IN HARPACTICOID COPEPOD NOMENCLATURE
Genus-group name (subgenera indented)
73
Type species (original binomen)
Type species (valid binomen)
Attheyella Brady, 1880a
Attheyella Brady, 1880a
Canthosella Chappuis, 1931
Chappuisiella Brehm, 1925
Delachauxiella Brehm, 1925
Neomrazekiella Özdikmen & Pesce, 2006
Ryloviella Borutzky, 1931d
Australocamptus Karanovic, 2004
Austrocletodes Pallares, 1979
Avdeevia Bresciani & Lützen, 1994
Balaenophilus Aurivillius, 1879
Balucopsylla Rao, 1972
Bathycamptus Huys & Thistle, 1989
Bathycletopsyllys Huys & Lee, 1999
Bathyesola Huys & Lee, 2000
Bathyidia Farran, 1926
Bathylaophonte Lee & Huys, 1999b
Bathypsammis Huys & Gee, 1993
Beatricella T. Scott, 1905b
Belemnopontia Huys, 1992
Bereraia Huys, 2009 mihi
Biameiropsis Karanovic, 2006
Biuncus Huys, 1996a
Bolbotelos Huys & Conroy-Dalton, 2006
Boreolimella Huys & Thistle, 1989
Boreopontia Willems, 1981
Boreovermis Huys & Conroy-Dalton, 2006
Bradya Boeck, 1873
Bradyellopsis Brian, 1925b
Brescianiana Avdeev, 1982
Breviconia Conroy-Dalton & Huys, 2000
Brevifrons Harris, 1994
Brianola Monard, 1927
Bryocamptus Chappuis, 1929a
Bryocamptus Chappuis, 1929a
Arcticocamptus Chappuis, 1929a
Echinocamptus Chappuis, 1929a
Rheocamptus Borutzky, 1952
Bulbamphiascus Lang, 1944
Caligopsyllus Kunz, 1975b
Canthocamptus crassus Sars, 1863
Attheyella crassa (Sars, 1863)
Canthocamptus crassus Sars, 1863
Attheyella crassa (Sars, 1863)
Canthocamptus muscicola Chappuis, 1928b
Attheyella muscicola (Chappuis, 1928b)
Canthocamptus crenulatus Mrázek, 1901
Attheyella crenulata (Mrázek, 1901)
Canthocamptus insignis Delachaux, 1918
Attheyella insignis (Delachaux, 1918)
Canthocamptus dentatus Poggenpol, 1874
Attheyella dentata (Poggenpol, 1874)
Attheyella (Ryloviella) baikalensis Borutzky, 1931d Attheyella baikalensis Borutzky, 1931d
Australocamptus hamondi Karanovic, 2004
Austrocletodes tricomatosum Pallares, 1979
Avdeevia antarctica Bresciani & Lützen, 1994
Balænophilus unisetus Aurivillius
Balaenophilus unisetus Aurivillius, 1879
Balucopsylla similis Rao, 1972
Bathycamptus eckmani Huys & Thistle, 1989
Bathycletopsyllus hexarthra Huys & Lee, 1999
Bathyesola compacta Huys & Lee, 2000
Bathyidia remota Farran, 1926
Bathylaophonte azorica Lee & Huys, 1999b
Psammis longifurca Bodin, 1968a
Bathypsammis longifurca (Bodin, 1968a)
Delavalia æmula T. Scott, 1893
Beatricella aemula (T. Scott, 1893)
Leptastacus dispinosus Mielke, 1982
Belemnopontia dispinosa (Mielke, 1982)
Ichnusella eione Cottarelli, 1971
Bereraia eione (Cottarelli, 1971)
Biameiropsis barrowi Karanovic, 2006
Singularia ingens Huys, 1995a
Biuncus ingens (Huys, 1995a)
Bolbotelos longisetosus Huys & Conroy-Dalton, 2006
Hemimesochra nympha Por, 1964b
Boreolimella nympha (Por, 1964b)
Boreopontia heipi Willems, 1981
Boreovermis bilobata Huys & Conroy-Dalton, 2006
Bradya typica Boeck, 1873
Bradyellopsis subniger Brian, 1925b
Brescianiana rotundata Avdeev, 1982
Arthropsyllus australis George, 1998a
Breviconia australis (George, 1998a)
Brevifrons faviolatum Harris, 1994
Brianella Stebleri Monard, 1926a
Brianola stebleri (Monard, 1926a)
Canthocamptus minutus Claus, 1863
Bryocamptus minutus (Claus, 1863)
Canthocamptus minutus Claus, 1863
Bryocamptus minutus (Claus, 1863)
Cyclopsine alpestris Vogt, 1845
Bryocamptus alpestris (Vogt, 1845)
Canthocamptus echinatus Mrázek, 1893
Bryocamptus echinatus (Mrázek, 1893)
Canthocamptus Zschokkei Schmeil, 1893
Bryocamptus zschokkei (Schmeil, 1893)
Bulbamphiascus imus (Brady, 1872)
Canthocamptus imus Brady, 1872
Caligopsyllus primus Kunz, 1975b
Type fixation
Notes
subsequent designation
subsequent designation
original designation
subsequent designation
subsequent designation
original designation
monotypy
original designation
monotypy
monotypy
original designation
original designation
original designation
original designation
original designation
monotypy
original designation
original designation
subsequent designation
original designation
original designation
original designation
original designation
original designation
original designation
original designation
original designation
indication
original designation
original designation
original designation
monotypy
monotypy
original designation
original designation
subsequent designation
original designation
original designation
original designation
monotypy
(15)
(15)
(16)
(16)
(17)
(18, 19)
(4)
(20)
(21)
(12, 22)
(4)
(4)
(4)
(4, 12)
TERMS OF USE
This pdf is provided by Magnolia Press for private/research use.
Commercial sale or deposition in a public library or website is prohibited.
74 · Zootaxa 2183 © 2009 Magnolia Press
Genus-group name (subgenera indented)
HUYS
Zootaxa 2183 © 2009 Magnolia Press ·
Type species (original binomen)
Type species (valid binomen)
Cancrincola Wilson, 1913
Canthocamptus Westwood, 1836
Canuella Scott & Scott, 1893c
Canuellina Gurney, 1927
Canuellopsis Lang, 1936a
Caribbula Huys & Gee, 1990
Carcinocaris Cottarelli, Bruno & Berrera, 2006
Carolinicola Huys & Thistle, 1989
Carraroenia McCormack, 2006
Ceratonotus Sars, 1909b
Cerconeotes Huys, 1992
Cervinia Norman in Brady, 1878
Cerviniella Smirnov, 1946
Cerviniopsis Sars, 1903
Ceuthonectes Chappuis, 1924b
Chappuisius Kiefer, 1938
Chaulionyx Kihara & Huys, 2009a
Chilaophonte Huys, 2009 mihi
Cholidya Farran, 1914
Cholidyella Avdeev, 1982
Ciplakastacus Sak, Karaytuğ & Huys, 2008
Cithadius Bowman, 1972
Cladorostrata Tai & Song, 1979
Clavigofera Harris & Iwasaki, 1996
Cletodes Brady, 1872
Cletopsyllus Willey, 1935
Clytemnestra Dana, 1847
Corallicletodes Soyer, 1966
Corbulaseta Huys & Lee, 2000
Cornylaophonte Willen, 1996
Coullana Por, 1984
Coullia Hamond, 1973a
Cubanocleta Petkovski, 1977
Cylindronannopus Coull, 1973a
Cylindropsyllus Brady, 1880b
Cylinula Coull, 1971a
Dactylopia Becker, 1974
Dactylopodamphiascopsis Lang, 1944
Dactylopodella Sars, 1905c
Dactylopodopsis Sars, 1911a
Cancrincola jamaicensis Wilson, 1913
Monoculus staphylinus Jurine, 1820
Canthocamptus staphylinus (Jurine, 1820)
Canuella perplexa Scott & Scott, 1893c
Canuellina insignis Gurney, 1927
Canuellopsis typica Lang, 1936a
Thompsonula hyaenae elongata Gee, 1988
Caribbula elongata (Gee, 1988)
Carcinocaris serrichelata Cottarelli, Bruno & Berera, 2006
Hemimesochra trisetosa Coull, 1973a
Carolinicola trisetosa (Coull, 1973a)
Carraroenia ruthae McCormack, 2006
Ceratonotus pectinatus Sars, 1909b
Leptastacus mozambicus Wells, 1967
Cerconeotes mozambicus (Wells, 1967)
Cervinia Bradyi Norman in Brady, 1878
Cervinia bradyi Norman in Brady, 1878
Cerviniella mirabilipes Smirnov, 1946
Cerviniopsis clavicornis Sars, 1903
Ceuthonectes serbicus Chappuis, 1924b
Chappuisius inopinus Kiefer, 1938
Chaulionyx paivacarvalhoi Kihara & Huys, 2009a
Chilaophonte maiquillahuensis Mielke, 1985
Chilaophonte maiquillahuensis (Mielke, 1985)
Cholidya polypi Farran, 1914
Cholydiella incisa Avdeev, 1982
Cholidyella incisa Avdeev, 1982
Ciplakastacus mersinensis Sak, Karaytuğ & Huys, 2008
Cithadius cyathurae Bowman, 1972
Cladorostrata brevipoda Shen & Tai, 1963
Cladorostrata brevipoda (Shen & Tai, 1963)
Clavigofera pacifica Harris & Iwasaki, 1996
Cletodes limicola Brady, 1872
Cletopsyllus papillifer Willey, 1935
Clytemnestra scutellata Dana, 1847
Corallicletodes boutierei Soyer, 1966
Laophonte bulligera Farran, 1913
Corbulaseta bulligera (Farran, 1913)
Cornylaophonte pleisteri Willen, 1996
Canuella canadensis Willey, 1923
Coullana canadensis (Willey, 1923)
Coullia heteropus Hamond, 1973a
Cubanocleta noodti Petkovski, 1977
Cylindronannopus primus Coull, 1973a
Cylindropsyllus lævis Brady, 1880b
Cylindropsyllus laevis Brady, 1880b
Cylinula proxima Coull, 1971a
Dactylopia peruana Becker, 1974
Amphiascus latifolius Sars, 1909c
Dactylopodamphiascopsis latifolius (Sars, 1909c)
Dactylopus flavus Claus, 1866
Dactylopodella flava (Claus, 1866)
Dactylopodopsis dilatata Sars, 1911a
Type fixation
original designation
monotypy
monotypy
monotypy
indication
original designation
original designation
original designation
monotypy
monotypy
original designation
monotypy
monotypy
subsequent designation
monotypy
monotypy
original designation
original designation
monotypy
original designation
original designation
original designation
original designation
original designation
monotypy
monotypy
monotypy
monotypy
original designation
original designation
monotypy
original designation
original designation
original designation
monotypy
original designation
monotypy
original designation
monotypy
monotypy
Notes
(23)
(12, 24)
(1, 5, 25)
(4)
(4)
(26)
(27)
(18, 28)
TERMS OF USE
This pdf is provided by Magnolia Press for private/research use.
Commercial sale or deposition in a public library or website is prohibited.
UNRESOLVED CASES IN HARPACTICOID COPEPOD NOMENCLATURE
Genus-group name (subgenera indented)
75
Type species (original binomen)
Type species (valid binomen)
Dactylopusia Norman, 1903a
Dactylopusioides Brian, 1928b
Dahlakocamptus Huys, 2009 mihi
Dahmsopottekina Özdikmen, 2009
Danielssenia Boeck, 1873
Darcythompsonia T. Scott, 1906
Delamarella Chappuis, 1954b
Delavalia Brady, 1869
Dendropsyllus Conroy-Dalton, 2003
Diagoniceps Willey, 1930
Diarthrodella Klie, 1949
Diarthrodes Thomson, 1883
Dilatatiocauda Harris, 2002
Diosaccopsis Brian, 1925a
Diosaccus Boeck, 1873
Discoharpacticus Noodt, 1954
Distioculus Huys & Böttger-Schnack, 1994
Dizahavia Por, 1979
Domnuia Willen, 2004
Donsiella Stephensen, 1936
Dorsiceratus Drzycimski, 1967b
Drescheriella Dahms & Dieckmann, 1987
Dyacrenhydrosoma Gee, 1999
Echinocletodes Lang, 1936b
Echinolaophonte Nicholls, 1941b
Echinopsyllus Sars, 1909b
Echinosunaristes Huys, 1995b
Ectinosoma Boeck, 1865
Ectinosomella Sars, 1910
Ectinosomoides Nicholls, 1945b
Elanella Por, 1984
Elaphoidella Chappuis, 1929a
Elapholaophonte Schizas & Shirley, 1994b
Ellucana Sewell, 1940
Emertonia Wilson, 1932
Enhydrosoma Boeck, 1873
Enhydrosomella Monard, 1935b
Eoschizopera Wells & Rao, 1976
Epactophanes Mrázek, 1893
Epactophanoides Borutzky, 1966
Esola Edwards, 1891
Dactylopus tisboides Claus, 1863
Dactylopusia tisboides (Claus, 1863)
Dactylopusioides stampaliae Brian, 1928b
Dactylopusioides macrolabris (Claus, 1866)
Cletocamptus xenuus Por, 1968
Dahlakia xenuus (Por, 1968)
Metahuntemannia curticauda Becker, 1979
Dahmsopottekina curticauda (Becker, 1979)
Danielssenia typica Boeck, 1873
Cylindropsyllus fairliensis T. Scott, 1899b
Darcythompsonia fairliensis (T. Scott, 1899b)
Delamarella arenicola Chappuis, 1954b
Delavalia palustris Brady, 1869
Dendropsyllus thomasi Conroy-Dalton, 2003
Diagoniceps lævis Willey, 1930
Diagoniceps laevis Willey, 1930
Diarthrodella orbiculata Klie, 1949
Diarthrodes novæ-zealandiæ Thomson, 1883
Diarthrodes novaezealandiae Thomson, 1883
Porcellidium dilatatum Hicks, 1971
Dilatatiocauda dilatata (Hicks, 1971)
Diosaccopsis rubeus Brian, 1925a
Dactylopus tenuicornis Claus, 1863
Diosaccus tenuicornis (Claus, 1863)
Discoharpacticus mirabilis Noodt, 1954
Miracia minor T. Scott, 1894a
Distioculus minor (T. Scott, 1894a)
Dizahavia halophila Por, 1979
Domnuia larsi Willen, 2004
Donsiella limnoriae Stephensen, 1936
Dorsiceratus octocornis Drzycimski, 1967b
Drescheriella glacialis Dahms & Dieckmann, 1987
Dyacrenhydrosoma breviseta Gee, 1999
Cletodes armata T. Scott, 1903b
Echinocletodes armatus (T. Scott, 1903b)
Cleta horrida Norman, 1876
Echinolaophonte horrida (Norman, 1876)
Echinopsyllus Normani Sars, 1909b
Echinopsyllus normani Sars, 1909b
Echinosunaristes bathyalis Huys, 1995b
Ectinosoma melaniceps Boeck, 1865
Ectinosomella nitidula Sars, 1910
Ectinosomoides longipes Nicholls, 1945b
Canuella elanitica Por, 1967
Elanella elanitica (Por, 1967)
Canthocamptus elaphoides Chappuis, 1924b
Elaphoidella elaphoides (Chappuis, 1924b)
Elapholaophonte decaceros Schizas & Shirley, 1994b
Canuella (Ellucana) longicauda Sewell, 1940
Ellucana longicauda (Sewell, 1940)
Emertonia gracilis Wilson, 1932
Enhydrosoma curticauda Boeck, 1873
Enhydrosomella staufferi Monard, 1935b
Schizopera syltensis Mielke, 1973
Eoschizopera syltensis (Mielke, 1973)
Epactophanes richardi Mrázek, 1893
Epactophanoides udegeicus Borutzky, 1966
Esola longicauda Edwards, 1891
Type fixation
Notes
designation by ICZN
monotypy
original designation
original designation
indication
original designation
monotypy
monotypy
original designation
monotypy
original designation
monotypy
original designation
subsequent designation
subsequent designation
original designation
original designation
original designation
original designation
monotypy
monotypy
original designation
original designation
original designation
original designation
monotypy
original designation
monotypy
monotypy
monotypy
monotypy
original designation
original designation
monotypy
original designation
subsequent designation
monotypy
original designation
monotypy
monotypy
monotypy
(29)
(30)
(4)
(115)
(31)
(32)
(18)
(18, 33)
(5, 34)
(35)
(36)
(37)
(4)
(38)
TERMS OF USE
This pdf is provided by Magnolia Press for private/research use.
Commercial sale or deposition in a public library or website is prohibited.
76 · Zootaxa 2183 © 2009 Magnolia Press
Genus-group name (subgenera indented)
HUYS
Zootaxa 2183 © 2009 Magnolia Press ·
Type species (original binomen)
Eucanuella T. Scott, 1901
Eudactylopus A. Scott, 1909
Eucanuella spinifera T. Scott, 1901
Dactylopus latipes T. Scott, 1894a nec Boeck
Eudactylopus andrewi Sewell, 1940
(1865)
Eupelte gracilis Claus, 1860
Cletodes laticauda Boeck, 1873
Eurycletodes laticauda (Boeck, 1873)
Cletodes laticauda Boeck, 1873
Eurycletodes laticauda (Boeck, 1873)
Cletodes lata T. Scott, 1892
Eurycletodes latus (T. Scott, 1892)
Euterpe gracilis Claus, 1863
Euterpina acutifrons (Dana, 1847)
Tetragoniceps incertus T. Scott, 1892
Evansula incerta (T. Scott, 1892)
Expansicervinia glacieria Montagna, 1981
Feregastes wellensi Fiers, 1986a
Fibulacamptus victorianus Hamond, 1988
Ameira longicaudata Nicholls, 1939a
Filexilia trisetosa Conroy-Dalton & Huys, 1997
Danielssenia robusta Sars, 1921
Fladenia robusta (Sars, 1921)
Folioquinpes mangalis Fiers & Rutledge, 1990
Parastenocaris forficata Noodt, 1963
Forficatocaris forficata (Noodt, 1963)
Parastenhelia anglica Norman & Scott, 1905
Foweya anglica (Norman & Scott, 1905)
Fultonia hirsuta T. Scott, 1902
Galapacanuella beckeri Mielke, 1979
Galapalaophonte pacifica Mielke, 1981
Tachidius incisipes Klie, 1913
Geeopsis incisipes (Klie, 1913)
Genesis vulcanoctopusi López-González, Bresciani
& Huys in López-González et al., 2000
Gideonia noncavernicola George & Martínez Arbizu, 2005
Ameira bengalensis Rao & Ganapati, 1969
Glabrameira bengalensis (Rao & Ganapati, 1969)
Hastigerella mehuinensis Mielke, 1986
Glabrotelson mehuinense (Mielke, 1986)
Glaciella yalensis Kikuchi, 1994
Godianiceps mayae Fiers, 1995
Goffinella stylifer Wilson, 1932
Goniopsyllus rostratus Brady, 1883
Gulcamptus uenoi Miura, 1969
Haifameira archibenthoica Por, 1964a
Ectinosoma sarsi Boeck, 1873
Halectinosoma sarsi (Boeck, 1873)
Alophytophilus fusiformis Brian, 1919
Halophytophilus fusiformis Brian, 1919
Stenhelia pygmæa Norman & Scott, 1905
Haloschizopera pygmaea (Norman & Scott, 1905)
Hamondia superba Huys, 1990b
Hemimesochra derketo Por, 1964a
Hanikraia derketo (Por, 1964a)
Harpacticella inopinata Sars, 1908c
Cyclops chelifer Müller, 1776
Harpacticus chelifer (Müller, 1776)
Eupelte Claus, 1860
Eurycletodes Sars, 1909a
Eurycletodes Sars, 1909a
Oligocletodes Lang, 1944
Euterpina Norman, 1903a
Evansula T. Scott, 1906b
Expansicervinia Montagna, 1981
Feregastes Fiers, 1986a
Fibulacamptus Hamond, 1988
Filexilia Conroy-Dalton & Huys, 1997
Fladenia Gee & Huys, 1990
Folioquinpes Fiers & Rutledge, 1990
Forficatocaris Jakobi, 1969
Foweya Gee, 2006
Fultonia T. Scott, 1902
Galapacanuella Mielke, 1979
Galapalaophonte Mielke, 1981
Geeopsis Huys in Huys et al., 1996
Genesis López-González, Bresciani & Huys
in López-González et al., 2000
Gideonia George & Martínez Arbizu, 2005
Glabrameira Conroy-Dalton & Huys, 1997
Glabrotelson Huys, 2009 mihi
Glaciella Kikuchi, 1994
Godianiceps Fiers, 1995
Goffinella Wilson, 1932
Goniopsyllus Brady, 1883
Gulcamptus Miura, 1969
Haifameira Por, 1964a
Halectinosoma Vervoort, 1962
Halophytophilus Brian, 1919
Haloschizopera Lang, 1944
Hamondia Huys, 1990b
Hanikraia Huys, 2009 mihi
Harpacticella Sars, 1908c
Harpacticus Milne-Edwards, 1840
Type species (valid binomen)
Type fixation
Notes
monotypy
original designation
(39)
monotypy
subsequent designation
subsequent designation
original designation
subsequent monotypy
original designation
original designation
original designation
original designation
original designation
original designation
original designation
original designation
original designation
monotypy
monotypy
original designation
monotypy
original designation
original designation
original designation
original designation
original designation
original designation
original designation
monotypy
original designation
monotypy
original designation
monotypy
original designation
original designation
original designation
monotypy
subsequent designation
(40)
(40)
(41)
(42)
(43)
(44)
(4)
(4)
(45)
(4)
(46)
TERMS OF USE
This pdf is provided by Magnolia Press for private/research use.
Commercial sale or deposition in a public library or website is prohibited.
UNRESOLVED CASES IN HARPACTICOID COPEPOD NOMENCLATURE
Genus-group name (subgenera indented)
77
Type species (original binomen)
Type species (valid binomen)
Harrietella T. Scott, 1906a
Helmutkunzia Wells & Rao, 1976
Hemicervinia Lang, 1935
Hemicletodes Lang, 1936a
Hemilaophonte Jakubisiak, 1932
Hemimesochra Sars, 1920b
Herdmaniopsis Brotskaya, 1963
Heterolaophonte Lang, 1948
Heteronychocamptus Lee & Huys, 1999b
Heteropsyllus T. Scott, 1894b
Hirtaleptomesochra Karanovic, 2004
Hoplolaophonte Hamond, 1973a
Huntemannia Poppe, 1884
Hypalocletodes Por, 1967
Hypocamptus Chappuis, 1929b
Ialysus Brian, 1927b
Idyanthe Sars, 1909c
Idyella Sars, 1905b
Idyellopsis Lang, 1948
Ifanella Vervoort, 1964
Indolaophonte Cottarelli, Saporito & Puccetti, 1986a
Inermipes Lee & Huys, 2002
Inermiphonte Huys & Lee, 2009
Intercanuella Becker & Schriever, 1979
Intercletodes Fiers, 1987
Intercrusia Huys, 1996b
Intersunaristes Huys, 1995b
Isocletopsyllus Huys & Lee, 1999
Isthmiocaris George & Schminke, 2003
Itunella Brady, 1896
Jamstecia Lee & Huys, 2000
Jonesiella Brady, 1880a
Juxtaramia Conroy-Dalton & Huys, 2000
Karllangia Noodt, 1964
Keraia Willen & Dittmar, 2009
Kinnecaris Jakobi, 1972
Klieonychocamptoides Noodt, 1958
Klieonychocamptus Noodt, 1958
Klieosoma Hicks & Schriever, 1985
Kollerua Gee, 1994
(?) Laophonte simulans T. Scott, 1894b
Harrietella simulans (T. Scott, 1894b)
Actopsyllus hartmannorum Kunz, 1971
Helmutkunzia hartmannorum (Kunz, 1971)
Herdmania stylifera Thompson, 1893
Hemicervinia stylifera (Thompson, 1893)
Hemicletodes typicus Lang, 1936a
Hemilaophonte Janinæ Jakubisiak, 1932
Hemilaophonte janinae Jakubisiak, 1932
Hemimesochra clavularis Sars, 1920b
Herdmaniopsis abyssicola Brotskaya, 1963
Cyclops Stromii Baird, 1837
Heterolaophonte stroemii (Baird, 1837)
Paronychocamptus connexus Pallares, 1979
Heteronychocamptus connexus (Pallares, 1979)
Heteropsyllus curticaudatus T. Scott, 1894b
Hirtaleptomesochra bispinosa Karanovic, 2004
Hoplolaophonte aculeata Hamond, 1973a
Huntemannia jadensis Poppe, 1884
Hypalocletodes salomonis Por, 1967
Maraenobiotus brehmi Douwe, 1922
Hypocamptus brehmi (Douwe, 1922)
Ialysus rufus Brian, 1927b
Idyopsis dilatata Sars, 1905b
Idyanthe dilatata (Sars, 1905b)
Idyella pallidula Sars, 1905b
Idyellopsis typica Lang, 1948
Ifanella chacei Vervoort, 1964
Indolaophonte ramai Cottarelli, Saporito & Puccetti, 1986a
Inermipes humphreysi Lee & Huys, 2002
Laophonte danversae Hamond, 1969
Inermiphonte danversae (Hamond, 1969)
Intercanuella lima Becker & Schriever, 1979
Intercletodes interita Fiers, 1987
Intercrusia problematica Huys, 1996b
Sunaristes dardani Humes & Ho, 1969a
Intersunaristes dardani (Humes & Ho, 1969a)
Cletopsyllus tertius Por, 1964a
Isocletopsyllus tertius (Por, 1964a)
Isthmiocaris longitelson George & Schminke, 2003
? Cletodes tenuiremis T. Scott, 1893
Itunella tenuiremis (T. Scott, 1893)
Jamstecia terazakii Lee & Huys, 2000
Jonesiella fusiformis Brady, 1880a
Juxtaramia polaris Conroy-Dalton & Huys, 2000
Karllangia arenicola Noodt, 1964
Pseudomesochra tamara Smirnov, 1946
Keraia tamara (Smirnov, 1946)
Parastenocaris forciculata Chappuis, 1953
Kinnecaris forficulata (Chappuis, 1953)
Klieonychocamptoides remanei Noodt, 1958
Laophonte kliei Monard, 1935a
Klieonychocamptus kliei (Monard, 1935a)
Halophytophilus ? triarticulatus Klie, 1949
Klieosoma triarticulatum (Klie, 1949)
Enhydrosoma radhakrishnai Ranga Reddy, 1979 Kollerua radhakrishnai (Ranga Reddy, 1979)
Type fixation
original designation
original designation
monotypy
indication
monotypy
monotypy
original designation
original designation
original designation
monotypy
original designation
monotypy
monotypy
monotypy
original designation
monotypy
subsequent designation
subsequent designation
indication
original designation
monotypy
original designation
original designation
original designation
original designation
original designation
original designation
original designation
original designation
monotypy
original designation
subsequent designation
original designation
original designation
original designation
original designation
original designation
original designation
original designation
original designation
Notes
(47)
(12, 18)
(4, 12, 48)
(49)
(1, 50)
(1, 51)
(4)
(52)
(53)
(54)
TERMS OF USE
This pdf is provided by Magnolia Press for private/research use.
Commercial sale or deposition in a public library or website is prohibited.
78 · Zootaxa 2183 © 2009 Magnolia Press
Genus-group name (subgenera indented)
HUYS
Zootaxa 2183 © 2009 Magnolia Press ·
Type species (original binomen)
Kristensenia Por, 1983b
Kunzia Wells, 1967
Kushia Harris & Iwasaki, 1996
Laophonte Philippi, 1840
Laophontina Norman & Scott, 1905
Laophontisochra George, 2002
Laophontodes T. Scott, 1894b
Laophontopsis Sars, 1908b
Laubieria Soyer, 1966
Leptameira Huys, 2009 mihi
Leptastacus T. Scott, 1906a
Leptocaris T. Scott, 1899b
Leptocletodes Sars, 1920c
Leptomesochra Sars, 1911b
Leptopontia T. Scott, 1902
Leptopsyllus T. Scott, 1894b
Leptopsyllus T. Scott, 1894b
Paraleptopsyllus Lang, 1944
Leptotachidia Becker, 1974
Lessinocamptus Stoch, 1997
Ligulocamptus Guo, 1998
Limameira Soyer, 1975b
Limnocletodes Borutzky, 1926
Lipomelum Fiers, 1986b
Lobitella Monard, 1934
Lobopleura Conroy-Dalton, 2004
Loefflerella Rouch, 1962
Longipedia Claus, 1862
Loureirophonte Jakobi, 1953
Lourinia Wilson, 1924
Lucayostratiotes Huys, 1990b
Macrosetella A. Scott, 1909
Malacopsyllus Sars, 1911b
Maiquilaophonte Mielke, 1985
Maraenobiotus Mrázek, 1893
Marbefia Huys & Lee, 2009
Marionobiotus Chappuis, 1940b
Marsteinia Drzycimski, 1968
Megistocletodes Por, 1986b
Meiopsyllus Cottarelli & Forniz, 1995
Kristensenia pallida Por, 1983b
Kunzia epacra Wells, 1967
Kushia zosteraphila Harris & Iwasaki, 1996
Laophonte cornuta Philippi, 1840
Laophontina dubia Norman & Scott, 1905
Laophontisochra maryamae George, 2002
Laophonte (Laophontodes) typicus T. Scott, 1894b
Cleta lamellifera Claus, 1863
Laubieria corallicola Soyer, 1966
Leptomesochra confluens Sars, 1911c
Tetragoniceps macronyx T. Scott, 1892
Leptocaris minutus T. Scott, 1899b
Leptocletodes debilis Sars, 1920c
Normanella attenuata A. Scott, 1896
Leptopontia curvicauda T. Scott, 1902
Leptopsyllus typicus T. Scott, 1894b
Leptopsyllus typicus T. Scott, 1894b
Paramesochra arctica Lang, 1936a
Leptotachidia iberica Becker, 1974
Lessinocamptus caoduroi Stoch, 1997
Ligulocamptus loffleri Guo, 1998
Limameira mediterranea Soyer, 1975b
Limnocletodes behningi Borutzky, 1926
Lipomelum heteromelum Fiers, 1986b
Lobitella apoda Monard, 1934
Lobopleura ambiducti Conroy-Dalton, 2004
Löfflerella dentata Rouch, 1962
Longipedia coronata Claus, 1863
Loureirophonte Catharinensis Jakobi, 1953
Jurinia armata Claus, 1866
Rhynchothalestris cornuta Geddes, 1969
Setella gracilis Dana, 1847
Malacopsyllus fragilis Sars, 1911b
Maiquilaophonte uachi Mielke, 1985
Maraenobiotus vejdovskýi Mrázek, 1893
Paronychocamptus carthyi Hamond, 1968
Marionobiotus Jeanneli Chappuis, 1940b
Marsteinia typica Drzycimski, 1968
Megistocletodes translucens Por, 1986b
Meiopsyllus marinae Cottarelli & Forniz, 1995
Type species (valid binomen)
Laophontodes typicus (T. Scott, 1894b)
Laophontopsis lamellifera (Claus, 1863)
Leptameira confluens (Sars, 1911c)
Leptastacus macronyx (T. Scott, 1892)
Leptocaris minuta T. Scott, 1899b
Leptomesochra attenuata (A. Scott, 1896)
Leptopsyllus arcticus (Lang, 1936a)
Loefflerella dentata Rouch, 1962
Loureirophonte catharinensis Jakobi, 1953
Lourinia armata (Claus, 1866)
Lucayostratiotes cornuta (Geddes, 1969)
Macrosetella gracilis (Dana, 1847)
Maraenobiotus vejdovskyi Mrázek, 1893
Lamelliphonte carthyi (Hamond, 1968)
Marionobiotus jeanneli Chappuis, 1940b
Type fixation
monotypy
original designation
original designation
monotypy
monotypy
original designation
indication
monotypy
monotypy
monotypy
original designation
monotypy
monotypy
subsequent designation
monotypy
indication
indication
original designation
original designation
original designation
original designation
monotypy
monotypy
original designation
monotypy
original designation
monotypy
subsequent monotypy
original designation
monotypy
original designation
subsequent designation
monotypy
monotypy
monotypy
original designation
monotypy
indication
original designation
monotypy
Notes
(55)
(56)
(57)
(4)
(58)
(1, 4)
(18, 59)
(60)
(12)
(61)
(62)
(18)
(12)
(63)
TERMS OF USE
This pdf is provided by Magnolia Press for private/research use.
Commercial sale or deposition in a public library or website is prohibited.
UNRESOLVED CASES IN HARPACTICOID COPEPOD NOMENCLATURE
Genus-group name (subgenera indented)
79
Type species (original binomen)
Type species (valid binomen)
Type fixation
Melima Por, 1964a
Meloriastacus Huys & Todaro, 1997
Membranastacus Huys, 1992
Mesochra Boeck, 1865
Mesocletodes Sars, 1909a
Mesopontia Sak, Huys & Karaytuğ, 2008
Mesopsyllus Por, 1960b
Metahuntemannia Smirnov, 1946
Metamphiascopsis Lang, 1944
Meteorina George, 2004b
Metis Philippi, 1843
Mexicolaophonte Cottarelli, 1977
Microarthridion Lang, 1944
Microcanuella Mielke, 1994a
Microchelonia Brady, 1918
Microlaophonte Vervoort, 1964
Micropsammis Gee & Huys, 1991
Microsetella Brady & Robertson, 1873
Mictyricola Nicholls, 1957
Mielkiella George, 1997
Minervella Cottarelli & Venanzetti, 1989
Miracia Dana, 1846
Miroslavia Apostolov, 1980
Miscegenus Wells, Hicks & Coull, 1982
Monardius Huys, 2009 mihi
Monocletodes Lang, 1936b
Monodicaris Schminke, 2009
Moraria Scott & Scott, 1893a
Morariopsis Borutzky, 1931b
Mourephonte Jakobi, 1953
Mucropedia Bouck, Thistle & Huys, 1999
Mucrosenia Gee & Huys, 1994
Muohuysia Özdikmen, 2009
Murunducaris Reid, 1994
Mwania Fiers & De Troch, 2000
Nannomesochra Gurney, 1932
Nannopus Brady, 1880a
Nathaniella Por, 1984
Navalonia Huys & Conroy-Dalton, 1994
Nematovorax Bröhldick, 2005
Neoacrenhydrosoma Gee & Mu, 2000
Melima caulerpae Por, 1964a
original designation
Meloriastacus ctenidis Huys & Todaro, 1997
original designation
Membranastacus inopinatus Huys, 1992
original designation
Mesochra Liljeborgii Boeck, 1865
Mesochra lilljeborgii Boeck, 1865*
subsequent designation
Cletodes irrasa Scott & Scott, 1894
Mesocletodes irrasus (Scott & Scott, 1894)
original designation
Arenopontia dillonbeachia Lang, 1965
Mesopontia dillonbeachia (Lang, 1965)
original designation
Mesopsyllus atargatis Por, 1960b
monotypy
Metahuntemannia gorbunovi Smirnov, 1946
monotypy
Dactylophusia hirsuta Thompson & Scott, 1903
Metamphiascopsis hirsutus (Thompson & Scott, 1903) original designation
Meteorina magnifica George, 2004b
monotypy
Metis ignea Philippi, 1843
monotypy
Mexicolaophonte arganoi Cottarelli, 1977
monotypy
Tachidius littoralis Poppe, 1881
Microarthridion littorale (Poppe, 1881)
original designation
Microcanuella bisetosa Mielke, 1994a
original designation
Microchelonia glacialis Brady, 1918
monotypy
Microlaophonte spongicola Vervoort, 1964
original designation
Micropsammis noodti Mielke, 1975
Micropsammis noodti (Mielke, 1975)
original designation
Microsetella atlantica Brady & Robertson, 1873
Microsetella norvegica (Boeck, 1865)
monotypy
Myctyricola typica Nicholls, 1957
Mictyricola typica Nicholls, 1957
indication
Mielkiella spinulosa George, 1997
monotypy
monotypy
Minervella baccettii Cottarelli & Venanzetti, 1989
Miracia efferata Dana, 1849
subsequent designation
Miroslavia longicaudata Apostolov, 1980
original designation
Miscegenus heretaunga Wells, Hicks & Coull, 1982
original designation
Teissierella salammboi Monard, 1935b
Monardius salammboi (Monard, 1935b)
original designation
Cletodes varians T. Scott, 1903a
Monocletodes varians (T. Scott, 1903a)
monotypy
Monodicaris larsi Schminke, 2009
original designation
Moraria Anderson-Smithi Scott & Scott, 1903b
Moraria brevipes (Sars, 1863)
monotypy
Morariopsis typica Borutzky, 1931b
indication
Mourephonte catharinensis Jakobi, 1953
Mourephonte longiseta (Nicholls, 1941a)
original designation
Mucropedia cookorum Bouck, Thistle & Huys, 1999
original designation
Mucrosenia kendalli Gee & Huys, 1994
original designation
Stenhelia xylophila Hicks, 1988a
Muohuysia xylophila (Hicks, 1988a)
original designation
Murunducaris juneae Reid, 1994
monotypy
Mwania phytocola Fiers & De Troch, 2000
original designation
Pseudomesochra parvula Gurney, 1927
Nannomesochra arupinensis (Brian, 1925a)
monotypy
Nannopus palustris Brady, 1880a
monotypy
Canuella reichi Por, 1964a
Nathaniella reichi (Por, 1964a)
monotypy
Stenocaris kerguelenensis Bodiou, 1977
Navalonia kerguelenensis (Bodiou, 1977)
original designation
Nematovorax gebkelinae Bröhldick, 2005
original designation
Neoacrenhydrosoma zhangi Gee & Mu, 2000
original designation
Notes
(12, 64)
(65)
(4)
(4)
(66)
(5, 67)
(4)
(68)
(69)
(70)
(116)
(71)
TERMS OF USE
This pdf is provided by Magnolia Press for private/research use.
Commercial sale or deposition in a public library or website is prohibited.
80 · Zootaxa 2183 © 2009 Magnolia Press
Genus-group name (subgenera indented)
HUYS
Zootaxa 2183 © 2009 Magnolia Press ·
Type species (original binomen)
Neoargestes Drzycimski, 1967a
Neobradya T. Scott, 1892
Neocancrincola Mañé-Garzón & Sobota, 1974
Neodactylopus Nicholls, 1945a
Neoechiniphora Huys, 1996b
Neogoniceps Fiers & De Troch, 2000
Neoleptastacus Nicholls, 1945b
Neomiscegenus Karanovic & Ranga Reddy, 2004
Neonitocrella Lee & Huys, 2002
Neopeltopsis Hicks, 1976
Neopsammastacus Cottarelli & Venanzetti, 1989
Neoscutellidium Zwerner, 1967
Neotachidius Shen & Tai, 1963
Neoargestes variabilis Drzycimski, 1967a
Neobradya pectinifer T. Scott, 1892
Neobradya pectinifera T. Scott, 1892
Neocancrincola platensis Mañé-Garzón & Sobota, 1974
Neodactylopus cyclopoides Nicholls, 1945a
Neoechinophora fosshageni Huys, 1996b
Neogoniceps martinezi Fiers & De Troch, 2000
Neoleptastacus spinicaudatus Nicholls, 1945b
Neomiscegenus indicus Karanovic & Ranga Reddy, 2004
Nitocrella insularis Miura, 1962
Neonitocrella insularis (Miura, 1962)
Neopeltopsis pectinipes Hicks, 1976
Psammastacus spinicaudus Wells, 1967
Neopsammastacus spinicauda (Wells, 1967)
Neoscutellidium yeatmani Zwerner, 1967
Tachidius (Neotachidius) triangularis Shen & Tai, Neotachidius triangularis (Shen & Tai, 1963)
1963
Neotisbella gigas Boxshall, 1979
Zausodes areolatus Geddes, 1968a
Neozausodes areolatus (Geddes, 1968a)
Nitokra typica Boeck, 1865
Nitocra typica Boeck, 1865
Nitocrella hirta Chappuis, 1924b
Nitocrellopsis rouchi Galassi, De Laurentiis & Dole-Olivier, 1999
Sigmatidium ? arenosetelloides Noodt, 1958
Noodtiella arenosetelloides (Noodt, 1958)
Noodtorthopsyllus psammophilus (Noodt, 1955)
Orthopsyllus psammophilus Noodt, 1955
Normanella dubia Brady, 1880a
Notopontia stephanieae Bodiou, 1977
Novanitocrella aboriginesi Karanovic, 2004
Novocrinia trifida Huys & Iliffe, 1998
Novolaophonte viatorum Cottarelli, Saporito & Puccetti, 1983
Nudivorax todai Lee & Huys, 2000
Nyxis rostrocularis Willen, 2009
Octopinella tenacis Avdeev, 1986
Octopinella tenax Avdeev, 1986
Miracia gracilis Dana, 1849
Oculosetella gracilis (Dana, 1849)
Odaginiceps clarkae Fiers, 1995
Odiliacletodes gracilis Soyer, 1965
Oikopus rostrilabrus Wells, 1967
Oligoxylora cooksoni Hicks, 1988b
Oniscopsis Pauliani Chappuis, 1955
Oniscopsis pauliani Chappuis, 1955
Onychocamptus heteropus Blanchard &
Onychocamptus mohammed (Blanchard &
Richard, 1891
Richard, 1891)
Onychopontia nichollsi Sak, Huys & Karaytuğ, 2008
Onychostenhelia falcifera Itô, 1979b
Lilljeborgia linearis Claus, 1866
Orthopsyllus linearis (Claus, 1866)
Neotisbella Boxshall, 1979
Neozausodes Bouck, Thistle & Huys, 1999
Nitocra Boeck, 1865
Nitocrella Chappuis, 1924b
Nitocrellopsis Galassi, De Laurentiis & Dole-Olivier, 1999
Noodtiella Wells, 1965b
Noodtorthopsyllus Lang, 1965
Normanella Brady, 1880a
Notopontia Bodiou, 1977
Novanitocrella Karanovic, 2004
Novocrinia Huys & Iliffe, 1998
Novolaophonte Cottarelli, Saporito & Puccetti, 1983
Nudivorax Lee & Huys, 2000
Nyxis Willen, 2009
Octopinella Avdeev, 1986
Oculosetella Dahl, 1895
Odaginiceps Fiers, 1995
Odiliacletodes Soyer, 1965
Oikopus Wells, 1967
Oligoxylora Hicks, 1988b
Oniscopsis Chappuis, 1955
Onychocamptus Daday, 1903
Onychopontia Sak, Huys & Karaytuğ, 2008
Onychostenhelia Itô, 1979b
Orthopsyllus Brady & Robertson, 1873
Type species (valid binomen)
Type fixation
monotypy
monotypy
original designation
monotypy
original designation
original designation
monotypy
original designation
original designation
original designation
original designation
original designation
monotypy
original designation
original designation
indication
monotypy
original designation
original designation
monotypy
monotypy
monotypy
original designation
original designation
monotypy
original designation
original designation
original designation
monotypy
original designation
monotypy
original designation
monotypy
monotypy
monotypy
original designation
original designation
monotypy
Notes
(72)
(73)
(74)
(75)
(76)
(77)
(12)
(78)
(79)
TERMS OF USE
This pdf is provided by Magnolia Press for private/research use.
Commercial sale or deposition in a public library or website is prohibited.
UNRESOLVED CASES IN HARPACTICOID COPEPOD NOMENCLATURE
Genus-group name (subgenera indented)
81
Type species (original binomen)
Pabellonia Gómez, 2000
Parabradya Lang, 1944
Paracerviniella Brotskaya, 1963
Paracrenhydrosoma Gee, 1999
Pabellonia olganoguerae Gómez, 2000
Bradya confluens Lang, 1936a
Parabradya confluens (Lang, 1936a)
Paracerviniella denticulata Brotskaya, 1963
Acrenhydrosoma maccalli Schizas & Shirley, 1994a Paracrenhydrosoma maccalli (Schizas & Shirley,
1994a)
Dactylopus latipes Boeck, 1865
Paradactylopodia latipes (Boeck, 1865)
Paradanielssenia kunzi Soyer, 1970b
Paraforficatocaris paranaensis Jakobi, 1972
Paraidya major Sewell, 1940
Paraidya major (Sewell, 1940)
Cleta brevirostris Claus, 1863
Paralaophonte brevirostris (Claus, 1863)
Laophonte echinata Willey, 1930
Paralaophontodes echinatus (Willey, 1930)
Paraleptastacus brevicaudatus Wilson, 1932
Paraleptomesochra minima Wells, 1967
Parameiropsis peruanus Becker, 1974
Paramenophia chilensis Lang, 1954
Paramesochra dubia T. Scott , 1892
Paramorariopsis anae Brancelj, 1991
Stenhelia hispida Brady, 1880a
Paramphiascella hispida (Brady, 1880a)
Paramphiascoides mixtus Wells, 1967
Paranaiara inajae Kihara & Huys, 2009b
Nannopus abyssi Sars, 1920c
Paranannopus abyssi (Sars, 1920c)
Paranitocrella bastiani Tang & Knott, 2009
Parapeltidium johnstoni A. Scott, 1909
Ameira trisetosa Krishnaswamy, 1957
Parapseudoleptomesochra trisetosa (Krishnaswamy,
1957)
Argestes tenuis Sars, 1921
Parargestes tenuis (Sars, 1921)
Dactylopus abyssi Boeck, 1873
Pararobertsonia abyssi (Boeck, 1873)
Paraschizopera beckeri Wells, 1981
Parasewellina prima Cottarelli, Saporito & Puccetti, 1986b
Parastenhelia hornelli Thompson & Scott, 1903
Parastenocaris brevipes Kessler, 1913
Parasunaristes cucullaris Fiers, 1982
Amymone sphaerica Claus, 1863
Parategastes sphaericus (Claus, 1863)
Thalestris Clausii Norman, 1869
Parathalestris clausii (Norman, 1869)*
Paratigriopus hoshidei Itô, 1969
Parbatocamptus jochenmartensi Dumont & Maas, 1988
Parevansula mediterranea Guille & Soyer, 1966
Tydemanella robusta Nicholls, 1941a
Parialysus robustus (Nicholls, 1941a)
Laophonte curticaudata Boeck, 1865
Paronychocamptus curticaudatus (Boeck, 1865)
Patagoniaella vervoorti Pallares, 1968c
Paradactylopodia Lang, 1944
Paradanielssenia Soyer, 1970b
Paraforficatocaris Jakobi, 1972
Paraidya Huys, 2009 mihi
Paralaophonte Lang, 1948
Paralaophontodes Lang, 1965
Paraleptastacus Wilson, 1932
Paraleptomesochra Wells, 1967
Parameiropsis Becker, 1974
Paramenophia Lang, 1954
Paramesochra T. Scott, 1892
Paramorariopsis Brancelj, 1991
Paramphiascella Lang, 1944
Paramphiascoides Wells, 1967
Paranaiara Kihara & Huys, 2009b
Paranannopus Lang, 1948
Paranitocrella Tang & Knott, 2009
Parapeltidium A. Scott, 1909
Parapseudoleptomesochra Lang, 1965
Parargestes Lang, 1944
Pararobertsonia Lang, 1944
Paraschizopera Wells, 1981
Parasewellina Cottarelli, Saporito & Puccetti, 1986b
Parastenhelia Thompson & Scott, 1903
Parastenocaris Kessler, 1913
Parasunaristes Fiers, 1982
Parategastes Sars, 1904b
Parathalestris Brady & Robertson, 1873
Paratigriopus Itô, 1969
Parbatocamptus Dumont & Maas, 1988
Parevansula Guille & Soyer, 1966
Parialysus Nicholls, 1941b
Paronychocamptus Lang, 1948
Patagoniaella Pallares, 1968c
Type species (valid binomen)
Type fixation
original designation
original designation
original designation
original designation
original designation
monotypy
original designation
original designation
original designation
monotypy
original designation
original designation
original designation
original designation
monotypy
original designation
original designation
original designation
original designation
original designation
original designation
monotypy
original designation
original designation
original designation
original designation
original designation
subsequent designation
monotypy
original designation
original designation
monotypy
original designation
original designation
monotypy
monotypy
original designation
original designation
Notes
(80)
(4)
(1, 4)
(1, 4)
(81)
(82)
(12)
TERMS OF USE
This pdf is provided by Magnolia Press for private/research use.
Commercial sale or deposition in a public library or website is prohibited.
82 · Zootaxa 2183 © 2009 Magnolia Press
Genus-group name (subgenera indented)
HUYS
Zootaxa 2183 © 2009 Magnolia Press ·
Type species (original binomen)
Type species (valid binomen)
Peltidiphonte Gheerardyn & Fiers in Gheerardyn et al., 2006a
Peltidium Philippi, 1839
Peltisenia Huys & Gee, 1996a
Peltthestris Monard, 1924
Peltobradya Médioni & Soyer, 1968
Peresime Dinet, 1974
Perissocope Brady, 1910
Perucamptus Huys & Thistle, 1989
Pesceus Özdikmen, 2008
Pholenota Vervoort, 1964
Pholetiscus Humes, 1947
Phyllognathopus Mrázek, 1893
Phyllopodopsyllus T. Scott, 1906a
Phyllothalestris Sars, 1905c
Pilifera Noodt, 1952
Pilocamptus Huys, 2009 mihi
Pindamoraria Reid & Rocha, 2003
Platychelipus Brady, 1880a
Platylaophonte Bodin, 1968b
Polyascophorus George, 1998b
Pontophonte Lee & Huys, 1999b
Pontopolites T. Scott, 1894b
Pontostratiotes Brady, 1883
Porcellidium Claus, 1860
Pordfus Özdikmen, 2008
Potamocaris Dussart, 1979
Praeleptomesochra Lang, 1965
Prionos Huys & Gee 1996b
Proameira Lang, 1944
Probosciphontodes Fiers, 1988
Proceropes Huys, 1990c
Propephonte Gheerardyn & Fiers in Gheerardyn et al., 2006b
Prosewellina Mielke, 1987
Protogoniceps Por, 1964a
Protolatiremus Itô, 1974
Protopsammotopa Geddes, 1968c
Psamathea Cottarelli & Venanzetti, 1989
Psammameira Noodt, 1952
Psammastacus Apostolov & Marinov, 1988
Peltidiphonte rostrata Gheerardyn & Fiers in Gheerardyn et al., 2006a
Peltidium purpureum Philippi, 1839
Idomene aberrans Por, 1964a
Peltisenia aberrans (Por, 1964a)
Peltthestris tripartita Monard, 1924
Peltobradya bryozoophila Médioni & Soyer, 1968
Peresime abyssalis Dinet, 1974
Perissocope typicus Brady, 1910
Hemimesochra rapiens Becker, 1979
Perucamptus rapiens (Becker, 1979)
Canthocamptus schmeilii Mrázek, 1893
Pesceus schmeilii (Mrázek, 1893)*
Pholenota spatulifera Vervoort, 1964
Cancrincola wilsoni Pearse, 1930
Pholetiscus wilsoni (Pearse, 1930)
Belisarius viguieri Maupas, 1892
Phyllognathopus viguieri (Maupas, 1892)
Tetragoniceps Bradyi T. Scott, 1892
Phyllopodopsyllus bradyi (T. Scott, 1892)
Thalestris mysis Claus, 1863
Phyllothalestris mysis (Claus, 1863)
Laophonte gracilis T. Scott, 1903c
Pilifera gracilis (T. Scott, 1903c)
Canthocamptus pilosus Douwe, 1910
Pilocamptus pilosus (Douwe, 1910)
Pindamoraria boraceiae Reid & da Rocha, 2003
Platychelipus littoralis Brady, 1880a
Platylaophonte delamarei Bodin, 1968b
Polyascophorus martinezi George, 1998b
Paronychocamptus leuke Por, 1959
Pontophonte leuke (Por, 1959)
Pontopolites typicus T. Scott, 1894b
Pontostratiotes abyssicola Brady, 1883
Thyone viridis Philippi, 1840
Porcellidium viride (Philippi, 1840)
Ophirion communis Por, 1967
Pordfus communis (Por, 1967)
Potamocaris bifida Dussart, 1979
Leptomesochra africana Kunz, 1951
Praeleptomesochra africana (Kunz, 1951)
Prionos ornata Huys & Gee, 1996b
Ameira simplex Norman & Scott, 1905
Proameira simplex (Norman & Scott, 1905)
Probosciphontodes stellata Fiers, 1988
Sarsocletodes secundus Smirnov, 1946
Proceropes secunda (Smirnov, 1946)
Propephonte duangitensis Gheerardyn & Fiers in Gheerardyn et al., 2006b
Prosewellina chilensis Mielke, 1987
Protogoniceps hebraeus Por, 1964a
Protolatiremus sakaguchii Itô, 1974
Protopsammotopa norvegica Geddes, 1968c
Psamathea nautarum Cottarelli & Venanzetti, 1989
Psammameira hyalina Noodt, 1952
Psammastacus confluens Nicholls, 1935
Psammastacus confluens (Nicholls, 1935)
Type fixation
original designation
monotypy
original designation
monotypy
monotypy
monotypy
indication
original designation
original designation
original designation
original designation
monotypy
original designation
original designation
original designation
original designation
original designation
monotypy
monotypy
original designation
original designation
indication
monotypy
monotypy
monotypy
monotypy
original designation
original designation
original designation
original designation
original designation
original designation
monotypy
original designation
original designation
monotypy
monotypy
original designation
original designation
Notes
(83)
(84)
(12)
(4)
(85)
(86)
(4)
TERMS OF USE
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UNRESOLVED CASES IN HARPACTICOID COPEPOD NOMENCLATURE
Genus-group name (subgenera indented)
83
Type species (original binomen)
Type species (valid binomen)
Type fixation
Psammis Sars, 1910
Psammocamptus Mielke, 1975
Psammolaophonte Wells, 1967
Psammoleptastacus Pennak, 1942
Psammoleptomesochra Mielke, 1994b
Psammonitocrella Huys, 2009 mihi
Psammoplatypus Lee & Huys, 1999b
Psammopsyllus Nicholls, 1945b
Psammotopa Pennak, 1942
Pseudameira Sars, 1911b
Pseudamphiascopsis Lang, 1944
Pseudechinopsyllus George, 2006a
Pseudectinosoma Kunz, 1935
Pseudoameiropsis Pallares, 1982b
Pseudobradya Sars, 1904a
Pseudodiosaccopsis Lang, 1944
Pseudodiosaccus T. Scott, 1906a
Pseudolaophonte A. Scott, 1896
Pseudoleptomesochra Lang, 1965
Pseudoleptomesochrella Lang, 1965
Pseudomesochra T. Scott, 1902
Pseudomoraria Brancelj, 1994
Pseudonsiella Hicks, 1988b
Pseudonychocamptus Lang, 1944
Pseudostenhelia Wells, 1967
Pseudotachidius T. Scott, 1898
Pseudozosime T. Scott, 1912
Psyllocamptus T. Scott, 1899b
Pteropsyllus T. Scott, 1906a
Pusillargillus Huys & Thistle, 1989
Quinquelaophonte Wells, Hicks & Coull, 1982
Rangabradya Karanovic & Pesce, 2001
Raoleptomesochra Karanovic, 2004
Raowellsia Özdikmen, 2008
Raptolaophonte Cottarelli & Forniz, 1989
Remanea Klie, 1929
Remaneicaris Jakobi, 1972
Retrocalcar Huys & Lee, 1999
Rhizothrix Sars, 1909a
Rhyncholagena Lang, 1944
Psammis longisetosa Sars, 1910
Psammocamptus axi Mielke, 1975
Psammolaophonte spinicauda Wells, 1967
Psammoleptastacus arenaridus Pennak, 1942
Psammoleptomesochra australis Mielke, 1994b
Psammonitocrella boultoni Rouch, 1992
Klieonychocamptus discipes Noodt, 1958
Psammopsyllus operculatus Nicholls, 1945b
Psammotopa vulgaris Pennak, 1942
Pseudameira crassicornis Sars, 1911b
Stenhelia herdmani A. Scott, 1896
Pseudechinopsyllus sindemarkae George, 2006a
Pseudectinosoma minor Kunz, 1935
Pseudoameiropsis argentinus Pallares, 1982b
Bradya minor (Scott & Scott, 1896)
Amphiascus rufescens Brian, 1925a
Diosaccus propinquus Scott & Scott, 1893b
Laophonte spinosa Thompson, 1893
Pseudoleptomesochra typica Lang, 1965
Nitocrella halophila Noodt, 1952
Pseudomesochra longifurcata T. Scott, 1892
Pseudomoraria triglavensis Brancelj, 1994
Pseudonsiella aotearoa Hicks, 1988b
Laophonte Koreni Boeck, 1873
Pseudostenhelia prima Wells, 1967
Pseudotachidius coronatus T. Scott, 1898
Pseudozosime browni T. Scott, 1912
Mesochra propinqua T. Scott, 1896
(?) Tetragoniceps consimilis T. Scott, 1894b
Hemimesochra nixe Por, 1964b
Laophonte quinquespinosa Sewell, 1924
Rangabradya indica Karanovic & Pesce, 2001
Parapseudoleptomesochra (?) reductus Rao, 1972
Langia maculata Wells & Rao, 1987
Raptolaophonte ardua Cottarelli & Forniz, 1989
Remanea arenicola Klie, 1929
Parastenocaris remanei Noodt, 1963
Cletopsyllus brattstroemi Geddes, 1981
Enhydrosoma curvatum Brady, 1880a
Amphiascus lagenirostris Sars, 1911a
monotypy
monotypy
original designation
monotypy
original designation
Psammonitocrella boultoni (Rouch, 1992)
original designation
Psammoplatypus discipes (Noodt, 1958)
original designation
monotypy
monotypy
subsequent designation
Pseudamphiascopsis herdmani (A. Scott, 1896)
original designation
original designation
monotypy
monotypy
Pseudobradya minor (Scott & Scott, 1896)
subsequent designation
Pseudodiosaccopsis rufescens (Brian, 1925a)
original designation
Pseudodiosaccus propinquus (Scott & Scott, 1893b) original designation
Pseudolaophonte spinosa (Thompson, 1893)
original designation
indication
Pseudoleptomesochrella halophila (Noodt, 1952)
original designation
monotypy
monotypy
monotypy
Pseudonychocamptus koreni (Boeck, 1873)
original designation
original designation
monotypy
monotypy
Psyllocamptus propinquus (T. Scott, 1896)
monotypy
Pteropsyllus consimilis (T. Scott, 1894b)
original designation
Pusillargillus nixe (Por, 1964b)
original designation
Quinquelaophonte quinquespinosa (Sewell, 1924)
original designation
original designation
Raoleptomesochra reducta (Rao, 1972)
original designation
Raowellsia maculata (Wells & Rao, 1987)
original designation
monotypy
monotypy
Remaneicaris remanei (Noodt, 1963)
original designation
Retrocalcar brattstromi (Geddes, 1981)
original designation
Rhizothrix curvata (Brady, 1880a)
monotypy
Rhyncholagena lagenirostris (Sars, 1911a)
original designation
Notes
(4)
(87)
(88)
(1, 5, 89)
(90)
(12)
(4, 91)
(92)
(93)
(94)
(95)
(4)
TERMS OF USE
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Genus-group name (subgenera indented)
HUYS
Zootaxa 2183 © 2009 Magnolia Press ·
Rhynchothalestris Sars, 1905c
Robertgurneya Lang, 1948
Robertsonia Brady, 1880a
Robustunguis Fiers, 1992
Romete Seifried & Schminke, 2003
Rosacletodes Wells, 1985
Rossopsyllus Soyer, 1975a
Rotundiclipeus Huys, 1988b
Sacodiscus Wilson, 1924
Sagamiella Lee & Huys, 1999a
Sarsameira Wilson, 1924
Sarsamphiascus Huys, 2009 mihi
Sarsocletodes Wilson, 1924
Scabrantenna Lee & Huys, 2000
Schizacron Gee & Huys, 1996
Schizopera Sars, 1905a
Schizoperoides Por, 1968
Schizothrix Huys, 1992
Scottolana Huys, 2009 mihi
Scutellidium Claus, 1866
Scutellopsis Wiborg, 1964
Selenopsyllus Moura & Pottek, 1998
Sentiropsis Huys & Gee, 1996a
Sewelliapusia Huys, 2009 mihi
Sewellina Krishnaswamy, 1956
Sextonis Huys, 1992
Sicameira Klie, 1950
Sigmatidium Giesbrecht, 1881
Simplicaris Galassi & De Laurentiis, 2004
Sinamphiascus Mu & Gee, 2000
Sinotachidius Huys, Ohtsuka, Conroy-Dalton & Kikuchi, 2004
Type species (original binomen)
Type species (valid binomen)
Thalestris helgolandica Claus, 1863
Rhynchothalestris helgolandica (Claus, 1863)
Stenhelia similis A. Scott, 1896
Robertgurneya similis (A. Scott, 1896)
Robertsonia tenuis Brady, 1880a
Robustunguis ungulatus Fiers, 1992
Romete bulbiseta Seifried & Schminke, 2003
Echinocletodes kuehnemanni Pallares, 1982a
Rosacletodes kuehnemanni (Pallares, 1982a)
Rossopsyllus kerguelensis Soyer, 1975a
Rotundiclipeus canariensis Huys, 1988b
Aspidiscus fasciatus Norman, 1869
Sacodiscus fasciatus (Norman, 1869)
Sagamiella latirostrata Lee & Huys, 1999a
Ameira parva Boeck, 1873
Sarsameira parva (Boeck, 1873)
Dactylopus minutus Claus, 1863
Sarsamphiascus minutus (Claus, 1863)
Pseudocletodes typicus Sars, 1920c
Sarsocletodes typicus (Sars, 1920c)
Scabrantenna yooi Lee & Huys, 2000
Enhydrosoma barnishi Wells, 1967
Schizacron barnishi (Wells, 1967)
Schizopera longicauda Sars, 1905a
Schizoperoides expeditionis Por, 1968
Leptastacus ctenatus Mielke, 1982
Schizothrix ctenata (Mielke, 1982)
Scottolana geei Mu & Huys, 2004
Scottolana geei (Mu & Huys, 2004)
Scutellidium longicauda (Philippi, 1840)
Psamathe longicauda Philippi, 1840
Scutellopsis armatus Wiborg, 1964
Selenopsyllus dahmsi Moura & Pottek, 1998
Danielssenia minuta Coull, 1969
Sentiropsis minuta (Coull, 1969)
Dactylopusia tropica Sewell, 1940
Sewelliapusia tropica (Sewell, 1940)
Sewellina reductus Krishnaswamy, 1956
Sewellina reducta Krishnaswamy, 1956
Leptastacus mehuinensis Mielke, 1985
Sextonis mehuinensis (Mielke, 1985)
Sicameira leptoderma Klie, 1950
Sigmatidium difficile Giesbrecht, 1881
Simplicaris lethaea Galassi & De Laurentiis, 2004
Sinamphiascus dominatus Mu & Gee, 2000
Tachidius (Tachidius) vicinospinalis Shen & Tai, Sinotachidius vicinospinalis (Shen & Tai, 1964b)
1964b
Smacigastes Ivanenko & Defaye, 2004
Smacigastes micheli Ivanenko & Defaye, 2004
Spelaeocamptus Chappuis, 1933b
Canthocamptus spelaeus Chappuis, 1925
Spelaeocamptus spelaeus (Chappuis, 1925)
Sphingothrix Fiers, 1997
Sphingothrix goldi Fiers, 1997
Spinaprecruris Gee, 2001
Cletodes curvirostris T. Scott, 1894b
Spinapecruris curvirostris (T. Scott, 1894b)
Spiniferaphonte Gheerardyn & Fiers in Gheerardyn et al., 2007 Spiniferaphonte ornata Gheerardyn & Fiers in Gheerardyn et al., 2007
Stenhelia Boeck, 1865
Stenhelia gibba Boeck, 1865
Stenocaris Sars, 1909b
Stenocaris gracilis Sars, 1909b
Stenocaris pristina Wells, 1968
Stenocaropsis pristina (Wells, 1968)
Stenocaropsis Apostolov, 1982
Type fixation
Notes
subsequent designation
original designation
monotypy
original designation
original designation
monotypy
monotypy
original designation
monotypy
original designation
original designation
original designation
indication
original designation
original designation
monotypy
monotypy
original designation
original designation
monotypy
monotypy
original designation
original designation
original designation
original designation
original designation
original designation
monotypy
original designation
original designation
original designation
(1, 96)
(1, 4)
original designation
monotypy
original designation
monotypy
original designation
monotypy
monotypy
original designation
(97)
(98)
(99)
(4)
(100)
(4)
(101)
(4)
(102)
TERMS OF USE
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UNRESOLVED CASES IN HARPACTICOID COPEPOD NOMENCLATURE
Genus-group name (subgenera indented)
85
Type species (original binomen)
Stenocopia Sars, 1907b
Stereoxiphos Huys & Conroy-Dalton, 2005
Stratiopontotes Soyer, 1970a
Strongylacron Gee & Huys, 1996
Stygepactophanes Moeschler & Rouch, 1984
Stygolaophonte Lang, 1965
Stygonitocrella Reid, Hunt & Stanley, 2003
Stygonitocrella Reid, Hunt & Stanley, 2003
Fiersiella Huys, 2009 mihi
Stylicletodes Lang, 1936b
Styracothorax Huys, 1993
Sunaristes Hesse, 1867
Superornatiremis Huys, 1996b
Syngastes Monard, 1924
Syrticola Willems & Claeys, 1982
Tachidiella Sars, 1909b
Tachidiopsis Sars, 1911c
Tachidius Lilljeborg, 1853
Ameira longicaudata T. Scott, 1892
Stenocopia longicaudata (T. Scott, 1892)
Stereoxiphos operculatus (Masry, 1970)
Stratiopontotes mediterranea Soyer, 1970a
Stratiopontotes mediterraneus Soyer, 1970a
Cletodes Buchholzii Boeck, 1873
Strongylacron buchholzii (Boeck, 1873)*
Stygepactophanes jurassicus Moeschler & Rouch, 1984
Stygolaophonte arenophila Lang, 1965
Nitocrella montana Noodt, 1965
Stygonitocrella montana (Noodt, 1965)
Nitocrella montana Noodt, 1965
Stygonitocrella montana (Noodt, 1965)
Stygonitocrella sequoyahi Reid, Hunt & Stanley, 2003
Cletodes longicaudata Brady, 1880a
Stylicletodes longicaudata (Brady, 1880a)
Styracothorax gladiator Huys, 1993
Sunaristes Paguri Hesse, 1867
Sunaristes paguri Hesse, 1867
Superornatiremis mysticus Huys, 1996b
Amymome [sic] clausii Thomson, 1883
Syngastes clausii (Thomson, 1883)*
Syrticola flandricus Willems & Claeys, 1982
Tachidiella minuta Sars, 1909b
Tachidiopsis cyclopoides Sars, 1911c
Cyclops brevicornis Müller, 1776 sensu
Tachidius discipes Giesbrecht, 1881
Lilljeborg (1853)
Tapholaophontodes rollandi Soyer, 1975a
Tapholeon ornatus Wells, 1967
Tectacingulum tumidum Harris, 1994
Tectacingulum tumidum (Harris, 1994)
Amymone Satyrus Claus, 1860
Tegastes satyrus (Claus, 1860)
Cleta secunda Sewell, 1924
Telodocus secundus (Sewell, 1924)
Micropsammis secunda Mielke, 1975
Telopsammis secunda (Mielke, 1975)
Tetanopsis typicus Brady, 1910
Tetragoniceps malleolata Brady, 1880a
Tetragoniceps malleolatus Brady, 1880a
Thalestris longimana Claus, 1863
Thermomesochra reducta Itô & Burton, 1980
Jonesiella hyænæ Thompson, 1889
Thompsonula hyaenae (Thompson, 1889)
Cyclops brevicornis Müller, 1776
Tigriopus brevicornis (Müller, 1776)
Cyclops furcatus Baird, 1837
Tisbe furcata (Baird, 1837)
Tisbella timsæ Gurney, 1927
Tisbella timsae Gurney, 1927
Tisbintra nankaurica Sewell, 1940
Tisbisoma spinisetum Božić, 1964
Tonpostratiotes tenuipedalis Itô, 1982
Polyascophorus schminkei George, 1998b
Touphapleura schminkei (George, 1998b)
Triathrix montagni Gee & Burgess, 1997
Tripartisoma trapezoidalis Avdeev, 1983
Troglophonte spelaea (Chappuis, 1938)
Laophonte spelaea Chappuis, 1938
Tapholaophontodes Soyer, 1975a
Tapholeon Wells, 1967
Tectacingulum Huys, 2009 mihi
Tegastes Norman, 1903a
Telodocus Huys & Willems, 1989
Telopsammis Gee & Huys, 1991
Tetanopsis Brady, 1910
Tetragoniceps Brady, 1880a
Thalestris Claus, 1862
Thermomesochra Itô & Burton, 1980
Thompsonula T. Scott, 1905b
Tigriopus Norman, 1869
Tisbe Lilljeborg, 1853
Tisbella Gurney, 1927
Tisbintra Sewell, 1940
Tisbisoma Božić, 1964
Tonpostratiotes Itô, 1982
Touphapleura Conroy-Dalton, 2001
Triathrix Gee & Burgess, 1997
Tripartisoma Avdeev, 1983
Troglophonte Huys & Lee, 2000
Type species (valid binomen)
Type fixation
Notes
subsequent designation
original designation
monotypy
original designation
monotypy
monotypy
original designation
original designation
original designation
monotypy
original designation
monotypy
original designation
subsequent designation
original designation
monotypy
monotypy
monotypy
(103)
monotypy
original designation
original designation
monotypy
original designation
original designation
indication
monotypy
subsequent designation
original designation
monotypy
monotypy
monotypy
monotypy
monotypy
monotypy
original designation
original designation
original designation
original designation
original designation
(12)
(104)
(4, 104)
(4)
(105)
(12)
(1, 106)
(107)
(4)
(12, 108)
(109)
(110)
(18)
(111)
(4)
(18)
TERMS OF USE
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Genus-group name (subgenera indented)
HUYS
Zootaxa 2183 © 2009 Magnolia Press ·
Type species (original binomen)
Type species (valid binomen)
Tryphoema Monard, 1926b
Tydemanella A. Scott, 1909
Typhlamphiascus Lang, 1944
Uptionyx Conroy-Dalton & Huys, 2000
Vermicaris Kornev & Chertoprud, 2008
Vibriopsyllus Kornev & Chertoprud, 2008
Volkmannia Boxshall, 1979
Weddellaophonte Willen, 1996
Wellsiphontina Fiers, 1991b
Wellsopsyllus Kunz, 1981
Wellsopsyllus Kunz, 1981
Scottopsyllus Apostolov & Marinov, 1988
Intermediopsyllus Huys, 2009 mihi
Willemsia Huys & Conroy-Dalton, 1994
Xanthilaophonte Fiers, 1991a
Xouthous Thomson, 1883
Xylora Hicks, 1988b
Yunona Avdeev, 1983
Zaus Goodsir, 1845
Zausodes Wilson, 1932
Zausopsis Lang, 1934
Zosime Boeck, 1873
Tryphoema porca Monard, 1926b
Tydemanella typica A. Scott, 1909
Amphiascus typhlops Sars, 1906b
Typhlamphiascus typhlops (Sars, 1906b)
Uptionyx verenae Conroy-Dalton & Huys, 2000
Stenocaris minuta Nicholls, 1935
Vermicaris minuta (Nicholls, 1935)
Vibriopsyllus curviseta Kornev & Chertoprud, 2008
Volkmannia forficula Boxshall, 1979
Weddellaophonte anyae Willen, 1996
Wellsiphontina striata Fiers, 1991b
Paramesochra gigas Wells, 1965a
Wellsopsyllus gigas (Wells, 1965a)
Paramesochra gigas Wells, 1965a
Wellsopsyllus gigas (Wells, 1965a)
Leptopsyllus minor Scott & Scott, 1895a
Wellsopsyllus minor (Scott & Scott, 1895a)
Leptopsyllus intermedius Scott & Scott, 1895b
Wellsopsyllus intermedius (Scott & Scott, 1895b)
Willemsia calceola Huys & Conroy-Dalton, 1993
Laophonte trispinosa Sewell, 1940
Xanthilaophonte trispinosa (Sewell, 1940)
Xouthous novæ-zealandiæ Thomson, 1883
Xouthous novaezealandiae Thomson, 1883
Xylora bathyalis Hicks, 1988b
Yunona marginata Avdeev, 1983
Zaus spinatus Goodsir, 1845
Zausodes arenicolus Wilson, 1932
Zausopsis mirabilis Lang, 1934
Zosime typica Boeck, 1873
Type fixation
monotypy
monotypy
original designation
original designation
original designation
original designation
original designation
original designation
original designation
original designation
original designation
original designation
original designation
original designation
original designation
monotypy
original designation
original designation
monotypy
original designation
monotypy
indication
Notes
(4)
(4)
(4, 18)
TERMS OF USE
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UNRESOLVED CASES IN HARPACTICOID COPEPOD NOMENCLATURE
Genus-group name (subgenera indented)
87
Actinocletodes Fiers, 1986c
Apolethon Wells, 1967
Argestoides Huys & Conroy-Dalton, 1997
Australonannopus Hamond, 1974
Barbaracletodes Huys, 2009 mihi
Bodinia George, 2004a
Cletocamptus Shmankewich, 1875
Laophontella Thompson & Scott, 1903
Leimia Willey, 1923
Nannopodella Monard, 1928
Parepactophanes Kunz, 1935
Pontocletodes Apostolov, 1980
Pseudocleta Lang, 1944
Pseudocletodes Scott & Scott, 1893b
Pyrocletodes Coull, 1973c
Scintis Por, 1986b
Taurocletodes Kunz, 1975a
Actinocletodes woutersi Fiers, 1986c
Apolethon fumator Wells, 1967
Argestoides prehensilis Huys & Conroy-Dalton, 1997
Australonannopus aestuarinus Hamond, 1974
Barbaracletodes barbara Becker, 1979
Barbaracletodes barbara (Becker, 1979)
Bodinia meteorensis George, 2004a
Cletocamptus retrogressus Shmankevich, 1875
Laophontella typica Thompson & Scott, 1903
Leimia vaga Willey, 1923
Nannopodella Denisi Monard, 1935
Nannopodella denisi Monard, 1935
Parepactophanes minuta Kunz, 1935
Pontocletodes ponticus Apostolov, 1980
Laophonte corbula Willey, 1935
Pseudocleta corbula (Willey, 1935)
Pseudocletodes vararensis Scott & Scott, 1893b
Pyrocletodes desuramus Coull, 1973c
Scintis variifurca Por, 1986b
Parepactophanes ? dubia Noodt, 1958
Taurocletodes dubius (Noodt, 1958)
original designation
original designation
original designation
original designation
original designation
(4)
original designation
monotypy
indication
monotypy
subsequent designation (5, 12, 112)
monotypy
original designation
original designation
monotypy
(113)
original designation
original designation
monotypy
Dactylopina villosa Brady, 1910
Eupeltidium glabrum A. Scott, 1909
Flavia crassicornis Brady, 1899
Marshia brevicaudata Herrick, 1895
Mawsonella typica Brady, 1918
Amphiascus parvus Sars, 1906b
Metaphroso gracilis Brady, 1910
Microcryobius nanus Brady, 1910
Phroso gracilis Brady, 1899
monotypy
monotypy
monotypy
subsequent designation (5, 113)
indication
designation dubious
(4)
monotypy
monotypy
monotypy
not yet designated
original designation
monotypy
(12)
monotypy
Genera inquirenda
Dactylopina Brady, 1910
Eupeltidium A. Scott, 1909
Flavia Brady, 1899
Marshia Herrick, 1895
Mawsonella Brady, 1918
Mesamphiascus Nicholls, 1941b
Metaphroso Brady, 1910
Microcryobius Brady, 1910
Phroso Brady, 1899
Portierella Labbé, 1926
Pseudocletopsyllus Vervoort, 1964
Reticulina Cleve, 1901
Tisemus Monard, 1928
Pseudocletopsyllus spiniger Vervoort, 1964
Reticulina Aurivilii Cleve, 1901
Tisemus pulchellus Monard, 1928
Mesamphiascus parvus (Sars, 1906b)
Reticulina aurivilii Cleve, 1901
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Genera incertae sedis
HUYS
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Notes
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
13.
14.
15.
In the introduction to his monograph, Lang (1948: 7) stated that whenever the type species of a genus was not
explicitly indicated by him, he considered the first species to have been described in that genus as the type (“Wenn
der Genotypus für eine Gattung nicht angegeben wird, betrachte ich die zuerst beschriebene Art als solcher.”). Such
a rule is not among the allowed criteria for ascertaining type fixation in the original publication (ICZN Art. 68), and
thus the status of being the “oldest species” assigned to one of Lang’s genera does not in and of itself confer type
status on that species. The term “designation” must be rigidly construed (ICZN Art. 67.5, which applies to Art. 69
for subsequent designation as well).
Subsequent designation by Boxshall (1979: 204). According to Seifried and Schminke (2003: 32) the family-group
name Aegisthidae Giesbrecht, 1893 [see Holthuis and Vervoort (2006) for correct publication date] is a senior
subjective synonym of Cerviniidae Sars, 1903.
Chappuis (1932a: 416) proposed the new generic name Eucamptus Chappuis, 1932 for Eucamptus uncinatus
Chappuis, 1932a (type by monotypy), overlooking that this name was preoccupied by four senior homonyms:
Eucamptus Guérin-Ménéville, 1832 (Coleoptera), Eucamptus Chevrolat, 1833 (Coleoptera), Eucamptus Dejean,
1834 (Coleoptera) and Eucamptus Dujardin, 1845 (Nematoda). Chappuis (1932b: 644) subsequently proposed the
new replacement name Afrocamptus which takes the same type species as its prior nominal taxon (ICZN Art. 67.8).
See present paper for explanation.
Originally included species designated herein (ICZN Art. 69.1).
According to Willen (2000: 202) the family-group names Hamondiidae and Ambunguipedidae proposed by Huys
(1990b: 45 and 93, respectively) are junior subjective synonyms of the name Rhynchothalestridae Lang, 1948.
Boxshall and Halsey (2004: 303) considered Hamondiidae the senior synonym of Ambunguipedidae, on the
grounds of page priority, and reinstated it as a valid family name.
In his redescription of Ameira longipes Boeck, 1865, Sars (1907a: 216) stated “... this is the form originally
recorded by Boeck as the type of the genus Ameira”. Although there is no such statement in Boeck’s (1865) paper,
Sars’s claim qualifies as a valid type designation according to ICZN Art. 69.1.1 and takes precedence over Lang’s
(1944: 24) more explicit type fixation. Contrary to previous authors (e.g. Lang 1948; Boxshall & Halsey 2004) who
attributed the family-group name Ameiridae to Monard (1927), Wells (2007: 88) pointed out that the authorship
and date must remain with Boeck (1865: 273) who established it under the name “Afdeling Ameirina”.
Subsequent designation by Apostolov and Marinov (1988: 238).
Nicholls (1941a: 415) proposed this genus for the debilis-group of Amphiascus Sars, 1905a but neglected type
fixation (published on 28 February). The name was made available in his revision of the Diosaccidae (= Miraciidae)
(Nicholls 1941b: 81; published on 27 October) when he fixed Dactylopus debilis Giesbrecht, 1881 as the type.
Subsequent designation by Nicholls (1941b: 75).
Sars (1909b: 312) changed the correct original spelling Ancorabolus to Anchorabolus and used the stem of this
incorrect subsequent spelling (ICZN Art. 33.3) for his new family-group name Anchorabolidae (Sars 1909b: 311).
Norman (1911: 139) pointed out Sars’s incorrect spelling of the type genus and Lang (1948: 1453) corrected the
spelling of the family to Ancorabolidae (ICZN Arts 32.5.3.3 and 35.4.1).
Species-group names are always to begin with a lower-case initial letter, regardless of how they were originally
published (ICZN Art. 28). In a species-group name first published with an initial upper-case letter the initial letter
must be replaced with a lower-case letter (ICZN Art. 32.5.2.5).
Bodin (1997: 193) considered Abyssameira Itô, 1983 (type by original designation: Abyssameira reducta Itô, 1983)
a potential junior subjective synonym of Argestes without giving any specific reasons for this; his course of action
however, has recently been endorsed by George (2008).
Atergopediidae Martínez Arbizu & Moura, 1998 (published 19 October) is a junior subjective synonym of
Novocriniidae Huys & Iliffe, 1998 (published 8 May).
Brady (1880a: 58) established the genus for two new species, Attheyella spinosa Brady, 1880a and A. cryptorum
Brady, 1880a. The former is a junior subjective synonym of Canthocamptus crassus Sars, 1863 (cf. Sars 1907a:
200) and was subsequently fixed as the type by Chappuis (1929a: 46).
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16. Both Chappuisiella Brehm, 1925 and Delachauxiella Brehm, 1925 were proposed as new genera by Brehm (1925:
314), not as subgenera as claimed by Lang (1948: 988, 996). Chappuis (1929a: 47–48) relegated them to subgenera
of Attheyella Brady, 1880a and fixed their respective type species by subsequent designation.
17. The original subgeneric name, Brehmiella Chappuis, 1929a, was preoccupied by Brehmiella Pascher, 1928
(Protozoa) and replaced by a new replacement name, Mrazekiella Brehm, 1949 (Brehm 1949: 515), which in itself
also proved to be a junior homonym of Mrazekiella Kijenskij, 1926 (Protozoa) (Özdikmen & Pesce 2006: 95). The
originally designated type was Canthocamptus northumbricus Brady, 1880a which is a junior subjective synonym
of Canthocamptus dentatus Poggenpol, 1874 (cf. Chappuis 1929a: 48); this type fixation also applies to the new
replacement name Neomrazekiella Özdikmen & Pesce, 2006 (ICZN Art. 67.8).
18. No diacritic or other mark (such as an apostrophe), or ligature of the letters a and e (æ) or o and e (œ) is to be used
in a scientific name (ICZN Art. 27). Names published with a diacritic or other mark, ligature, apostrophe, or
hyphen, are incorrect original spellings that must be corrected according to ICZN Art. 32.5.2.
19. Subsequent designation by Mu and Huys (2002: 203).
20. Huys (1996a: 1261) proposed the new replacement name Biuncus Huys, 1996a for the preoccupied Singularia
Huys, 1995a [junior homonym of Singularia Arenberger, 1988 (Lepidoptera)]. Huys’s (1995a: 674) fixation by
original designation also applies to Biuncus (ICZN Art. 67.8).
21. Brian (1923: 129) first cited the genus as part of a binominal nomen nudum “Bradyellopsis subniger n. gen. n. sp.
mihi” and subsequently formally proposed the genus for two species in a later paper (Brian 1925b: 116). Since the
expression “n. g. n. sp.” in this latter work was only applied to B. subniger (p. 122) this is deemed to be an original
type designation (ICZN Art. 68.2.1).
22. A new replacement name Brianola Monard, 1927 was proposed for Brianella Monard, 1926a [junior homonym of
Brianella Wilson, 1915 (Copepoda: Lernaeopodidae)] by Monard (1927: 145). Monard’s (1926a: 46) fixation by
monotypy consequently also applies to Brianola (ICZN Art. 67.8).
23. Westwood (1836: 228) stated that species belonging to “… our genus Canthocampus, having for its type the C.
staphylinus, have the abdomen of the females recurved with a spine beneath at the base…”. Stebbing (1910: 546)
claimed “… since the premier genus is Canthocampus (not Canthocamptus) Westwood, 1836, the spelling of the
family should be arranged to correspond” (i.e. “Canthocampidae”). Baird (1850: 204), however, pointed out that
the name was derived from the Greek ακανθα (a spine) and καμπτος (flexible) and should be spelled
Canthocamptus instead of Westwood’s misprinted incorrect original spelling Canthocampus. The subsequent
spellings Canthocarpus (Baird 1834: 97) and Cantocamptus (Sars 1909c: 32) are also to be considered lapsus
calami. Contrary to previous authors (e.g. Lang 1948; Boxshall & Halsey 2004) who attributed the family-group
name Canthocamptidae to Sars (1906c), Wells (2007: 90) correctly pointed out that the authorship and date must
remain with Brady (1880a: 47) who established it under the name “Canthocamptinæ”. Borutzky (1931c: 281–283)
proposed the genus Baikalocamptus (type by monotypy: B. verestschagini Borutzky, 1931c) as the type of a new
family Baikalocamptidae. Chappuis’s (1935: 284) course of action to relegate Baikalocamptus to a junior
subjective synonym of Canthocamptus (and hence the family name Baikalocamptidae to a junior subjective
synonym of Canthocamptidae) was endorsed by Lang (1948: 920).
24. Since Norman was alone responsible both for the name Cervinia and for satisfying the criteria of availability other
than actual publication, he takes the authorship of the generic name and of its type species (ICZN Art. 50.1.1;
Recommendation 51E). The same rule applies to the type species of Ambunguipes Huys, 1990b (Thalestris
rufocincta Norman in Brady, 1880a).
25. Sars (1903: 22) proposed the genus Cerviniopsis for two new species (C. clavicornis Sars, 1903 and C.
longicaudata Sars, 1903) but neither he nor any subsequent worker fixed a type. Cerviniopsis clavicornis is here
fixed as the type by subsequent designation.
26. T. Scott (1905a: 146) listed the family-group name “Cletodeidae” without accompanying description or definition
but included under it the description of the new species Cletodes sarsi (incorrect original spelling C. Sarsi – now
placed in the genus Enhydrosoma Boeck, 1873). Since the name was published before 1931 and based on the then
(and still) valid generic name Cletodes (ICZN Art. 12.2.4) it satisfies all criteria required for availability. Sars
(1909a: 281) was the first author to amend it to “Cletodidae”, using the correct stem of the type genus.
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27. Goniopelte Claus, 1891 (type species by monotypy: Goniopelte gracilis Claus, 1891) is a junior subjective
synonym of Clytemnestra Dana, 1847. Huys and Conroy-Dalton (2000: 4) pointed out that the unused family name
Goniopeltidinae proposed by Claus (1891: 430) loses in priority from its junior synonym Clytemnestridae A. Scott,
1909 since reversal of precedence applies (ICZN Art. 23.9). Poppe’s (1891) family-group name, with its alternative
spellings Pseudo-Peltididae (Poppe 1891: 141), Pseudo-Peltidiidae (Lang 1944: 11) and Pseudopeltidiidae (Wells
1976: 6, 11), was rejected by Boxshall (1979: 232) because its usage contravenes ICZN Art. 11.7.1.1. Seifried
(2003: 139) recently synonymised Clytemnestridae with Peltidiidae Boeck, 1873 but Wells (2007: 101) reinstated it
as a subfamily within the latter.
28. Brady (1878: 31) cited the generic name Cylindrosoma in a list but subsequently (Brady 1880b: 23, 30) remarked
that this name was preoccupied [being a junior homonym of Cylindrosoma Tschüdi, 1839 (Amphibia),
Cylindrosoma Gray in Jones, 1843 (Myriapoda) and Cylindrosoma Rondani, 1856 (Diptera)]. Cylindrosoma Brady,
1878 is however a nomen nudum since it does not satisfy the provisions of ICZN Art. 12.1.
29. Vervoort and Holthuis (1983) submitted a revised version of Vervoort's (1963) application, asking the Commission
to use its plenary powers to set aside all previous type fixations for Dactylopusia and to designate Dactylopus
tisboides as type species of the genus. This proposal was adopted under the plenary powers as recorded and
published in Opinion 1356 (Melville 1985).
30. Brian (1928b: 338) designated Dactylopusioides stampaliae Brian, 1928b [first cited as a nomen nudum in Brian
(1928a: 37)], which is a junior subjective synonym of Dactylopus macrolabris Claus, 1866 (cf. Lang 1936c:
34–35), as the type species.
31. Wells (1978: 1) changed the incorrect original spelling D’Arcythompsonia T. Scott, 1906a to Darcythompsonia and
the family name (Darcythompsoniidae) accordingly.
32. Latiremus Božić, 1969, the type genus of the nominal family-group taxon Latiremidae Božić, 1969, was considered
to be a junior synonym of Delamarella Chappuis, 1954b by Huys et al. (2005), however, according to ICZN Art.
40.1 the family name is not to be replaced.
33. According to Lang (1936c: 14) Diarthrodes Thomson, 1883 is a junior subjective synonym of Westwoodia Dana,
1854 [type: Harpacticus nobilis Baird, 1846b (incorrect original spelling Arpacticus)], however Sharpe (1910) had
already shown that the latter is an invalid name, being preoccupied by Westwoodia Brullé, 1846 (Hymenoptera).
Another subjective synonym, Pseudothalestris Brady, 1883 (type by monotypy: Pseudothalestris imbricata Brady,
1883), although published in the same year, loses in priority to Diarthrodes. Thomson’s (1883) work was published
in May 1883 while the exact date of Brady’s (1883) H.M.S. Challenger report is incompletely specified – the
publication date of the latter to be adopted is 31 December 1883 (ICZN Art. 21.3.2). Being based on a junior
homonym (ICZN Art. 39), the family-group name “Westwoodina” introduced by Boeck (1865: 257) becomes a
permanently invalid synonym of Dactylopusiidae Lang, 1936c.
34. Brian (1925a: 15) proposed the genus without type fixation for two new species, Diosaccopsis rubeus Brian, 1925a
and D. amphiasculus Brian, 1925a, but subsequently (Brian 1928b: 8) discovered that the former was the male of
the latter. He maintained D. rubeus as the only species but this course of action does not qualify as a valid type
fixation (ICZN Art. 69.4). Diosaccopsis rubeus is here formally fixed as the type of Diosaccopsis Brian, 1925a.
35. Sars’s (1906a: 145) claim that Brady (1880a) described “… the first-named species [Dactylopus tenuicornis] as the
type of the genus Diosaccus” can be considered a subsequent type designation according to ICZN Art. 69.1.1 even
though there is no such explicit statement in Brady’s (1880a: 68) diagnosis of the genus.
36. The spelling of the family-group name Ectinosomidae (Sars 1903: 28) was emended to Ectinosomatidae by Moore
(1978: 111).
37. Fixed as type by monotypy in the subgenus Canuella (Ellucana) by Sewell (1940: 136) (ICZN Art. 61.2.2). The
subgenus was upgraded to generic level by Coull (1971a: 203).
38. Subsequent designation by Sars (1909a: 298).
39. Vervoort (1964: 152) pointed out that Dactylopus latipes T. Scott, 1894a is a primary homonym of D. latipes
Boeck, 1865 (= type of Paradactylopodia Lang, 1944) and replaced it by the next available name, Eudactylopus
latipes f. andrewi Sewell, 1940 which he raised to species level. Since Sewell’s (1940) infrasubspecific name was
adopted as the valid name of a species before 1985 (Vervoort 1964: 154), the subspecific name andrewi is deemed
to be available from its original publication (ICZN Art. 45.6.4.1).
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40. Sars (1909a: 294) stated that “… this species [Cletodes laticauda], being the first recorded, may be regarded as the
type of the present genus…” but this is not an acceptable designation under the Code (Art. 67.5). Lang (1944: 30)
fixed this species as the type of the nominotypical subgenus Eurycletodes Sars, 1909a and hence also of the genus
(ICZN Art. 61.2.2).
41. Although generally (cf. Lang 1948) attributed to Claus (1863) who fixed Euterpe gracilis Claus, 1863 as the type
species and provided the first illustrated description, the genus Euterpe was first diagnosed the year before (Claus
1862: 87) but without any reference to species included under it. New names published before 1931 which are
accompanied by a description or a definition of the taxon (as in Claus 1862) that it denotes (ICZN Art. 12.1) satisfy
the criteria of availability and hence type fixation is not mandatory nor is the use of one or more available specific
names in combination with it, or clearly included under it. Norman (1903a: 368) proposed Euterpina as a new
replacement name for the preoccupied Euterpe Claus, 1862, which is a junior homonym of Euterpe Swainson, 1831
(Lepidoptera). Claus’s (1863: 110) fixation of Euterpe gracilis by subsequent monotypy also applies to Euterpina
Norman, 1903a (ICZN Art. 67.8). The type species is a junior subjective synonym of Harpacticus acutifrons Dana,
1847 (cf. Giesbrecht 1893: 555). According to Seifried (2003: 108) Brian’s (1921: 58) family name Euterpidinae
[Lang (1948: 285) and subsequent authors but not Gurney (1932: 17) cited this name as “Euterpinidae” – it is
uncertain whether Brian’s spelling is a lapsus calami] is a junior subjective synonym of the name Tachidiidae
Boeck, 1865.
42. T. Scott’s (1906a: 461) original type designation for Evansia T. Scott, 1906a [a junior homonym of Evansia
Pickard-Cambridge, 1900 (Arachnida)] also applies to its new replacement name Evansula T. Scott, 1906b (ICZN
Art. 67.8).
43. Conroy-Dalton and Huys (1997: 322) pointed out that Ameira longicaudata Nicholls, 1939a is a primary homonym
of A. longicaudata T. Scott, 1892 (= type of Stenocopia Sars, 1907b) and replaced it by the new binomen Filexilia
trisetosa Conroy-Dalton & Huys, 1997 which they designated as the type species of Filexilia Conroy-Dalton &
Huys, 1997.
44. The Code (Art. 13.3) does not require “designation” of a type species of a new genus, only “fixation”. Huys et al.’s
(1996: 228) explicit inclusion of Tachidius incisipes Klie, 1913 in Geeopsis Huys in Huys et al., 1996 resulted in
the fixation of the type species by monotypy. The second, unnamed, species referred to in Huys et al. (1996: 228)
pertains to Tachidius longicornis Olofsson, 1917. This was synonymised with the type species by Lang (1948:
289), who suspected that Olofsson’s (1917) description was deficient. Re-examination of material from the type
region Spitzbergen (provided by Dr W. Mielke) has confirmed the differences between T. incisipes and T.
longicornis and established the latter as a distinct species. Consequently, the genus Geeopsis now contains G.
incisipes (Klie, 1913) and G. longicornis (Olofsson, 1917) comb. nov.
45. The incorrect original spelling Alophytophilus by Brian (1919: 72) was corrected to Halophytophilus by Sars
(1920a: 48).
46. Milne-Edwards (1840) established the genus Harpacticus Milne-Edwards, 1840 (incorrect original spelling
Arpacticus) to include Cyclops chaelifer Müller, 1776 (incorrect original spelling C. chælifer), Cyclops armatus
Tilesius, 1815, and a new species Harpacticus chauseica (incorrect original spelling Arpacticus Chauseica). The
latter species has not been mentioned by any subsequent workers (including Lang (1948)) although it is listed as a
species in Sherborn (1925: 1216). Baird (1846a: 416) explicitly designated Cyclops chelifer as the type of
Harpacticus. Huys and Song (2004: 38) relegated Ismardis Leigh-Sharpe, 1936 (type by monotypy: Ismardis
spartacus Leigh-Sharpe, 1936) to a junior subjective synonym of Harpacticus. Hence, Leigh-Sharpe’s (1936: 68)
family-group name Ismardiidae is likewise a junior subjective synonym of the name Harpacticidae Dana, 1846
[Dana’s (1846: 226) incorrect original spelling Arpacticidae was corrected by Dana (1847: 152) although this did
not gain universal approval (cf. Stebbing 1910: 542)]
47. Thompson (1893: 185) proposed the genus Herdmania for Herdmania stylifera Thompson, 1893 (type by
monotypy). The placement of this genus in the Misophriidae [as defined under Brady’s (1878) broader concept]
made Lang (1935: 262) overlook it when he proposed Hemicervinia Lang, 1935 for a new species Hemicervinia
ryforsi (type by monotypy). Lang (1948: 183) subsequently recognized this oversight, pointed out the
conspecificity of Herdmania stylifera and Hemicervinia ryforsi, and relegated Hemicervinia to a junior objective
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48.
49.
50.
51.
52.
53.
54.
55.
56.
57.
synonym of Herdmania. Por (1964a: 65) eventually pointed out that Thompson’s genus was a junior homonym of
Herdmania Lahille, 1888 (Urochordata) and reinstated Hemicervinia.
Since Baird (1837: 330) had named the species after the prominent Norwegian zoologist Hans Ström (also spelled
Strøm), under ICZN Art. 32.5.2.1 the correct spelling is stromii as originally proposed by Baird (1837). It should,
however, be noted that Vervoort and Holthuis (1983: 56) had changed the spelling of stromii to stroemii, and the
latter spelling was accepted by the Commission. In order to ensure consistency with the ruling in Opinion 1356
(Melville 1985) the incorrect subsequent spelling stroemii is to be preserved unaltered and no Official Correction
(ICZN Art. 80.4) is required. Some authors (Apostolov & Marinov 1988; Bodin 1997; Wells 2007) have also
erroneously attributed Cyclops stroemii to Baird (1834) but that paper only quoted the species as Cyclops
brevicornis O.F. Müller, 1776 (p. 97) with which it had originally, but incorrectly, been identified. The original
description of Cyclops stroemii (as ‘Cyclops Stromii’) was given in Baird (1837: 330).
Designated by Kornev and Chertoprud (2008: 233) as the type genus of their recently proposed family
Heteropsyllidae.
Sars (1905b: 97) established the genus Idyopsis Sars, 1905b [a junior homonym of Idyopsis Agassiz, 1860
(Ctenophora)] for two new species, I. dilatata Sars, 1905b and I. pusilla Sars, 1905b, but did not fix a type.
Consequently, this condition also applies to its new replacement name Idyanthe proposed by Sars (1909c: 23) (see
also Sars 1911a: 369) (ICZN Art. 67.8). Vervoort (1964: 103) was the first author to validly fix Idyopsis dilatata as
the type species by subsequent designation. Seifried’s (2003: 92) subsequent fixation of the same type is invalid.
Subsequent designation by Vervoort (1964: 103).
Brady (1896: 24) designated Itunella subsalsa Brady, 1896 (by monotypy), a junior subjective synonym of
Cletodes tenuiremis T. Scott, 1893 (cf. T. Scott 1897: 151).
Subsequent designation by Huys and Gee (1993: 62).
Hicks and Schriever’s (1983: 2) original type designation for Kliella Hicks & Schriever, 1983 [a junior homonym
of Kliella Schäfer, 1945 (Ostracoda)] also applies to its new replacement name Klieosoma Hicks & Schriever, 1985
(ICZN Art. 67.8).
T. Scott (1905a: 145) listed the family-group name “Laophontidae” without accompanying description or definition
but included under it the description of the new species Laophonte longiremis (now placed in the genus Archesola
Huys & Lee, 2000). Since the name was published before 1931 and based on the then (and still) valid generic name
Laophonte (ICZN Art. 12.2.4) it satisfies all criteria required for availability. Although Stebbing (1910: 546)
pointed out that T. Scott (1907: 209) himself had attributed the family name to G.O. Sars, the date and authorship
must remain with T. Scott (1905a). The family-group name Namakosiramiidae (type genus: Namakosiramia Ho &
Perkins 1977 = Microchelonia Brady, 1918) proposed by Ho & Perkins (1977: 371) is a junior subjective synonym
of the name Laophontidae T. Scott, 1905a (Huys 1988a: 1520).
T. Scott (1894b: 249) designated “Laophontodes typicus (T. Scott, 1894b)” as type (by monotypy) for the new
subgenus Laophontodes T. Scott, 1894b in Laophonte Philippi, 1840 (ICZN Art. 61.2.2). The correct interpolated
combination should have been Laophonte (Laophontodes) typicus T. Scott, 1894b (ICZN Art. 6.1). Merope
Thomson, 1883 (type species: Merope hamata Thomson, 1883) is a junior homonym of Merope Newman, 1838
(Mecoptera), Merope Adams & Adams, 1856 (Mollusca) and Merope Albers, 1860 (Mollusca). Wilson (1932: 562)
proposed the new substitute name Meropia, which is preceded by its junior subjective synonym Laophontodes T.
Scott 1894b.
Although generally attributed to Claus (1863: 123), who included in it Harpacticus fortificationis Fischer, 1860,
Cleta brevirostris Claus, 1863, Cleta lamellifera Claus, 1863 and Cleta serrata Claus, 1863, the genus Cleta was
first diagnosed the year before (Claus 1862: 87) but without any reference to species included under it. According
to ICZN Art. 12.1 Cleta Claus, 1862 satisfies the criteria of availability and hence type fixation is not mandatory
nor is the use of one or more available specific names in combination with it, or clearly included under it. Sars
(1908b: 265) established the genus Laophontopsis (but not as a nomen novum) for Cleta lamellifera Claus, 1863
(type by monotypy) and transferred Claus’s (1863) other Cleta species to Laophonte Philippi, 1840 (Sars 1907b:
237; 1908a: 256). Sewell (1924: 834) pointed out that Cleta lamellifera should be designated as the type of Cleta,
implying the relegation of Laophontopsis to a junior objective synonym of the latter. However, Cleta Claus, 1862 is
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58.
59.
60.
61.
62.
63.
64.
65.
a junior homonym of Cleta Duponchel, 1844 (Lepidoptera) and Cleta Mulsant, 1850 (Coleoptera) and hence
Laophontopsis is the valid name.
Leptocaris T. Scott, 1899b permanently takes precedence (as a nomen protectum) over the earlier homonym
Leptocaris Aurivillius, 1898 (Mysida) (nomen oblitum) (Anonymous 2000). According to ICZN Art. 30.1.2, names
ending in -caris are feminine; hence, the specific epithet should agree in gender (minuta) [see editorial postscript in
Apostolov (2007: 1169)].
Wells (2007: 94) discussed the confusion surrounding the correct spelling of this genus.
Although generally attributed to Claus (1863), who fixed L. coronata as the type species by subsequent monotypy
and provided the first illustrated description, the genus was first diagnosed the year before (Claus 1862) but without
any reference to species included under it. New names published before 1931 that are accompanied by a description
or a definition of the taxon (as in Claus 1862) that it denotes (ICZN Art. 12.1) satisfy the criteria of availability and
hence type fixation is not mandatory nor is the use of one or more available specific names in combination with it,
or clearly included under it. Contrary to previous authors (e.g. Lang 1948; Boxshall & Halsey 2004) who attributed
the family-group name Longipediidae to Sars (1903), Wells (2007: 83) pointed out that the authorship and date
must remain with Boeck (1865: 252) who established it under the name “Afdeling Longipedina”.
Wilson (1924: 14) proposed the new replacement name Ceyloniella for the preoccupied generic name Ceylonia
Thompson & Scott, 1903 (type by monotypy: Ceylonia aculeata Thompson & Scott, 1903), a junior homonym of
Ceylonia Buckton in Cotes, 1891 (Hemiptera). In the same work, Wilson (1924: 15) also proposed the new
replacement name Lourinia for the preoccupied generic name Jurinia Claus, 1866 (type by monotypy: Jurinia
armata Claus, 1866), which is a junior homonym of Jurinia Robineau-Desvoidy, 1830 (Diptera) and Jurinia Costa,
1839 (Hymenoptera). Jurinia Claus, 1866 and Ceylonia Thompson & Scott, 1903 (and hence Lourinia and
Ceyloniella) are objective synonyms since their respective type species (although having different names) are based
on the same name-bearing type (ICZN Art. 61.3.3). Monard (1927: 173), acting as the first reviser (ICZN Art.
24.2.1), not only pointed out the synonymy but also fixed the precedence by selecting Lourinia as the senior name
[see also Vervoort (1964: 304)]. Claus’s (1866: 25) fixation by monotypy for Jurinia consequently also applies to
Lourinia (ICZN Art. 67.8). The family name Ceyloniidae proposed by A. Scott (1909: 227) is invalid because it
was based on a junior homonym (ICZN Art. 39); Monard’s (1937: 83) Ceyloniellidae is a junior subjective
synonym of the family name Louriniidae Monard, 1927.
Dana (1846, 1847, 1849, 1854) did not designate a type species for Setella Dana, 1846, but Giesbrecht (1893: 560 –
see Holthuis and Vervoort (2006) for correct publication date of this work), being the first person to fix a type
species, subsequently designated Setella gracilis Dana, 1847. Both A. Scott (1909: 230) and Wilson (1924: 16)
pointed out that Setella Dana, 1846 is a junior homonym of Setella Schrank, 1802 (Lepidoptera), and replaced it by
the new replacement names, Macrosetella and Dwightia, respectively, the former of which takes priority (Huys &
Böttger-Schnack 1994: 230). Giesbrecht’s (1893) type fixation also applies to Macrosetella (ICZN Art. 67.8). The
family-group name Macrosetellidae proposed by A. Scott (1909: 230) is a junior subjective synonym of Miraciinae
Dana, 1846. The family names “Setellina” proposed by Boeck (1865: 281) and “Setellidae” proposed
independently as intentionally new by Brian (1921: 57) are invalid because they were based on a junior homonym
(ICZN Art. 39).
According to Huys et al. (1996: 198) the family-group name Marsteiniidae proposed by Drzycimski (1969) is a
junior subjective synonym of Neobradyidae Olofsson, 1917.
Boeck (1865: 275) proposed the genus Mesochra for three species: Mesochra lilljeborgii Boeck, 1865 (incorrect
original spelling Liljeborgii), M. kroeyeri Boeck, 1865 (incorrect original spelling Krøyeri) and Dactylopus
pygmaeus Claus, 1863. Sars (1905a: 388) subsequently designated the first species as the type.
Wells (2007: 88) pointed out that the authorship and date of the family name Metidae should rest with Boeck (1873:
59) who first coined the name as “Metinae” and not Sars (1910) as generally accepted (e.g. Lang 1948; Boxshall &
Halsey 2004). The family-group names Abacolidae Edwards, 1891 (p. 92) and Ilyopsyllidae A. Scott, 1909 (p. 228)
are junior subjective synonyms of Metidae Boeck, 1873, being based on two junior subjective synonyms of Metis
Philippi, 1839: Abacola Edwards, 1891 (type by monotypy: Abacola holothuriae Edwards, 1891) and Ilyopsyllus
Brady & Robertson, 1873 (type by monotypy: Ilyopsyllus coraceus Brady & Robertson, 1873), respectively.
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66. Brady and Robertson (1873: 130) designated Microsetella atlantica Brady & Robertson, 1873, which is a junior
subjective synonym of Setella norvegica Boeck, 1865 (cf. Sars 1904a: 44).
67. Dana diagnosed the genus in 1846; however the two species (Miracia efferata Dana, 1849; M. gracilis Dana, 1849)
first assigned to it were not specified until 1849. According to ICZN Art. 11.4.1, a work containing genus-group
names without associated nominal species is accepted as consistent with the Principle of Binominal Nomenclature.
Huys and Böttger-Schnack (1994: 210) stated that Miracia efferata became the type and only species when Dahl
(1895) proposed M. gracilis as the type of a new genus Oculosetella, however, according to ICZN Art. 69.4,
elimination of all but one of the originally included species does not in itself constitute type fixation. Unless an
earlier type fixation has been overlooked, Boxshall’s (1979: 234) citation of M. efferata as the type species is here
accepted as the first type fixation by subsequent designation. Willen (2000: 214) relegated the family Miraciidae
Dana, 1846 to a subfamily in the family Diosaccidae Sars, 1906a but subsequently recognized the name Miraciidae
has priority over Diosaccidae (Willen 2002: 40).
68. Scott and Scott (1893a: 213) fixed Moraria andersonsmithi Scott & Scott, 1893a (original incorrect spelling
Anderson-Smithi), which is a junior subjective synonym of Canthocamptus brevipes Sars, 1863 (cf. Lang 1948:
1029), as the type species by monotypy. The generic name Ophiocamptus Mrázek, 1893, proposed for
Ophiocamptus sarsii Mrázek, 1893 (type by original designation), O. poppei Mrázek, 1893 and Canthocamptus
brevipes was published on 20 May 1893 (Mrázek 1893: 112) and hence is a junior subjective synonym of Moraria
(published March 1893). Borutzky (1931b: 263) subdivided the genus by proposing a new subgenus
Baikalomoraria Borutzky, 1931b [subsequent incorrect spelling Baicalomoraria by Borutzky (1949: 873)] for five
new species: Moraria baikalensis Borutzky, 1931b; M. dentata Borutzky, 1931b; M. intermedia Borutzky, 1931b;
M. laticauda Borutzky, 1931b; and M. tenuicauda Borutzky, 1931b. None of these originally included species was
fixed as the type by Borutzky (1931b) or subsequently by another author, consequently rendering the subgeneric
name unavailable (ICZN Art. 13.3). Instead of making the name available by fixing a type and providing a
description of the taxon it denotes, I have elected to follow Wells’s (2007: 92) judgement in that a comprehensive
revision of the entire genus is required before the validity of these subgenera can be assessed. Löffler (1962: 195)
added a third subgenus Kuehneltiella Löffler, 1962 (incorrect original spelling Kühneltiella) (type by monotypy: K.
neotropica Löffler, 1962). Cicchino and Ringuelet (1977: 226) invalidated this subgenus because the type belongs
to Antarctobiotus Chappuis, 1930.
69. Borutzky (1931b: 271) originally designated the type Morariopsis typica Borutzky, 1931b by indication. In the
Russian summary (Borutzky 1932: 20) of his earlier descriptions in German (Borutzky 1931a–c) the author
proposed the genus under a different name, Morariopus g. n., containing Morariopus typica sp. n. and Moriariopus
latifurcata sp. n.; “Morariopus” is here regarded as an incorrect subsequent spelling (ICZN Art. 33.3).
70. Jakobi (1953: 56) designated Mourephonte catharinensis Jakobi, 1953, which is a junior subjective synonym (cf.
Lang 1965: 446) of Laophonte longiseta Nicholls, 1941a, as the type species.
71. Gurney’s (1927: 543) type fixation by monotypy for Pseudomesochra Gurney, 1927 [a junior homonym of
Pseudomesochra T. Scott 1902 (family Pseudotachidiidae)] also applies to its new replacement name
Nannomesochra Gurney, 1932 (ICZN Art. 67.8). The type species Pseudomesochra parvula Gurney, 1927 is a
junior subjective synonym of Mesochra arupinensis Brian, 1925a (cf. Monard 1935b: 54; Lang 1936d: 451, 457).
72. It is clear from Nicholls’s (1945a: 2) heading “Neodactylopus cyclopoides gen. et sp. nov.” and his statement
“Since there is only the single species no generic diagnosis is given” that he had established the new nominal genusgroup taxon for a single species. Fixation by this means is deemed to be fixation by monotypy, regardless of the
author’s claim in an addendum to his paper (p. 15) that the species denoted by the name ? Eudactylopus anomala
Sewell, 1940 should also be included in Neodactylopus (ICZN Art. 68.3).
73. The type was fixed by monotypy in the subgenus Tachidius (Neotachidius) by Shen and Tai (1963: 417) (ICZN
Art. 61.2.2). The subgenus was upgraded to full generic rank by Huys et al. (2005: 134).
74. Dussart’s (1982: 102) subsequent type fixation of N. spinipes Boeck, 1865 is invalid.
75. The genus-group name Nitocrellopsis Petkovski, 1976 (no original type fixation) was made available by Galassi et
al. (1999: 178) by fixing N. rouchi Galassi, De Laurentiis & Dole-Olivier, 1999 as the type species and by
providing an amended generic diagnosis (ICZN Arts 13.1 and 13.3).
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76. Both Noodtiella Wells, 1965b and Noodtia Lang, 1965 have the same name-bearing type, Sigmatidium ?
arenosetelloides Noodt, 1958, and are therefore objective synonyms (Lang 1965: 12; Wells 1965b: 30). The former
takes precedence over the latter [see postscript in Lang (1965: 547)] and has page priority over Lineosoma Wells,
1965b (Wells 1967; Kihara & Huys 2009a).
77. Cristacoxa Huys, 1990a, the type genus of the nominal family-group taxon Cristacoxidae Huys, 1990a, was
considered to be a junior synonym of Noodtorthopsyllus Lang, 1965 by Huys and Kihara (2009), however,
according to ICZN Art. 40.1 the family name is not to be replaced.
78. Daday (1903: 157) fixed Onychocamptus heteropus Daday, 1903, which is a junior subjective synonym of
Laophonte mohammed Blanchard & Richard, 1891 (cf. Zykoff 1904: 247), as the type species by monotypy.
79. Claus (1866: 22) proposed the genus Liljeborgia (an incorrect original spelling as it was expressly intended as a
patronym in honour of Wilhelm Lilljeborg) for which he fixed Liljeborgia linearis Claus, 1866 as the type species
by monotypy. The correction of incorrect original spellings resulting from inadvertent errors is mandatory and
hence Liljeborgia has to be emended to Lilljeborgia in accordance with ICZN Art. 32.5.1, retaining the authorship
and date of the original spelling. Lilljeborgia Claus, 1866 is a junior homonym of Lilljeborgia Spence Bate, 1862
(Amphipoda) and was replaced by a new replacement name Orthopsyllus Brady & Robertson, 1873 (Brady &
Robertson 1873: 138). Consequently, Claus’s (1866) type fixation for Lilljeborgia also applies to its nomen novum,
Orthopsyllus (ICZN Art. 67.8).
80. The subgeneric name Bradya (Parabradya) Lang, 1944 was attributed full generic rank by Seifried et al. (2007: 3).
81. Thompson and Scott (1903: 263) proposed the genus for two new species, Parastenhelia hornelli and P. similis but
did not fix the type of the genus. Lang (1934: 24) reduced the latter to a synonym of P. hornelli which he declared
as the type of the genus. For some inexplicable reason Lang (1944: 13) designated Harpacticus spinosus Fischer,
1860 as the type species, and his course of action has been perpetuated in the literature [e.g. Vervoort (1964: 180),
Apostolov & Marinov (1988: 134)]. This subsequent fixation is invalid since this species was not originally
included in the genus (ICZN Art. 69.2).
82. Chappuis (1933b: 48) fixed Parastenocaris as the type of the family Parastenocaridae and this date was adopted by
most subsequent authors. However, as recognised by Damian-Georgescu (1970: 219) and Reid (1994: 775) the
name did not become available until 1940, when a brief diagnosis was provided by Chappuis (1940a: 294). The
spelling of the name was corrected by Noodt (1965: 101) who changed it to Parastenocarididae. H.K. Schminke in
a paper read to the 5th International Conference on Copepoda, Baltimore, 1993 (but not published in the
Proceedings) divided the family into two subfamilies – Parastenocaridinae and Fontinalicaridinae – but the latter
name remains unavailable until the provisions of ICZN Arts 13.1, 13.2, 16.1 and 16.2 are satisfied (Galassi & De
Laurentiis 2004: 426; Wells 2004: 96; Ranga Reddy & Defaye 2007: 17).
83. Chappuis (1929b: 475) proposed the new generic name Paracamptus Chappuis, 1929b (type species by original
designation: Canthocamptus schmeilii Mrázek, 1893) which proved a junior homonym of Paracamptus Casey,
1895 (Coleoptera). Özdikmen (2008: 268) replaced the junior homonym by Pesceus Özdikmen, 2008 (nomen
novum) which takes the same type species as its prior nominal taxon (ICZN Art. 67.8).
84. Mrázek (1893: 97) fixed Phyllognathopus paludosus Mrázek, 1893 as the type by monotypy but Hartwig (1896:
320) claimed that this species was identical with Belisarius viguieri Maupas, 1892 (type by monotypy of Belisarius
Maupas, 1892). Wilson (1924: 14) pointed out that Phyllognathopus Mrázek, 1893 takes priority over the
preoccupied Belisarius [a junior homonym of Belisarius Simon, 1879 (Arachnida)]. The generic name Viguierella
Maupas, 1892 is an unavailable name, being an unjustified manuscript correction in a reprint which was mistakenly
adopted by Perrier (1893: 995) [see also (Gurney 1932: 6)]. Being based on a nomen nudum, the family-group name
Viguierellidae proposed by Gurney (1928: 331) is likewise unavailable. Also note that P. laticauda Por, 1964a (as
P. labicauda [sic]) and P. medius Por, 1964a were erroneously assigned to Phyllognathopus (family
Phyllognathopodidae) instead of Phyllopodopsyllus T. Scott, 1906a (family Tetragonicipitidae) by Dussart and
Defaye (1990: 16).
85. Philippi (1840: 190) proposed the genus Thyone Philippi, 1840 for three species of which he named only one
(Thyone viridis Philippi, 1840). According to ICZN Art. 68.3 T. viridis is deemed to be fixed by monotypy,
regardless of whether the author considered the nominal genus-group taxon to contain other species which he did
not cite by name. Both names being objective synonyms, Philippi’s (1840: 190) type fixation by monotypy for the
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86.
87.
88.
89.
90.
91.
92.
93.
latter also applies to its new replacement name Porcellidium Claus, 1860 (ICZN Art. 67.8). Claus (1860: 6) pointed
out that Philippi’s (1840) generic name was a junior homonym of Thyone Oken, 1815 (Holothuroidea; type
Holothuria fusus Müller, 1776) and proposed the new replacement name Porcellidum Claus, 1860. A report
prepared by Hemming (1954) concluded that Oken’s (1815, 1816) Lehrbuch der Naturgeschichte was nonbinominal and therefore that new names published in it were unavailable. The work was rejected by the
International Commission on Zoological Nomenclature and placed on the Official Index (Opinion 417, September
1956), and consequently, as an unavailable name Thyone Oken, 1815 cannot enter into homonymy with Thyone
Philippi, 1840 (ICZN Art. 54.2). Although Oken’s (1815) genus is still cited – even in the recent holothuroid
literature (e.g. Solís-Marín et al. 2005; Thandar 2006) – this does not change its status of a previously unavailable
name (ICZN Art. 11.5.2). The first author who published the name Thyone in a way that satisfied the criteria of
availability was Goldfuß (1820: 177) who provided a definition and included two previously described species
(Holothuria fusus and Fistularia impatiens Forsskål, 1775). Although Oken was mentioned as the author, it was
Goldfuß who made the name available (the name of the author does not form part of the name of a taxon; ICZN Art.
51.1). Consequently, Thyone Philippi, 1840 becomes a permanently invalid name as a junior homonym of Thyone
Goldfuß, 1820. Contrary to previous authors who attributed the family-group name Porcellidiidae to either Sars
(1904b) (e.g. Lang 1948; Bodin 1997) or Brady (1880a) (e.g. Huys & Boxshall 1991; Huys et al. 1996), WalkerSmith (2001: 653) pointed out that the authorship and date should rest with Boeck (1865: 279) who coined the name
as “Afdeling Porcellidina”.
Por (1967: 143) proposed the generic name Ophirion Por, 1967 for Ophirion communis Por, 1967 (type by
monotypy) but this generic name proved to be preoccupied by its senior homonym Ophirion Townsend, 1911
(Diptera). Özdikmen (2008: 267) proposed the new replacement name Pordfus which takes the same type species
as its prior nominal taxon (ICZN Art. 67.8).
Chappuis (1953: 146), who had overlooked Nicholls’s (1945b: 21) proposal of the genus Psammopsyllus Nicholls,
1945b, established the same genus under a different name, Paulianicaris Chappuis, 1953. In a later paper Chappuis
(1954c: 273) classified it as incertae sedis in the Harpacticoida but was of the opinion that a new family may be
proposed, which he named “Paulianicaridae” (p. 275). However, since Chappuis (1954c) refrained from presenting
an explicit statement or formal diagnosis, this family name remained a nomen nudum in the harpacticoid
nomenclature and cannot be considered a synonym of the family-group name Cylindropsyllidae Sars, 1909b as
claimed by Boxshall and Halsey (2004: 851).
Sars (1911b: 400) proposed the new generic name Pseudameira Sars, 1911b for two new species (Pseudameira
crassicornis and P. furcata) but neither was fixed as the type. Lang’s (1948: 835) claim that Sars (1911b) had
designated P. crassicornis as “… Typus seiner neuen Gattung Pseudameira …” can be considered a subsequent
type designation according to ICZN Art. 69.1.1 even though there is no such explicit statement in Sars’s (1911b:
400–402) description of the genus.
Apostolov and Marinov (1988: 72) listed Pseudobradya brevicornis (T. Scott, 1894a), the earliest described species
included in Lang (1948), as the type but this species was not even originally included. Sars (1904a: 40) had created
the genus for six species of which he named five: Bradya minor Scott & Scott, 1896; Bradya elegans Scott & Scott,
1896; Bradya hirsuta Scott & Scott, 1896; Bradya similis Scott & Scott, 1896 and Pseudobradya acuta Sars, 1904a.
Sars (1910: 356; 1920a: 31) added a further 12 new species but the type issue remained unresolved. Bradya minor
is here fixed as the type by subsequent designation.
A. Scott (1896: 144) designated Pseudolaophonte aculeata A. Scott, 1896, which is a junior subjective synonym of
Laophonte spinosa Thompson, 1893 (cf. Sars 1911c: 429), as the type species.
T. Scott (1899b: 254) designated Psyllocamptus fairliensis T. Scott, 1899b, which is a junior subjective synonym of
Mesochra propinqua T. Scott, 1896 (cf. Lang 1948: 826), as the type species.
According to Lang (1948: 1621) Nicholls’s (August 1944: 492) family-group name Pteropsyllidae (type genus:
Pteropsyllus T. Scott, 1906a) is a junior subjective synonym of Lang’s (19 February 1944: 27) name
Tetragonicipitidae [Lang’s (1944) incorrect original spelling Tetragonicepsidae was emended by Wells (1967:
310)].
Wells and Rao (1987: 182) proposed the generic name Langia Wells & Rao, 1987 for its only originally included
species Langia maculata Wells & Rao, 1987 (type by original designation). Langia Wells & Rao, 1987 is a junior
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94.
95.
96.
97.
98.
99.
100.
101.
102.
103.
104.
105.
homonym of Langia Moore, 1872 (Lepidoptera) and was recently replaced by the new replacement name
Raowellsia Özdikmen, 2008 by Özdikmen (2008: 269), which takes the same type species as its prior nominal
taxon (ICZN Art. 67.8).
According to Lang (1948: 1621) Nicholls’s (1945c: 96) family-group name Remaneidae (type genus: Remanea
Klie, 1929) is a junior subjective synonym of the name Paramesochridae Lang, 1944.
The type species was named in honour of the Swedish zoologist Hans Brattström. According to ICZN Art. 32.5.2.1
the specific name brattstroemi is an incorrect original spelling that must be corrected to brattstromi.
Subsequent designation by Apostolov and Marinov (1988: 117) [see also Huys (1990b: 80)].
Wells (1985: 13) proposed Rosacletodes as a new replacement name for the preoccupied Echinocletodes Pallares,
1982a (family Huntemanniidae), which is a junior homonym of Echinocletodes Lang, 1936b (family
Ancorabolidae). Pallares’s (1982a: 26) type fixation by monotypy takes precedence over Wells’s (1985: 13) more
explicit fixation by original designation.
Wilson (1924: 14) proposed the new replacement name Sacodiscus for the preoccupied generic name Aspidiscus
Norman, 1869. Norman’s (1869: 298) fixation by monotypy consequently also applies to Sacodiscus (ICZN Art.
67.8). Aspidiscifera Strand, 1929, which was also proposed as a new replacement name for Aspidiscus (Strand
1929: 10), is a junior objective synonym of Sacodiscus.
Wilson (1924: 16) proposed Sarsameira as a new replacement name for the preoccupied Parameira Sars, 1907b,
which is a junior homonym of Parameira Seidlitz, 1868 (Coleoptera). Parameira Sars, 1907b was established to
accommodate Ameira parva Boeck, 1873 and a new species Parameira major Sars, 1907b. Sars’s (1907b: 222)
statement “… Boeck’s species [A. parva], which may be regarded as the type of the present genus” does not qualify
as a rigidly construed type designation (ICZN Art. 67.5.3). The type of Sarsameira was formally fixed by Wilson
(1924: 16) who designated A. parva – as Parameira parva Sars [sic]; hence this type fixation also applies to the
prior nominal taxon Parameira (ICZN Art. 67.8).
Wilson (1924: 22) proposed the new replacement name Sarsocletodes for the preoccupied generic name
Pseudocletodes Sars, 1920c. Sars’s (1920c: 89) original designation (by indication; ICZN Art. 68.2.2) consequently
also applies to Sarsocletodes (ICZN Art. 67.8).
Psamathe Philippi, 1840 and Scutellidium Claus, 1866 are objective synonyms. Claus (1866: 20) fixed Scutellidium
tisboides Claus, 1866 (by monotypy) as the type of Scutellidium; this species is a junior subjective synonym of the
type species of Psamathe (Psamathe longicauda Philippi, 1840, fixed by monotypy) (cf. Lang 1948: 387). Wilson
(1924: 16) pointed out that the generic name Psamathe Philippi, 1840 is preoccupied twice by Psamathe
Rafinesque, 1814 (Isopoda) and Psamathe Johnston, 1836 (Polychaeta) and hence Scutellidium is the valid name
for the taxon it denotes.
Sars (1909b: 323) established the genus Stenocaris to accommodate a new species S. gracilis Sars, 1909b and
further remarked on the striking similarity with the species described by T. Scott (1892) as Cylindropsyllus minor.
From the original diagnosis it is not clear whether Sars meant to include C. minor and, if so, which species he
designated as the type. In a later paper redescribing S. minor (T. Scott, 1892), Sars (1911c: 434) reiterated the close
relationship between both species by stating that Scott's species “... is evidently referable to the genus Stenocaris,
...., agreeing with the type species, S. gracilis, in all essential characters”. Since C. minor was doubtfully included it
does not qualify as an originally included species (ICZN Art. 67.2.5) and, consequently, Sars’s (1911c) statement
cannot be regarded as a formal subsequent designation. Sars’s (1909b) type fixation of S. gracilis was by
monotypy, rendering Apostolov and Marinov's (1988: 278) fixation of S. minor as the type species invalid. The
family-group name Stenocaridae proposed by Monard (1927: 171) is a junior subjective synonym of
Cylindropsyllidae Sars, 1909b.
Subsequent designation by Sars (1920b: 63).
The genus-group name Stygonitocrella Petkovski, 1976 (no original type fixation) was made available by Reid et
al. (2003: 997) by fixing S. montana Reid, Hunt & Stanley, 2003 as the type species and by providing an amended
generic diagnosis (ICZN Art. 13.1 and 13.3).
Huys (1993: 769) proposed the family-group name Styracothoracidae but this is now considered a junior subjective
synonym of Idyanthidae (Moura & Martínez Arbizu 2003: 178; Seifried 2003: 98).
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106. Subsequent designation by Lang (1948: 482). Thomson’s (1883: 98) spelling of the generic name is a lapsus calami
of Amymone Claus, 1860.
107. As pointed out by Giesbrecht (1881: 255), Lilljeborg’s (1853: 196) type and only species included in Tachidius was
misidentified as Cyclops brevicornis Müller, 1776 [= type of Tigriopus Norman, 1903a), and hence the incorrectly
applied original binomen, Tachidius brevicornis (Müller, 1776), had to be rejected and replaced by a new name,
Tachidius discipes Giesbrecht, 1881. Contrary to previous authors (e.g. Lang 1948; Seifried 2003) who attributed
the family-group name Tachidiidae to Sars (1909b), Wells (2007: 88) pointed out that the authorship and date must
remain with Boeck (1865: 256) who established it under the name “Afdeling Tachidina”.
108. Norman (1903a: 368) proposed the generic name Tegastes as a new replacement name for the pre-occupied
Amymone Claus, 1860, which is a junior homonym of Amymone Müller, 1776 (Crustacea), Amymone Savigny,
1822 (Polychaeta) and Amymone Agassiz, 1846 (Mollusca). Claus’s (1860: 11) type fixation by monotypy also
applies to Tegastes Norman, 1903a (ICZN Art. 67.8). Being based on a junior homonym (ICZN Art. 39), the
family-group names Amyonea Boeck, 1865 (based on the incorrect subsequent spelling Amyone; Boeck 1865: 255)
and its alternative “Amymoninae” used by Brady (1880a: 27) are invalid synonyms of the name Tegastidae Sars,
1904b.
109. Agreement in gender between genus and species-group names proposed by Lang (1948: 882).
110. Although generally attributed to Claus (1863: 128), who provided the first illustrated species descriptions, the genus
Thalestris was first diagnosed the year before (Claus 1862) but without any reference to originally included species.
New generic names published before 1931 which are accompanied by a description or a definition of the taxon (as
in Claus 1862: 88) that they denote (ICZN Art. 12.1) satisfy the criteria of availability and hence type fixation is not
mandatory nor is the use of one or more available specific names in combination with them, or clearly included
under them. Claus (1863) created the genus for seven new species (Thalestris robusta, T. mysis, T. microphylla, T.
helgolandica, T. longimana, T. forficula, T. harpacticoides) and three Harpacticus species described by Fischer
(1860) (Harpacticus fulvus = Tigriopus brevicornis, H. aquilinus = Thalestridae incertae sedis, H. spinosus =
Parastenhelia spinosa). Sars (1905b: 105) subsequently designated Thalestris longimana Claus, 1863 as the type.
111. Norman (1869: 296) designated Tigriopus Lilljeborgii Norman, 1869 (by monotypy), which is a junior subjective
synonym of Cyclops brevicornis Müller, 1776 (cf. Lang 1948: 311), as the type species.
112. Monard (1928: 438) proposed Nannopodella for Nannopodella denisi (incorrect original spelling Denisi) but also
remarked “… l’espèce Enhydrosoma minutum de Scott doit certainement rentrer dans notre nouveau genre”; neither
of these species was fixed as the type. Klie (1929: 374) and Lang (1936b: 477) remarked that the latter species
should be transferred to Rhizothrix Sars, 1909a as R. minuta (T. Scott, 1903a). Since elimination of all but one of
the originally included nominal species from a nominal genus does not in itself constitute type fixation (ICZN Art.
69.4), Nannopodella denisi Monard, 1928 is here fixed as the type species.
113. Fixed as type by monotypy in the subgenus Cletodes (Pseudocletodes) by Scott and Scott (1893b: 239) (ICZN Art.
61.2.2).
114. Herrick (1895: 41) proposed the genus Marshia for two new species, M. albuquerquensis Herrick, 1895 and M.
brevicaudata Herrick, 1895. The latter is here fixed as the type species by subsequent designation; the former was
previously transferred to the genus Cletocamptus Shmankevich, 1875 (Lang 1936b: 473).
115. Dahms and Pottek's (1992: 28) original type designation for Talpina Dahms & Pottek, 1992 [a junior homonym of
Talpina Hagenow, 1840 (Phoronidea)] also applies to its new replacement name Dahmsopottekina Özdikmen, 2009
(ICZN Art. 67.8).
116. Mu and Huys's (2002: 205) original type designation for Hicksia Mu & Huys, 2002 [a junior homonym of Hicksia
Delgado, 1904 (Trilobita)] also applies to its new replacement name Muohuysia Özdikmen, 2009 (ICZN Art. 67.8).
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