Zootaxa,A revision of the family Typhlotanaidae Sieg ...

ZOOTAXA 1598

A revision of the family Typhlotanaidae Sieg 1984 (Crustacea: Tanaidacea) with the remarks on the Nototanaidae Sieg, 1976 MAGDALENA BŁAŻEWICZ-PASZKOWYCZ

Magnolia Press Auckland, New Zealand

MAGDALENA BŁAŻEWICZ-PASZKOWYCZ A revision of the family Typhlotanaidae Sieg 1984 (Crustacea: Tanaidacea) with the remarks on the Nototanaidae Sieg, 1976 (Zootaxa 1598) 141 pp.; 30 cm. 28 Sept. 2007 ISBN 978-1-86977-163-8 (paperback) ISBN 978-1-86977-164-5 (Online edition)

FIRST PUBLISHED IN 2007 BY Magnolia Press P.O. Box 41-383 Auckland 1346 New Zealand e-mail: [email protected] http://www.mapress.com/zootaxa/

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BŁAŻEWICZ-PASZKOWYCZ

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ZOOTAXA

A revision of the family Typhlotanaidae Sieg 1984 (Crustacea: Tanaidacea) with the remarks on the Nototanaidae Sieg, 1976 MAGDALENA BŁAŻEWICZ-PASZKOWYCZ Department of Polar Biology and Oceanobiology, University of ºódï, Banacha 12/16, 90-237 POLAND, [email protected]

Table of contents Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Material and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Family: Nototanaidae sensu lato . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Genus: Meromonakantha Sieg, 1986 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Meromonakantha natatoris n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Genus: Paratyphlotanais Kudinova-Pasternak & Pasternak, 1978 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 Family: Typhlotanaidae Sieg, 1984 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 Key for Typhlotanaidae genera and morpho-groups . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 Genus: Typhlotanais G.O. Sars, 1882 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 Typhlotanais aequiremis (Lilljeborg, 1864) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 Genus: Hamatipeda n. gen. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35 Hamatipeda longa (Kudinova-Pasternak, 1975) n. comb. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35 Hamatipeda trapezoida n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37 Genus: Larsenotanais gen. n. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41 Larsenotanais amabilis n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42 Genus Peraeospinosus Sieg, 1986 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46 Peraeospinosus acruxi n. sp.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47 Genus: Pulcherella n. gen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51 Key for determination of Pulcherella females . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52 Pulcherella filatovae (Kudinova-Pasternak, 1975) n. comb. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52 Pulcherella pulcher (Hansen, 1913) n. comb. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56 Pulcherella spiniventris (Dollfus, 1897) n. comb. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60 Pulcherella juraszi n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61 ? Pulcherella pulcher Hansen, 1913 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65 Genus: Torquella n. gen. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65 Key for identification of Torquella females . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66 Torquella angularis (Kudinova-Pasternak, 1966) n. comb. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66 Torquella elegans (Kudinova-Pasternak, 1978) n. comb. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67 Torquella grandis (Hansen, 1913) n. comb. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71 Torquella longisetosa (Kudinova-Pasternak, 1990) n. comb. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73 Torquella parangularis (Kudinova-Pasternak 1975) n. comb. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78 Torquella rotundirostris (Lang, 1970) n. comb . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78 Torquella eltaninae n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80 Torquella galatheae n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83 Genus: Typhlamia n. gen. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86 Key for identification of Typhlamia females . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87

Accepted by Kim Larsen: 3 Sept. 2007; published: 28 Sept. 2007

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Typhlamia mucronata (Hansen, 1913) n. comb. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87 Typhlamia sandersi (Kudinova-Pasternak, 1985) n. comb. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90 Typhlamia bella n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94 Typhlotanais sensu lato . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98 ‘greenwichensis’ group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98 Typhlotanais greenwichensis Shiino, 1970 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98 Typhlotanais messinensis G.O. Sars 1882 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103 ? Typhlotanais greenwichensis Shiino, 1970 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107 ‘mixtus’ group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108 Typhlotanais mixtus Hansen, 1913 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108 Typhlotanais mimosis n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112 ‘spinicauda’ group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116 Typhlotanais spinicauda Hansen, 1913 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117 Typhlotanais squamiger n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121 ‘cornutus’ group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 126 Typhlotanais andeepae n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 126 ‘plicatus’ group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131 Typhlotanais plicatus Kudinova-Pasternak, 1993 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131 ‘eximius’ group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132 Typhlotanais sp. A . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132 Tube building ability of nototanaids and typhlotanaids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136 Remarks on world-wide Tanaidacea distribution in deep-water . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136 Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137 Literature cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137

Abstract Recent tanaidacean material collected from Antarctic waters, primarily during the ANDEEP expeditions of 2002 and 2005, includes a number of new taxa attributable to the families Nototanaidae and Typhlotanaidae sensu Sieg. Analysis of this material has exposed a problem with the recent contention of the two families, and has revealed consistent morphological trends which support the distinction of these two families. In the present paper, examination of both museum specimens and newly-collected material, has allowed a re-analysis based on a series of detailed morphological observations, resulting in a new definition of the families Typhlotanaidae Sieg, 1984 with the establishment of five new genera (Hamatipeda n. gen., Larsenotanais n. gen., Pulcherella n. gen., Torquella n. gen., Typhlamia n. gen.), a the description of thirteen new species, the redescription of fifteen species, and the construction of keys for the determination of typhlotanaid genera and of the species of three newly-erected genera. Key words: Tanaidacea, Nototanaidae, Typhlotanaidae, ANDEEP, Antarctic, abyssal, Hamatipeda, Larsenotanais, Meromonakantha, Paratyphlotanais, Peraeospinosus, Torquella, Typhlamia, Typhlotanais, Pulcherella

Introduction Recent tanaidacean material collected from Antarctic waters, primarily during the ANDEEP expeditions of 2002 and 2005, includes a number of new taxa attributable to the families Nototanaidae and Typhlotanaidae sensu Sieg (see below). Analysis of this material has exposed problems with recent suggestions that the two families should be united, and has revealed consistent morphological trends which support the distinction of these two families. In order to undertake a sensible classification of these taxa, it has been necessary to reexamine the genera attributed to these families in detail, based on as much material, including types, as is presently feasible.



In their preliminary cladistic analysis of the Paratanaoidea, Larsen & Wilson (2002) combined the two families Nototanaidae Sieg, 1976 and Typhlotanaidae Sieg 1984 into one family, uniting eleven genera into their redefined Nototanaidae: Bathytanaissus Bird & Holdich, 1989, Metatanais Shiino, 1952, Nesotanais Shiino, 1968, Nototanais Richardson, 1906, Paratyphlotanais Kudinova-Pasternak & Pasternak, 1978, Peraeospinosus Sieg, 1986, Protanaissus Sieg, 1982, Tanaissus Norman & Scott, 1906, Teleotanais Lang, 1956, Typhlotanais G.O. Sars, 1882, and Typhlotanaoides Sieg, 1983a. The other genus included by Sieg (1986b) in the Typhlotanaidae, Meromonakantha, was classed as incertae sedis by Larsen & Wilson (2002) and subsequently moved into the Nototanaidae by Larsen (2005). As has been pointed out by Bamber (2005), the conclusions of Larsen & Wilson’s (2002) cladistic analysis, which failed to resolve the familial associations of numerous genera, must be regarded only as a preliminary hypothesis. Owing to synapomorphies within the Nototanaidae-Typhlotanaidae, at present a clear phylogenetic analysis is not feasible based purely on the available morphological information, and it is therefore appropriate to keep with the earlier classification of these families (e.g. sensu Sieg, loc. cit.). The Nototanaidae sensu Larsen & Wilson (2002) aggregated forms with a three-articled antennule and six-articled antenna, a labium without medial spiniform setae, no reduction of pereonite-1, a marsupium formed of four pairs of oostegites, no articulated setae on the pleonites, and with a fused coxa and with the dactylus fused with the unguis on pereopods 4 to 6. The remaining characters given in the diagnosis of Larsen and Wilson (ibid.) are ambiguous and have only minor relevance in discrimination of family level (eyes present or absent, antenna article-3 with or without dorsal spiniform setae, molar process broad or pointed, maxilliped endites fused or not fused). It is apparent that the inconclusive results obtained by Larsen and Wilson (2002) were a consequence of including into their analysis a number of poorly described taxa, e.g. Paratyphlotanais (see Bird, 2004a) or Peraeospinosus (see Błażewicz-Paszkowycz, 2005), as well as inappropriate character-states obtained from the scant literature. One such character largely responsible for forcing the typhlotanaids into a group with the nototanaids was the fusion of the dactylus-unguis. The classification of these two families sensu Larsen & Wilson (2002) clearly requires re-analysis, through careful examination and redefinition of all the included taxa. The severity of the task is emphasized by the existence of nearly a hundred species that need to be redescribed, and by the difficulty in obtaining the type material. Many of those species were described in the early 20th century from only a single specimen, and their superficial descriptions are inadequate for taxonomical analysis. When available, some holotypes are in poor condition or covered by crystalline deposits that make further observation impossible. Many typhlotanaid species which were described by Kudinova-Pasternak between the years 1966 and 1993 now remain only as incomplete descriptions and as type specimens either in bad condition or lost. In such circumstances the detailed phylogenetic analysis proposed is unfeasible for the present. Despite this, the present study, on both museum material and newly-collected material, has undertaken a reanalysis based on a series of detailed morphological observations, resulting in a new definition of the family Typhlotanaidae Sieg, 1984, distinguishing it from the family Nototanaidae Sieg, 1976, in the establishment of five new genera, the description of thirteen new species, the redescription of fifteen species, and the construction of keys for the determination of typhlotanaid genera and of the species of three newly-erected genera, Pulcherella n. gen. Typhlamia n. gen. and Torquella n. gen., all of which are presented below. Species sharing a set of unique characters are provisionally gruped in ‘morpho-groups’. It is assumed they may constitute valid new generae and it will be tested when more new species are described. In the result of the present studies most typhlotanaids have got a new generic range or were classified to one of the stated a morpho-group. Some others (T. angusticheles Kudinova-Pasternak, Typhlotanais assimilis G.O. Sars, 1882, T. brevicornis Lilljeborg, 1864, T. kussakini Kudinova-Pasternak 1970, T. longicephala Kudinova-Pasternak 1970, T. proctagon Tattersal, 1904 T. compactus Kudinova-Pasternak, 1966, T. plebejus, Hansen 1913, T. profundus Hansen,1913, T. simplex, Kudinova-Pasternak 1984, T. williamsae Dojiri & Sieg, 1997) weren't classified to any of the established genera or perceived morpho-groups usually due to unavailability of the material. The summary of this results are presented in Table 1.

A REVISION OF FAMILY TYPHLOTANAIDAE SIEG 1984

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TABLE 1. Classification of the nototanaids and typhlotanais. genus/morpho–group

species NOTOTANAIDAE

Bathytanaissus

B. spinulosus

Meromonakantha

M. guilei; M. irregularis; M. macrocephala sensu (Hansen, 1913); M. macrocephala sensu (Sieg, 1986); M. setosa; M. natatoris n. sp.; M. nutae n. sp.

Nesotanais

N. lacustris; N. maclaughlinae; N. rugula

Nototanais

N. antarcticus; N. dimorphus

Paratyphlotanais

P. armatus; P. gracilipes; P. japonicus; P. mananensis; P. microcheles; P. pectinatus; P. richardi; P. typicus; P. alveolus n. sp.

Protanaissus

P. alvesi; P. floridensis; P. longidactylus; P macrotrichos

Tanaissus

T. danica; T. lilljeborgi; T. psammophilus TYPHLOTANAIDAE

Hamatipeda

H. longa; H. trapezoida n. sp.

Larsenotanais

L. amabilis n. sp.

Obesutanais

O. sigridae

Peraeospinosus

P. brachyurus; P. emergensis; P. exiliremis; P. kerguelenensis; P. magnificus; P. magnus; P. peculiaris, P. peculiaroides; P. pushkini; P. rectus; P. subtigaleatus; P. acruxi n. sp.

Pulcherella

P. pulcher; P. filatovae; P. spiniventris; P. juraszi n. sp.

Typhlotanais

T. aequiremis

Typhlotanaoides

T. insolitus; T. rostralis

Torquella

T. angularis; T. elegans; T. grandis; T. longisetosa; T. parangularis; T. rotundirostris; T. eltaninae n. sp.; T. galatheae n. sp.

Typhlamia

T. mucronata; T. sandersi; T. bella n. sp.

‘greenwichensis’ group

Typhlotanais greenwichensis; T. messinensis

‘mixtus’ group

Typhlotanais mixtus; T. mimosis n. sp.

‘spinicauda’ group

Typhlotanais spinicauda, Typhlotanais squamiger n. sp.

‘cornutus’ group

Typhlotanais cornutus; ?T. adipatus sensu Tzareva, 1982; ? T. crassus, T. andeepae n. sp.

‘plicatus’ group

Typhlotanais longimanus; T. plicatus; ?T. variabilis

‘eximius’ group

Typhlotanais eximius; T. spinipes; Typhlotanais sp. A.

‘trispinosus’ group

Typhloatnais trispinosus; T. tenuicornis

non classified

T. assimilis; T. compactus; T. finmarchicus; T. inaequipes; T. inermis; T. kussakini; T. longicephala; T. plebejus; T. proctagon; T. profundus; T. simplex; T. solidus; T. williamse

Material and methods Material (including types) analyzed for this study has been loaned from the Zoological Museum University Copenhagen (Denmark), the Zoological Museum, University of Lomonosow (Moscow, Russia), the Zoologisches Institut und Museum der Universität Hamburg (Germany), the National Museum of Natural History, the Smithsonian Institute (Washington DC), Icelandic Museum of Natural History (Reykjavik), and the Museum of Oceanography (Monaco). Most of the new species described in this paper were collected during ANDEEP Expedition I and II in the austral summer of 2002 and the following ANDEEP III from 21 January to 6 April 2005. The fieldwork was conducted on board the German icebreaker Polarstern using various zoobenthic sampling gears including a



Multibox corer, Agassiz trawl and epibenthic sledge. The samples were taken at numerous transects in the Scotia Sea, the Weddell Sea, along the Antarctic Peninsula and along the Scotia Arc (http://www.biologie.unihamburg.de/zim/niedere2/cruise_report.pdf). Specimens were examined with Nikon SMZ 800 dissecting and Zeiss Axiolab compound microscopes. Dissected appendages were stained with chlorazol black and mounted on slides in glycerin jelly. Body length was measured from the tip of the rostrum to the end of the pleotelson. Typhlotanaids of arbitrary length: width ratio 6 are called short and long respectively. In case of those taxa with a simple pereon outline (e.g. rectangular pereonites in Typhlotanais spinicauda Hansen, 1913) there was no problem in defining the average width. In the case of those with a complex pereon outline (convex/concave lateral margins) (e.g. Meromonakantha natatoris n. sp.) the average body width was subjectively discriminated. Length of the pleon was measured from the anterior of pleomere-1 to the tip of the pleotelson. The length and width of particular articles in the cheliped were measured as shown in the figure 1A. The morphologic terminology generally follows that proposed by Larsen (2003) and Błażewicz-Paszkowycz (2005). The term “clinging pereopods” (Fig. 1D–H) is used for the last three pairs of thoracic appendages when they are adapted for holding the animals inside tubes or burrows: they have a reduced coxa, widened basis, short merus and carpus, and are armored with specialized structures such as ‘prickly tubercles’ or serrated/setulated hooks on the carpus (Plate 1A). The term ‘microtrichia’ (‘comb of setae’ in Błażewicz-Paszkowycz, 2005) is adopted after Bird (2004b) and refers to the small, scale-like, cuticular structures with a distal fringe of spines or small setae (Plate 1B). The round, small distal structures on the maxilliped endites are called tubercles (Fig. 1C); it is not known whether they are homologous with the “flat setae” found in the Leptocheliidae (Fig. 1B), but because of their different morphology a different name is proposed. The terms ‘manca I, II and III’ are accepted after Larsen (2003) while the term ‘neuter’ is used according to the definition given by Bird (2004b). Type material of the species described in this paper has been deposited in Zoological Museum University Copenhagen (Denmark), Zoologisches Institut und Museum der Universität Hamburg (Germany), and National Museum of Natural History, Smithsonian Institute (Washington DC). The literature list cited through the paper has been prepared with help of two web pages by Anderson et al. (2006) and Anderson (2005). The synonymy of the species is based mostly on Sieg (1983), Bird (2004a) and Larsen (2005).

Results From the eleven genera assigned by Larsen & Wilson (2002) to the Nototanaidae (Bathytanaissus, Metatanais, Nesotanais, Nototanais, Paratyphlotanais, Peraeospinosus, Protanaissus, Tanaissus, Teleotanais, Typhlotanais, and Typhlotanaoides, only three (Typhlotanais sensu lato, Typhlotanaoides and Peraeospinosus) have prickly tubercles (Fig. 1D–G) on the clinging pereopods 4–6. Natatory males have been identified a few times in Typhlotanais and Peraeospinosus (Sieg, 1986a; Błażewicz-Paszkowycz, 2004; 2005), although they have not yet been found in Typhlotanaoides. Natatory males sharing no specific character with females have not been recognized as the same species and in consequence might have been omitted in many taxonomical studies. The other hypothesis assumes a lack of full terminal males (natatory/swimming), instead the existence of a form called ‘preparatory males’. This form closely resembles the female but for a thicker antennule and sometimes better developed pleopods. It is assumed that ‘preparatory’ males are sexually mature although dissection or sectioning to look for testes is required for confirmation (Bird pers. comm.).



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FIGURE 1. A) The measurements of the cheliped; B) Maxilliped endites in the Leptocheliidae with flat setae; C) Maxilliped endites in the Typhlotanaidae with tubercles; D–H) Variety of prickly tubercles in pereopods 4–6 of Typhlotanaidae.



PLATE 1. A) Peraeospinosus kerguelenensis, pereopod-6 carpus with clinging apparatus; B) Microtrichia (scale-like, cuticular structures with a distal fringe spines or small setae) on maxilliped endites of Typhlotanais greenwichensis.

Based on the presence of prickly tubercles on pereopods 4–6 and a combination of other consistent features (see diagnosis of Typhlotanaidae below) the family Typhlotanaidae is re-erected here, to include three existing genera (Typhlotanais, Typhlotanaoides, and Peraeospinosus), three new genera derived from Typhlotanais sensu lato (present work), Obesutanais Larsen, Błażewicz-Paszkowycz & Cunha, 2006, and Hamatipeda n. gen. These last two genera have specialized hooks on pereopods 4–6 replacing the prickly tubercles; as both of these taxa share more characters with the Typhlotanaidae than with the Nototanaidae, they are placed in the Typhlotanaidae, although a new family may be established for them in the future. As a consequence, the rest of the ‘nototanaid’ genera are included in a revised Nototanaidae. Charbeitanais Bamber & Bird, 1997 that was preliminary included to the family was suggested as a possible synonym of Pseudozeuxo and transferred to the Pseudozeuxidae Bird & Bamber (2000).

Family: Nototanaidae sensu lato Diagnosis (modified after Larsen &Wilson, 2002). Female: Antennule three-articled. Antenna six-articled. Cheliped with side piece. Pereopods 4-6 of the walking type. Marsupium formed from four pairs of oostegites. Male (where known): terminal (non-feeding) type. Genera included: Bathytanaissus; Nesotanais; Nototanais; Protanaissus; Tanaissus; Paratyphlotanais, and Meromonakantha. Remarks: Undoubtedly the Nototanaidae is a polyphyletic taxon and perhaps two clear phylogenetic lines can be observed. The monotypic Teleotanais gerlachi Lang, 1956, with a four-articled antennule, has been probably wrongly assigned to the family and, following the family definition, cannot be included. The first phylogenetic line embraces nototanaid genera with the cheliped attached via a sclerite and with a bifid



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right mandible incisor with crenulated upper margin (Bathytanaissus, Nesotanais, Nototanais, Protanaissus, Tanaissus). Noreworthy if that three of those five genera have ‘terminal males’, with reduced mouthparts and strongly modified chelipeds (adult males Bathytanaissus and Protanaissus are currently unknown). The ‘typhlotanaid-like’ Paratyphlotanais and Meromonakantha may form another phylogenetic line, being species with a cheliped lacking a sclerite and with a simple incisor. There is scarce information about males within those genera; Bird (2004a) suggested that they may be morphologically similar to the females and often overlooked. The most evident characters distinguishing true Nototanaidae (first phylogenetic line) from true Typhlotanaidae is the attachment of the cheliped (via a sclerite in the Nototanaidae and without a sclerite in the Typhlotanaidae) and the walking morphology of the last three pair of pereopods in the Nototanaidae versus the clinging type in the Typhlotanaidae. Two further differences between these families regard the structure of the mouthparts. The upper part of the mandible incisor is bifid and serrated in the Nototanaidae, but in the Typhlotanaidae it is simple and smooth. The other character applies to the distal part of maxillule endite: in the Nototanaidae this appendage is bent almost at a right angle distally and usually its outer margin is covered by dense setae, but in the Typhlotanaidae it is almost straight and naked. The shape and setation of the maxillule and associated mouthparts, although difficult for discrimination or observation (the maxilla is often damaged or lost during dissection), are probably important features which should be considered more when attempting to reconstruct tanaidomorph phylogeny.

Genus: Meromonakantha Sieg, 1986 Meromonakantha: Sieg (1986b) 2(4): 72; Larsen (2005) 5: 202–203.

Diagnosis (modified after Larsen, 2005): Female. Body almost completely cylindrical in cross-section. Carapace wider than pereonites. Pereonites 2–5 hexagonal or round, never rectangular or square; well separated from one another. Pleon slightly wider than pereon in dorsal view. Antennule three-articled, article-1 stout (twice as long as wide). Antenna with long setae on articles 2 and 3. Molar process of mandible crenulated, with several blunt tubercles. Maxillule with seven or eight spiniform terminal setae. Maxilliped bases partially fused, endites not fused, each with two tubercles distally. Chelipeds not attached via a sclerite, not reaching the anterior edge of pereonite-1. Pereopods 1–3 with coxa; merus and carpus with simple setae only. Pereopods 4–6 of walking type, with coxa; merus and carpus with at least one strong curved spiniform seta; dactylus and unguis (semi-) fused. Pleopods present. Uropods biramous; endopod and exopod with two articles (occasionally semifused). Male: Unknown. Gender of generic name: Feminine. Type species: Typhlotanais macrocephala Hansen 1913. Species included: Meromonakantha guilei Larsen, 2005; M. irregularis (ex Typhlotanais) (Hansen, 1913); M. (ex Typhlotanais) macrocephala sensu (Hansen, 1913); M. (ex Typhlotanais) macrocephala sensu (Sieg, 1986b); M. (ex Typhlotanais) setosa (Kudinova-Pasternak, 1966); M. natatoris n. sp.; and M. nutae n. sp. Remarks: As with the other nototanaids, Meromonakantha shows conservative morphology within the genus and strongly reduced setation. While the conservative morphology seems to be convenient in recognition of genera, the reduced setation is responsible for the scarcity of discernible species characteristics that enhance the difficulty in creating diagnoses. A full revision of this genus is impossible at present owing to only having a single specimen of Meromonakantha (=Typhlotanais) macrocephala sensu (Hansen, 1913) and a lack of types of M. setosa. The same reasons preclude constructing a key for identification of the Mero-



monakantha species. In spite of this, it can be assumed that the characteristic body habitus (i.e. carapace and pleon wider than pereonites, hexagonal or oval pereonites 3–5), stout antennule, characteristic spiniform (usually hook-like) setae on pereopods 4–6, and (semi-) fused unguis and dactylus of pereopods 4–6 are characters distinctive enough to consider Meromonakantha a valid genus. The distribution of M. macrocephala in both Antarctic and North Atlantic is controversial. A large carapace, as long as the first two and half of the third pereonites, and the serrated pereonites of M. macrocephala sensu Hansen are distinctive characters that suggest separating the species from M. macrocephala sensu Sieg. This Antarctic taxon was used by Sieg (op.cit) to redescribe the species, the type locality of which is actually off south-west Iceland. Because only a single type specimen Hansen’s species exists, I have avoided redescribing these two taxa and seeking characters to unequivocally distinguish them. If M. macrocephala is a complex of two species, the modified genus Meromonakantha presently contains the following species: M. guilei, M. irregularis, M. macrocephala (sensu Hansen), M. macrocephala (sensu Sieg), M. setosa, M. natatoris n. sp. and M. nutae n. sp. From those seven (nominally six) species the most distinctive is M. natatoris (see remarks under species description). An unusual character is observed in the maxilliped palp of M. guilei: it has two, three and four setae on articles 2–4 respectively while the other Meromonakantha species (as well as all other nototanaids except Protanaissus) have three, four and six setae respectively. Reduced setation in M. guilei is also found in the maxillule which has only six distal spiniform setae instead of the eight or nine commonly occurring in nototanaids.

Meromonakantha natatoris n. sp. (Figs 2–4) Material examined: Holotype: female, (K 41350), ANT XXII/3, PS 67/81-8-E, 70°32.02'-70°32.19'S, 14°35.05'–14°35.13'W, depth 4392–4385 m, epibenthic sledge, 24 Feb 2005; Paratypes: one female (K 41351), the same locality; one female dissected on slides, (K 41352), ANT XXII/3, PS 67/81-9, 70°32.94'– 70°33.15'S, 14°34.40'–14°32.74'W, depth 4390–4392 m, Agassiz trawl, 24 Feb 2005. Diagnosis: Body long, 7.5 times as long as wide. Carapace 1.4 times as long as wide. Pereonite-1 shorter than pereonite-2, with proximal seta on each side. Pleon as long as pereonites 4–5 combined, each pleomere with single lateral seta. Antennule article-3 three times as long as article-2. Chela clearly shorter than cheliped carpus. Pereopod-1 propodus longer than merus and carpus combined. Pleopod exopod normal or reduced to oval squama. Uropod exopod clearly shorter than endopod. Etymology: Natator (Latin) = ‘swimmer’. Named after its “swimming” habitus. Description: Non-ovigerous female. Body length 1.5 mm (Fig. 2A, B), 7–8 times as long as wide. Cephalothorax 20% of overall length, 1.4 times as long as wide. Pereonite-1 three times as wide as long, with single lateral seta and with strong sternal hook; pereonites 2–6 of similar size, 1.2 times as long as wide; pereonite-2 trapezoidal, narrower caudally; pereonites 3–4 hexagonal; pereonites 5–6 trapezoidal, narrower proximally. Pleon almost 20% of body length, each pleomere with single lateral seta. Antennule (Fig. 3A): Little shorter than carapace; article-1stout, twice as long as wide, with two long setae medially, one long seta distally and with three pinnate setae distally; article-2 with two long (reaching beyond end of article-3) simple setae and one pinnate seta distally; article-3 three times as long as article-2, with terminal spur and four setae distally. Antenna (Fig. 3B): Article-1 short; article-2 twice as long as article-3, with one long (reaching beyond half of article-4) and one short simple setae; article-3 with long (reaching end of article-4) simple seta; article4 1.5 times as long as article-5, with two short simple setae and three pinnate setae distally; article-5 with single distal seta; article-6 with five simple setae.



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FIGURE 2. Meromonakantha natatoris n. sp., holotype, adult female. A) lateral view; B) dorsal view. Scale = 0.1 mm.

Mouth parts: Labrum (Fig. 3C) hood-shaped, covered by numerous minute setae. Mandible (Fig. 3D, E) stout; molar process well-developed, with crenulated, well-calcified edges; lacinia mobilis well-developed, crenulated. Maxillule and maxilla lost during dissection. Maxilliped (Fig. 3G) coxa absent; bases fused, without seta; endite with seta and two tubercles distally; palp four-articled, article-1 naked; article-2 wedge-shaped with three strong short setae on inner margin and one minute seta on outer margin; article-3 elongated, with four strong setae on inner margin, article-4 slender (over twice as long as wide) with six strong setae terminally. Cheliped (Fig. 3H): Basis 1.6 times as long as wide; merus triangular, with long ventral seta; carpus almost three times as long as broad, with two dorsal setae and two setae ventrally, one of which is twice as long as the other; propodus and fixed finger shorter than carpus, about half as broad as long; dactylus about



half length of whole; fixed finger inner margin well-calcified, with three setae distally and with two simple setae ventrally.

FIGURE 3. Meromonakantha natatoris n. sp., female paratype. A) Antennule; B) Antenna; C) Labrum; D) Left mandible; E) Right mandible; F) Labium; G) Maxilliped; H) Cheliped. Scale: A, B, H = 0.1 mm; C–G = 0.01 mm.



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Pereopod-1 (Fig. 4A): Of walking type; basis shorter than rest of articles combined, with simple seta proximally; ischium short with one seta; merus as long as carpus, with one seta distally; carpus with four setae distally; propodus longer than merus and carpus combined length, with one dorsal and one ventrally setae subdistally; dactylus and unguis combined 0.7 times as long as propodus; unguis almost twice as long as dactylus.

FIGURE 4. Meromonakantha natatoris n. sp., female paratype. A) Pereopod-1; B) Pereopod-2; C) Pereopod-3; D) Pereopod-5; E) Pereopod-6; F) Pleopod with rudimental exopod; G) Pleopod with regular exopod; H) Uropod. Scale = 0.1 mm.



Pereopod-2 (Fig. 4B): Of walking type; coxa with one simple seta; basis shorter than rest of articles combined, naked; ischium with one seta; merus as long as carpus, with one simple and one spiniform setae distally; carpus with three spiniform setae distally; propodus longer than merus and carpus combined, with one short ventral seta and one long dorsal seta sub-distally; dactylus and unguis combined half as long as propodus. Pereopod-3 (Fig. 4C): Similar to pereopod-2, but propodus a little longer than carpus, both propodal setae short. Pereopod-4 broken. Pereopod-5 (Fig. 4D): Basis walking type, 2.5 times as long as wide, with two pinnate setae ventrally; ischium with two setae; merus almost as long as carpus, with strong spiniform seta distally; carpus with two spiniform setae (small and large) and three setae (two long, one short) distally; propodus a little longer than carpus with one spiniform seta ventrally; dactylus broken. Pereopod-6 (Fig. 4E): Similar to pereopod-4; dactylus four times as long as unguis. Pleopods 1–5 (Fig. 4G): Basal article naked. Endopod with eighteen setae. Exopod four times as long as wide, with seven pinnate outer setae and one inner seta. Large gap between most proximal seta and the others in both rami. Uropod (Fig. 4H): Endopod two-articled; proximal article 0.6 times length of ramus, with seta distally; distal article tipped by four long seta. Exopod two-articled, a little longer than endopod proximal article with one seta distally; distal article tipped by one long and one short setae. Distribution: Antarctic: Eastern Weddell Sea, at abyssal depths of 4385–4392 m. Remarks: One of three specimens of Meromonakantha natatoris n. sp. had the exopod of the pleopods vestigial (Fig. 4F) while the two others had them normally developed (Fig. 4G). As all studied specimens had the same body length, the presence of residual pleopods in the only one of them cannot be explained by age of the animals. Also no gonopores or cones could be observed in any of those three specimens so the character cannot be explained by sexual dimorphism. Either these are three females, one of which has incompletely developed pleopods, or one is an immature animal at a moult stage just before maturity. The slender, relatively long (slightly shorter than carapace) antennule, narrow pereonite-1, long pereopod1 propodus, and relatively short uropod exopod, distinguish Meromonakantha natatoris n. sp. from the other members of the genus. A controversial character is the spiniform setae on merus and carpus of pereopods 2 and 3 in the new species, where the other species usually have only simple setae. The peculiar body habitus, hook-like setae on merus and carpus of pereopods 4–6, presence (in specific stages) of large hyposphaenium (pereonal hook) and slender uropods support including the new species in Meromonakantha.

Meromonakantha nutae n. sp. (Figs 5–7) Material examined: Holotype: one female, (K 41353), ANT XXII/3, PS 67/151-7-E, 61°45.52'–61°45.42'S, 47°7.68'–47°8.04'W, depth 1182–1185 m, epibenthic sledge, 21 Mar 2005; Paratypes three females (one damaged, one dissected on slides), (K 41354), the same locality; one female, (K 41355), ANT XXII/3, PS 67/1506-E, 61°48.70'–61°48.57' S, 47°28.04'–47°28.19' W, depth 1996–1993 m, epibenthic sledge, 20 Mar 2005. Diagnosis: Carapace as long as pereonite-1 and a quarter of pereonite-2 combined. Antennule article-3 less than twice as long as article-2. Pereopods 1 and 2 propodus shorter than merus and carpus combined. Pleopod endopod with four-five, exopod with six-seven distal setae. Uropod exopod a little shorter than endopod, articles semifused. Etymology: Nut is the Egyptian Goddess presented as an elongated woman bending over the Earth and



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touching the horizons with her toes and finger tips. The name reflects the bending body habitus of the preserved animal. Description: Female 1.2 mm long (Fig. 5A,B). Body long, about eight times as long as wide; carapace as long as pereonite-1 and a quarter of pereonite-2 combined; pereonite-1 as long as pereonite-2; pereonite-2 subequal to pereonite-4; pereonite-3 subequal to pereonite-6, pereonite-5 longest; pleon as long as pereonites 5 and 6 combined.

FIGURE 5. Meromonakantha nutae n. sp., holotype, adult female. A) lateral view. Scale 0.1 mm.

Antennule (Fig. 6A): 0.6 times as long as carapace; article-1 stout, twice as long as wide, with long simple setae in middle and distal, and four pinnate setae along article; article-2 with two long simple setae distally; article-3 less than twice as long as article-2, with terminal small spur and three setae distally. Antenna (Fig. 6B): Article-1 short; article-2 twice as long as article-3, with one long seta, article-3 with short simple seta; article-4 three times as long as wide, twice as long as article-5 with one simple and two pinnate setae distally; article-5 with single seta distally; article-6 short, with four simple setae. Mouth parts: Labrum (Fig. 6C) hood-shaped, covered by numerous minute setae. Mandible (Fig. 6D,E) stout; molar process well-developed, with crenulated, well-calcified edges and with blunt teeth in lower margin; lacinia mobilis well-developed, crenulated. Maxillule (Fig. 6F) with eight terminal spiniform setae, palp (Fig. 6F’) with two terminal setae. Maxilla (Fig. 6G) subtriangular, naked. Maxilliped (Fig. 6I): coxa reduced; bases fused, a little longer than wide, with seta just reaching end of endites; endite with two setae and two tubercles distally; palp 4-articled: article-1 naked; article-2 with three setae on inner margin and one seta on outer margin; article-3 with four simple setae on inner margin; article-4 slender with five inner setae and small outer seta. Labium (Fig. 6H) 2-lobed, outer and inner lobe distal corner setose. Cheliped (Fig. 7A): Basis 1.5 times as long as wide; merus with ventral seta; carpus almost 1.5 times as long as broad, with two dorsal setae and two long ventral setae; propodus and fixed finger little longer than carpus, about 2.5 times as long as broad; dactylus about half length of whole; fixed finger with three calcified teeth and three setae on inner margin, and two simple setae ventrally. Pereopod-1 (Fig. 7B): Of walking type; coxa present; basis a little shorter than rest of articles combined, naked; ischium with one seta; merus equal to carpus, with one seta distally; carpus with four setae (one very small) distally; propodus little shorter than merus and carpus combined, with setae subdistal on dorsal and minute seta ventrally.



Pereopod-2 (Fig. 7C): Of walking type; coxa present; basis shorter than rest of articles combined, naked; ischium short, with one seta; merus as long as carpus with one seta distally; carpus with three setae distally; propodus little shorter than merus and carpus combined, with one seta ventrally and subdistal seta dorsally; dactylus and unguis combined as long as propodus.

FIGURE 6. Meromonakantha nutae n. sp., paratype. A) Antennule; B) Antenna; C) Labrum; D) Left mandible; E) Right mandible; F) Maxillule; F’) Palp; G) Maxilla; H) Labium; I) Maxilliped. Scale: A–C = 0.1 mm; D–H = 0.01 mm.



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FIGURE 7. Meromonakantha nutae n. sp., paratype. A) Cheliped; B) Pereopod-1; C) Pereopod-2; D) Pereopod-3; E) Pereopod-4; F) Pereopod-5; G) Pereopod-6; H) Pleopod; I) Uropod. Scale = 0.1 mm.



Pereopod-3 (Fig. 7D): Similar to pereopod-2. Pereopod-4 (Fig. 7E): Of walking type; basis shorter than rest of articles combined, three times as long as wide, naked; ischium with one seta; merus as long as carpus with two spiniform seta distally; carpus with two spiniform setae, one hook and one seta distally; propodus a little shorter than merus and carpus combined, with two spiniform setae ventrally, one pinnate seta dorsally and one dorso-distal seta; dactylus semifused with unguis, half as long as propodus. Pereopod-5 (Fig. 7F): Similar to pereopod-4. Pereopod-6 (Fig. 7G): Similar to pereopod-4 with three short setae on propodus distally (absent in drawn specimen). Pleopods 1–5 (Fig. 7H): Basal article naked. Exopod with five setae on outer margin and one seta on inner margin. Endopod with eight setae. Both rami with most proximal seta separated by large gap from the others. Uropod (Fig. 7I): Endopod two-articled, proximal article 0.6 times length of ramus, with five distal and subdistal setae. Exopod two-articled, articles semifused, a little shorter than endopod, with one outer seta near fusion line; distal article tipped by long seta. Distribution: Antarctic: East off South Orkney Islands, at bathyal depths of 1182–1996 m. Remarks: Meromonakantha nutae n. sp. is most similar to the North-Pacific M. setosa, having a relatively short carapace little longer than pereonite-1. The poor description of M. setosa and its lost types make any detailed comparison of the species impossible. The new species however has distinctly less slender chelae which are only twice as long as wide and semifused articles in the exopod of uropod.

Genus: Paratyphlotanais Kudinova-Pasternak & Pasternak, 1978 Paratyphlotanais: Sieg (1986a) 45: 73–74; Sieg (1986b) 2: 76; Larsen (2005) 5: 201, 203, 208, 261, 302; Bird (2004a) 38: 1359–1363, 1374, 1381. Paratyphlotandis: Guţu & Sieg (1999): 365, lapsus calami

Diagnosis (modified after Bird, 2004a): Cephalothorax longer than broad, distinctly conical; rostrum usually prominent. At least pereonites 1–2 with sternal spurs. Antennule article-3 nearly as long as article-1; article-3 with terminal spur. Antennal articles 2 and 3 with dorsal spiniform seta. Mandible with broad molar. Cheliped without side piece. Pereopods 2 and 3 with one meral and three serrate carpal setae. Pereopods 4–6 walking type; merus with two serrate setae; carpus with three (or four) serrate setae; dactylus and unguis unfused, unguis bifurcated. Pleopods with proximal seta on endopod and exopod separated from remainder by gap. Uropods slender, endopod two-articled, distal article much shorter than proximal, exopod one or two-articled. Male: Terminal or swimming males unknown. See Bird (2004a) for ‘preparatory’ males (perhaps the mature male form). Gender of generic name: Masculine. Type species: Paratyphlotanais typicus Kudinova-Pasternak & Pasternak, 1978. Species included: P. armatus (Vanhffen, 1914); P. gracilipes (Hansen, 1913); P. japonicus (KudinovaPasternak, 1984); P. mananensis (Wallace, 1919); P. microcheles (G.O. Sars, 1882); P. pectinatus Bird, 2004; P. richardi (Dollfus, 1897); Paratyphlotanais typicus Kudinova-Pasternak & Pasternak, 1978; Paratyphlotanais alveolus n. sp. Remarks: Paratyphlotanais has been revised by Bird (2004a). He assigned to the genus eight species and noted the similarity of T. mananensis to the genus. My observations on the ANDEEP material, as well Hansen’s, Kudinova-Pasternak’s, Sars’s and Dolffus’ type specimens confirm that the characters chosen by Bird define well this monophyletic genus.



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Paratyphlotanais alveolus n. sp. (Figs 8–10) Material examined: Holotype: one female, (K 41390), ANT XXII-3, PS 67/74-6-E, 71°18.35’–71°18.28’S, 13°57.71’–13°57.31’W, depth 1030–1040 m, 20 Feb 2005; Paratypes: two females (one dissected on slides), (K 41387), ANT XXII-3, PS67/133-2-S, 62°46.49’–62°46.38’S, 53°3.50’–53°3.98’W, depth 1584–1579 m, 16 Mar 2005; eight females, (K 41389), ANT XXII-3, PS 67/133-2-Epi, the same locality; one female, (K 41 391), ANT XXII-3, PS 67/80-9-Epi, 70°39.07’–70°39.22’S, 14°43.36’–14° 43.39’W, depth 3102–3102 m, 23 Feb 2005; one female, (K 41449), ANT XXII-3, PS 67/80-9-S, the same locality; three mancae, (K 41388), ANT XXII-3, PS 67/74-6-S, 71°18.35’–71°18.28’S, 13°57.71’–13°57.31’W, depth 1030–1040 m, 20 Feb 2005. Diagnosis (neuter/female): Paratyphlotanais with complex pereon outline; pereonite-1 depressed dorsally, with large hyposphaenium; pereonites 2 and 3 with hyposphaenium prominent; pereonites 4 and 5 with small hyposphaenium. Pereopods 1–2 coxa with large spur. Chelipeds gracile with one ventral carpal seta exceeding carpal width and with only two setae dorsally; meral and carpal spiniform setae of pereopods 1–3 only weakly serrate. Uropod endopod up to ten times as long as broad. Etymology: Alveolus [Lat.] = cavity, tray, river-bed. The name alludes to the dorso-ventrally depressed form of pereonite-1. Description: Non-ovigerous female body length 2.8 mm (Fig. 8A,B). Cephalothorax 15% of total length, 1.5 times as long as basal width; rostrum pointed, overlapping base of antennules. Pereonites 1–3 narrower caudally; pereonites 4 and 5 rounded; pereonite-6 narrowest rostrally; pereonite-1 four times wider than long, depressed at middle; pereonites width: length ratios: 3.5, 1.5, 1.2, 1.2, 1.0, 1.4 respectively; large sternal hyposphaenium on pereonite-1, prominent hook on each of pereonites 2 and 3, and small hook on each of pereonites 4 and 5. Pleon 22% of body length, little longer than carapace. Pleotelson (Fig. 10H) rectangular, just longer than two preceding pleonites, with modest apex. Antennule (Fig. 9A): Almost 0.6 times length of cephalothorax; article-1 twice as long as wide, 1.4 times as long as articles 2 and 3 combined, with four setae along margin and one long (longer than article-2) simple seta and two pinnate setae distally; article-2 half as long as article-3, with one long (just as article-3) distal seta; article-3 with terminal spur and four simple setae distally. Antenna (Fig. 9B): Article-2 three times as long as article-3, with dorsal seta; article-3 naked; article-4 with two long, one short and one pinnate setae distally, 4.5 times as long as wide; article-5 with one terminal seta; article-6 with four setae. Mouth parts: Labrum (Fig. 9C) with rounded, setose distal margin. Mandible (Fig. 9D,E) molar broad with crenulated margin, and crenulated incisor; left mandible with broad lacinia mobilis. Maxillule endite (Fig. 9F) with eight distal setae; palp (Fig. 9F’) with two terminal setae. Maxilla (Fig. 9G) oval. Labium (Fig. 9H) with inner lobe setose distally and with minute setae along margin on outer lobe. Maxilliped (Fig. 9I) basis 1.5 times as long as wide, with long seta reaching over endites distally; endites with one tubercle and one seta on distal margin; palp four-articled: article-1 naked; article-2 wedge-shaped, with three setae on inner margin and one relatively long seta on outer margin; article-3 with three long and one short setae on inner margin; article-4 shorter than article-3 with four long and two short setae distally. Cheliped (Fig. 10A): Basis over twice as long as wide; merus triangular with long ventral seta; carpus three times as long as broad, with two small setae dorsally and two subequal, long setae and one minute seta ventrally; propodus and fixed finger slender, about three times as long as broad; dactylus about 0.7 times as long as propodus, with one seta near insertion of dactylus; fixed finger with three setae on strongly calcified inner margin and two setae on ventral margin; dactylus little shorter than fixed finger, with small setae in proximal part of dorsal margin.



FIGURE 8. Paratyphlotanais alveolus n. sp., holotype, adult female. A) dorsal view; B) lateral view. Scale = 0.1 mm.

Pereopod-1 (Fig. 10B): Coxa with one seta on spur; basis slender 6.5 times as long as broad, a little longer than merus and carpus combined, with proximal seta ventrally; ischium with small seta; merus about twice as long as broad, with spiniform ventral seta and simple dorsal seta; carpus little longer than merus, 2.3 times as long as broad, with spiniform dorsal seta and four minute setae distally; propodus about five times as long as broad, with one short serrate and one simple seta distally; dactylus and unguis about as long as propodus, unguis longer than dactylus. Pereopod-2 (Fig. 10C): Coxa with one seta; basis slender, about six times as long as broad, with proximal and distal seta dorsally; ischium with small seta; merus about twice as long as broad, with ventral spine and two setae distally; carpus as long as merus, with two spiniform and five minute setae distally; propodus about 1.5 times as long as carpus, with one disto-dorsal spiniform seta and one minute seta ventrally; dactylus and unguis together as long as propodus.



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FIGURE 9. Paratyphlotanais alveolus n. sp., female paratype. A) Antennule; B) Antenna; C) Labrum; D) Left mandible; E) Right mandible; F) Maxillule; F’) Palp; G) Maxilla; H) Labium; I) Maxilliped. Scale = A, B, I = 0.1 mm; C-H = 0.01 mm.



FIGURE 10. Paratyphlotanais alveolus n. sp., female paratype. A) Cheliped; B) Pereopod-1; C) Pereopod-2; D) Pereopod-3; E) Pereopod-4; F) Pereopod-6; G) Pleopod; H) Uropod. Scale = 0.1 mm A REVISION OF FAMILY TYPHLOTANAIDAE SIEG 1984


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Pereopod-3 (Fig. 10D): Similar to pereopod-2, but coxa without prominent spur, basis with one seta ventrally. Pereopod-4 (Fig. 10E): More robust than pereopods 1–3; basis three times as long as wide, as long as merus, carpus and propodus combined, naked; ischium with one seta; merus, with two spiniform ventral serrate setae; carpus almost as long as merus, three times as long as broad, with four spiniform setae (one longer than the others); propodus little longer than carpus, with two spiniform setae on ventral margin and one spiniform seta distally; unguis 0.3 times dactylus length, combined 0.6 times propodus length. Pereopod-5: similar to pereopod-4. Pereopod-6 (Fig. 10F): Basis about three times as long as wide and almost as long as merus, carpus and propodus combined, with one pinnate seta ventrally; ischium with one seta; merus with two spiniform serrated setae ventrally; carpus almost as long as merus, twice as long as broad, with four spiniform setae and two short simple setae distally; propodus longer than carpus, with one spiniform seta subdistally on ventral margin and three spiniform setae distally reaching half of dactylus; unguis and dactylus 0.3 times as long as of propodus; unguis 0.3 times as long as dactylus. Pleopods (Fig. 10G): Both rami subovate, with proximal seta separated from the others; endopod with ten plumose setae, exopod with twenty plumose setae. Uropods (Fig. 10H): Very slender, nearly twice as long as pleotelson; endopod two-articled, proximal article 0.7 times length of ramus, ten times as long as broad; exopod one-articled almost as long as proximal article of endopod; setation as figured. Distribution: Antarctic: Eastern Weddell Sea and line between tip of Antarctic Peninsula and Southern Orkney Island, at depths of 1030–3102 m. Remarks: Paratyphlotanais alveolus n. sp. has a transverse depression on first pereonites. This unique character allows immediate recognition of the species; equally it is the only species with a coxal spur present only on the first pereopod.

Family: Typhlotanaidae Sieg, 1976 Diagnosis (new): Blind. Antennule three-articled. Antenna six-articled. Molar process wide. Maxilliped distal endites with two tubercles. Cheliped without side piece. Pereopods 1–3 with coxa, pereopods 4–6 clinging type, without coxa, with prickly tubercles on carpus (except Obesutanais and Hamatipeda), dactylus and unguis not fused (if fused than modified to hook), pereopod-6 propodus with three terminal setae distally. Uropod rami one- or two-articled. Marsupium formed from four pairs of oostegites. Male (where known): swimming type. Genera included: Typhlotanais G.O. Sars, 1882 sensu stricto; Typhlotanais sensu lato; Peraeospinosus Sieg, 1986; Typhlotanaoides Sieg, 1983; Obesutanais Larsen et al. 2006; Hamatipeda n. gen.; Larsenotanais n. gen.; Pulcherella n. gen.; Torquella n. gen.; Typhlamia n. gen. Remarks: Two species, Typhlotanais dubius and T. gruzovi, do not have the clinging form of pereopods 4–6. T. dubius almost certainly belongs in Pseudoparatanais, while T. gruzovi may belong to Paratyphlotanais or a similar genus (Bird, pers. comm.). As this material is not available for study the problem cannot be resolved for a while, but it is quite probable that these species cannot be included within the Typhlotanaidae as defined here.



Key for Typhlotanaidae genera and morpho-groups 1. Body short, five or less times as long as wide (Fig. 11A) ............................................................................2 - Body long, six or more times as long as wide (Fig. 11A’)............................................................................8 2. Antennule twice as long as carapace (Fig. 11B) ............................................................................................ ..................................................‘eximus’ group (T. penicillatus, Typhlotanais sp. A, T. eximus, T spinipes) - Antennule about as long as carapace (Fig. 11B’) ........................................................................................3 3. Pereopods 4–6 dactylus and unguis fused and modified to hook (Fig. 11I).......................... Typhlotanoides - Pereopods 4–6 dactylus not fused with unguis, both normally developed (Fig. 11I’) ................................4 4. Pereopods 4–6 with specialized hooks on carpus (Fig. 11C) .....................................................Obesutanais - Pereopods 4–6 with prickly tubercles on carpus (Figs 11D, D’) ................................................................ 5 5. Pereopods 4–5 unguis bifurcated (Fig. 11D’), propodus distal seta longer than dactylus unguis combined (Fig. 11D’) ...... ‘cornutus’ group (T. cornutus, T. crassus, T. andeepae n. sp., T. adipatus sensu Tzareva) - Pereopods 4–5 unguis simple (Fig. 11D), propodus distal seta shorter than dactylus (Fig. 11D)................6 6. Uropod exopod 2-articled ............................................................................................................. T. grahami - Uropod exopod 1-articled .............................................................................................................................7 7. Uropod exopod minute, less than half endopod length ...................................................................T. parvus - Uropod exopod longer than half endopod length ....................................................... Larsenotanais n. gen. 8. Pereonites 1–3 corrugated (Fig. 11E) ............... ‘plicatus’ group (T. plicatus, T. longimanus, T. variabilis) - Pereonites 1–3 smooth (not corrugated) .......................................................................................................9 9. Uropod exopod equal or subequal to endopod (Fig. 11F); pereonites 2–5 semi-annulated proximally and distally (Fig. 11G) ................................................................................................................ Peraeospinosus - Uropod exopod shorter than endopod (Fig. 11H); pereonites 2–5 smooth (without rings)........................10 10. Pleotelson distal projection with two strong spiniform setae (Fig. 11J) ........................................................ ............................................................................... ‘spinicauda’ group (T. spinicauda, T. squamiger n. sp.) - Pleotelson distal projection without strong setae (Fig. 11H)......................................................................11 11. Pereopod coxa 1–3 with spur (Fig. 11K) ................................................................................................... 12 - Pereopod coxa 1–3 without spur (Figs 11L).............................................................................................. 13 12. Pereonite-1 with long setae dorsodistally (Fig. 12A) .........................................................T. mimosis n. sp. - Pereonite-1 without long setae dorsodistally ... ‘greenwichensis’ group (T. messinensis, T. greenwichensis) 13. Pereopods 1–3 ischium with long seta (Fig. 11M); antenna article-2 and 3 with ventral hooks (Fig. 11N’). ....................................................................................... ‘trispinosus’ group (T. trispinosus, T. tenuicornis) - Pereopod-1-3 ischium with short seta (Figs 11K, L); antenna article-2 and 3 without teeth (Fig. 11N) ...14 14. Pereopods 4–6 carpus with specialized hooks (Fig 12B) ...............................................Hamatipeda n. gen. - Pereopod 4–6 carpus with prickly tubercles (Fig. 12G–I)..........................................................................15 15. Pereonite-1 almost as long as carapace (Fig. 12D)..........................................................Pulcherella n. gen. - Pereonite-1 clearly shorter than carapace (Figs 12C, E) ............................................................................16 16. Antennule over twice as long as carapace (Fig. 12 C); prickly tubercles on pereopods 4–6 minute (Fig. 12J) ....................................................................................................................................Typhlamia n. gen. - Antennule shorter than double length of carapace (Figs 12D, E, K)..........................................................17 17. Prickly tubercles on pereopods 4–6 surrounded by spines (Figs 12G, G’) ........................ Torquella n. gen. - Prickly tubercles on pereopods 4–6 simple, not surrounded by spines (Figs 12 H–J) ...............................18 18. Large gap between cheliped bases and pereonite-1 ventrally (Fig. 12L) ...................................................... ................................................................................................. ’mixtus’ group (T. mixtus, T. mimosis n. sp.) - Small (or no) gap between cheliped basis and pereonite-1 ventrally (Figs 12 K, L’) ...............................19 19. A few setae on pereopod-1 basis and carpus dorsal edge (Fig. 12M) ............................................................ .......................................................................................................... T. aequiremis (Typhlotanais sensu str.) - No setae on pereopod-1 basis and carpus dorsal edge (Fig. 12M’).......................... Typhlotanais sensu lato A REVISION OF FAMILY TYPHLOTANAIDAE SIEG 1984


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FIGURE 11. Morphological characters used in the key to Typhlotanaidae. A) Typhlotanais sp. A, dorsal view; A’) T. spiniventris, dorsal view; B) Typhlotanais sp. A, proximal part of body dorsal view; B’) L. amabilis n. sp., proximal part of body dorsal view; C) O. sigridae, pereopod-5; D) T. mimosis n. sp., pereopod-5; D’) T. andeepae n. sp. pereopod-5; E) T. plicatus, lateral view; F) P. kerguelenensis, uropod; G) P. kerguelenensis, lateral view; H) T. messinensis, pleotelson and uropod; K) T. greenwichensis, pereopod-1; L) T. juraszi n. sp., pereopod-1; M) T. tenuicornis, pereopod-2; N’) P. acruxi n. sp., antenna; N’) T. trispinosus, antenna; J) T. spinicauda, pleotelson and uropod.



FIGURE 12. Morphological characters used in the key to Typhlotanaidae (continued). A) T. mimosis n. sp., proximal part of body dorsal view; B) H. trapezoida n. sp. pereopod-6; C) T. mucronatus, lateral view of proximal body part; D) P. pulcher, lateral view of proximal body part; E) T. variabilis n. sp., lateral view of proximal body part; F) T. spinicauda, dorsal view; F’) T. mucronatus, dorsal view; G, G’) P. kerguelenensis, pereopod-4; H) T. mimosis, pereopod-5; I) P. filatovae, pereopod-6; J) P. pulcher; K) T. andeepae; L) T. mixtus, carapace ventral view; L’) T. aequiremis, carapace ventral view; M) T. aequiremis, pereopod-2; M’) T. inaequipes, pereopod-2.



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Genus: Typhlotanais G.O. Sars, 1882 Tanais: Lilljeborg (1864) (in part) : 12. Typhlotanais: G.O. Sars (1882a) 7: 52; Sieg (1986a) 94–95; Larsen (2005) 209–210.

Diagnosis (new): Body long (seven times as long as wide). All pereonites wider than long, lateral edges parallel (not rounded in dorsal view). Antennule shorter than carapace. Labrum hood-shaped, sparsely setose. Mandible lacinia mobilis delicately crenulated, molar with numerous, regular, blunt denticles. Maxilla with nine spines. Maxilliped basis as long as wide, with short seta; endites with two setae and two tubercles. Labium bilobed, sparsely setose distally. Cheliped slender; chela almost as long as carpus, three times as long as wide. Pereopods 1–3 coxa without spurs; pereopod-1 with regular setae (eight, two, and five on basis, merus and carpus respectively) on dorsal margin; pereopods 2 and 3 with spiniform setae on carpus and propodus; pereopods 4–6 carpus with prickly tubercles of moderate size (less than half of carpus length), unguis with bifid tip, propodus distal seta (-ae) reaching end of dactylus. Uropod rami one-articled; exopod little shorter than endopod, tipped with two simple setae (one seta robust). Male: Unknown. Gender of generic name: Masculine. Type species: Typhlotanais (=Tanais) aequiremis (Lilljeborg, 1864). Species included: Typhlotanais aequiremis (see remarks). Remarks: The first step toward rationalizing typhlotanaid taxonomy was taken by Sieg (1984). In considering the ornamentation of the first three pairs of pereopods, he separated typhlotanaids from leptognathiids, primarily as a subfamily (op. cit.) and later as a family (Sieg, 1986b) judging them as an apomorphic group. His conclusions were based mainly on the presence of fully developed ‘spines’ (‘spiniform’ setae in presently accepted terminology) on the anterior pereopods in Leptognathiidae, while pereopod spines are strongly reduced or completely absent in typhlotanaids. In contradiction to the diagnosis given by Sieg 1986a, there are minute spiniform setae occuring on pereopods 2–3 of most typhlotanaids, including the type species—T. aequiremis. They are absent in a few species only. Two of these (Typhlotanais parvus Sieg, 1986 and T. grahami Błażewicz-Paszkowycz, 2004) have reduced setation on all pereopods and indeed the carpus of their pereopods 2 and 3 is armed with two regular setae only. This feature differs in the new genus Pulcherella described below, which has 3–5 setae on the pereopod 2–3 carpus. If Sieg’s concept that spiniform setae on these pereopods is an apomorphic character is correct, the regular and relatively numerous setae in Pulcherella should be considered plesiomorphic, while reduced setation, including the reduction of spiniform setae, should be an apomorphy. Sieg (1986a) suggested two phylogenetic trends within Typhlotanais and created the subgenera Typhlotanais and Monosmerinx. His preliminary system was based on the number of terminal seta on the uropodal exopod (one in Monosmerinx and two in Typhlotanais) and the absence or presence of a proximal seta in the endopod of the pleopod. However, these two characters need to be entirely re-assessed. My initial observations reveal that there is only one seta on exopod of the uropods in just two typhlotanaids (T. parvus and T. grahami), while the lack of the proximal setae on the pleopods may be an artifact due to frangible character of pleopod setation. At the moment Typhlotanais sensu stricto comprises only the type species (T. aequiremis). Consequently all other species must remain Typhlotanais sensu lato. It is recognized that further studies may place other Typhlotanais sensu lato species into Typhlotanais sensu stricto, but such a transfer is not possible now owing to the scarcity or poor condition of the material available for the study. Morphological studies of Typhlotanais sensu lato have shown a variety of characters: the proportionate length of pereonite-1, the setation of the dorsal edge of the cheliped carpus, the morphology of the mandible



molar process, the shape of the basis of pereopods 1–3, and the character of the clinging apparatus of pereopods 4–6. These characters can be used to provisionally split Typhlotanais into coherent morpho-groups and that may provide the basis for distinguishing new genera in the future.

Typhlotanais aequiremis (Lilljeborg, 1864) (Figs 13–15) Tanais aequiremis: Lilljeborg (1864) 1864: 12, 21; Lilljeborg (1865) 1: 14, 25; Gerstaecker (1888) 5(1): 540; Sieg (1980a) 537: 11–12. Tanais æquiremis: G.O. Sars (1866) 15: 122. Tanais depressus: G.O. Sars (1866) 15: 121–122; G.O. Sars (1871(1872)) 275, 285; G.O. Sars (1877) 2: 246; G.O. Sars, (1882a) 7: 34; Gerstaecker (1888) 5(2): 540; G.O. Sars (1896): 21–22; Norman (1899) 3: 340; Sieg (1980a) 537: 11– 12. Typhlotanais æquiremis: G.O. Sars (1882a) 7: 34–35; G.O. Sars ((1882)1883): 14; G.O. Sars (1896): 21–22; Norman (1899) 3: 340; Norman (1905) 16: 78–95; Hansen (1909) 1909: 229; Hansen (1913) 3(3): 56; Grieg (1914): 105; Stephensen (1937) 2(24): 19; Stephensen (1943) 121(10): 73; Stephensen (1948) 53: 164–165. Typhlotanais aequiremis: Zirwas (1911) 12: 105; Nierstrasz (1913) 32(a): 35; Nierstrasz & Schuurmans Stekhoven (1930) 10: 162–164; Stephensen (1932) 6: 349; Stephensen (1937) 3(27): 23; Greve (1965b) 1(27): 5; Greve (1965c) 20: 15; Greve (1965a) 20: 44, 50, 53; Greve (1965d) 38: 142; Greve (1967) 29: 295–297; Greve (1968) 36: 77–78, 82; Kudinova-Pasternak (1969) 48: 1737; Lang (1970) 23: 267–270, 288; Morino (1971) 18(5): 349; Brattegard & Vader (1972) 49: 36; Greve (1972) 48: 35; Sieg (1980a) 537: 12; Holdich & Jones (1983a) 60–61; Holdich & Jones (1983b) 17: 158, 170–172, 176, 178, 180. Hassack & Holdich (1987) 16(3): 223, 226–227, 229. Typhlotanais aeqviremis: Kudinova-Pasternak (1966) 45: 528. Typhlotanais aequiremis: Wallace (1919) 18: 7–8. Lang (1970) 23: 270. ? Typhlotanais aequiremis: Lang (1957) 52: 2.

Material examined: 32 females, CRU 3887, (one female dissected on slides), Norway, material determined by G.O. Sars, 1898; three females, CRU 3888, Norway, material determined by G.O. Sars, 1898. Diagnosis: As for the genus. Complementary description: Female 3.5 mm (Fig. 13A, B) body long, 7.5 times as long as wide; carapace smooth, 1.3 times as long as wide, lateral margins slightly rounded; rostrum pointed; pereonites smooth, all wider than long; pereonite-1 shortest, twice as wide as long; pereonites 2, 4 and 5 subequal, 1.5 times as long as pereonite-1; pereonite-3 longest; pereonite-6 slightly longer than pereonite-1. Pleon about as long as carapace, pleonites 1–5 similar in size; pleotelson rounded. Antennule (Fig. 14A): Article-1 slender, about four times as long as wide, with two groups of pinnate and simple setae along article; article-2 with three simple setae distally; article-3 four times as long as article-2, with five apical setae. Antenna (Fig. 14B): Article-2 as long as wide, twice as long as article-3, both naked; article-4 four times as long as article-5, with two simple and three pinnate setae distally; article-5 with one simple seta distally; article-6 very short, with six terminal setae. Mouth parts: Labrum (Fig. 14C) hood-shaped, with minute setae. Mandible (Figs 14D,E) molar welldeveloped, longer than incisor with regular, blunt teeth; incisor blunt, crenulated; lacinia mobilis well-developed, crenulated. Maxillule (Fig. 14F) endite with nine apical spiniform setae, palp with two terminal setae. Both lobes of labium (Fig. 14G) poorly separated and sparsely setose distally. Epignath (Fig. 14I) curved, simple distally. Maxilliped (Fig. 14H): coxa reduced; basis fused, heart-shaped, as long as wide, with short seta; endite with two simple setae and two tubercles distally; palp article-1 unarmed; article-2 wedge-shaped, with three setae (one robust) on inner margin and one minute seta on outer margin; article-3 trapezoidal, with four weakly-serrated setae on inner margin; article-4 slender, with one simple seta on outer margin and five weakly-serrated distal setae (one broken off).



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Cheliped (Fig. 15A): Basis slightly rounded, twice as long as wide; merus wedge shaped, with one seta ventrally; carpus with two long and one short setae ventrally and two setae dorsally; propodus with distal seta on inner side; fixed finger (propodus projection) tipped with strong spine, with three setae on well-calcified inner margin and two setae ventrally; dactylus slightly curved with one short, rod seta proximally on dorsal margin.

FIGURE 13. Typhlotanais aequiremis, female. A), dorsal view; B) lateral view; C) carapace ventral side. Scale 1 mm.



FIGURE 14. Typhlotanais aequiremis, female. A) Antennule; B) Antenna; C) Labrum; D) Left mandible; E) Right mandible; F) Maxillule; G) Labium; H) Maxilliped; I) epignath. Scale: A, B = 0.1 mm; C-I = 0.01 mm.



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FIGURE 15. Typhlotanais aequiremis. A) Cheliped; B) Pereopod-1; C) Pereopod-2; D) Pereopod-3; E) Pereopod-4; F) Pereopod-6; G) Pleopod; H) Uropod. Scale = 0.1 mm.

Pereopod-1 (Fig. 15B): Slender (walking type); coxa with one seta; basis as long as merus, carpus and half of propodus combined, with eight short simple setae dorsally and two setae ventrally; ischium short, with



one seta; merus shorter than carpus with two simple setae ventro-distally and two setae dorso-distally; carpus with two setae ventro-distally and five setae on dorsal margin; propodus a little longer than carpus, with two setae distally; dactylus half as long as unguis; both slightly shorter than propodus. Pereopod-2 (Fig. 15C): Slender (walking type); coxa with one seta; basis slightly shorter than rest of articles combined, with three minute setae proximally on dorsal margin and two short setae on ventral margin; ischium with simple seta; merus a little shorter than carpus, with three simple setae distally; carpus slightly longer than merus, with four simple setae and one spiniform distally; propodus a little longer than merus and carpus combined length, with spiniform seta ventrally and two setae dorsally; dactylus half as long as unguis, with one simple seta. Pereopod-3 (Fig. 15D): Similar to pereopod-2, but basis with only one seta. Pereopod-4 (Fig. 15E): Clinging type; basis robust, less than twice as long as wide, with two setae ventrally and one seta dorsally; ischium with two setae; merus with two spiniform setae ventrally; carpus with prickly tubercles of moderate size distal hooks and one setae dorso-distally; propodus with two ventro-distal spiniform setae and one dorso-distal seta as long as dactylus; unguis bifid tipped, half as long as dactylus; both almost as long as propodus. Pereopod-5 similar to pereopod-4. Pereopod-6 (Fig. 15F): Similar to pereopod-5, but propodus with three dorso-distal setae. Pleopod (Fig. 15G): Basal article naked; exopod with one seta on inner margin and fourteen plumose setae on outer margin; endopod with eighteen plumose setae on outer margin; no clear gap between proximal seta and other setae. Uropods (Fig. 15H): Basal article as long as wide; both rami one-articled (see comment in generic diagnosis); exopod 0.7 times as long as endopod, with one strong seta and one regular seta distally; endopod tipped by one strong, four regular and one pinnate setae. Distribution: North Sea, The Skagerrak, Sweden, Norwegian south and west coast up far north of Tromso, Spitsbergen, British waters, western and eastern Iceland, at the depth 22–430 m (Holdich and Jones, 1983a). Remarks: Pragmatically, all typhlotanaids with prickly tubercles (i.e. all except Obesutanaids and Hamatipedis) can be divided into short body (length: width ratio 6). The first group includes species of two morpho-groups—the ‘eximus’ group, i.e. T. penicillatus, T. eximus, T. spinipes and Typhlotanais sp. A (present work), and the ‘cornutus’ group, i.e. T. cornutus, T. crassus, T. andeepae n. sp., and T. adipatus sensu Tzareva—as well as Larsenotanais amabilis n. sp. and three species incertae sedis T. parvus, T. grahami and T. solidus. Typhlotanais aequiremis, at present the only member of Typhlotanais sensu stricto, is representative of the long form (length: width ratio >6) with pereonites wider than long or square, but never longer than wide. The smooth cuticle covering the pereonites distinguishes Typhlotanais sensu stricto from Peraeospinosus and the ‘plicatus’ group. The first pereonite that is shorter than the carapace allows immediate distinction from Pulcherella n. gen and Hamatipeda n. gen., while the unarmed pleotelson allows distinction from members of the ‘spinicauda’ group. T. sensu stricto is distinguished from members of genus Typhlamia n. gen. by having the antennule shorter than the carapace, and from species presently combined in the ‘trispinosus’ group—T. trispinosus Hansen, 1913 and T. tenuicornis G.O. Sars, 1882—by the lack of teeth on the second and third antennal articles. Mouthparts are conservative in their morphology within the family, although some variability occurs in the character of the molar and the ornamentation of the maxilliped endites. The mandible molar of the type species is armed by a row of blunt and regular teeth. The same blunt and regular molar teeth occur in members of Pulcherella, Torquella and Typhlamia, although they are less densely arranged. The short seta on the maxilliped basis distinguishes T. aequiremis from the genera above. In addition, the distal edges of the maxilliped endites are armed with two setae and two tubercles of regular size. Tubercles of similar character occur gener-



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ally in most typhlotanaids, although they are clearly smaller in Peraeospinosus, and especially large in T. inaequipes Hansen, 1913 and T. williamsae Dojiri & Sieg, 1997 (Typhlotanais sensu lato). Unique characters of T. aequiremis are the relatively numerous setae on the dorsal margin of the basis (eight) and carpus (five) of pereopod-1. The presence of spiniform setae on the carpus of the second and third pereopods is a character shared by T. aequiremis and most typhlotanaids (including the genera Peraeospinosus, Torquella, Typhlamia), but they absent in Pulcherella, T. parvus and T. grahami (see remarks under Typhlotanais s. str In T. aequiremis the prickly tubercles are well embossed but relatively small (not longer half of carpus), in the ‘spinicauda’ group they are large (as long as carpus), in Pulcherella they are large but almost flat, in Peraeospinosus and Torquella they are surrounded by a row of spines, while in Typhlamia they are minute. The gap between the most proximal and the other setae on the pleopods is so small in T. aequiremis that the pleopod setation in this species is considered to be continuous. The uni-articled uropod rami of T. aequiremis is a rather common character among typhlotanaids and thus far has been observed in most members of Peraeospinosus, Typhlotanais adipatus (both sensu Tzareva and sensu Sieg), T. crassus Dojiri & Sieg, 1997, T. finmarchicus G.O. Sars 1882, and in the ‘plicatus’ group. Articulation of the uropod is a convenient character unless the articles are partially fused (semi-fused). The fusion line is often difficult to observe under the light microscope, and may become even less clear in adult and in preadult stages (pers. obs.). Typhlotanais aequiremis can be distinguished by various characters from those species not included to any of proposed morpho-groups. Most difficult for comparison at the moment is the poorly-described T. simplex, of which type material is not available for study. T. brachyurus Beddard, 1886 and T. longicephala Kudinova-Pasternak 1970 can be immediately distinguished: the first species has pereonite-1 just as long as the carapace, while the second has an extremely long carapace that is about twice as long as wide. The relatively large ventral gap between the distal edge of the cheliped basis and pereonite-1 allows the distinction of T. mixtus from T. aequiremis. The same large gap probably exists in T. plebejus Hansen, 1913, but because its cheliped distal edge is distally expanded, this gap is not visible. Further comparison of T. plebejus with the type species is not possible owing to the insufficiency of material available for study (the holotype only). Details of the first three pairs of pereopods allows the distinction of three other Typhlotanais sensu lato species from T. aequiremis: these details are large spines on the merus of pereopod-3 in T. proctagon Tattersall, 1904, the long setae on the carpus of pereopods-1 and –2 in T. inermis Hansen, 1913, and the long seta far exceeding the unguis on the propodus of pereopods 4-6 in T. kussakini Kudinova-Pasternak, 1970. Six other species—T. assimilis G.O. Sars, 1882, T. compactus Kudinova-Pasternak, 1966, T. finmarchicus G.O. Sars, 1882, T. inaequipes Hansen, 1913 T. profundus Hansen, 1913, T. williamsae Dojiri & Sieg, 1997 – share a similar body habitus with T. aequiremis. Two of them, T. inaequipes and T. williamsae, can be distinguished by their large tubercles on the maxilliped endites and long setae (just reaching the distal edge of the endites) on the maxilliped basis. Except for T. profundus, all the other species share with T. aequiremis the characters of prickly tubercles, small spiniform setae on the carpus of the second and third pair of pereopods, the relatively short distal seta on the propodus of the fourth and fifth pereopods, and the short setae on the maxilliped basis. Apart from that, T. compactus has numerous setae on the basis, merus and carpus of the first pereopod. According to observations by Hassack & Holdich (1987), tubes made by non-ovigerous females of T. aequiremis are incrusted mostly by sand grains, with a higher sand concentration towards the anterior part of tube. The posterior part of the tube is sac-like and is made of a mucous lining which can adjust to pleon. The anterior part of tube is open to allow the animal to emerge from the tube, for example for grazing for food particles.



Genus: Hamatipeda n. gen. Diagnosis (female): Pereonite-1 long (subequal or longer than carapace), pereonites 1–5 longer than wide (pereonites 2 and 3 over 1.5 times as long as wide). Mandible molar process with irregular edge but without tubercles and teeth. Maxillipedal endites lack distal tubercles. Chelae smaller than carpus, tapering distally. Pereopods 4–6 merus and carpus with two large specialized hooks, propodus with two long spiniform setae in the distal part of ventral margin, unguis tip trifurcate. Etymology: Named after the large spiniform-setae on pereopods 4–6: hamatus [lat.] = bearing hooks; pedes [lat.] = legs. Type species: Hamatipeda trapezoida n. sp. Gender of generic name: Feminine. Species included: Hamatipeda (= Typhlotanais) longa (Kudinova-Pasternak, 1975); H. trapezoida n. sp. Remarks: Hamatipeda n. gen. lacks prickly tubercles in pereopods 4–6 but, because it also lacks a cheliped ‘side piece’ and the climbing character of pereopods 4–6 is evident, it is included into the family Typhlotanaidae as was Obesutanais.

Hamatipeda longa (Kudinova-Pasternak, 1975) n. comb. (Fig. 16) Typhlotanais longus: Kudinova-Pasternak (1975) 103: 212–213, 215.

Material examined: Non-ovigerous female holotype (MC-978), Sta. 929, r/v Akademik Kurtchatov, Falkland Islands, 52º08.5'S, 57º16.1'W, depth 720 m. Diagnosis (female): Pereonite-3 twice as long as wide. Pereonites 1–3 lateral margins parallel. Cheliped fixed finger ventral setae extending beyond the article. Uropod exopod clearly shorter than endopod. Complementary description: Ovigerous female. Body length 3.5 mm (Fig. 16A,B), body very long, about twelve times as long as wide; carapace smooth, narrowed anteriorly, little longer than wide. Pereonites smooth with parallel lateral margins; pereonite-1 little shorter than carapace; pereonites 2 and 3 the longest; pereonite-2 1.5 times as long as pereonite-1; pereonite-6 shortest. Pleonites 1–5 subequal; pleotelson with distal projection. Antennule (Fig. 16C): Article-1 about 1.5 times as long as wide, with two distal and medial pinnate setae and one simple distal seta; article-2 0.3 times as long as article-3, with one simple seta; article-3 with three terminal simple setae. Antenna (Fig. 16D): Article-1 broken; article-2 longer than article-3; articles 2 and 3 with one simple seta; article-4 twice as long as article-5, with three simple and two pinnate setae distally; article-5 with one simple seta distally; article-6 with four setae. Mouth parts: Labrum missing. Mandible (Fig. 16E) stout; molar process well-developed, with crenulated, well-calcified edge; lacinia mobilis well-developed with five teeth. Maxillule and maxilla missing. Maxilliped (Fig. 16G) basis elongate and fused; endites separated, each with one seta only on distal margin; palp with four articles: article-1 naked; article-2 wedge-shaped, with one seta on outer margin; article-3 trapezoidal in shape with five setae on inner margin; article-4 slender (1.5 times as long as wide) with six terminal setae. Labium (Fig. 16F) inner lobe with a few setules. Cheliped (Fig. 16H): Basis twice as long as wide; merus wedge-shaped, with one seta on ventral margin; carpus over 1.6 times as long as wide, with one long seta ventrally and two short setae dorsally (proximally and distally); chela narrower than carpus, 1.5 times as long as wide, with one seta at insertion of dactylus; inner margin of fixed finger with two, or three teeth and three setae in distal part, ventral margin with two long simple setae; dactylus slightly curved. A REVISION OF FAMILY TYPHLOTANAIDAE SIEG 1984


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FIGURE 16. Hamatipeda longa. A) Holotype, female, dorsal view; B) Holotype, female, present state; C) Antennule; D) Antenna; E) Right mandible; F) Labium; G) Maxilliped; H) Cheliped; I) Pereopod-1; J) Pereopod-2; K) Pereopod-3; L) Pereopod-4; M) Pleopod; N) Pleotelson with uropods. Scale: A, B = 1 mm; C, H-M = 0.1 mm; E-G = 0.1 mm; D, F = 0.05 mm; E = 0.03 mm. A, C–G after Kudinova-Pasternak (1975).

Pereopod-1 (Fig. 16I): Of walking type; basis naked and shorter than merus, carpus and propodus combined length; ischium with one seta; merus almost as long as carpus, with one distal seta; carpus with four short setae distally; propodus with two, subdistal setae on dorsal margin and one small seta ventrally; unguis twice as long as dactylus; together as long as propodus. Pereopod-2 (Fig. 16J): Of walking type; basis almost as long as carpus and propodus combined; ischium with one seta ventrally; merus almost as long as carpus with one distal seta; carpus with two simple seta and



one spiniform seta; propodus 1.5 times as long as carpus, dorsal margin with one seta distally and one spiniform seta ventrally; dactylus almost as long as unguis. Pereopod-3 (Fig. 16K): Similar to pereopod-2. Pereopod-4 (Fig. 16L): Of clinging type; basis narrow about three times as long as wide; ischium with two setae ventrally; carpus little longer than merus, each with two large hook-like setae, carpus additionally with one seta dorsally; propodus as long as carpus with two setae ventrally and one dorsal terminal seta; unguis tip trifurcated. Pereopod-5 missing. Pereopod-6 similar to pereopod-4, but propodus with three terminal setae dorsally (according to Kudinova-Pasternak, 1975: 215). Pleopods 1–5 (Fig. 16M): All pleopods similar; basal article naked, most proximal seta of both exopod and endopod separated from others by gap; exopod outer margin with eight plumose setae, inner margin with one plumose seta; endopod with twelve plumose setae on outer margin. Uropod (Fig. 16N): Uropod exopod with one article, endopod two-articled; exopod about as long as proximal article of endopod. Male: Unknown. Distribution : Species known only from the type locality: Scotia Arch, Falkland Islands (52º08.5'S, 57º16.1'W), depth 720 m. Remarks: This complementary description has been prepared after studying the holotype that lacks the head, pleotelson and pereopods 5 and 6, so it has had to be supported by the description and drawing made by Kudinova-Pasternak (1975). In the illustrations made by Kudinova-Pasternak there are some inconsistencies in the antennule and antenna setation. It is thought that the plumose seta drawn for antennule article-2 belongs to article-1 and the proximal seta drawn for the antenna article-5 belongs to article-4.

Hamatipeda trapezoida n. sp. (Figs 17–19) Material examined: Holotype: non-ovigerous female, (NMNH 1100137), r/v Eltanin Cr 4, Sta. 129., 61º46'– 61º48'S, 61º35'–61º25'W, depth 3678–3861 m, Menzies Trawl, 2 Aug 1962. Paratype: non-ovigerous female, (K 41447), r/v Polarstern, ANT XIX/3-4, PS 61/41-3, Sta. 41-3, 59° 22.24'-59° 22.40' S, 60° 4.06'–60° 3.99' W, depth 2375–2372, epibenthic sledge, 26 Jan 2002; (K 41465) PS 61/129-7, 59°52.30'S, 59°57.63'W, depth 3614 m, Multi Corer, 23 Feb 2002. Diagnosis: Pereonites 1–3 trapezoidal in shape (lateral margins of pereonites 1–3 not parallel). Setae on ventral margin of cheliped fixed finger not extending beyond the article. Uropod exopod almost as long as endopod. Etymology: Named after the trapezoidal shape of pereonites 1–3. Description: Female, with body length 5.7 mm (Fig. 17 B, C), body long, about ten times as long as wide. Carapace glabrous, narrow, 1.5 times as long as wide. Pereonites smooth; pereonites 1–3 wider anteriotly than posteriorly (trapezoidal shape); pereonite-1 0.75 times as long as pereonite-2; pereonites 2 and 3 subequal in length; pereonite-4 slightly shorter than pereonite-3, little longer than wide (Fig. 17B) or 1.7 as long as wide (Fig. 17 A) pereonite-5 slightly shorter than 4; pereonite-6 shortest. Pleonites 1–5 similar in size. Pleotelson rectangular, slightly tapering distally. Antennule (Fig. 18A): Conical, three-articled; article-1 robust, about twice as wide as long, one simple medial seta, and two groups of pinnate setae distally and medially; article-2 0.25 times length of article-3, two simple setae and one pinnate seta distally; article-3 with four simple terminal setae. Antenna (Fig. 18B): Article-1 damaged during dissection; article-2 with two simple setae; article-3 with A REVISION OF FAMILY TYPHLOTANAIDAE SIEG 1984


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one simple seta; article-4 almost 1.7 times as long as article-5, with three long simple setae and two distal, pinnate setae; article-5 with one simple distal seta; article-6 with six terminal setae.

FIGURE 17. Hamatipeda trapezoida n. sp. A) Paratype, dorsal view (Polarstern Sta. 41-3); B, C) Holotype, lateral view (Eltanin Cr 4 Sta. 129). Scale = 1 mm.



FIGURE 18. Hamatipeda trapezoida n. sp., female paratype. A) Antennule; B) Antenna; C) Left mandible; D) Right mandible; E) Maxillule; F) Maxilla; G) Labium; H) Maxilliped; I) Epignath; J) Cheliped. Scale: A, B, J = 0.1 mm; C-H = 0.01 mm.

Mouthparts: Labrum not found. Mandible (Figs. 18C,D) large, molar process well-developed, with crenulated edge; lacinia mobilis well-developed, with five teeth. Maxillule (Fig. 18E) with eight distal spiniform setae. Maxilla (Fig. 18F) oval. Labium (Fig. 18G) two-lobed; inner and outer lobe with a few setules distally. Maxilliped (Fig. 18H) bases elongated and fused, with one distal seta; each endite armed with two setae on



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distal margin; palp article-1 naked; article-2 wedge-shaped with one seta on outer margin and three setae on inner margin; article-3 with four setae on inner margin; article-4 slender (1.5 times as long as wide), with six terminal setae. Cheliped (Fig. 18J) robust; basis slightly longer than wide; merus wedge-shaped, with one seta ventrally; carpus over 1.4 times as long as wide, with two long setae ventrally and two short setae dorsally (proximally and distally); chela narrower and shorter than carpus, twice as long as wide, with one seta at dactylus insertion; inner edge of fixed finger with three teeth and three setae in distal part; ventral edge with two long simple setae not extending beyond the article; dactylus slightly curved with one proximal seta.

FIGURE 19. Hamatipeda trapezoida n. sp., female paratype. A) Pereopod-1; B) Pereopod-2; C) Pereopod-3; D) Pereopod-4; E) Pereopod-5; F) Pereopod-6; G) Pleopod; H) Uropod. Scale = 0.1 mm.



Pereopod-1 (Fig. 19A): Of walking type; coxa present; basis with one pinnate seta proximally; about as long as merus, carpus and half of propodus combined; ischium short, with one seta; merus as long as carpus with two distal setae; carpus with four short setae distally; propodus with three subdistal setae. Pereopod-2 (Fig. 19B): Of walking type; coxa present; basis almost as long as merus, carpus and propodus combined; ischium with one seta ventrally; merus almost as long as carpus, with three setae distally; carpus with two simple, one bipinnate, and one spiniform seta; propodus 1.5 times as long as carpus; dorsal edge with two setae distally and one seta ventrally; unguis 1.5 times as long as dactylus. Pereopod-3 (Fig. 19C): Similar to pereopod-2, but merus with two simple and one spiniform setae. Pereopod-4 (Fig. 19D) Of clinging type; basis robust, about twice as long as wide; ischium with two setae ventrally (one shorter than other); carpus slightly longer than merus, each with two large hook-like setae; propodus as long as carpus, with two ventral spiniform setae, one terminal seta and one pinnate seta on dorsal edge; unguis trifurcated. Pereopod-5 (Fig. 19E): Similar to pereopod-4. Pereopod-6 (Fig. 19F): Similar to pereopod-4, but propodus with three dorso-distal setae. Pleopods 1–5 (Fig. 19G): All pleopods similar; basal article naked; exopod and endopod each with proximal seta separated from the others by gap; exopod outer edge with nine plumose setae, inner edge with one pinnate seta; endopod armed with 17 plumose setae on outer edge. Uropod (Fig. 19H): Both rami one-articled. Endopod with pinnate seta in the middle and one pinnate and four simple setae distally; exopod just as long as endopod, with one seta proximally, one robust seta and one simple seta terminally. Male: Unknown. Distribution: West Antarctic, north of South Shetland Islands, at depths of 2372–3876 m. Remarks: Hamatipeda trapezoida n. sp. is distinguished from H. longa by the trapezoidal shape of pereonites 1–3 in contrast to H. longa in which the pereonite lateral edges are parallel. The other diagnostic feature is the length of the uropodal rami; the exopod of the uropod is subequal in H. trapezoida but it is clearly shorter than the endopod in H. longa.

Genus: Larsenotanais gen. n. Diagnosis (female): Body short, four times as long as wide, all pereonites wider than long, pereonites 1–3 subequal. Antennule slightly longer than carapace; article-1 with about five simple setae along article. Mandible molar with regular tubercles and three spines ventrally. Maxilliped basis with seta longer than endites; endites with two tubercles and one seta distally. Cheliped compact, basis reaching the edge of pereonite-1, carpus three times as long as wide. Pereopods 1–3 basis without acute projection, merus, carpus and propodus without spiniform seta. Pereopods 2 and 3 carpus and propodus with spiniform seta; pereopods 4–6 carpus with moderate prickly tubercles (less than half of carpus length), unguis simple, propodus distal setae shorter than dactylus. Both rami of pleopods with proximal seta separated from others by gap. Uropods rami one-articled, exopod 0.8 times as endopod. Etymology: Named after Dr Kim Larsen in recognition of his contribution to knowledge of the Tanaidacea. Type species: Larsenotanais amabilis n. sp. Gender of generic name: Masculinae. Species included: Larsenotanais amabilis n. sp. Remarks: Larsonotanais n. gen. shows a few features in common with members of the ‘cornutus’ group (see remarks under that group) sharing a similar body habitus, setation on antennule article-1, mandible ornamentation, proportion of cheliped articles, character of prickly tubercles on pereopods 4–6, and a short distal



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seta on the propodus of pereopods 4–6. In contrast to members of the ‘cornutus group' the present species has a simple unguis and uni-articled uropod rami, which preclude it from the ‘cornutus’ group. A short body (less than five times as long as wide) is also characteristic of members of the ‘eximus’ group, and species unclassified at the moment—T. solidus Hansen (1913) (Fig. 80). The new species cannot be included to the ‘eximus’ group as it has a short antennule. Distinction from T. solidus is impossible at present, as that species is known only from a single specimen (holotype) that is covered by crystals, a condition which precludes detailed observation of its morphology.

Larsenotanais amabilis n. sp. (Figs 20–22) Material examined: Holotype: non-ovigerous female, (K 41345), ANT XXII/2, 60°38.35'–60° 38.12'S, 53°57.36'–53° 57.49'W, depth 2893–2893 m, EBS, 30 Jan 2002; Paratypes: two females (one dissected on slides), (K 41346), the same locality; one female, (K 41438), ANT XXII/2, PS 61/42-2, 59°40.29'– 59°40.42'S, 57°35.43'–57°35,27'W, depth 3683–3680 m, EBS, 27 Jan 2002. Diagnosis: as for the genus Etymology: Amabilis (Lat.) = pleasant. Description: Non-ovigerous female (Figs 20A, B). Body short, 4.4 times as long as wide. Carapace 0.8 times as long as wide, tapering proximally, rounded laterally; all pereonites wider than long, clearly rounded laterally; pereonites 1–3 subequal, three times as wide as long; pereonites 4–5 2.4 times as wide as long; pereonite-6 a little shorter than pereonite-5, 2.7 times as wide as long; pereonites 1 and 2 with small lateral seta; pleon 1.5 times as long as carapace; pleotelson gently rounded; caudal projection well-developed, with two setae. Antennule (Fig. 21A): Article-1 about three times as long as wide, 1.4 times as long as article-2 and 3 combined, with five simple setae on inner margin, outer margin with one simple and two pinnate setae at middle and distally; article-2 0.6 times as long as article-3, with one long, simple seta and two pinnate setae distally; article-3 with three long and two short, terminal setae. Antenna (Fig. 21B): Article-1 damaged during dissection; article-2 with one distal seta; article-3 naked; article-4 twice as long as article-5, with two simple and two pinnate setae distally; article-5 with simple seta distally; article-6 with four three terminal setae. Mouth parts: Labrum lost during dissection. Mandible (Fig. 21D,E) stout; molar well-developed (Fig. E’), with regular tubercles on distal edges and three spines on lower margin; lacinia mobilis well-developed, crenulated. Maxillule (Fig. 21F) inner endite with eight terminal setae; palp lost during dissection. Maxilla (Fig. 21G) semi-oval. Labium (Fig. 21H) with bunch of setae on outer corner of inner lobe; outer lobe with small setae. Maxilliped (Fig. 21I) bases with simple setae reaching over endite; each endite armed with two middle setae and two tubercles on distal margin; palp article-1 naked; article-2 wedge-shaped, with three setae (one strongly serrated) on inner margin and one short, simple seta on outer margin; article-3 trapezoidal, with four setae on inner margin; article-4 with one simple seta on outer margin and five terminal setae (one shorter than others). Cheliped (Fig. 22A): Basis 1.8 times as long as wide; merus wedge-shaped with seta ventrally; carpus three times as long as wide, with two simple setae ventrally and two short setae dorsally; carpal shield poorly developed; chela as long as carpus, three times as long as wide; fixed finger with three setae on inner margin and two simple setae ventrally; dactylus almost straight, with one short seta proximally. Pereopod-1 (Fig. 22B): Of walking type; coxa present; basis as long as merus, carpus and half of propodus combined, with one medial seta dorsally; ischium with one simple seta; merus slightly shorter than carpus with three setae distally; carpus a little longer than merus with three setae distally and one minute seta subdis-



tally on ventral margin; propodus slightly longer than carpus with one seta dorsally and one ventral seta subdistally; unguis 1.5 times as long as dactylus; unguis and dactylus combined 0.6 times length of propodus. Pereopod-2 (Fig. 22C): Of walking type; coxa with one seta; basis little shorter than rest of articles combined, with one proximal setae dorsally; ischium with one seta; merus as long as carpus, with three setae distally; carpus as long as merus, with two simple setae, two minute and one spiniform setae distally; propodus with two subdistal setae dorsally (one longer than unguis) and one spiniform seta ventrally; dactylus shorter than unguis. Pereopod-3 (Fig. 22D): Similar to pereopod-2. Pereopod-4 (Fig. 22E): Similar to pereopod-5 (basis damaged).

FIGURE 20. Larsenotanais amabilis n. sp., holotype, female. A) dorsal view; B) lateral view. Scale = 0.1 mm



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FIGURE 21. Larsenotanais amabilis n. sp., female paratype. A) Antennule; B) Antenna; C) Left mandible incisor; D) Right mandible; E) Molar details F) Maxillule; G) Maxilla H) Labium; I) Maxilliped. Scale: A, B = 0.1 mm; C-I = 0.01 mm.

Pereopod-5 (Fig. 22F): Of clinging type; basis wide, with two pinnate setae disto-ventrally; ischium with two setae; merus almost as long as carpus, with two subdistal spiniform setae and numerous combs of spines on ventrally; carpus with hooks distally, one sensory seta dorsally, and with rounded, small (less than half as long as article) prickly tubercles ventrally; propodus five times as long as wide, with one pinnate seta dorsally



and two spiniform setae ventrally, one seta reaching half dactylus; dactylus tipped by simple unguis; dactylus and unguis 0.7 times as long as propodus.

FIGURE 22. Larsenotanais amabilis n. sp., female paratype. A) Cheliped; B) Pereopod-1; C) Pereopod-2; D) Pereopod-3; E) Pereopod-4; F) Pereopod-5; G) Pereopod-6; H) Pleopod; Uropod. Scale = 0.1 mm.



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Pereopod-6 (Fig. 22G): Similar to pereopod-5, but propodus with three terminal setae not reaching half length of unguis. Pleopods 1–5 (Fig. 22H): All pleopods similar; exopod outer margin with eight plumose setae, innermargin with one plumose seta; endopod armed with fourteen plumose setae on outer margin; both rami with gap between proximal seta and others. Uropod (Fig.22I): Basal article less than half as long as endopod; both rami uni-articled; endopod 1.5 times as long as exopod, with pinnate setae on inner margin and six setae terminally; exopod and endopod two-articled; exopod tipped by short and long seta. Distribution: West Antarctic, North Weddell Sea and North of South Shetland Island at depths ranging from 2893 to 3683 m.

Genus Peraeospinosus Sieg, 1986 Diagnosis (after Błażewicz-Paszkowycz, 2005): Body robust, elongate, usually 6–10 times as long as wide; pereonites 1–5 with collar-like grooves, usually anteriorly or posteriorly. Carapace oval or rounded, rarely narrow, rostrum well-developed. Dorsal margin of cheliped carpus with row of minute setae. Labrum hoodshaped, with upper part covered with needle-like setae. Molar process with row of spines on lower margin. Maxillule with nine spiniform setae distally, two semi-fused. Labium external lobes with row of minute setae on edges. Maxilliped bases wide, heart-shaped, endites with two small flat setae (tubercles) on distal margins. Epignath sharply tipped, often bifurcated. Distal seta on propodus of pereopods 4 and 5 longer than unguis; prickly tubercles on pereopods 4–6 surrounded by blunt, calcified spines; unguis of pereopods 4–6 bifurcated. Exopod of pleopods semi-elliptical. Uropod rami usually uni-articled, equal or subequal. Uropod exopod tipped by diminutive and robust seta (at basis almost as wide as exopod). Male: Terminal/swimming type. Gender of generic name: Masculine. Type species: Typhlotanais kerguelenensis Beddard, 1886. Species included: Peraeospinosus brachyurus (Beddard, 1886); Peraeospinosus emergensis BłażewiczPaszkowycz, 2005; P. exiliremis Błażewicz-Paszkowycz, 2005; P. kerguelenensis; P. magnificus (KudinovaPasternak, 1969); P. magnus (Kudinova-Pasternak, 1990); P. peculiaris (Lang, 1968), P. peculiaroides Błażewicz-Paszkowycz, 2005; P. pushkini (Tzareva, 1982); P. rectus (Kudinova-Pasternak, 1968); P. subtigaleatus Błażewicz-Paszkowycz, 2005; P. acruxi n. sp. Remarks: Peraeopsinosus brachyurus (Beddard,1886), originally placeded in Typhlotanais, is transferred to the genus Peraeopsinosus. The type specimens are mounted in Canada Balsam, precluding detailed analysis, but the subequal rami of the uropods, the long setae on the propodus of pereopods 4 and 5 and the minute setae on the cheliped carpus support the reallocation. The species has a long seta on the propodus of pereopod-6. Based on the definition of Peraeospinosus by Błażewicz-Paszkowycz (2005), three species previously included in this genus are now excluded. Typhlotanais adipatus Tzareva, 1982 (originally described in Typhlotanais and included to Peraeospinosus by Sieg, 1986 a) and Typhlotanais angusticheles (KudinovaPasternak, 1987) (originally described in Peraeospinosus) are transferred to Typhlotanais sensu lato. The third species, Typhlotanais (=Peraeospinosus) mixtus Hansen, 1913, probably represents a new genus of Typhlotanaidae (see remarks page 108). As a result, Peraeospinosus currently includes twelve species.



Peraeospinosus acruxi n. sp. (Figs 23-25) Material examined: Holotype: one female, (K 41347), ANT XXII /3, PS 67/74-5, 71°18.35'S–71°18.28'S, 13°57.71'–13°57.31'W, depth 1030–1040 m, epibenthos sledge, 20 Feb 2005; Paratypes: two females, two mancae, (K 41349), the same locality; one manca, (K 41348), ANT XXII/3, PS 67/153-7, 63°19.31'S– 63°19.15'S, 64°36.94'–64°37.18' W, depth 2092–2118 m, epibenthic sledge, 29 Mar 2005. Diagnosis: Body long, about ten times as long as wide; carapace smooth, not swollen, 1.2 times as long as wide, 1.5 times as long as pereonite-1. Pereonites 2–3 rectangular; pereonite-6 short (1.7 times as wide as long). Pleon a little wider than pereonites, as long as carapace, over twice as long as pereonite-6; pleotelson rectangular. Antennule article-3 twice as long as article-2. Propodus of pereopod-2 length:width ratio 4.8: pereopods 4–6 merus and carpus with microtrichae; distal setae on pereopod-6 propodus well-calcified, just as long as dactylus. Pleopods normally developed. Uropod exopod slightly longer than endopod. Etymology: Acrux is the biggest star in the constellation ‘The Southern Cross’. Description: Female holotype body length 5.7 mm. Body ten times as long as wide. Pereonite-1 about as long as wide, tapering distally; pereonites 2–4 longer than wide (Fig. 23A,B), narrower in the middle and wider at proximal and distal margin; pereonite-5 as long as wide; pereonite-6 1.7 as wide as long. Pleon as long as carapace, a little wider than pereonites; all pleonites the same size; pleotelson rectangular. Antennule (Fig. 24A): three-articled; article-1 stout, about 1.3 times as long as articles 2 and 3 combined, with two groups of pinnate setae and two simple setae along article; article-2 half as long as article-3, with two simple and one pinnate setae distally; article-3 with three apical setae. Antenna (Fig. 24B): Article-2 twice as long as article-3, with one simple seta; article-4 eight times as long as wide, less than twice as long as article-5, with three minute setae distally; article-5 with one simple seta distally; article-6 very short, with six terminal setae. Mouth parts: Labrum lost during dissection. Mandible (Fig. 24C,D) stout; molar process well-developed, with strongly crenulated edges, bunch of small spines at “lower” margin; lacinia mobilis well-developed, crenulated. Maxillule (Fig. 24E) endite longer than palp, with eight apical spiniform setae; two of them are semi-fused; palp with two terminal setae. Maxilla (Fig. 24F) semi-triangular. Lobes of labium (Fig. 24G) poorly separated and setose in distal parts; outer lobe with row of short setae along outer margin. Maxilliped (Fig. 24H): coxa reduced; basis fused into heart-shaped plate, as long as wide, with seta reaching half of endites; endite with two simple setae and two small tubercles distally; palp article-1 naked; article-2 wedgeshaped, with two weakly serrated and one strongly serrated setae on inner margin and one minute seta on outer margin; article-3 trapezoidal, with two weakly serrated and two simple setae on inner margin; article-4 slender, with one simple seta on outer margin and five weakly serrated setae distally. Epignath (Fig. 24I) curved, distally simple. Cheliped (Fig. 25A): Basis robust, slightly rounded, twice as long as wide; merus wedge-shaped, with one rod seta; carpus with row of eight small rod setae dorsally and two rod setae ventrally; propodus with seta near fixed finger insertion; fixed finger (propodus projection) tipped with a strong spine, with well-calcified inner margin, and with three setae dorsally and two setae ventrally; dactylus slightly curved, with one short rod seta proximally on dorsal margin. Pereopod-1 (Fig. 25B): Slender (walking type); basis with five short setae along article; ischium short with one seta; merus 1.25 times as long as propodus, with two setae distally; carpus with one simple and two rod setae (dorsal seta long); merus and carpus combined longer than propodus; propodus length: width ratio 7, with one rod setae distally and one minute spiniform seta ventrally; dactylus less than half of unguis length, with long seta. Pereopod-2 (Fig. 25C): Slender (walking type); basis little shorter than rest of articles combined, with three minute setae along article; ischium with short seta ventrally; merus little longer than carpus, each with



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two simple setae and thick spiniform seta distally; carpus with microtrichia; propodus little longer than merus and carpus combined length, with spiniform seta ventrally and one simple and one thick rod seta dorsally; propodus length: width ratio 4.8; dactylus with one simple seta. Pereopod-3 (Fig. 25D): Similar to pereopod-2, but merus and carpus subequal, merus with only one simple seta distally.

FIGURE 23. Peraeospinosus acruxi n. sp., holotype, female. A) dorsal view; B) lateral view. Scale =1 mm.



FIGURE 24. Peraeospinosus acruxi n. sp., female paratype. A) Antennule; B) Antenna; C) Left mandible; D) Right mandible; E) Maxillule; F) Maxilla; G) Labium; H) Maxilliped; I) Epignath. Scale = 0.1 mm.

Pereopod-4 (Fig. 25E): Clinging type; basis twice as long as wide, with one simple and two pinnate setae dorsally and one pinnate seta ventrally; ischium with two setae; merus almost as long as to carpus, with two strong spiniform setae on ventral margin; carpus with two distal tooth, one seta distally and large prickly tubercle covered by little spines and surrounded by dense row of well-calcified blunt spines ventrally; propoA REVISION OF FAMILY TYPHLOTANAIDAE SIEG 1984


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dus with two spiniform setae ventrally and with distal seta twice as long as propodus and dactylus combined length; dactylus tipped by weakly bifurcated unguis.

FIGURE 25. Peraeospinosus acruxi n. sp., female paratype. A) Cheliped; B) Pereopod-1; C) Pereopod-2; D) Pereopod3; E) Pereopod-4; F) Pereopod-5; G) Pereopod-6; G’) ; H) Pleopod; I) Uropod. Scale = 0.1 mm.



Pereopod-5 (Fig. 25F): Similar to pereopod-4. Pereopod-6 (Fig. 25G): Similar to pereopod-5; propodus tipped by three terminal setae just as long as dactylus (two coarsely, one finely serrated). Pleopods 1–5 (Fig. 25H): Rami similar in structure; endopod with row of eighteen setae and exopod with row of six plumose setae on outer margin (length: width ratio of both exopod and endopod 2.7); clear gap between the most proximal and other setae in both rami. Uropod (Fig. 25I): Both rami one-articled. Exopod little longer than endopod; endopod with one pinnate seta at middle, tipped by two simple terminal setae and one pinnate seta; exopod with pinnate seta on outer margin, tipped by strong simple setae. Distribution: Antarctic: Eastern part of the Weddell Sea and off Palmer Archipelago, at depths of 1030– 2118 m. Remarks: Peraeospinosus acruxi n. sp. is the seventh species of Peraeospinosus recorded in the Antarctic. It can be distinguished from the other six species (Peraeospinosus kerguelenensis, P. pushkini, P. emergensis, P. exiliremis, P. magnificus, P. subtigaleatus) by its narrow carapace (longer than wide), by its pereonites 2–4 being longer than wide and by the long dorso-distal rod seta on the carpus of pereopod-1. From the list above, only P. emergensis has elongate pereonites and a similar rod seta on the propodus of pereopod-1, but it has also a round carapace, a more compact carpus of the cheliped (1.25 times as long as wide), a narrower propodus on pereopod-2 (width: length 8), and sixteen setae on the pleopod exopod. In contrast, P. acruxi has a cheliped carpus twice as long as wide, a compact propodus on pereopod-2 (width: length 4.8) and only six setae on the pleopod exopod. Similar elongate pereonites can be observed in P. magnificus, although that species has one long distal seta on the propodus of pereopod-6, while P. acruxi has all three setae shorter than the dactylus. Kudinova-Pasternak (1969; 1970; 1973; 1993) recorded P. magnificus at various localities in the North Pacific (off California, off Japan and Alaska), the South Atlantic (Argentina), and the West Antarctic (Bransfield Strait) as well (see Błażewicz-Paszkowycz, 2005). That material is not available for study, so the apparently wide distribution of the species cannot be confirmed. However, it seems likely that Kudinova-Pasternak dealt with two (or more) species with distinct areas of distribution (North Pacific and West Antarctic).

Genus: Pulcherella n. gen Diagnosis: Carapace almost as long as pereonite-1. Pereonites 1–3 longer than wide; pereonites 5 and 6 wider than long. Mandible molar with teeth or tubercles on margin. Maxilliped bases as wide as long, with long setae (reaching end of endites); endites with two tubercles and with one seta distally. Cheliped bases separated from pereonite-1 by gap ventrally. Pereopod-1 carpus with long seta (over half of propodus length) and unguis with spatulate apex. Pereopods 4–6 carpus with large (about as long as half of article) but flat prickly tubercles, merus with one spiniform and one simple seta; unguis bifurcated. Pleopod rami with gap between most proximal and other setae. Uropod endopod two-articled, exopod one-articled. Exopod little longer than endopod proximal article. Male: Unknown. Etymology: The generic name is derived from the name of the type species Typhlotanais pulcher Hansen, 1913. Type species: Typhlotanais pulcher Hansen, 1913. Gender of generic name: Feminine. Species included: Pulcherella (= Typhlotanais) pulcher; P. (= Typhlotanais) filatovae (Kudinova-Pasternak, 1975); P. (= Typhlotanais) spiniventris (Dollfus, 1897); Pulcherella juraszi n. sp. Remarks: Pulcherella n. gen. is characterized by having a carapace shorter than pereonite-1, pereonites 1–3 longer than wide and flat prickly tubercles on pereopods 4–6. The combination of these three character



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allows clear separation of the new genus from the other typhlotanaids. The long first three pereonites, which are almost as long as the carapace, are elsewhwere found only in the genus Hamatipeda, although in contrast to Pulcherella its members have a trifurcated unguis and strong spines instead of prickly tubercles on the carpus of pereopods 4–6. Further differences can be observed in the mouthparts, where the maxilliped endites have two tubercles distally in Pulcherella and none in Hamatipeda.

Key for determination of Pulcherella females* * Pulcherella spiniventris (Dollfus, 1897) was omitted from the key owing to the poor condition of the examined lectotype and paralectotypes.

1. Pereopods 2 and 3 carpus distal setae short (shorter than half of propodus)............................... P. filatovae - Pereopods 2 and 3 carpus distal setae long (almost as long as propodus) ...................................................2 2. Pereopods 2 and 3 propodus distal seta three times as long as dactylus and unguis combined, pereopod-6 propodus four times as long as wide, pereopods 4–6 unguis half as long as dactylus ................. P. pulcher - Pereopod 2 and 3 propodus distal seta just reaching end of unguis, pereopod-6 propodus twice as long as wide, pereopods 4–6 unguis almost as long as dactylus........................................................ P. juraszi n. sp.

Pulcherella filatovae (Kudinova-Pasternak, 1975) n. comb. (Figs 26–28) Typhlotanais filatovae: Kudinova-Pasternak (1975) 103: 214, 217, 226. Larsen (2005): 209.

Material examined: Holotype: female, Mc-979, Akademik Kurchatov Sta. 880(D1), 57°07.4'S, 26°40'W, depth 497 m. Other material examined: one female, (K 41415), ANT XIX/3, PS 61/43-8, 60°27.12'– 60°27.24'S, 56°5.10'–56°05.25'W, depth 3961.0–3963.4 m, 04 Feb 2002; one female, (K 41416), ANT XIX/3, PS 61/46-7, 60°38.35'–60°38.12'S, 53° 57.36'–53° 57.49'W, depth 2893.2–2892.8 m, 30 Jan 2002; one female in tube, (K 41417), ANT XXII/3, PS 67/151-7, 61°45.52'–61°45.42'S, 47°07.68'–47°08.04'W, depth 1182– 1185 m, EBS, 21 Mar 2005; one female, (USNM 1100139), Eltanin 9/724, 54°05'–53°04'S, 033°43'– 033°37'W, depth 2714–2727 m, 9 Sep 1963; one female, (USNM 1100140), Eltanin 6/339, 53°05'–53°08'S, 059°31'–059°24'W, depth unknown, 3 Dec 1962. Diagnosis: Pereopods 2 and 3 merus and carpus with short setae (clearly shorter than half of following article), propodus with two dorsal seta not reaching beyond unguis; pereopods 4 and 5 propodus with distal seta as long as dactylus and unguis combined. Description: Female body (Figs 26A, 27 A, B) cylindrical, body long, about ten times as long as wide, 1.8–2.8 mm long (according to Kudinova-Pasternak, 1975). Cephalothorax 0.8 times as long as pereonite-1. Pereonites 1–3 longer than wide, pereonite-1 shorter than pereonite-2, pereonites 2 and 3 subequal, almost twice as long as wide; pereonite-4 as long as wide; pereonite-5 little wider than long; pereonite-6 half as long as pereonite-5, almost twice as wide as long; pleon short (including pleotelson only about 12 % of total body length); pleonites subequal, carrying pleopods; pleotelson as long as combined length of two pleonites. Antennule (Fig. 27C): Almost as long as cephalothorax; article-1 with one simple distal setae; article-3 1.5 times as long as article-2 that has two long (as long as article-3) simple distal setae; article-3 five times as long as wide, with three simple distal setae and one aesthetasc. Antenna (Fig. 27D): Article-2 twice as long as article-3, with one dorsal seta; article-3 with one dorsal seta; article-4 twice as long as article-5, with one pinnate seta, two long and one short simple distal setae; article-5 with one distal seta; article-6 minute, with four distal setae.



FIGURE 26. Pulcherella filiatovae. A) Holotype, female, dorsal view, present stage; B) Pereopod-1; C) Pereopod-2; D) Pereopod-3; E) Pereopod-4; F) Pereopod-5; G) Pereopod-6. Scale = 0.1 mm.

Mouthparts: Labrum (Fig. 27E) flat, with setules on lateral margin. Mandibular molar process broad, longer than incisor, with sharp denticles on distal edge. Left mandible (Fig. 27F) lacinia mobilis longer than incisor, crenulated, with four denticles. Labium (Fig. 27I) with inner and minute outer processes, both with a few setules. Maxillule (Fig. 27G) endite with eight distal spiniform setae; palp shorter than endite, with two terminal setae. Maxilla (Fig. 27H) oval, naked. Maxilliped (Fig. 28A) basis short (as long as wide), with one long seta at palp insertion; endites with two tubercles and long inner seta on distal margin; palp article-1 A REVISION OF FAMILY TYPHLOTANAIDAE SIEG 1984


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naked; article-2 with one outer and three inner setae; article-3 with four bipinnated inner setae; article-4 with one outer and five bipinnate inner setae. Epignath (Fig. 27J) longer than maxillule endite, rounded distally.

FIGURE 27. Pulcherella filiatovae (ANDEEP material). A) Female dorsal view; B) Female lateral view; C) Antennule; D) Antenna; E) Labrum; F) Left mandible; G) Maxillule; G’) Palp; H) Maxilla; I) Labium; J) Epignath. Scale A-D, I, J = 0.1 mm; E-H, 0.01 mm.



FIGURE 28. Pulcherella filiatovae (ANDEEP material). A) Maxilliped; B) Cheliped; C) Pereopod-1; D) Pereopod-2; E) Pereopod-3; F) Pereopod-4; G) Pereopod-5; H) Pereopod-6; I) Pleopod; J) Uropod. Scale = 0.1 mm.

Cheliped (Fig. 28B): Basis less than twice as long as wide, 0.7 times as long as carpus; merus with one seta ventrally; carpus shorter than propodus including fixed finger, with two ventral and two small dorsal setae; propodus with one seta at dactylus insertion; fixed finger with two ventral setae and three on inner margin, with three prominent denticles on inner margin; dactylus as long as fixed finger. Pereopod-1 (Fig. 28C): Coxa present; basis shorter than merus, carpus, and propodus combined, with two dorsal setae; ischium with one seta ventrally; merus shorter than carpus; carpus shorter than propodus, with two short and one long (longer than half of propodus) dorsal setae and one minute subterminal seta on ventral margin; propodus as long as half of basis, with three dorso-distal setae, and one ventral seta; dactylus and unguis longer than propodus; dactylus shorter than unguis; unguis with spatulate tip. A REVISION OF FAMILY TYPHLOTANAIDAE SIEG 1984


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Pereopod-2 (Fig. 28D): As pereopod-1 except: basis as long as merus, carpus and propodus combined, with one dorsal seta. Carpus with five short distal setae. Propodus with two long dorso-distal setae reaching half length of unguis and one ventral seta. Dactylus and unguis combined shorter than propodus. Pereopod-3 (Fig. 28E): As pereopod-2 except: basis with three dorsal seta. Carpus with four simple distal setae. Pereopod-4 (Fig. 28F): Clinging type; basis with two bipinnate ventro-medial setae; ischium with one seta; merus with one simple and one spiniform seta; carpus with flat, large (as long as half of article) prickly tubercles, one simple dorso-distal seta and two spiniform ventro-distal hooks; propodus with two ventro-distal spiniform seta and one dorso-distal setae as long as dactylus and unguis combined; unguis bifurcated. Pereopod-5 (Fig. 28G): Similar to pereopod-4. Pereopod-6 (Fig. 28H): Similar to as pereopod-5 but propodus with three dorso-distal setae. Pleopod (Fig. 28I): Basal article naked; exopod with one inner and seven outer plumose setae; endopod with 13 inner plumose setae, large gap between proximal seta and other setae. Uropod (Fig. 28J): Basal article about quarter as long as endopod proximal article; endopod two-articled; proximal article-1 with simple seta distally; distal article with five four setae; exopod one-articled as long as endopod proximal article, tipped with short and long setae. Distribution: West Antarctic: vicinity of the Falkland Islands and North-East of Elephant Island, at depths of 1182 to 3963 m. Remarks: The material of Pulcherella filatovae studied by Kudinova-Pasternak (1975) consisted of five females and one manca-III. From that collection only the holotype was available for study (Figs 26A–G). It lacked the carapace, the pleotelson and most appendages. The collection of a few specimens of P. filatovae during the ANDEEP Program enabled the redescription of the species. P. filatovae can be distinguished from the other members of the genus by having short setae on the merus, carpus and propodus of pereopods 2 and 3, and relatively long distal setae on the propodus of pereopods 4–6 (see remarks under P. pulcher).

Pulcherella pulcher (Hansen, 1913) n. comb. (Figs 29–31) Typhlotanais pulcher: Hansen (1913) 3(3): 39–40; Nierstrasz (1913) 32 (a): 36; Stephensen (1913) 22: 267–268, 418; Stephensen (1932) 6: 350; Stephensen (1936) Medd. Gronl. 80(2): 29; Lang (1970) (2), 23 (4): 276, 287; Holdich & Jones (1983a): 62–63; Holdich & Jones (1983b) 17: 158, 172, 175, 177–179; Holdich & Bird (1985): 443; Hassack & Holdich (1987) 16(3): 224 225, 227; Larsen (2005): 210.

Material examined: Holotype: Ingolf Station, South of Davis Strait, St. 38, 59°12'N, 5l°05'W, depth 3420 m (1870 fm); two females (one dissected on slides), BIOICE 2836, Shipek Grab, 26 Aug 1995, 63º16.64’N, 16.52º00’W, depth 615 m, bottom temp. 6.74ºC, salinity 35.12, silty sand. Diagnosis: Pereopods 2 and 3 with long setae on merus and carpus, almost as long as following article respectively; pereopods 4 and 5 propodus distal seta not reaching the end of dactylus. Description: Female (Figs 29A,B). Body 2.2 mm long, cylindrical, about nine times as long as wide; cephalothorax shorter than pereonite-1. Pereonites 1–4 longer than wide. Pereonites 5 and 6 wider than long; pereonite-5 1.5 times as long as pereonite-6; pereonite-6 twice as wider as long. Pleon short (including pleotelson only about 12 % of total body length); pleonites subequal, carrying pleopods; pleotelson as long as combined length of two pleonites. Antennule (Fig. 30A): Article-1 longer than distal two articles combined, four times as long as wide, with two medial and two distal simple setae; article-2 less than half as long as article-3, with two simple distal setae; article-3 about six times as long as wide, less than half of article-1 length, with three simple distal setae.



FIGURE 29. Pulcherella pulcher, holotype, female. A) lateral view; B) dorsal view; C) Pulcherella (=Typhlotanais) spiniventris. Scale: 1 mm.

Antenna (Fig. 30B): Article-1 not broader than following articles, naked; article-2 little longer than article-3, with one dorsal seta; article-3 with one dorsal seta; article-4 about seven times as long as wide, three times as long as article-5, with two pinnate setae, one short and two long simple setae distally; article-5 with one distal seta; article-6 minute, with four distal setae. Mouthparts: Labrum (Fig. 30C) flat and with many setules on distal margin. Mandibular molar process broad, as long as incisor, with terminal tubercles on distal edge. Left mandible (Fig. 30D) lacinia mobilis longer than incisor, crenulated; incisor shaped like a blunt spine. Right mandible (Figs 30E,E’) incisor tapering into two denticles. Labium (Fig. 30I) with inner and outer lobe covered with a few setules. Maxillule (Fig. 30F) endite with eight distal spiniform setae; palp longer than endite, with two terminal setae. Maxilla not recovered. Maxilliped (Fig. 30G) basis with long seta; endites as wide as basis, with two distal tubercles and one middle seta distally. Palp article-1 with microtrichae on outer margin; article-2 with one small outer and three inner setae; article-3 with four inner setae; article-4 with one outer and five inner pinnate setae. Epignath (Fig. 30H) rounded distally. Cheliped (Fig. 31A): Basis twice as long as wide, half as long as carpus, not reaching pereonite-1 ventrally; merus with one ventral seta; carpus marginally shorter than propodus including the fixed finger, with



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two ventral and two small dorsal setae; propodus with one seta at dactylus insertion; fixed finger with two ventral setae and three on inner margin, well-calcified inner margin; dactylus marginally shorter than fixed finger, with two strong spiniform setae on inner margin (Fig. 28A’).

FIGURE 30. Pulcherella pulcher. A) Antennule; B) Antenna; C) Labrum; D) Left mandible; E) Right mandible; E') Right mandible incisor details; F) Maxillule; F’) Palp; G) Maxilliped; H) Epignath; I) Labium. Scale: A, B,G-I = 0.1 mm; C-F = 0.01 mm.



FIGURE 31. Pulcherella pulcher, female. A) Cheliped; A’) Cheliped dactylus detalis; B) Pereopod-1; C) Pereopod-2; D) Pereopod-3; E) Pereopod-4; F) Pereopod-5; G) Pereopod-6; H) Pleopod; I) Uropod. Scale = 0.1 mm.



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Pereopod-1 (Fig. 31B): Coxa with one seta; basis shorter than merus, carpus, and propodus combined, with three simple ventral and two small pinnate setae dorsally; ischium with one seta; merus as long as carpus, with two distal setae; carpus little shorter than propodus, with three short and one long (longer than half of propodus) setae distally; propodus longer than half of basis, with three dorso-distal setae and one ventral seta; dactylus and unguis combined shorter than propodus; dactylus half as long as unguis, with one proximal seta; unguis with spatulate tip. Pereopod-2 (Fig. 31C): Coxa with one seta; basis as long as merus, carpus and propodus combined, with one dorsal seta; ischium with one ventral seta; merus with one short and one long (as long as carpus) distal setae; carpus with one short and two long (almost as long as propodus) setae; propodus with one short and one long setae (three times as long as dactylus and unguis combined length); dactylus naked. Pereopod-3 (Fig. 31D): As pereopod-2, but merus with only one seta. Pereopod-4 (Fig. 31E): Clinging type; basis with plumose ventro-medial seta and small dorso-proximal seta; ischium with two setae; merus little longer than carpus, with one simple and one spiniform ventro-distal seta; carpus with one distal seta and one large and flat prickly tubercle; propodus with two ventro-distal spiniform setae, and distal seta reaching about half length of dactylus; dactylus and unguis together shorter than propodus, unguis with bifid tip. Pereopod-5 (Fig. 31F): As pereopod-4. Pereopod-6 (Fig. 31G): As pereopod-5 except basis naked; carpus with one pinnate dorso-distal seta. Propodus with three dorso-distal setae. Pleopod (Fig. 31H): Basal article naked; exopod with one inner and 14 outer plumose setae, with gap between proximal seta and other setae; endopod with 17 outer plumose setae, large gap between proximal seta and other setae. Uropod (Fig. 31I): Basal article 0.3 times as long as endopod, naked; endopod 2-articled; proximal article with one simple and two pinnate medial setae; distal article with one pinnate and four simple setae; exopod 1articled, with one medial seta and one short simple and one long thick setae distally. Distribution: The species is known from the North Atlantic: Davis Strait (holotype) at a depth of 3422 m (Hansen 1913), British waters (Lynn of Lorn, Western Scotland) at depth 38 m (muddy clay) and widely in the Rockall-Biscay area of the NE Atlantic (Bird pers. comm.). Remarks: Pulcherella pulcher can be immediately distinguished from P. filatovae by presence of one long dorsal seta on the propodus of the second and third pereopods reaching far over the unguis. In P. filatovae and P. juraszi n. sp. there are two setae there, both shorter than or subequal to unguis. According to observations by Hassack & Holdich (1987), P. pulcher constructs tubes little longer than the specimen (5 mm) covered by a fine layer of mud. The tube is sealed in front and distally deflected ventrally in the same manner as the pleotelson.

Pulcherella spiniventris (Dollfus, 1897) n. comb. (Fig. 29C) Typhlotanais spiniventris: Dollfus (1897) 22: 210-212; Norman (1899) (7), 3: 340; Tattersall (1904): 601; Tattersall (1905) 2:5; Nierstrasz (1913): 34; Vanhoffen (1914) 15: 472; Dollfus (1938) 97: 94; Belloc (1960) 1188: 2; Lang (1970) (2), 23 (4):277-278, 287.

Material examined: Lectotype: female, 365104, CAM, Hirondelle Station 112, between Pico and Sao-Jorge (Azores), depth 1287 m, fond sable vaseux, 1 Jul 1887; Paralectotypes: two females (broken), 365168, one dissected on slides, the same locality; one female, 365158, Station 226, Pico Faya Strait, depth 130 m, gravel, sand, shell grit.



Distribution: NE Atlantic, Azores, depth 130–1287m. Remarks: The lectotype and paratypes of Pulcherella spiniventris are covered by crystalline deposits which preclude detailed morphological observations for defining character states and differences from the other members of the genus. The carapace being shorter than pereonite-1, the long dorso-distal seta on pereopod-1 and the flat prickly tubercles on pereopods 4 and 5 undoubtedly classify the species in the genus Pulcherella. Additional material from, or close to, the type locality would help to clarify the taxonomic status of this species versus the similar congener P. pulcher.

Pulcherella juraszi n. sp. (Figs 32–34) Material examined: Holotype: female, (K 41342), ANT XXII/3, PS 78-9, 71°09.39'–71°09.36'S 13°59.30'– 13°58.81'W, depth 2156–2147 m, epibenthic sledge, 21 Feb 2005; Paratypes: two females, (K 41343), ANT XXII/3, PS 121-11-E, 63°37.73'-63°37.55'S, 50°38.09'–50°38.37'W, depth 2663–2659 m, epibenthic sledge, 15 Mar 2005; two females, (K 41344), ANT XXII/3, PS 133-2, 62°46.49'–62°46.38'S, 53°03.50'–53°03.98', depth 1584–1579 m, epibenthic sledge, 16 Mar 2005. Diagnosis: Pereopod-1 carpus with long seta (little shorter than propodus), dactylus and unguis combined longer than propodus; pereopods 2 and 3 carpus with seta (-ae) about half as long as following article; pereopods 4–6 with long unguis (over half as long as dactylus); pereopod-6 compact (propodus twice as long as wide). Uropod exopod little longer than endopod proximal article. Description: Non-ovigerous female. Body length 1.7 mm (Figs 32A, B), long, ten times as long as broad. Carapace smooth, rounded laterally, 1.5 times as long as wide, 18 % of total body length; rostrum blunt. Pereonite-1 trapezoid, tapering distally as long as wide; pereonites 2 and 3 slightly rounded 1.5 times as long as wide; pereonites 4 and 5 little shorter than pereonite-3; pereonite-6 shortest, almost twice as wide as long. Pleon little longer than carapace, tapering distally; pleonites 1–5 similar in size; pleotelson semi-rounded; caudal process poorly developed. Antennule (Fig. 33A): Shorter than carapace; article-1 half as long as article-2, with one simple and two pinnate setae at middle and one seta distally; article-2 with one seta; article-3 five times as long as wide, 2.5 times as long as article-2, tipped by two short and three long setae. Antenna (Fig. 33B): Article-2 about half as long as article-3; article-3 with one small seta; article-4 thick, three times as long as wide, just twice as long as article-5, with two long, two short and two pinnate setae distally; article-5 with one distal setae; article-6 tipped by four setae distally. Mouth parts: Labrum (Fig. 33C) covered by numerous small setae. Mandible (Figs 33D, D’,E) stout; molar process well-developed, distal margin with about eight tubercle-like structures; lacinia mobilis welldeveloped, with four teeth distally. Maxillule (Fig. 33F) endite with eight spiniform setae distally. Maxilla (Fig. 33G) suboval. Maxilliped (Fig. 33I): bases fused, about as long as wide, with simple setae reaching end of endites; endite with one tubercle and one seta in mid-distal corner; palp four-articled: article-1 naked; article-2 wedge-shaped, with three setae on inner margin and one seta on outer margin; article-3 trapezoidal, with three bipinnate and one short simple setae on inner margin; article-4 with one simple seta on outer margin and five bipinnate terminal setae. Epignath lost during dissection. Labium (Fig. 33H) bilobed, outer corner of inner lobe and outer lobe with minute setae.



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FIGURE 32. Pulcherella juraszi n. sp. A) Holotype, female, dorsal view; B) Holotype, female, lateral view. Scale = 0.1 m.



FIGURE 33. Pulcherella juraszi n. sp., female paratype. A) Antennule; B) Antenna; C) Labrum; D) Left mandible; D’) Molar process; E) Right mandible; F) Maxillule; G) Maxilla; H) Labium; I) Maxilliped; J) Cheliped. Scale: A, B, J = 0.1 mm, C-H = 0.01 mm.

Cheliped (Fig. 33J): Basis twice as long as wide, with one seta long on dorsal margin; merus wedgeshaped, with one long seta ventrally; carpus twice as long as broad, with two dorsal setae and with two long setae ventrally; propodus and fixed finger little longer than carpus, about 2.5 times as long as wide; fixed finger with three setae and three calcified teeth on inner margin and two setae on ventral margin; dactylus with one seta in proximal part of dorsal margin.



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Pereopod-1 (Fig. 34A): Coxa with seta; basis as long as merus, carpus and half of propodus combined; ischium with simple seta; merus little shorter than carpus, with simple seta distally; carpus with one short seta ventrally, two setae dorso-distally (one little shorter than propodus); propodus 1.4 times as long as carpus, with one subdistal seta ventrally and two subdistal setae dorsally; dactylus little shorter than propodus, dactylus and unguis combined longer than propodus.

FIGURE 34. Pulcherella juraszi n. sp., female paratype. A) Pereopod-1; B) Pereopod-2; C) Pereopod-3; D) Pereopod-4; E) Pereopod-5; F) Pereopod-6; G) Uropod. Scale = 0.1 mm.



Pereopod-2 (Fig. 34B): Coxa without seta; basis as long as merus, carpus and half of propodus combined with one pinnate seta dorsally; ischium with simple seta; merus little shorter than carpus, with simple seta distally; carpus with two long and two short setae distally; propodus 1.2 times as long as carpus, with one subdistal seta ventrally and two subdistal setae dorsally; dactylus little shorter than unguis; dactylus and unguis combined a little shorter than propodus. Pereopod-3 (Fig. 34C): Similar to pereopod-2, but merus with two seta and one of propodus dorsal seta shorter than the others. Pereopod-4 (Fig. 34D): Clinging type; basis with one pinnate seta proximally on dorsal margin; ischium with one seta ventrally; merus with two spiniform setae distally; carpus with spiniform seta distally and flat prickly tubercle as long as half of carpus; propodus over three times as long as wide, with two spiniform ventro-distal setae and with distal seta reaching dactylus; unguis simple, little shorter than dactylus. Pereopod-5 (Fig. 34E): Similar to pereopod-4. Pereopod-6 (Fig. 34F): Similar to pereopod-4, but propodus compact, twice as long as wide, with three distal setae, little shorter than dactylus. Pleopod: Both exopod and endopod without setae. Uropod (Fig. 34G): Endopod two-articled, 2.5 times as long as basal article; proximal article almost as long as distal article, with one distal seta; distal article tipped by five simple setae; exopod one-articled, a little longer than endopod proximal article, tipped by strong, long and short simple setae. Distribution: Antarctic: Weddell Sea, in the vicinity of Elephant Island at depths of 1579–2663 m. Remarks: The inclusion of Pulcherella juraszi n. sp. in the genus Pulcherella may be doubtful owing to the lack of setation on the pleopods, and the presence of a simple and relatively long unguis on pereopods 4– 6. Also it has two spiniform setae on the merus of pereopods 4–6 instead of one simple and one spiniform seta as in the other species of Pulcherella. Although these characters clearly distinguish P. juraszi from the other Pulcherella species, the new species has been included in the genus as it shares more characters in common with its members than with the other typhlotanaids.

? Pulcherella pulcher Hansen, 1913 Material examined: two females, ANT XXII/3, PS 67/80-9-Epi, 70°39.07'–70° 39.22'S, 14°43.36'– 14°43.39'W, depth 3103–3102 m, epibenthic sledge, 23 Feb 2005. Remarks: Two females of the new genus Pulcherella found in the ANDEEP material are so closely similar to North Atlantic Pulcherella pulcher (Hansen, 1913) that I could not find any clear differences. Such observation suggests a wide distribution of P. pulcher unless North Atlantic and Antarctic species represent cryptic taxa (see remarks on tanaid distribution).

Genus: Torquella n. gen. Diagnosis: Body long, about 6–7 times as long as wide. Pleon short, less than 20 % of total body length. Pereonite-1 anterior edge is deeply concave. Mandible molar process with well-calcified and crenulated edge, occasionally with teeth. Maxilliped basis with seta at least as long as three-quarter of endites; endite distal margin with two tubercles and usually one seta. Cheliped basis posterior margin separated from pereonite-1 by gap. Chela slender, little longer than cheliped carpus. Pereopods 2 and 3 with spiniform seta on carpus and with long seta on propodus; occasionally with rod setae. Pereopods 4–6 unguis simple, with row of spines ventrally; prickly tubercles large (half as long as carpus), surrounded by row of spines. Uropod rami two-articled; exopod usually reaching three-quarters of endopod; exopod distal article twice as long as proximal article. A REVISION OF FAMILY TYPHLOTANAIDAE SIEG 1984


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Male: Unknown. Etymology: Torquis (Latin) = collar. The name reflects the collar-like character of the pereonite-1. Gender of generic name: Feminine. Type species: Typhlotanais longisetosus Kudinova-Pasternak, 1990. Species included: Torquella (= Typhlotanais) angularis (Kudinova-Pasternak, 1966); T. (=Typhlotanais) elegans (Kudinova-Pasternak, 1978); T. (=Typhlotanais) grandis (Hansen, 1913); T. (=Typhlotanais) longisetosa; T. (= Typhlotanais) parangularis (Kudinova-Pasternak, 1975); T. (= Typhlotanais) rotundirostris (Lang, 1970); T. eltaninae n. sp.; T. galatheae n. sp. Remarks: The genus is established to designate species previously included in Typhlotanais sensu lato sharing the cluster of characters listed in diagnosis above. The collar-shape of pereonite-1, characteristic for Torquella, can be observed (to some extent) in T. variabilis Hansen, 1913; however, that species has prickly tubercles that are not surrounded by rows of spines and the uropod exopod is a little longer than the endopod. Again, a collar-shape of pereonite-1 can be observed in the genus Typhlamia (= Typhlotanais) but it differs in having small prickly tubercles and an antennule twice as long as the carapace. Prickly tubercles surrounded by spines characteristic for Torquella occur also in pereopods 4–6 of Peraeospinosus (see Błażewicz-Paszkowycz, 2005). Except for this single feature no other similarity between Torquella n. gen. and Peraeospinosus Sieg, 1986 can be observed.

Key for identification of Torquella females 1. Pereonite-2 twice as long as pereonite-1 ......................................................................................................2 - Pereonite-2 clearly less than twice as long as pereonite-1............................................................................3 2. Carapace round, as long as wide; pereopod-2 merus with two setae shorter than a quarter of the carpus, carpus with only two long setae and three short setae................................................................ T. angularis - Carapace longer than wide; pereopod-2 merus with four long setae reaching half of carpus, carpus with six setae not reaching half of propodus ......................................................................................... T. elegans 3. Uropod exopod as long as proximal article of endopod ........................................................T. parangularis - Uropod exopod longer than endopod proximal article .................................................................................4 4. Pereonite-5 about as long as wide.................................................................................................................5 - Pereonite-5 longer than wide ........................................................................................................................6 5. Pereopod-6 propodus with three short setae distally ........................................................... T. rotundirostris - Pereopod-6 propodus with two short and one long setae distally......................................T. eltaninae n. sp. 6. Pereopods 2 and 3 merus and carpus with distal setae reaching half of following article, propodus with distal seta reaching over unguis.............................................................................................. T. longisetosa - Pereopods 2 and 3 merus and carpus with distal setae shorter than half of following article, propodus with distal seta not reaching over unguis..............................................................................................................8 7. Pereopod-1 basis shorter than merus and carpus combined, propodus with long seta ventrally and two rod setae dorsally ................................................................................................................................. T. grandis - Pereopod-1 basis longer than merus and carpus combined, propodus with spiniform seta ventrally, and regular seta dorsally ......................................................................................................... T. galatheae n. sp.

Torquella angularis (Kudinova-Pasternak, 1966) n. comb. (Fig. 35) Typhlotanais sp. A: Belyaev (1966): 88



Typhlotanais angularis: Kudinova-Pasternak (1966) 45: 529–531; Kudinova-Pasternak (1968) 2: 73; Lang (1970) (2) 23: 270, 287; Shiino (1970) 1: 98; Morino (1971) 18(5): 354; Kudinova-Pasternak (1975) 103: 213; Kudinova-Pasternak (1977 (1978)) 108: 127.

Material examined: Holotype: female, Mc-940; Vitjaz Station 4074, 40°19.7'N, 175°45.3'E, depth 6065 m.

FIGURE 35. Torquella angularis. A) Holotype, female, dorsal view, present stage; B) Pereopod-1; C) Pereopod-2. Scale = 0.1 mm

Diagnosis: Carapace round, as long as wide; pereonite-2 twice as long as pereonite-1. Pereonite-3 over 1.3 times as long as wide. Chela 3.7 times as long as wide. Antennule article-3 about three times as long as article-2. Pereopod-1 merus and carpus little shorter than basis, merus with short setae, propodus with one long simple seta and one rod seta dorsally; merus with two short setae, carpus with two long setae just reaching half of propodus; pereopods 4–5 propodus distal seta reaches end of unguis. Distribution: Known only from the type locality. Remarks: Torquella angularis is known from only one specimen recorded from the North Pacific. The poor condition of the holotype excludes redescription of the species (Fig 31). Based on observation of the holotype remains and information gained from Kudinova-Pasternak’s drawings and description (pages 529– 531) the diagnosis of the species could be presented above.

Torquella elegans (Kudinova-Pasternak, 1978) n. comb. (Fig. 36–37) Typhlotanais elegans: Kudinova-Pasternak (1977 (1978)) 108: 127–130, 131.

Material examined: Holotype Mc- 986, Vitjaz Sta. 7391, 24°06.8'N, 143°47.1'E, depth 6330 m. A REVISION OF FAMILY TYPHLOTANAIDAE SIEG 1984


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Diagnosis: Carapace 1.2 times as long as wide, 2.5 times as long as pereonite-1. Pereonite-2 over twice as long as pereonite-1; pereonite-3 over 1.3 times as long as wide. Antennule article-3 three times as long as article-2. Chela 2.8 times as long as wide. Pereopod-1 merus and carpus little shorter than basis, merus with short setae, propodus with three relatively long simple setae; pereopod-2 propodus with two long setae dorsally, merus with four long setae (just half of carpus), carpus with six long setae just reach half of propodus; pereopods 4–5 propodus with distal seta reaches end of dactylus; pereopods 4–6 with numerous spines around prickly tubercles; pereopod-6 propodus distal setae subequal. Uropod exopod reaching half of endopod distal article. Complementary description: Female (Fig. 36K). Body length 2.2–5.4 mm (according to Kudinova-Pasternak, 1978), 7.4 times as long as wide; carapace little longer than wide, smooth, 15% of body length, narrowing rostrally, rounded laterally, rostrum acute; pereonite-1 twice as wide as long, latero-proximal corners expanded forward. Pereonites 2, 3 and 5 similar size, little longer than wide; pereonite-4 1.3 times as long as wide; pereonite-6 1.3 times as wide as long. Pleon 15% of body length, pleonites 1–5 similar in size; pleotelson rectangular. Antennule (Fig. 36A): Article-1 slender, more than three times as long as wide, with simple and pinnate setae in two groups medially and distally on outer margin and simple proximal seta on inner margin; article-2 with two setae distally, article-3 just three times as long as article-2 with five setae distally. Antenna (Fig. 36B): Article-2 long, three times as long as article-3, with one serrated seta; article-3 with one seta; article-4 is 2.5 times as long as article-5, with two long, one short simple and four pinnate setae distally; article-5 with one simple seta distally; article-6 very short, with four terminal setae. Mouth parts: Labrum (Fig. 36C) hood-shaped with numerous setae. Mandible (Figs 36D,E) stout; molar process well-developed, edges supported with tubercle-like structures; lacinia mobilis well-developed, crenulated. Maxillule (Fig. 36F) endite with eight apical spiniform setae; palp lost during dissection. Maxilla (Fig. 36G) triangular. Labium (Fig. 36H) bilobed, outer lobe and outer corner of inner lobe setose. Maxilliped (Fig. 36I): bases fused, 2-lobed with simple setae reaching half of endites; endite armed with plumose seta in distolateral margin and two tubercles on distal margin; palp 4-articled: article-1 naked; article-2 wedge-shaped with three strong serrated setae on inner margin and one short seta on outer margin; article-3 with three strong serrated setae and one long simple setae on inner margin; article-4 with four strong serrated setae and two short setae. Epignath (Fig. 36J) tip rounded. Cheliped (Fig. 37A): Basis 1.8 times as long as wide; merus wedge-shaped with one seta ventrally; carpus about 2.5 times as long as broad, with two dorsal setae and with two long setae ventrally; propodus and fixed finger little longer than carpus, about three times as long as wide, with one seta on inner side and one seta on outer side near insertion of dactylus; fixed finger with three setae on inner margin and with two setae on ventral margin. Pereopod-1 (Fig. 37B): Of walking type; coxa with one seta; basis shorter than ischium, merus and carpus combined, with three simple setae dorsally and one seta ventrally; ischium short with one seta; merus little shorter than carpus, with two setae distally; carpus with four setae distally and one subdistal short seta ventrally; propodus 1.3 times as long as carpus with one seta ventrally and three setae dorsally; dactylus with long seta; unguis 1.5 times as long as dactylus. Pereopod-2 (Fig. 37C): Of walking type; coxa with one seta; basis with one ventral and one simple, dorsal setae, as long as merus, carpus and propodus combined; ischium with one seta; merus almost as long as carpus, with four long simple setae distally; carpus with six simple and one spiniform setae distally; propodus a little shorter than merus and carpus combined, with spiniform seta ventrally and two setae dorsally reaching half length of unguis; dactylus with one seta, little shorter than unguis. Pereopod-3 (Fig. 37D): Similar to pereopod-2, but merus with two setae only.



FIGURE 36. Torquella elegans, female holotype. A) Antennule; B) Antenna; C) Labrum; D) Left mandible; E) Right mandible; F) Maxillule; G) Maxilla; H) Labium; I) Maxilliped; J) Epignath; K) Female dorsal view. Scale: K = 1 mm, A, B, H-J = 0.1 mm, C-G = 0.01 mm.



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FIGURE 37. Torquella elegans, female holotype. A) Cheliped, B) Pereopod-1; C) Pereopod-2; D) Pereopod-3; E) Pereopod-4; F) Pereopod-5. Scale = 0.1 mm.

Pereopod-4 (Fig. 37E): Basis with one seta ventrally and one seta dorsally; ischium with two seta ventrally; merus with two spiniform setae and microtrichae on ventral margin; carpus with hooks distally and



prickly tubercles surrounded by dense spines; propodus over seven times as long as wide with two spiniform ventro-distal setae and with distal setae just reaching end of dactylus; propodus twice as long as dactylus and unguis combined; unguis distally simple, with row spines on ventral margin. Pereopod-5 (Fig. 37F): Similar to pereopod-4. Pereopod-6: Similar to pereopod-4, but propodus with three setae distally (according to Kudinova-Pasternak, 1978, Fig 8, page 129). Pleopods missing from specimen. Uropod (according to Kudinova-Pasternak, 1978, Fig. 8, page 129): Endopod with two articles, proximal and distal article subequal; proximal with two pinnate setae distally; distal article with five terminal setae. Exopod with two articles, reaching half of endopod distal article. Distribution: The species is known from two locations in the North Pacific: 24°06.8'N 143°47.l'E, at a depth of 6330m and 29°18.3'N, 143°15.2'E, at the depth range from 6770 to 6850m (Kudinova-Pasternak, 1978). Remarks: Kudinova-Pasternak (1978) measured 19 females of lengths between 2.2 and 5.4 mm and four mancae of lengths 1.8–2.9 mm. From that collection only the holotype was available for study. The holotype (Fig. 36K) differs from the specimen illustrated by Kudinova-Pasternak (1977 (1978)) in the proportion of pereonites. For example, the holotype’s pereonite-2 is twice as long as pereonite-1 and pereonite-4 is the longest, but in figure 7 by Kudinova-Pasternak (page: 127–130) pereonite-2 is 1.5 times as long as pereonite-1 while pereonite-3 is the longest. The holotype also has a shorter antennule, longer setae on the carpus of pereopod-1, no setae on the merus of pereopod-2 and a generally more compact pereopod-2. It is also interesting that the nineteen females and four mancae measured by Kudinova-Pasternak have overlapping body length ranges. It may be that the material studied by Kudinova-Pasternak embraced two sympatric hadal species of Torquella, but because the material was lost it is impossible to resolve this problem.

Torquella grandis (Hansen, 1913) n. comb. (Fig. 38) Typhlotanais grandis: Hansen (1913) 3(3): 52, 54–55; Nierstrasz (1913) 32(a): 37; Kudinova-Pasternak (1970) 86: 348; Lang (1970) 23(4): 273, 288; Morino (1971) 18(5): 353; Kudinova-Pasternak (1973) 91: 153; Kudinova-Pasternak (1977 (1978)) 108: 130; Holdich & Bird (1985) ): 443; Larsen (2005): 210. Typhlotanais sp. C: Belyaev (1966) 88; Kudinova-Pasternak (1975b) 103: 213, 216, 226. Typhlotanais grandis (?): Kudinova-Pasternak (1966) 45: 531; Kudinova-Pasternak (1968) 2: 73.

Material examined: Holotype: female, Ingolf Station 54, South of Iceland, 63°08'N, 15°40'W, depth 691 fm (= 1749 m); length 4.18 mm. Diagnosis: Carapace 1.4 times as long as wide, over twice as long as pereonite-1. Pereonite-2 1.5 times as long as pereonite-1; pereonite-3 0.75 times as long as wide. Antennule article-3 about three times as long as article-2. Pereopod-1 merus and carpus longer than basis, merus with short seta (-ae), propodus with two short rod setae dorsally; pereopod-2 propodus with one seta just reaching half of dactylus, merus with short setae (about quarter of carpus), carpus with two simple setae reaching quarter of propodus; pereopods 1 and 2 with rod setae on merus, carpus and propodus; pereopods 4–5 propodus distal seta little shorter than dactylus; pereopods 4–6 with numerous spines around prickly tubercles; pereopod-6 propodus distal setae subequal. Uropod exopod about as long as half of endopod distal article. Distribution: The type locality of T. grandis is situated in the Iceland Basin, 63°08’N 15°40’W at a depth of 1749 m. The species has also been reported several times from the North Pacific (Kudinova-Pasternak 1970, 1973) at depths of 3800–6135 m.



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FIGURE 38. Torquella grandis. A) Holotype, female, lateral view; B) carapace ventral side; C) Pereopod-1; D) Pereopod-2; E) Pereopod-4; F) Pereopod-6 distally; G) uropod. Scale 0.1 mm.

Remarks: A complementary description of T. grandis is impossible owing to the existence of only the single type specimen (holotype) (Fig. 38 A–G). The holotype has an elongated pereopod-1 with merus and carpus combined longer than the basis, that distinguishes T. grandis from the other members of the genus Torquella. The North-Pacific material studied by Kudinova-Pasternak is not available for study and it is impossible to make a comparison between Hansen’s and Kudinova-Pasternak’s material. The distribution of T. grandis in both the North Atlantic and the North-East Pacific is doubtful; since about ten putative (and undescribed) spe-



cies of Torquella are known from the northeast Atlantic between Iceland and the Bay of Biscay (Bird per. comm.), the likelihood of finding new species in the North Pacific is high.

Torquella longisetosa (Kudinova-Pasternak, 1990) n. comb. (Fig. 39–42) Typhlotanais longisetosus: Kudinova-Pasternak (1990a) 126: 96–97; Kudinova-Pasternak (1993) 127: 141; Larsen (2005): 210.

Material examined: Holotype: female (Mh 10), Akademik Kurchatov Sta. 4923, 25°17’S, 12°24’E, depth 3175m; one female, three mancae, (K 41411), ANT XXII/3, PS 67/81-8-E, 70°32.02'–70°30.3l'S, 14°35.05– 14°35.13'W, depth 4392–4385 m, epibenthic sledge, 24 Feb 2005; (K 41412), ANT XXII/2, PS 67/43-8, 60°27.12'–60°27.24' S, 56°05.10'–56°05.25'W, depth 3961.0–3962.4 m, epibenthic sledge, 4 Feb 2002.

FIGURE 39. Torquella (=Typhlotanais) longisetosa. A) Holotype, female, dorsal view. B) Uropod. Scale: A = 1 mm; B = 0.1 mm.

Diagnosis: Carapace 1.2 times as long as wide; pereonite-2 1.6 times as long as pereonite-1; pereonites 2 and 3 subequal; pereonite-5 longest, 1.3 times as long as wide; chela about four times as long as wide; anten-



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nule article-3 less than three times as long as article-2; pereopod-1 merus and carpus more than half of basis length, merus with spiniform and simple seta, propodus two short and one long rod setae dorsally; pereopod-2 propodus with two setae dorsally (one short and one reaching over unguis), merus with spine and simple seta (half length of carpus), carpus with three spiniform setae and two simple setae reaching half length of propodus; pereopods 4 and 5 propodus distal seta barely reaching half length of dactylus; pereopods 4–6 with few spines around prickly tubercles; pereopod-6 propodus distal setae subequal; uropod rami two-articled; exopod reaching to about half length of endopod distal article. Description: Ovigerous female. Body length 2.9 mm (Fig. 40), 7.5 times as long as wide. Carapace smooth, 15% of body length, a little longer than wide, slightly narrowing rostrally, rounded laterally, rostrum acute. Pereonite-1 three times as wide as long, latero-proximal corners expanded forward; pereonite-2 and 3 subequal, both about 1.3 times as long as wide; pereonite-4 as long as wide; pereonite-5 longest, as long as carapace; pereonite-6 rhomboidal. Pleon 15% of body length, pleonites 1–5 similar in size; pleotelson rectangular. Antennule (Fig. 41A): Article-1 stout, three times as long as wide, with three setae along article; article-2 with one seta distally; article-3 three times as long as article-2. Antenna (Fig. 41B): Article-2 twice as long as article-3, with one seta; article-3 naked; article-4 is 2.4 times as long as article-5, with three simple and one pinnate setae distally; article-5 with one simple seta distally; article-6 very short, with five terminal setae. Mouth parts: Labrum (Fig. 41C) hood-shaped, covered by numerous minute setae. Mandible (Fig. 41D,E) stout; molar process well-developed, with crenulated edges; lacinia mobilis well-developed, crenulated. Maxillule (Fig. 41F) endite with eight apical spiniform setae; palp (Fig. 41F’) with two setae. Maxilla (Fig. 41G) semi-oval, naked. Maxilliped (Fig. 41H) bases fused about as wide as long, with two simple setae reaching end of endites; endite armed with pinnate seta and two tubercles on distal margin; palp four-articled: article-1 naked, article-2 wedge-shaped with one bipinnate seta and two simple setae on inner margin, article-3 with three serrated setae and one simple seta on inner margin, article-4 slender (twice as long as wide) with one simple seta on outer margin and five terminal setae. Labium (Fig. 41I) bilobed, outer corner of inner lobe and outer lobe with minute setae. Cheliped (Fig. 42A) slender. Basis twice as long as wide; merus wedge-shaped with one seta ventrally; carpus about three times as long as broad, with two dorsal setae and with two long and one short ventral setae; propodus and fixed finger little longer than carpus, about four times as long as broad; dactylus little longer than whole; fixed finger with well-calcified inner margin, only one seta visible, ventral margin with two simple setae. Pereopod-1 (Fig. 42B): Of walking type; coxa present; basis 1.4 times as long as merus and carpus combined, with two small setae on ventral margin and two simple setae and one pinnate seta on dorsal margin; ischium short with one seta; merus almost as long as carpus, with one rod and one simple setae distally; carpus with three simple and three rod setae distally; propodus 1.3 times as long as carpus with one setae ventrally and two (short and long) rod setae dorsally; dactylus and unguis combined almost as long as propodus, dactylus 0.6 times as long as unguis. Pereopod-2 (Fig. 42C): Of walking type; coxa present; basis with one simple and one pinnate seta proximally, longer than ischium, merus and carpus combined; ischium with one seta; merus almost as long as carpus, with one simple and one spiniform setae distally, some microtrichae on ventral margin; carpus with three spiniform, and two simple setae distally; propodus 1.3 times as long as carpus with spiniform seta ventrally and with long simple and spiniform setae dorsally; dactylus and unguis combined 0.6 times length of propodus, dactylus almost as long as unguis. Pereopod-3 (Fig. 42D): Similar to pereopod-2, but microtrichae on ventral margin well-calcified. Pereopod-4 (Fig. 42E): Clinging type; coxa absent; basis with one seta ventrally; ischium with two setae ventrally; merus with two spiniform setae distally; carpus with two spiniform setae distally and with prickly



tubercles surrounded by rare spines; propodus six times as long as wide, with pinnate seta mid-dorsally, two spiniform ventro-distal setae and one distal seta just reaching half of dactylus; unguis simple, with spines ventrally.

FIGURE 40. Torquella longisetosa female (ANDEEP material). A) lateral view; B) dorsal view. Scale = 1 mm.



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FIGURE 41. Torquella longisetosa (ANDEEP material). A) Antennule; B) Antenna; C) Labrum; D) Left mandible; E) Right mandible; F) Maxillule; F’) Palp; G) Maxilla; H) Maxilliped; I) Labium. Scale: A, B, I = 0.1 mm; C-H = 0.01 mm.



FIGURE 42. Torquella (=Typhlotanais) longisetosa (ANDEEP material). A) Cheliped; B) Pereopod-1; C) Pereopod-2; D) Pereopod-3; E) Pereopod-4; F) Pereopod-5; G) Uropod. Scale = 0.1 mm.



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Pereopod-5 missing from specimen. Pereopod-6 (Fig. 42F): Similar to pereopod-4, but propodus with three distal setae a little longer than half of dactylus. Pleopods 1–5 damaged. Uropod (Fig. 42G): Endopod with two articles, proximal and distal article subequal; proximal with two pinnate setae distally; distal article with five terminal setae. Exopod with two articles; distal article 1.6 times as long as proximal, tipped by long simple seta. Distribution: South-eastern Atlantic (25°17’S, 12°24’E), Eastern Weddell Sea, south of Falkland Islands, depth range 3060–6100 m. Remarks: The holotype provided by Kudinova-Pasternak lacks a carapace and most appendages, the remains of which are shown in figure 39. The finding of T. longisetosa in ANDEEP samples allows the drawing of its morphological details and redescription. It can be immediately distinguished from its congeners by the long seta on the propodus of pereopods 2 and 3.

Torquella parangularis (Kudinova-Pasternak 1975) n. comb. (Fig. 43) Typhlotanais parangularis: Kudinova-Pasternak (1975) 103: 213, 216, 226; Kudinova-Pasternak & Pasternak (1981) 115: 122, 123; Kudinova-Pasternak (1985) 120: 55; Kudinova-Pasternak (1990) 126: 96; Larsen (2005): 210.

Material examined: Holotype: female, Mc 981, Akademik Kurchatov, Station 916, 56°29.4'S 50°51'W, depth 4664–5631m. Diagnosis: Carapace 1.2 times as long as wide. Pereonite-2 1.6 times as long as pereonite-1; pereonite-3 longest, 1.25 times as long as wide. Chela three times as long as wide. Antennule article-3 twice as long as article-2. Pereopod-1 merus and carpus more than half as long as basis, merus with two short setae, propodus with three subdistal setae dorsally; pereopod-2 propodus with one seta reaching half of dactylus, merus with one short and one long seta (about half of carpus), carpus with three short simple setae dorso-distally; pereopod-4 and 5 propodus distal seta reaches end of dactylus; uropod exopod as long as endopod proximal article. Distribution: Species known only from type locality: Scotia Sea at the depth range from 4664 to 5631 m. Remarks: Torquella parangularis is distinguished from the other species of the genus by the short uropod exopod that is as long as the endopod proximal article. A relatively short uropod exopod occurs in T. eltaninae n. sp., but it has a much shorter antennule article-3 and one long distal seta on the pereopod-6 propodus.

Torquella rotundirostris (Lang, 1970) n. comb Typhlotanais rotundirostris: Lang (1970) (2) 23(4): 277, 279–286.

Diagnosis: Carapace 1.5 times as long as wide. Pereonite-2 about 1.4 times as long as pereonite-1; pereonite3 wider (1.3 times) than long, and pereonite-5 about as long as wide. Chela three times as long as wide. Antennule article-3 twice as long as article-2. Pereopod-1 basis 1.5 times as long as merus and carpus combined, merus with short setae (one rod seta), propodus with short and long setae on dorsal margin; pereopod-2 propodus with two setae (one simple seta reaching half of dactylus), merus with one spiniform seta, carpus with two spiniform setae and two short simple setae; pereopods 4 and 5 propodus distal seta reaches end of unguis; pereopods 4–6 with numerous spines around prickly tubercles; pereopod-6 propodus with three sub-



equal setae distally; uropod endopod twice as long as basal article, uropod exopod reaches about half of endopod proximal article.

FIGURE 43. Torquella parangularis. A) Female; dorsal view. E) Pleopod; B–D) Pereopods 1–3, respectivelly; F) Uropod. Scale = A = 1 mm; B-E = 0.1 mm. A REVISION OF FAMILY TYPHLOTANAIDAE SIEG 1984


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Distribution: Species known only from type locality: Southern-East Atlantic (30°14.09'S, 13°03'E) at 3073 m depth. Remarks: T. elegans, T. galatheae and T. rotundirostris are the only three members of Torquella having two long dorso-distal setae on the propodus of the second and third pereopods. Of these three species only T. galatheae has a spiniform dorso-ventral seta on first pair of pereopod while two others species have regular seta there. The presence of four and two setae on the merus of pereopods 2 and 3 respectively in T. elegans allows its distinction from T. rotundirostris which has only one spiniform and one regular seta on each.

Torquella eltaninae n. sp. (Figs 44–45) Diagnosis: Carapace 1.3 times as long as wide. Pereonite-2 about 1.3 times as long as pereonite-1; pereonites 3 and 5 about as long as wide. Chela 2.5 times as long as wide. Antennule article-3 1.6 times as long as article2. Pereopod-1 merus with short seta; pereopod-2 propodus with one simple seta reaching half of unguis, merus with one spiniform seta, carpus with spiniform setae ventrally and two short simple seta dorsally; pereopods 4 and 5 propodus distal seta reaching end of dactylus; pereopods 4–6 with numerous spines around prickly tubercles; pereopod-6 propodus with one long and two short setae distally. Uropod endopod twice as long as basal article; uropod exopod about half of endopod proximal article. Material examined: Holotype: one female, (K 41454), ANT XXII/3, PS67/078-9, 71°09.39'– 71°09.36'S, 13°59.30'–13°58.81'W, depth 2156–2147 m, epibenthic sledge, 21 Feb 2005; Paratypes: one female, (USNM 1100142), Eltanin 9, Sta.724, 54°05'–54°04'S, 033°43'–033°37'W, depth 2714–2727 m, 9 Sep 1963; two females (one damaged), (USNM 100141) Eltanin 4, Sta.127, 61°45’S–61°14’W, depth 4758 m, 1 Aug 1962. Etymology: The name is given after the research vessel Eltanin from which the species was collected. Description: Female. Body length 2.3 mm (Figs 44A,B), 7.2 times as long as wide. Carapace smooth, 20% of body length, 1.3 times as long as wide, slightly narrowing rostrally, rounded laterally, rostrum acute. Pereonite-1 twice as wide as long, antero-lateral corners expanded forward; pereonite-2 is 1.2 times as wide as long; pereonites 4 and 5 subequal, a little wider than long; pereonite-6 twice as wide as long. Pleon 18% of body length, pleonites 1–5 similar in size; pleotelson rectangular. Antennule (Fig. 44D): Article-1 stout, over twice as long as wide, with few pinnate setae distally; article2 with two setae distally; article-3 1.6 times as long as article-2, with six setae distally. Antenna (Fig. 44E): Article-2 twice as long as article-3, with one seta; article-3 naked; article-4 is 2.6 times as long as article-5, with two simple and five pinnate setae distally; article-5 with one simple seta distally; article-6 very short, with five terminal setae. Mouth parts: Labrum lost during dissection. Mandible (Fig. 44F,G) stout; molar process (Fig. 44G’) welldeveloped, edges supported with tubercle-like structures, lower edge with spines; lacinia mobilis well-developed, crenulated. Maxillule (Fig. 44H) inner endite with eight apical spiniform setae; palp with two distal setae. Maxilla lost during dissection. Maxilliped (Fig. 45A): bases fused, bilobed with simple setae reaching half of endites; endite armed with seta in dorso-lateral corner and two tubercles on distal margin; palp fourarticled: article-1 naked; article-2 wedge-shaped, with three setae on inner margin and one short seta on outer margin; article-3 with three biserrated setae, one simple setae on inner margin; article-4 with one bipinnate seta on outer margin and five terminal setae. Cheliped (Fig. 45B): Basis 1.5 times as long as wide, with one seta; merus wedge-shaped with one seta ventrally; carpus over twice as long as broad, with two dorsal setae and with two long setae ventrally; propodus and fixed finger 1.2 times as long as carpus, about three times as long as wide, with one setae near insertion of dactylus; fixed finger with three setae on inner margin and with two setae on ventral margin.



Pereopod-1 (Fig. 45C): Of walking type; coxa present: basis longer than merus and carpus combined, with one simple and pinnate setae proximally and one seta ventrally; ischium with one seta; merus little shorter than carpus, with one seta distally; carpus with three setae distally; propodus 1.6 times as long as carpus, with one seta ventrally and one spiniform seta dorsally; dactylus with seta; unguis broken.

FIGURE 44. Torquella eltaninae n. sp. A) Holotype, female, dorsal view. B) Holotype, female, lateral view; C) Carapace ventral side; D) Antennule; E) Antenna; F) Right mandible; G) Left mandible; G’) Molar process; H) Maxillule; H’ detail of terminals setae. Scale = 0.1 mm.



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FIGURE 45. Torquella eltaninae n. sp., female paratype. A) Maxilliped; B) Cheliped; C) Pereopod-1; D) Pereopod-2; E) Pereopod-3; F) Pereopod-4; G) Pereopod-5; H) Pereopod-6; I) Pleopod; J) Uropod. Scale 0.1 mm.

Pereopod-2 (Fig. 45D): Of walking type; coxa present; basis with one seta proximally and one seta on ventral margin; longer than merus and carpus combined; ischium with one seta; merus almost as long as car-



pus, with one spiniform setae distally; carpus with three spiniform setae ventrally and two (short and long) dorso-distal setae; propodus a little longer than merus and carpus combined, with spiniform seta ventrally and one long seta (reaching half length of unguis) dorsally; dactylus half as long as unguis. Pereopod-3 (Fig. 45E): Similar to pereopod-2, but merus with spiniform and simple seta. Pereopod-4 (Fig. 45F): Coxa absent: Basis clearly shorter than rest of articles combined, with two pinnate setae on ventral margin; ischium with two setae ventrally; merus with two spiniform setae ventrally and small spines arranged in combs on ventral margin; carpus with two spiniform setae and prickly tubercles surrounded by dense spines; propodus five times as long as wide, with two spiniform ventral setae and with distal setae reaching end of dactylus; dactylus and unguis combined half as long as propodus; unguis distally simple with row spines on ventral margin. Pereopod-5 (Fig. 45G): Similar to pereopod-4. Pereopod-6 (Fig. 45H): Similar to pereopod-4, but propodus with two short setae (as long as half of dactylus) and one long seta (reaching over unguis). Pleopods 1–5 (Fig. 45I): Endopod with 16 plumose setae on outer margin; exopod with eleven setae on outer margin and one seta on inner margin; large gap between most proximal seta and others. Uropod (Fig. 45J): Exopod and endopod two-articled; endopod twice as long as peduncle; exopod twothirds length of endopod. Distribution: Vicinity of South Georgia, South Shetland Islands and Eastern Weddell Sea and at depths from 2147 to 4758 m. Remarks: From the other Torquella species T. eltaninae n. sp. can be distinguished by its more compact body, with pereonites 2–4 about as wide as long. The main diagnostic character is the long seta on the propodus of pereopod-6.

Torquella galatheae n. sp. (Figs 46–47) Material examined: Holotype: female, (CRU ###), partially dissected on slides, Galathea Station 716, 9°23'N, 89°32'W, depth 3570 m, herring otter trawl, bottom dark, muddy clay, bottom temp.: c. 1.9°C, 6 May 1952. Diagnosis: Carapace 1.2 times as long as wide. Pereonite-2 1.5 times as long as pereonite-1; pereonites 3 and 5 1.4 times as long as wide. Chela three times as long as wide. Antennule article-3 less than three times as long as article-2. Pereopod-1 merus and carpus shorter than basis, merus with two short setae; pereopod-2 propodus with two long setae dorsally, merus with one spiniform and one simple seta, carpus with three spiniform and two short simple setae; pereopods 4–5 propodus distal seta reaches end of unguis; pereopods 4–6 with numerous spines around prickly tubercles; pereopod-6 propodus with distal setae subequal; uropod rami two-articled, exopod 0.8 times length of endopod. Etymology: The name is given after the research vessel r/v Galathea from which the species was collected. Description: Female. Body length 3.7 mm (Fig. 46A), almost ten times as long as wide; carapace smooth, 15% of body length, 1.3 times as long as wide, slightly narrowing rostrally, rounded laterally, rostrum acute. Pereonite-1 twice as wide as long, antero-lateral corners expanded forward; pereonite-2 little wider than long, 1.5 times as long as pereonite-1; pereonites 3–5 subequal, 1.4 times as long as wide; pereonite-6 1.5 as wide as long. Pleon 15% of body length, pleonites 1–5 similar in size; pleotelson semi-rectangular. Antennule (Fig. 46B): Article-1 over three times as long as wide, with three groups of pinnate and simple setae; article-2 with two simple and one pinnate setae distally; article-3 over twice as long as article-2, with five simple and one pinnate setae distally. A REVISION OF FAMILY TYPHLOTANAIDAE SIEG 1984


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Antenna (Fig. 46C): Article-2 little longer than article-3; article-3 with one seta; article-4 is 2.2 times as long as article-5, with two simple and two pinnate setae distally; article-5 with one simple seta distally; article-6 very short, with five terminal setae.

FIGURE 46. Torquella galatheae n. sp., female paratype. A) Holotype, female, dorsal view; B) Antennule; C) Antenna; D) Labrum; E) Left mandible; F) Right mandible; G) Maxillule; H) Maxilla; I) Labium; J) Maxilliped. Scale A = 1 mm; B, C, I = 0.1 mm, D-J = 0.01mm.



FIGURE 47. Torquella galatheae n. sp., female paratype. A) Cheliped; B) Pereopod-1; C) Pereopod-2; D) Pereopod-3; E) Pereopod-4; F) Pereopod-5; G) Pereopod-6; H) Pleopod; I) Uropod. Scale = 0.1 mm.

Mouth parts: Labrum (Fig. 46D) hood-shaped with numerous small setae. Mandible (Fig. 46E,F) stout; incisor with row of teeth on upper margin; molar process well-developed, edges supported with teeth-like structures, lower edge with spines; lacinia mobilis well-developed, crenulated. Maxillule (Fig. 46G) endite A REVISION OF FAMILY TYPHLOTANAIDAE SIEG 1984


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with eight apical spiniform setae. Maxilla (Fig. 46H) oval, naked. Maxilliped (Fig. 46J) bases fused, bilobed, with simple setae reaching over half length of endites; endite with two setae and two tubercles on distal margin; palp four-articled: article-1 naked; article-2 wedge-shaped, with three setae on inner margin and one short seta on outer margin; article-3 with three serrated setae and one simple setae on inner margin; article-4 with one serrated seta on outer margin and five terminal setae. Labium (Fig. 46I) bilobed; outer corner of inner lobe and outer lobe with minute setae. Cheliped (Fig. 47A): Basis twice as long as wide; merus wedge-shaped with one seta ventrally; carpus about 2.8 times as long as broad, with one dorsal seta and with two long setae ventrally; propodus and fixed finger as long as carpus, about three times as long as wide, with one long setae near insertion of dactylus; fixed finger with three setae on inner margin and with two simple setae on ventral margin. Pereopod-1 (Fig. 47B): Of walking type; coxa present; basis a little longer than merus and carpus combined, with three setae on dorsal margin; ischium with one seta; merus as long as carpus, with one spiniform and one simple seta distally; carpus with one spiniform, two simple, one pinnate setae distally and one small spiniform setae subventrally; propodus 1.3 times as long as carpus with one seta ventrally and three setae dorsally; dactylus and unguis combined 0.75 times length of propodus. Pereopod-2 (Fig. 47C): Of walking type; coxa present; basis with three setae dorsally, shorter than ischium, merus and carpus combined; merus a little shorter than carpus, with one simple and one spiniform setae distally; carpus with three spiniform setae and two simple setae distally; propodus a little longer than merus and carpus combined, with spiniform seta ventrally and two long setae (reaching end of unguis) dorsally; dactylus shorter than half of unguis. Pereopod-3 (Fig. 47D): Similar to pereopod-2, but carpus with four spiniform setae. Pereopod-4 (Fig. 47E): Clinging type; coxa absent; basis with one simple seta dorsally and one pinnate seta ventrally; ischium with two setae ventrally; merus with two spiniform setae and microtrichiae on ventral margin; carpus with two spiniform setae and prickly tubercles surrounded by dense spines; propodus five times as long as wide, with two spiniform ventro-distal setae and with distal setae reaching the end of unguis; dactylus and unguis combined as long as two-thirds of propodus; unguis distally simple with row spines on ventral margin. Pereopod-5 (Fig. 47F): Similar to pereopod-4. Pereopod-6 (Fig. 47G): Similar to pereopod-4, but propodus with three subequal setae distally. Pleopods 1-5 (Fig. 47H): Endopod with 25 plumose setae on outer margin; exopod with 16 setae on outer margin and one seta on inner margin. Uropod (Fig. 47I): Endopod two-articled; proximal and distal article subequal; proximal with one seta distally, distal article with five simple and two pinnate terminal setae. Exopod with two articles; distal article 1.7 times as long as proximal article, tipped by long and short simple seta. Distribution. Species known only from the type locality: Acapulco-Panama: 9°23'N, 89°32'W, at a depth of 3570 m. Remarks: From most of its congeners T. galatheae n. sp. can be distinguished by having pereonites 3–5 longer than wide. This character is shared with T. elegans, but T. elegans lacks the spiniform setae on the merus of pereopods 2 and 3 that are present in T. galatheae.

Genus: Typhlamia n. gen. Diagnosis: Body long, about 8–9 times as long as wide. Carapace about as wide as long, tapering proximally, rounded marginally. Pereonites clearly rounded marginally. Antennule twice as long as carapace; antennule article-1 over six times as long as wide (occasionally with suture in the middle of article); antennule article-3 slender, about ten times as long as wide, with distal setae as long as half of whole appendage. Maxilliped basis



with seta reaching far beyond endites. Cheliped bases not reaching the pereonite-1 ventrally. Pereopods 1 and 3 coxa without spur, pereopod-1 with long seta on carpus; pereopods 4–6 prickly tubercles small; unguis with bifid tip. Pleopod rami with gap between most proximal seta and others. Uropod endopod two-articled, exopod one-articled. Male: Unknown. Etymology: Typhlos (Gr.) = blind, Lamia – the goddess with long hair. The name reflects long setae on the antennule. Gender of generic name: Feminine. Type species: Typhlamia bella n. sp. Species included: Typhlamia (= Typhlotanais) mucronata (Hansen, 1913); Typhlamia (= Typhlotanais) sandersi (Kudinova-Pasternak, 1985); Typhlamia bella n. sp. Remarks: The genus can immediately be distinguished by having a long antennule bearing long setae. In general body habitus (rounded pereonites in dorsal view, long cheliped with basis not reaching pereonite-1 ventrally) Typhlamia resembles Typhlotanais variabilis, but it differs in having a uni-articulated exopod of uropods, a bifurcated unguis and small prickly tubercles on pereopods 4–6.

Key for identification of Typhlamia females 1. 2. -

Pereopod-2 with short setae on carpus (less than half length of propodus) ............................ T. mucronata Pereopod-2 with long setae on carpus (as long as propodus) ..................................................................... 2 Pereopod-2 merus with short seta (as long as half of carpus) ..................................................... T. sandersi Pereopod-2 merus with long seta (as long as carpus) ..............................................................T. bella n. sp.

Typhlamia mucronata (Hansen, 1913) n. comb. (Figs 48–49) Typhlotanais mucronatus: Hansen (1913) 3 (3): 37, 42–44; Stephensen, (1932) 6: 350; Kudinova-Pasternak (1970) 86: 348; Lang (1970) (2), 23 (4): 275, 289; Kudinova-Pasternak (1973) 91: 153; Kudinova-Pasternak (1985) 120: 63; Kudinova-Pasternak (1990) 126: 94; Holdich & Bird (1985): 443; Larsen (2005): 210.

Material examined: Syntype: female, CRU 3916, Ingolf-Station, St. 120, North-East of Iceland, 67°29'N, 11°32'W, depth 885 fm (1618 m); syntype: female, CRU 7423, Ingolf-Station, Sta 119, North-East of Iceland, 67°53'N, 10°19'W, depth 1010 fm (1947 m); syntype: female dissected on slides, CRU 3885, the same locality. Diagnosis: Carapace a little longer than wide; pereopod-1 carpus with seta as long as propodus length. Pereopod-2 merus and carpus with short setae (not longer than half of following article respectively). Uropod exopod little longer than endopod proximal article. Description: Nonovigerous female. Body length 2.1 mm (Figs 48A,B), 7.8 times as long as broad. Carapace smooth, rounded laterally, as long as wide, 13 % of total body length; rostrum blunt. Pereonites smooth, slightly rounded lateral margins; pereonites 1, 2 and 6 subequal, 1.7 times as wide as long, pereonite-1 0.7 times as long as carapace; pereonite-3 almost as long as pereonite-4; pereonite-5 a little shorter than pereonite-4, as long as wide. Pleon 20% of body length, as long as pereonites 5, 6 plus 0.3 of pereonite-4 combined; pleonites 1–5 similar in size; pleotelson semi-circular; caudal projection reduced. Antennule (Fig. 48D): Twice as long as carapace; article-1 with fusion line, seven times as long as wide, with a few setae on outer and inner margin, two submedial setae reaching over end of article; article-2 with two simple setae distally; article-3 nine times as long as wide, tipped by six long simple setae.



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Antenna (Fig. 48E): Article-2 more than half as long as article-3; article-3 with one small seta; article-4 twice as long as article-5, with six simple setae distally; article-5 with one distal seta; article-6 tipped by four setae.

FIGURE 48. Typhlamia mucronata. A) Holotype, female, dorsal view; B) Holotype, female, lateral view; C) Carapace, ventral view ; D) Antennule; E) Antenna; F) Left mandible; G) Right mandible; H) Maxillule endite distally I) Epignath; J) Labium. Scale A, B = 1 mm; C–E, J= 0.1 mm; F–I = 0.01 mm.



FIGURE 49. Typhlamia mucronata. A) Maxilliped; B) Cheliped; C) Pereopod-1; D) Pereopod-2; E) Pereopod-3; F) Pereopod-4; G) Pereopod-5; H) Pereopod-6 distal part; I) Pleopod; J) Uropod. Scale 0.1 mm.

Mouth parts: Labrum lost during dissection. Mandible (Figs 48F,G) stout; molar process well-developed, with crenulated distal margin; lacinia mobilis well-developed, crenulated. Maxillule (Fig. 48H) endite with nine spiniform setae distally; palp lost during dissection. Maxilla lost during dissection. Maxilliped (Fig. 49A): bases fused little longer than wide; each with simple setae reaching far over endites; endite with two tubercles on distal margin; palp four-articled: article-1 naked; article-2 wedge-shaped, with three setae on inner margin; article-3 trapezoidal, with three bipinnate and one simple setae on inner margin, article-4 slender (2.5 times as long as wide), with one simple seta on outer margin and five bipinnate terminal setae. Epignath (Fig. 48I) tip not rounded. Labium (Fig. 48J) bi-lobed, outer corner of inner lobe and outer lobe with minute setae.



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Cheliped (Fig. 49B): Basis 1.7 times as long as wide, with one seta on dorsal margin; merus wedgeshaped, with one short seta ventrally; carpus about 2.7 times as long as broad, with two dorsal setae and with two long setae ventrally; propodus and fixed finger just as long as carpus, about four times as long as wide; fixed finger with three setae on inner margin and two setae on ventral margin. Pereopod-1 (Fig. 49C): Coxa with seta; basis little shorter than merus, carpus and propodus combined; ischium with simple seta; merus little shorter than carpus, with simple seta distally; carpus with four short and one long (longer than half of propodus) setae distally; propodus 1.4 times as long as carpus, with one ventral and two dorsal setae; dactylus almost as long as unguis, together as long as half of propodus. Pereopod-2 (Fig. 49D): Coxa with seta; basis a little longer than merus and carpus combined; ischium with simple seta; merus a little shorter than carpus, with one simple seta and one thick seta distally; carpus with four short and one spiniform setae distally; propodus a little longer than carpus, with one spiniform seta ventrally and two simple setae on dorsal margin; dactylus almost as long as unguis. Pereopod-3 (Fig. 49E): Similar to pereopod-2. Pereopod-4 (Fig. 49F): Clinging type; basis with one seta ventrally; ischium with one seta ventrally; merus with two spiniform setae; carpus with spiniform seta distally and small prickly tubercles; propodus over three times as long as wide, with two spiniform ventro-distal setae and with distal setae just reaching end of dactylus; unguis with bifid tip. Pereopod-5 (Fig. 49G): Similar to pereopod-4. Pereopod-6 (Fig. 49H): Similar to pereopod-4, but propodus with three distal setae as long as dactylus. Pleopods 1–5 (Fig. 49I): Basal article naked; exopod with nine plumose setae on outer margin and one seta on inner margin; endopod with 17 pinnate setae on outer margin. Gap between most proximal setae and the other setae in both rami. Uropod (Fig. 49J): Endopod two-articled; proximal article almost as long as distal article, with one distal seta; distal article tipped by five simple setae; exopod one-articled, little longer than endopod proximal article, tipped by strong long and short simple setae. Distribution: From published sources, the species has been found at a few places in the North Atlantic: North-East off Iceland (type locality), South of Jan Mayen, and the Bay of Biscay at depths of 1618–1847 m; and it was also recorded in the North Pacific (45°32'N, 153°46'E, depth 6675–6710 m; 44°17'N 149°37'E, depth 4840 m; 56°14'N,139°44'W, depth 3450 m) by Kudinova-Pasternak (1970; 1973). Remarks: The wide distribution of Typhlamia (= Typhlotanais) mucronata (Hansen, 1913) in both the North Atlantic and Pacific is questionable, but because material studied by Kudinova-Pasternak is not available for study the problem cannot be solved at the moment. T. mucronata can be distinguished from the two other species (T. sandersi and T. bella n. sp.) by its having relatively short setae on the carpus of the second pereopod while in the two other species there are two long setae reaching the end of the propodus.

Typhlamia sandersi (Kudinova-Pasternak, 1985) n. comb. (Figs 50–51) Typhlotanais sandersi: Kudinova-Pasternak (1985) 120: 52–64; Larsen (2005): 210.

Material examined: Holotype: female (Mh 4), Vitjaz Station, Great Meteor, 29°50'N, 28°08'6E, depth 3080– 3140 m. NE Atlantic. Diagnosis: Carapace longer than wide. Pereopod-1 carpus seta almost as long as propodus; pereopod-2 merus with seta not reaching the end of carpus, carpus with two setae reaching the end of propodus. Uropod exopod as long as endopod proximal article.



Description: Nonovigerous female. Body length 4 mm (Figs 50A), long, 8.2 times as long as broad. Carapace smooth, rounded laterally, 1.2 times as long as wide, 14 % of total body length; rostrum blunt. Pereonites smooth, slightly rounded lateral margins, pereonite-1 shortest, a little longer than pereonite-6; pereonite2 almost as long as pereonite-5, 1.5 times as long as pereonite-1; pereonites 3 and 4 subequal, as long as wide. Pleon 21% of body length, as long as pereonites 5 and 6 combined; pleonites 1–5 similar in size; pleotelson semi-rounded; caudal projection prominent.

FIGURE 50. Typhlamia sandersi. A) Holotype, female, dorsal view; B) Antennule; C) Antenna; D) Labrum; E) Left mandible; F) Right mandible; G) Maxillule; G’) Palp; H) Maxilliped; I) Labium. Scale: A = 1 mm; B, C = 0.1 mm; E-I = 0.01 mm.

Antennule (Fig. 50B): Twice as long as carapace; article-1 almost seven times as long as wide, with a few simple and pinnate setae on outer, inner and distal margin; two distal setae exceeding tip of article-2; article-2 with two simple setae and one pinnate seta distally; article-3 twelve times as long as wide, tipped by five long simple setae. A REVISION OF FAMILY TYPHLOTANAIDAE SIEG 1984


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Antenna (Fig. 50C): Article-2 more than 1.5 as long as article-3, with one seta; article-3 naked; article-4 twice as long as article-5, with two simple and two pinnate setae distally; article-5 with one distal setae; article-6 tipped by six setae distally. Mouth parts: Labrum (Fig. 50D) flat, covered by minute setae. Mandible (Figs 50 E,F) stout; molar process well-developed, with crenulated distal margin; lacinia mobilis on left mandible well-developed, crenulated. Maxillule (Fig. 50G) endite with eight spiniform setae distally; palp (Fig. 50G’) with two setae distally. Maxilla lost during dissection. Maxilliped (Fig. 50H): bases fused little longer than wide; each with simple setae reaching far over endites; endite with two tubercles on distal margin; palp four-articled: article-1 naked; article-2 wedge-shaped, with three setae on inner margin and one seta on outer margin; article-3 trapezoidal, with four setae on inner margin; article-4 slender (twice as long as wide), with one simple seta on outer margin and five terminal setae. Epignath lost during dissection. Labium (Fig. 50I) bi-lobed, outer corner of inner lobe and outer lobe with minute setae. Cheliped (Fig. 51A): Basis a little longer than wide, with one seta on dorsal margin; merus wedgeshaped, with one long seta ventrally; carpus less than three times as long as broad, with two dorsal setae and with two long setae ventrally; propodus and fixed finger as long as carpus, about three times as long as wide; fixed finger with three setae on inner margin and two setae on ventral margin. Pereopod-1 (Fig. 51B): Coxa with seta; basis as long as merus, carpus and half of propodus combined; ischium with simple seta; merus as long as carpus, with simple seta distally; carpus with four short and one long (as long as propodus) setae distally; propodus 1.3 times as long as carpus, with one long seta subdistally on ventral margin and two short setae on dorsal margin; dactylus as long as half of unguis, together 0.7 times length of propodus. Pereopod-2 (Fig. 51C): Coxa present; basis as long as merus, carpus and half of propodus combined; ischium with simple seta; merus little shorter than carpus, with one simple seta and one thick seta distally; carpus with one short, two long and two spiniform setae distally; propodus a little longer than carpus, with one spiniform seta ventrally and two subdistal simple setae on dorsal margin; dactylus almost as long as unguis. Pereopod-3 broken. Pereopod-4 (Fig. 51D): Clinging type; basis with one seta ventrally; ischium with two setae ventrally; merus with one simple and one spiniform seta; carpus with hooks distally and small prickly tubercles; propodus six times as long as wide, with two spiniform ventrodistal setae and with distal setae reaching over half dactylus; unguis with bifid tip. Pereopod-5 (Fig. 51E): Similar to pereopod-4. Pereopod-6 (Fig. 51F): Similar to pereopod-4, but propodus with three distal setae, as long as dactylus. Pleopods 1–5 (Fig. 51G): Basal article naked; exopod with ten plumose setae on outer margin and one seta on inner margin; endopod with 19 pinnate setae on outer margin. Gap between most proximal setae and the other setae in both rami. Uropod (Fig. 51H): Endopod two-articled; proximal article 1.3 times as long as distal, with two distal seta (one longer than proximal article), distal article tipped by four simple setae; exopod one-articled, as long as proximal endopod article, tipped by one strong, long and one short simple setae. Distribution: This species is known only from the type locality: North-eastern Atlantic, west of Tenerife at the depth range of 3080–3140 m. Remarks: The relatively good condition of the holotype allowed partial dissection of the specimen. T. sandersi is most similar to T. bella n. sp., but it can be distinguished by the setation of the cheliped, pereopod 1 and 2. On the carpus of pereopod-1 of T. sandersi one of the distal setae reaches the end of the propodus, while the longest seta on the same article in T. bella reaches to just half the length of the propodus. The dorsal setae on the pereopod-2 propodus are subequal in T. sandersi while in T. bella there is one short and one long; finally, the cheliped merus ventral seta is much longer in T sandersi than T. bella.



FIGURE 51. Typhlamia sandersi, female paratype. A) Cheliped; B) Pereopod-1; C) Pereopod-2; D) Pereopod-3; E) Pereopod-4; F) Pereopod-6; G) Pleopod; H) Uropod. Scale = 0.1 mm.



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Typhlamia bella n. sp. (Figs 52–54) Material examined: Holotype, female (K 41413), ANT XXII-3, PS67/81-8-E 70°32.02'–70°32.19'S, 14°35.05'–14°35.13'W, depth 4392-4385, 24 Feb 2005; Paratypes: ten juveniles females, three females (one dissected on slides, one half, one broken) (K 41414), ANT XXII-3, PS 67/81-8-E, 70°32.02'-70°32.19'S, 14°35.05'–14°35.13'W, depth 4392–4385 m, EBS, 24 Feb 05; ?Typhlamia bella one female (K 41435), ANT XXII-3, PS 67/154-9, 62°31.47'–62° 31.36'S, 64°39.45'–64° 39.25'W, depth 3804–3808 m, 30 Mar 2005. Diagnosis: Carapace as long as wide. Pereopod-1 carpus seta just half of propodus length; pereopod-2 merus and carpus with setae reaching end of following article. Uropod exopod as long as endopod proximal article. Etymology: Bella (Lat.) = pretty, neat. The name reflects the author’s personal impression of the habitus of the animal. Description: Non-ovigerous female (Fig. 52A,B). Body length 3.6 mm, 7.8 times as long as broad. Carapace smooth, rounded laterally, as long as wide, 13% of body length, rostrum blunt. Pereonites smooth, slightly rounded lateral margins; pereonite-6 trapezoidal; pereonite-1 is 0.7 times length of carapace; pereonites 2, 3 and 6 of similar size; pereonite-2 a little longer than pereonite-1; pereonite-4 a little longer than wide; pereonite-5 as long as wide. Pleon 20% of body length, a little shorter than pereonites 5 and 6 combined; pleonites 1–5 similar in size; pleotelson semi-rounded; caudal projection weak. Antennule (Fig. 53A): Twice as long as carapace; article-1 seven times as long as wide, with five simple setae and five pinnate setae arranged in groups on inner and outer margin; article-2 with two distal setae; article-3 2.5 times as long as article-2, over ten times as long as wide, tipped by four long simple setae. Antenna (Fig. 53B): Article-2 three times as long as article-3; article-4 twice as long as article-5, with three simple and two pinnate setae distally; article-5 with one distal setae; article-6 tipped by four setae. Mouth parts: Labrum (Fig. 53C) covered by minute setae. Mandible (Figs 53D,E) stout; molar process (Fig. 53D’) well-developed, with crenulated distal margin; lacinia mobilis well-developed, crenulated. Maxillule (Fig. 53F) endite with eight spiniform setae distally; palp lost during dissection. Maxilla (Fig. 53G) semitriangular. Maxilliped (Fig. 53I) bases fused little longer than wide; each with simple seta reaching far over endites; endite armed with two setae and two tubercles on distal margin; palp four-articled: article-1 naked; article-2 wedge-shaped, with three setae on inner margin; article-3 trapezoidal, with three bipinnate and one simple setae on inner margin; article-4 slender (1.5 times as long as wide), with one simple seta on outer margin and five bipinnate terminal setae. Epignath lost during dissection. Labium (Fig. 53H) bilobed, outer corner of inner lobe and outer lobe with minute setae. Cheliped (Fig. 54A): Basis 1.3 times as long as wide; merus wedge-shaped with one short seta ventrally; carpus about 2.5 times as long as broad with two dorsal setae and with two long setae ventrally; propodus and fixed finger about as long carpus, about three times as long as wide; fixed finger with three setae on inner margin and two setae on ventral margin. Pereopod-1 (Fig. 54B): Coxa with seta; basis a little shorter than merus, carpus and propodus combined; ischium with simple seta; merus slightly shorter than carpus, with simple seta distally; carpus with four short and two long (longer than half of propodus) setae distally; propodus 1.2 times as long as carpus, with one ventral and one dorsal setae subdistally; dactylus almost as long as unguis, with seta reaching end of unguis, together 0.6 times length of propodus. Pereopod-2 (Fig. 54C): Coxa with seta; basis slightly longer than merus and carpus combined; ischium with simple seta; merus a little shorter than carpus, with one short and long (as long as carpus) seta distally; carpus with two short, two long and two spiniform setae distally; propodus slightly longer than carpus, with one spiniform seta ventrally and two subdistal simple setae on dorsal margin; dactylus almost as long as unguis, with short seta. Pereopod-3 broken.



FIGURE 52. Typhlamia bella n. sp., holotype, female A), dorsal view; B) lateral view. Scale 1 mm.

Pereopod-4 (Fig. 54D): Clinging type; basis with two pinnate setae; ischium with two setae ventrally; merus with one spiniform seta and one short simple seta; carpus with two spiniform setae, small prickly tubercle, and one seta distally; propodus six times as long as wide, with two spiniform ventro-distal setae and with distal setae just reaching end of dactylus; unguis broken. Pereopod-5 (Fig. 54F): Similar to pereopod-4; unguis with bifid tip. Pereopod-6 (Fig. 54E): Similar to pereopod-4, but propodus with three distal setae, little shorter than dactylus..



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FIGURE 53. Typhlamia bella n. sp., female paratype. A) Antennule; B) Antenna; C) Labrum; D) Right mandible; D’) Molar process; E) Left mandible; F) Maxillule; G) Maxilla; H) Labium; I) Maxilliped. Scale: A, B = 0.1 mm; C-I = 0.01 mm.



FIGURE 54. Typhlamia bella n. sp., female paratype. A) Cheliped; B) Pereopod-1; C) Pereopod-2; D) Pereopod-4; E) Pereopod-6; F) Pereopod-5; G) Pleopod; H) Uropod. Scale = 0.1 mm.

Pleopods 1–5 (Fig. 54G): Basal article naked; exopod with twelve plumose setae on outer margin andone seta on inner margin; endopod with 20 plumose setae on outer margin. Gap between most proximal seta and the other setae in both rami.



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Uropod (Fig. 54H): Endopod two-articled; proximal article 1.5 times length of distal article, with one distal seta; distal article tipped by three simple setae. Exopod one-articled, as long as endopod proximal article; tipped by one short and one long setae. Distribution: The species occurs in the Antarctic Ocean: Eastern part of Weddell Sea, at depths of 43924385 m, and in the vicinity of the South Shetland Islands at depths of 3804–3808 m.

Typhlotanais sensu lato ‘greenwichensis’ group Diagnosis: Body elongate, over seven times as long as wide. Pereonite margins parallel. Mandible molar process with irregular edges. Maxilliped basis with long setae (longer than endites); endites edge with two tubercles and two setae. Cheliped basis reaching pereonite-1 ventrally. Chela slender, carpus longer than propodus, with minute setae on dorsal margin. Pereopods 1–3 coxa with spur. Pereopods 4–6 prickly tubercles surrounded by minute spines. Uropod rami one- or two-articled. Species included: Typhlotanais greenwichensis Shiino, 1970; Typhlotanais messinensis G.O. Sars, 1882 Remarks: The ‘greenwichensis’ morpho-group is well defined by the combined presence of minute setae on the dorsal margin of the cheliped, an anteriorly-directed spur on the coxa of pereopods 1–3, and prickly tubercles on pereopods 4–6 surrounded by blunt spines. The minute setae on the cheliped carpus also occur in the genus Peraeospinosus, while prickly tubercles surrounded by blunt spines are found in both Peraeospinosus and Torquella n. gen. The first genus is well defined by the long setae on propodus of pereopods 4–5 and the subequal uropod rami, while Torquella by the characteristic body habitus and shape of pereonite-1. Spurs on the coxa of pereopods 1–3 are characteristic for all described members Paratyphlotanais and for Typhlotanais mimosis n. sp. that is included in this paper with the ‘mixtus’ group (see remarks under ‘mixtus’ group).

Typhlotanais greenwichensis Shiino, 1970 (Figs 55–57) Typhlotanais greenwichensis: Shiino (1970) 1: 95–98; Kudinova-Pasternak (1975) 103: 211, 226; Shiino ((1978)1979) 5: 85–86; Larsen (2005): 209. Peraeospinosus adipatus: Błażewicz & Jażdżewski (1996) 17(3–4): 215–216, 219; Błażewicz-Paszkowycz & Jażdżewski (2000): 179, non Peraeospinosus adipatus (Tzareva, 1982).

Material examined: Admiralty Bay – central basin: 12 females, OC-336, 24 Mar 1981, depth 21 m; one female, C-734, 10 Jan 1988, depth 400–600 m; two females, OC-736, 17 Dec 1988, depth 480m; Ezcurra Inlet: one female, OC-366, 19 Oct 1983, depth 226m; one female, OC-411, 11 Jan 1985, depth 81 m; Section I: one female, OC-270, 21 Jan 1980, depth 175 m; one female, OC-452, 15 Mar 1985, depth 37 m; one female, OC-431, 8 Feb 1985, depth 120 m; one female, OC-444, 21 Feb 1985, depth 175 m; one female, OC496, 9 Sep 1985, depth 187 m ; 3 females, OC-485, 10 Oct 1985, depth 232 m; two females, OC-548, 8 Jan 1985, depth 296 m; one female, OC-447, 9 Sep, depth 317 m; one female, OC-520, 3 Nov 1985, depth 335 m; one female, OC-449, 9 Mar 1985, depth 349 m; one female, OC-448, 9 Mar 1985, depth 352 m; Section II: 14 females OC-321, 17 Mar 1980, depth 290 m; one female OC-252 Jan 1980, depth 240 m,; Section III: four females, OC-303, 7 Mar 1980, depth 12 m. Diagnosis: Antennule longer than carapace; article-2 half as long as article-3. Antenna article-4 seven times as long as wide. Cheliped carpus three times as long as wide. Pereopods 2–3 merus with long setae (longer than half of carpus); propodus with simple seta ventrally; uropod rami one-articled.



FIGURE 55. Typhlotanais greenwichensis, female. A), lateral view; B) dorsal view. Scale 1 mm.



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Complementary description: Female 4.6 mm (Fig. 55A, B), body long, 6.8 times as long as wide; carapace smooth, slightly longer than wide, lateral margins slightly rounded; rostrum pointed; pereonites smooth, all wider than long; pereonite-1 shortest, almost twice as wide as long, half as long as pereonite-3; pereonites 2, 4 and 5 subequal, 1.5 times as long as pereonite-1; pereonite-3 longest; pereonite-6 0.6 times as long as pereonite-5. Pleon 1.5 times as long as carapace; pleonites 1–5 similar in size; pleotelson rounded. Antennule (Fig. 56A): Article-1 slender, almost four times as long as wide, with two simple setae along article and five simple setae distally; article-2 with one simple seta distally; article-3 twice as long as article-2, with apical spur and four distal setae. Antenna (Fig. 56B): Article-2 with two setae distally, just longer than wide, almost twice as long as article-3; article-4 7.5 times as long as wide, twice as long as article-5, with one long and six short setae distally; article-5 with one simple seta distally; article-6 very short, with three terminal setae. Mouth parts: Labrum (Fig. 56C) hood-shaped, with minute setae. Mandible (Figs 56D,E) molar welldeveloped, longer than incisor with irregular edges; incisor blunt; lacinia mobilis well-developed, crenulated. Maxillule (Fig. 56F) endite with eight apical spiniform setae, palp with two terminal setae (not drawn). Labium both lobes (Fig. 56G) poorly separated and sparsely setose distally. Epignath (Fig. 56I) curved, simple distally. Maxilliped (Fig. 56H): coxa reduced; basis fused into heart-shaped plate, 1.5 times as long as wide, with long seta; endite with two simple setae and two tubercles distally; palp article-1 unarmed; article-2 wedge-shaped, with three setae on inner margin and one minute seta on outer margin; article-3 trapezoidal, with four serrated setae on inner margin; article-4 slender, with one simple seta on outer margin and five serrated distal setae. Cheliped (Fig. 57A): Basis slightly rounded, twice as long as wide; merus wedge shaped, with one seta ventrally; carpus with one long, one short and one minute seta ventrally and with seven minute setae along dorsal margin; propodus slightly shorter than carpus, with distal seta on inner side; fixed finger (propodus projection) tipped by strong spine, with three setae on well-calcified inner margin and two setae ventrally. Pereopod-1 (Fig. 57B): Slender (walking type); coxa with spur; basis six times as long as wide, almost as long as carpus and propodus combined, with four setae dorsally; ischium short, with one seta; merus almost as long as carpus, with two simple setae distally; carpus with six simple setae distally; propodus 1.3 times as long as carpus, with three setae subdistally and one seta ventrally; dactylus half as long as unguis; both together little shorter than propodus. Pereopod-2 (Fig. 57C): Slender (walking type); coxa with spur; basis 4.5 times as long as wide, with four setae dorsally and one ventrally; ischium with simple seta; merus a little shorter than carpus, with three simple setae distally; carpus with four simple setae and one spiniform distally; propodus almost as long as merus and carpus combined, with one simple seta ventrally and two setae dorsally; dactylus almost as long as unguis, with one simple seta. Pereopod-3 (Fig. 57D): Similar to pereopod-2. Pereopod-4 (Fig. 57E): Clinging type; basis robust, twice as long as wide, with three setae ventrally; ischium with two setae; merus with two spiniform seta ventrally; carpus with spine distally and with prickly tubercles surrounded by row of minute spines; propodus with two spiniform ventrally and one dorso-distal setae little shorter than dactylus; dactylus three times as long as unguis; both subequal to propodus. Pereopod-5 (Fig. 57F): Similar to pereopod-4. Pereopod-6 (Fig. 57G): Similar to as pereopod-5, but propodus with three dorso-distal setae. Pleopod (Fig. 57H): Basal article naked; exopod with one seta on inner margin and twelve plumose setae on outer margin; endopod with eighteen plumose setae on outer margin; clear gap between proximal seta and other setae. Uropod (Fig. 57I): Basal article as long as wide; both rami one-articled; exopod 0.7 times as long as endopod, with one strong seta and one regular seta distally; endopod tipped by one strong, four regular and one pinnate setae.



FIGURE 56. Typhlotanais greenwichensis, female. A) Antennule; B) Antenna; C) Labrum; D) Left mandible; E) Right mandible incisor; F) Maxillule; G) Labium; H) Maxilliped; I) Epignath. Scale: A, B = 0.1 mm; C-I = 0.01 mm.



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FIGURE 57. Typhlotanais greenwichensis, female. A) Cheliped; B) Pereopod-1; C) Pereopod-2; D) Pereopod-3; E) Pereopod-4; F) Pereopod-5; G) Pereopod-6; H) Pleopod; I) Uropod. Scale = 0.1 mm.



Distribution: Antarctic: Antarctic Peninsula, South Shetland Islands, South Sandwich Islands at the depth range of 12–757 m. Remarks: Typhlotanais greenwichensis can be distinguished from the Mediterranean T. messinensis by lacking a spiniform seta on the merus of pereopods 2 and 3 and in having relatively long setae on the merus, carpus and propodus of the same appendages (those on the merus reach half the length of the carpus). T. messinensis also has spiniform setae on the ventral side of the propodus.

Typhlotanais messinensis G.O. Sars 1882 (Figs. 58–60) Typhlotanais messinensis: G.O. Sars (1882a) 7: 39–40; Carus (1885) 1(2): 431; Norman & Stebbing (1886) 12: 109; G.O. Sars (1886) 11: 346–349; G.O. Sars, (1896): 18; Dollfus (1898) 11: 36; Norman (1899) 3: 340; Nierstrasz (1913) 32 (a): 34; Lang (1970) 23 (4): 274–275, 288; Larsen (2005): 209.

Material examined: CRU 3896, Messina 1-20/5. Diagnosis: Antennule shorter than carapace; article-2 0.3 times as long as article-3. Antenna article-4 four times as long as wide. Cheliped carpus 2.5 times as long as wide wide. Pereopod-2-3 merus with short setae (shorter than half of carpus); propodus with spiniform seta ventrally; uropod rami two-articled. Complementary description: Female 4.6 mm (Fig. 58A,B), body long, 7.1 times as long as wide; carapace smooth, 1.3 times as long as wide, lateral margins slightly rounded; rostrum slightly pointed; pereonites smooth, all wider than long; pereonite-1 shortest, almost twice as wide as long, half as long as pereonite-3; pereonites 2, 4 and 5 subequal, 1.5 times as long as pereonite-1; pereonite-3 longest; pereonite-6 0.6 times as long as pereonite-5. Pleon 1.5 times as long as carapace, pleonites 1–5 similar in size; pleotelson rounded. Antennule (Fig. 59A): Article-1 slender, three times as long as wide, with four short setae along article on inner margin and one simple and three pinnate setae distally; article-2 with simple seta distally; article-3 almost four times as long as article-2, with apical spur and four distal setae. Antenna (Fig. 59B): Article-2 a little longer than wide, with two setae distally, almost twice as long as article-3; article-4 five times as long as wide, with two setae distally; article-5 with one long seta distally; article-6 very short, with five terminal setae. Mouth parts: Labrum (Fig. 59C) hood-shaped, covered by minute setae. Mandible (Figs 59D,E) molar well-developed, longer than incisor with irregular edges; incisor blunt; lacinia mobilis well-developed, crenulated. Maxillule (Fig. 59F) endite with eight apical spiniform setae, palp lost during dissection. Labium (Fig. 59G) withboth lobes poorly separated and sparsely setose distally. Epignath (Fig. 59I) curved, simple distally. Maxilliped (Fig. 59H): coxa reduced; basis fused in heart-shaped plate, 1.5 times as long as wide, with long seta; endite with two tubercles distally; palp article-1 unarmed; article-2 wedge-shaped, with three setae on inner margin and one minute seta on outer margin; article-3 trapezoidal, with four setae on inner margin; article-4 slender, with one simple seta on outer margin and five distal setae. Cheliped (Fig. 60A): Basis slightly rounded, 1.8 times as long as wide; merus wedge-shaped, with one seta ventrally; carpus 2.5 times as long as wide, with two setae ventrally and with six minute setae along dorsal margin; propodus slightly shorter than carpus, with distal seta on inner side; fixed finger (propodus projection) tipped by strong spine, with three setae on well-calcified inner margin and two setae ventrally. Pereopod-1 (Fig. 60B): Coxa with spur; ischium short, with one seta; merus almost as long as carpus; carpus with three simple setae distally; propodus 1.3 times as long as carpus, with three setae subdistally; dactylus half as long as unguis with one seta; dactylus and unguis combined 0.7 times as long as propodus. Pereopod-2 (Fig. 60C): Coxa with spur; basis with three setae along article; ischium with simple seta; merus slightly shorter than carpus, with two simple setae and one spiniform seta distally; carpus a little longer



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than merus, with three simple setae and one spiniform distally; propodus almost as long as merus and carpus combined, with one spiniform seta ventrally and two setae dorsally; dactylus almost as long as unguis. Pereopod-3 (Fig. 60D): Similar to pereopod-2. Pereopod-4 (Fig. 60E): Clinging type; basis robust, twice as long as wide, ith two pinnate setae ventrodistally; ischium with two setae; merus with two spiniform setae ventrally; carpus with spine distally and with prickly tubercles; propodus with two spiniform setae ventrally and one dorso-distal seta, a little shorter than dactylus; dactylus three times as long as simple unguis; both almost as long as propodus.

FIGURE 58. Typhlotanais messinensis G.O. Sars, 1882, female. A), lateral view; B) dorsal view. Scale = 1 mm.



FIGURE 59. Typhlotanais messinensis G.O. Sars, 1882, female. A) Antennule; B) Antenna; C) Labrum; D) Left mandible; E) Right mandible; F) Maxillule; G) Labium; H) Maxilliped; I) Epignath. Scale: A, B = 0.1 mm; C–I = 0.01 mm.



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FIGURE 60. Typhlotanais messinensis, female. A) Cheliped; B) Pereopod-1; C) Pereopod-2; D) Pereopod-3; E) Pereopod-4; F) Pereopod-5; G) Pereopod-6; H) Pleopod; I) Uropod. Scale = 0.1 mm.

Pereopod-6 (Fig. 60F): Similar to as pereopod-5, but propodus with three dorso-distal setae. Pleopod (Fig. 60G): Basal article naked; exopod with one seta on inner margin and twelve plumose setae on outer margin; endopod with sixteen plumose setae on outer margin; clear gap between proximal seta and other setae.



Uropods (Fig. 60H): Basal article little longer than wide; both rami two-articled; exopod 0.6 times as long as endopod; proximal article little shorter than distal with one seta distally; distal article with one strong and one small seta distally; endopod proximal article 0.3 times as long as distal article; endopod distal article with one simple and two pinnate seta at middle, tipped by two long, two short and one pinnate setae distally. Distribution: Mediterranean Sea, Sicily, Messina at a depth range of 37–50 m.

? Typhlotanais greenwichensis Shiino, 1970 (Fig. 61) Material examined: one female, two mancae, (K 41392), ANT XXII-2, PS 61/46-7, 60°38.35'–60°38.12'S, 53°57.36'–53°57.49'W, depth 2893.2–2893.2 m, 30 Jan 2002; one female, (K 41394), ANT XXII-2, PS 61/ 129-7, 59°52.41'S, 59°57.69'W, depth 3601.0 m, LBC, 23 Feb 2002; one female, (K 41397), ANT XXII-3, 746, 71°18.35'–71°18.28'S, 13°57.71'–13°57.31'W, depth 1030–1040 m, 20 Feb 2005; one female, (K 41395), ANT XXII-3, PS 67/151-7-E, 61°45.52'–61°45.42'S, 47°7.68'–47°8.04'W, depth 1182.0–1185.0 m, 21 Mar 2005.

FIGURE 61. ?Typhlotanais greenwichensis Shiino, 1970, female. A) Antennule; B) Antenna; C) Cheliped; D) Pleopod ; E) Uropod. Scale = 0.1 mm.

Distribution: West Antarctic, vicinity of the Falkland Islands, Drake Passage, Eastern Weddell Sea at depths of 1030–3601 m. Remarks: The specimens found deeper than 1030 m in the ANDEEP samples were very closely similar to those occurring in shallow waters, although they had a more slender antennule, chelae and uropods (Fig. 61). In such circumstances it is not clear if there is one eurybathic species exhibiting the phenomenon of polar emergence or whether there are two (shallow- and deep-water) cryptic or sibling species. A REVISION OF FAMILY TYPHLOTANAIDAE SIEG 1984


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‘mixtus’ group Diagnosis: Body elongate (over seven times as long as wide). Carapace elongate (over 1.5 times as long as wide). Cheliped basis separated from pereonite-1 by a gap ventrally. Pereonite-1 shorter then the others, all pereonites wider than long or square; pereonite margins delicately rounded; pereonites 1–3 carpus and propodus with spiniform seta; pereopods 4–5 merus with large prickly tubercles (larger than half of carpus length), propodus distal seta as long as half of dactylus; unguis simple; dactylus and unguis combined almost as long as propodus. Both pleopod rami with proximal seta separated from the others gap. Both uropod rami two-articled. Male: Unknown. Species included: Typhlotanais mixtus Hansen, 1913; Typhlotanais mimosis n. sp. Remarks: The ‘mixtus’ group is well defined by the large gap between the cheliped basis and pereonite1 ventrally, large prickly tubercles on the carpus of pereopods 4–6 and a long dactylus on pereopods 4–6 that, combined with the unguis, is longer than the propodus. These characters are also valid for the provisional ‘spinicauda’ group, although they have pair of large terminal spines on the pleotelson. Typhlotanais mimosis n. sp. has a distinctive projection on the coxa of pereopods 1 to 3 characteristic also of T. greenwichensis and T. messinensis. The last two species constitute a separate ‘greenwichensis’ group defined by a number of other characters such as a lack of a gap between the cheliped basis and pereonite-1 ventrally, a row of short seta on cheliped carpus, and prickly tubercles surrounded by a row of spines. Since T. mimosis has the specific coxal projection on pereopods 1–3 the assignment of the species to the ‘mixtus’ group may be ambiguous, but because the new species shares more characters with T. mixtus than with any other ‘typhlotanaid’ it is placed in the ‘mixtus’ group at the moment.

Typhlotanais mixtus Hansen, 1913 (Figs 62–64) Typhlotanais mixtus: Hansen (1913) 3 (3): 53, 59–61; Nierstrasz (1913) 32 (a): 38; Stephensen (1913) 22: 270, 421; Stephensen (1932) 6: 350; Stephensen (1936) 80 (2): 29; Lang (1970) 23 (4): 275, 288; Morino (1971) 18 (5): 353. Peraeospinosus mixtus: Sieg (1986a) 45: 156–163; Błażewicz-Paszkowycz & Sekulska-Nalewajko (2004) 27: 224, 226.

Material examined: Syntype: one female, CRU 7351, Ingolf 125, Arctic, N. of Iceland, 68º08'N, 016º02'W, depth 1373m, 29 Jul 1896; Other material: two females (one dissected on slides), BIOICE 2773, Iceland Plateau, 69º24.8'N, 14º27.53'W, depth 1629 m, RP, 3 Aug 1995; bottom temp -0.86ºC, salinity 34.90 ‰. Diagnosis: Carapace narrow (1.5 times as long as wide). Cheliped carpus 2.5 times as long as wide, propodus as long as carpus. Pereopods 1–3 coxa without projection; pereopod-1 merus and carpus about as wide as long; pereopods 2–3 propodus with two simple setae on dorsal margin. Description: Female (Fig. 62). 1.7 mm body length, body long, 7.3 times as long as wide. Carapace 1.5 times as long as wide, almost as long as pereonite-1 and 2 combined, 20% of total body length. All pereonites rectangular, with lateral margin parallel; pereonites 2 and 6 subequal, 1.5 times as long as pereonite-1; pereonite-3 slightly longer than pereonite-2, as long as wide; pereonites 4 and 5 subequal, as long as wide. Pleon a little longer than carapace; pleonites 1–5 subequal; pleotelson with well-developed distal process. Antennule (Fig. 63A): Article-1 2.75 times as long as wide, 1.75 times as long as articles 2 and 3 combined, with two groups of simple and pinnate setae at middle and distally; article-2 with two short setae distally; article-3 twice as long as article-2, with two four setae distally.



FIGURE 62. Typhlotanais mixtus Hansen, 1913, female. A), lateral view; B) dorsal view. Scale = 0.1 mm.



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FIGURE 63. Typhlotanais mixtus, female. A) Antennule; B) Antenna; C) Left mandible; D) Right mandible incisor; E) Maxillule; F) Labium. Scale: A, B = 0.1 mm; C–F = 0.01 mm.

Antenna (Fig. 63B): Article-2 slightly longer than article-3, with two setae almost as long as article-3; article-3 with one seta distally; article-4 1.75 times as long as article-5 with two simple and two pinnate setae distally; article 5 with one seta distally; article-6 small with four setae distally. Mouth parts: Labrum, maxilla, maxilliped and epignath destroyed during dissection. Mandible (Fig. 63C, D) stout; molar process well-developed, with well-calcified distal margin supported with tubercle-like structures; lacinia mobilis well-developed, crenulated. Maxillule (Fig. 63E) endite with eight apical spiniform setae with numerous delicate short seta along margins; palp lost during dissection. Both lobes of labium (Fig. 63F) poorly separated, hirsute in distal parts. Cheliped (Fig. 64A): Basis 1.9 times as long as wide; merus wedge-shaped, with one short seta ventrally; carpus over 2.3 times as long as broad, with two dorsal setae and with two short setae ventrally; propodus and fixed finger as long as carpus, over three times as long as wide; fixed finger with three teeth and three setae on inner margin and two setae on ventral margin. Pereopod-1 (Fig. 64B): Slender (walking type); coxa present; basis a little shorter than merus, carpus and propodus combined, with two setae on dorsal margin and one seta on ventral margin; ischium with simple seta; merus almost as long as carpus, with two setae distally; carpus with two short, simple and one spiniform setae distally; propodus with three subdistal seta dorsally; dactylus 0.3 times length of propodus. Pereopod-2 (Fig. 64C): Slender (walking type); basis as long as merus, carpus and half of propodus combined, with one seta ventrally and one seta dorsally; ischium with simple seta; merus almost as long as carpus with two setae distally; carpus with four short simple setae and one spiniform seta distally; propodus with one spiniform seta ventrally and two simple setae dorsally reaching end of dactylus.



FIGURE 64. Typhlotanais mixtus, female. A) Cheliped; B) Pereopod-1; C) Pereopod-2; D) Pereopod-3; E) Pereopod-4; F) Pleopod; G) Uropod. Scale = 0.1 mm. A REVISION OF FAMILY TYPHLOTANAIDAE SIEG 1984


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Pereopod-3 (Fig. 64D): Similar to pereopod-2. Pereopod-4 (Fig. 64E): Clinging type; basis three times as long as wide; ischium with two setae; merus with two spiniform setae; carpus with spiniform hooks and large prickly tubercles, as long as carpus; propodus almost four times as long as wide, with two spiniform ventro-distal setae and with distal seta just reaching half of dactylus; unguis simple, with dactylus almost as long as propodus. Pereopods 5 and 6 lost during dissection. Pleopods 1–5 (Fig. 64F): Basal article naked; exopod with eight plumose setae on outer margin and one seta on inner margin; endopod with twelve pinnate setae on outer margin. Gap between most proximal seta and other setae in both rami. Uropod (Fig. 64G): Both rami two-articled. Endopod proximal article little longer than distal, with one pinnate seta distally; distal article tipped by five simple setae. Exopod two-articled; proximal article little longer than distal article, with two simple seta distally; distal article with two (short and long) terminal setae. Distribution: North Atlantic Davis Strait, North of the Faroes, East and North of Iceland, south Jan Mayen at the depth range 906–2626m (temp. 0.6–1.5°C) and Iceland Plateau (69º24.8'N, 14º27.53'W) at depths of 1373–1629 m.

Typhlotanais mimosis n. sp. (Figs 65–67) Material examined: Holotype: female (K 41420), ANT XXII-3, PS 67/59-5-E, 67°29.74'–67°29.61'S, 0°01.93'–0°02.19'W, depth 4655–4655 m, epibenthic sledge, 14 Feb 2005. Paratypes: three females (two dissected on slides) (K 41421), ANT XXII-3, PS 121-11-E, 63°37.73'–63°37.55'S, 50°38.09'–50°38.37'W, depth 2663–2659 m, epibenthic sledge, 15 Mar 2005. Diagnosis: Carapace narrow (1.85 times as long as wide). Cheliped slender, carpus over three times as long as wide, propodus shorter than carpus. Pereopods 1–3 coxa with projection; pereopod-1 merus and carpus about three times as long as wide; pereopod propodus 2–3 with strong seta on dorsal margin. Etymology: Mimosa is one of four stars in the constellation of ‘The Southern Cross’. Description: Ovigerous female length 1.8 mm (Fig. 65A,B), body long, 8.3 times as long as wide. Carapace 1.8 times as long as wide, almost as long as pereonites 1 and 2 combined, 20% of total body length. All pereonites rectangular, with lateral margin slightly rounded; pereonite-2 almost 1.5 times as long as pereonite1; pereonites 2 and 4 subequal; pereonite-3 slightly longer than pereonite-2, as long as wide; pereonite-5 1.25 times as wide as long; pereonite-6 shortest, over twice wider than long, pleon as long as carapace; pleonites 1– 5 subequal; pleotelson with well-developed distal process. Antennule (Fig. 66A): Article-1 over three times as long as wide, longer than articles 2 and 3 combined, with one medial seta and one simple and two pinnate setae distally; article-2 with one short distal seta; article3 three times as long as article-2, with two short and three long setae distally. Antenna (Fig. 66B): Article-2 a little longer than article-3, with two setae almost as long as article-3; article-3 with one seta distally; article-4 1.6 times as long as article-5, with one seta distally, article-6 short with one short and three long setae distally. Mouth parts: Labrum lost during dissection. Mandible (Figs 66C, D) stout; molar process well-developed, with well-calcified distal margin supported with tubercle-like structures; lacinia mobilis well-developed, crenulated. Maxillule (Fig. 66E) endite with eight apical spiniform setae, with numerous delicate setae along margins; palp lost during dissection. Maxilla lost during dissection. Both lobes of labium (Fig. 66F) poorly separated, setose in distal parts. Maxilliped (Fig. 66G): basis longer than wide, with short seta on distal margin; endites distal margin with two tubercles and one seta in disto-lateral corner; palp four-articled: article1 naked; article-2 wedge-shaped, with three setae on inner margin and one small seta on outer margin; article-



3 with three simple setae on inner margin; article-4 slender (over twice as long as wide), with one simple seta on outer margin and five (two bipinnate and three simple) terminal setae.

FIGURE 65. Typhlotanais mimosis n. sp., holotype, female. A) dorsal view; B) lateral view. Scale 0.1 mm. A REVISION OF FAMILY TYPHLOTANAIDAE SIEG 1984


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FIGURE 66. Typhlotanais mimosis n. sp., female paratype. A) Antennule; B) Antenna; C) Left mandible; D) Right mandible; E) Maxillule; F) Labium; G) Maxilliped; H) Cheliped. Scale: A, B, H = 0.1 mm, C–G = 0.01 mm.



FIGURE 67. Typhlotanais mimosis n. sp., female paratype. A) Pereopod-1; B) Pereopod-2; C) Pereopod-3; D) Pereopod-4; E) Pereopod-5; F) Pereopod-6; G) Pleopod; H) Uropod. Scale = 0.1 mm. A REVISION OF FAMILY TYPHLOTANAIDAE SIEG 1984


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Cheliped (Fig. 66H): Basis 2.5 times as long as wide; merus wedge-shaped, with one short seta ventrally; carpus over three times as long as broad, with two dorsal setae and with two short setae ventrally; propodus and fixed finger shorter than carpus, over three times as long as wide; fixed finger with three teeth and three setae on inner margin and two rod setae on ventral margin. Pereopod-1 (Fig. 67A): Slender (walking type); coxa with spur tipped with strong seta; basis as long as carpus and propodus combined, with two setae; ischium with simple seta; merus and carpus subequal, with one distal seta; carpus with five short setae; propodus with short subdistal seta ventrally and three subdistal seta dorsally; dactylus a little shorter than unguis; dactylus and unguis combined almost as long as propodus. Pereopod-2 (Fig. 67B): Slender (walking type); coxa with spur tipped with strong seta; basis as long as merus, carpus and propodus combined, with two setae along article; ischium with simple seta; merus and carpus subequal, with two ventro-distal setae; carpus with three short simple setae and one spiniform seta distally; propodus with one spiniform, subdistal seta ventrally and strong seta dorsally reaching end of dactylus; dactylus little shorter than unguis, with one seta; dactylus and unguis combined about half as long as propodus. Pereopod-3 (Fig. 67C): Similar to pereopod-2, but merus with one simple and one spiniform setae. Pereopod-4 (Fig. 67D): Clinging type; basis with one seta dorsally; ischium without seta; merus with two spiniform setae; carpus with spiniform hooks and one dorso-distal seta, prickly tubercles, as long as carpus; propodus four times as long as wide, with two spiniform ventro-distal setae and with distal setae just reaching half of dactylus; unguis simple; unguis and dactylus almost as long as propodus. Pereopod-5 (Fig. 67E): Similar to pereopod-4, but basis ventral margin with simple and pinnate setae, propodus with pinnate seta on dorsal margin. Pereopod-6 (Fig. 67F): Similar to pereopod-4, but propodus with three distal setae as long as half of propodus. Pleopods 1-5 (Fig. 67G): Basal article naked; exopod with eight plumose setae on outer margin and one seta on inner margin; endopod with eleven pinnate setae on outer margin. Gap between most proximal seta and the other setae in both rami. Uropod (Fig. 67H): Endopod two-articled; proximal article less than twice as long as distal article, with one distal seta; exopod two-articled, longer than endopod proximal article; proximal article as long as half of distal article, with one subdistal seta. Distribution: The species is known from two localities in the Southern Ocean: North of Martha Coast and between the tip of the Antarctic Peninsula and South Orkney Island, at depths of 2659–4655 m. Remarks: T. mimosis n. sp. can be distinguished from all typhlotanaids by two long proximall setae on pereonite 1. From T. mixtus that is puitative congeneris species, it differs having relative long carapace that is almost twice as long as wide (only 1.5 as long as wide in T. mixtus). The well developed spur on basis of pereopods 1–3 in the new species is character commonly observed in “greenwichenis” group. The lack of the minute setation along cheliped carpus and large prickly tubercles in pereopods 4–6 of the new species exclude it from “greeenwichensis” group. At the persent stage of knowledge it is assumed that some features (e.g. spur on basis) are consistent within particular morpho-group while can vary within the others groups. A new look on this question could bring a description of more new species while a phylogenetic analysis would test a real realation between ‘mixtus’ and ‘greenwichensis’ group.

‘spinicauda’ group Diagnosis: Body over seven times as long as wide. Carapace slightly tapering proximally. Pereonites rectangular, carapace and pereonites lateral margin parallel. Pleotelson apex supporting two strong spines; small gap between cheliped basis and proximal margin of pereonite-1 ventrally. Antennule sparsely setose; labrum with



relatively long setae on upper margin; mandible molar with crenulated margin with blunt spines or tubercles. Maxilliped basis longer than wide, without seta; endites with two large tubercles and two setae distally. Cheliped relatively narrow (chela three times as long as wide), merus and carpus dorsally well-calcified; pereopod-1 articles without spiniform setae. Pereopods 2 and 3 merus and carpus with spiniform setae; pereopods 4–6 with large prickly tubercle (longer than half of carpus length), propodus dorso-distal setae reaching half of dactylus, dactylus and unguis combined just as long as propodus. Both uropod rami two-articled; endopod proximal article with spiniform setae on inner margin, exopod shorter than endopod proximal article. Species included: Typhlotanais spinicauda Hansen 1913, Typhlotanais squamiger n. sp. Remarks: A pleotelson with the apex supported by two strong terminal spiniform setae, and spiniform setae on the inner margin of the uropod endopod proximal article in Typhlotanais spinicauda and T. squamiger n. sp. are unique characters among known ‘typhlotanaids’ and the potential for erecting a new genus to which to assign these two species is fairly high. Because only a few specimens were available for study, resolution of a generic status is premature; instead, a ‘spinicauda’ morpho-group is proposed to emphasize the close similarity between T. spinicauda and T. squamiger n. sp. T. spinicauda and T. squamiger share large prickly tubercles and a general body habitus with T. mixtus possibly indicating a phylogenic affinity. The last taxon, however, lacks spiniform setae on the merus of pereopods 2 and 3, and it has a large gap ventrally between the cheliped basis and the proximal margin of pereonite-1 (the gap is much smaller in the ‘spinicauda’ group). The presence of strong terminal spines on the pleotelson and a row of spiniform setae on the inner margin of the uropod endopods is characteristic for Tanabnormia cornicauda Guţu, 1986. This character suggested to Guţu the possibility of a predatory potential of the species, although no specific evidence for this kind of feeding was given. During the dissection of the one specimen of T. squamiger n. sp. the gut was found to contain only fine particles resembling fine benthic sediments, similar to the gut contents found in most shallow water tanaids (Błażewicz-Paszkowycz and Ligowski, 2002). This feature may have more to do with defense of the posterior of the animal when in its tube or burrow.

Typhlotanais spinicauda Hansen, 1913 (Figs 68–70) Typhlotanais spinicauda: Hansen (1913) 3 (3): 53–54; Niestrasz (1913) 32: 37; Stephensen (1913) 22: 268–269, 418; Stephensen (1936) 6: 35; Lang (1970) (2), 23 (4): 277, 288; Morino (1971) 18(5): 353; Hassak & Holdich (1987) 16(3): 224, 226–229; Kudinova-Pasternak (1990b) 69: 139; Larsen (2005): 210.

Material examined: Holotype: non-ovigerous female, Davis Strait, Ingolf St. 28, 65°14’N, 55°42’W; depth 420 fm (= 768 m); Other material: two females BIOICE 2257 (one dissected on slides), Irminger Basin, 5 Sep 1992, 63o 14.61'N 26o 29.14'W, depth 1209 m, epibenthic sled (Rothlisberg-Pearcy); bottom temp. 4.09ºC, salinity 34.94o/oo, clay with shell sand. Diagnosis: Pereopods 2 and 3 carpus with spiniform setae only; propodus with two regular dorso-distal seta (one more robust then the other). Complementary description: Non-ovigerous female. Body length 2.3 mm (Fig. 68A,B), body long, 7.5 times as long as wide; carapace smooth, 1.4 times as long as wide, margins almost parallel, tapering proximally; rostrum weakly pointed; pereonites smooth, rectangular, with parallel lateral margins: pereonite-1 shortest, 2.2 times as wide as long; pereonites 2 and 6 0.7 times as long as wide; pereonite-3 square; pereonites 4 and 5 subequal, slightly shorter than wide. Pleon about as long as carapace and pereonite-1 combined; pleonites 1–5 similar in size; pleotelson rounded; caudal projection with two strong spines (Fig. 68D). Antennule (Fig. 69A): Article-1 about twice as long as wide and twice as long as article-3, strongly taper-



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ing distally, with one simple seta and two pinnate setae distally and three short setae in row on outer margin; article-2 as long as wide with small simple and one pinnate setae distally; article-3 three times as long as article-2, with six distal setae. Antenna (Fig. 69B): Articles 1 and 2 broken; article-2 with one simple seta distally; article-3 naked; article-4 twice as long as article-5, with four setae distally; article-5 with simple seta; article-6 with six simple subterminal and terminal setae.

FIGURE 68. Typhlotanais spinicauda Hansen, 1913, holotype, female. A) dorsal view; B) Holotype, female, lateral view; C) Carapace, ventral view. D) Pleotelson with uropod. Scale: A, B = 1 mm; C, D = 0.1 mm.



FIGURE 69. Typhlotanais spinicauda, female paratype. A) Antennule; B) Antenna; C) Labrum; D) Left mandible; D’) Molar process; E) Right mandible; E’) Molar process; F) Maxillule; G) Maxilla; H) Labium; I) Maxilliped. Scale: A, B = 0.1 mm; C–I = 0.01 mm.

Mouth parts: Labrum (Fig. 69C) hood-shaped, covered by numerous relatively long setae. Mandible (Figs 69D,E) stout; molar process well-developed, with strongly crenulated edges (Figs 69D’,E’); lacinia mobilis



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well-developed, crenulated. Maxillule (Fig. 69F) endite with eight terminal spiniform setae, three short simple setae on outer margin as well as on inner margin; palp lost during dissection. Maxilla elongated (Fig. 69G). Maxilliped (Fig. 69I) bases nearly twice as long as wide, without simple setae distally; each endite with two setae and two large tubercles on distal margin; palp article-1 naked; article-2 wedge-shaped, with three simple setae on inner margin and one short, simple seta on outer margin; article-3 trapezoidal; with four setae on inner margin; article-4 with one simple seta on outer margin and five strong terminal setae. Labium (Fig. 69H) with group of setae on outer corner of inner lobe; outer lobe with small setae. Cheliped (Fig. 70A): Basis less than twice as long as wide, naked, not reaching pereonite-1 ventrally (Fig. 68C); merus with one seta; carpus twice as long as wide, covered by numerous scales, with two simple setae and one minute ventrally and one short setae dorsally; carpal shield poorly developed; chela almost as long as carpus, three times as long as wide and with three setae on inner margin, with two simple setae ventrally; dactylus curved distally, slightly longer than fixed finger, with one short seta proximally. Pereopod-1 (Fig. 70B): Of walking type; coxa naked; basis just as long as propodus and carpus combined, with two setae dorsally and two setae ventrally; ischium short with one simple seta; merus almost as long as carpus, with one short seta dorso-distally; carpus with four short setae distally; propodus with two subdistal setae dorsally and minute subdistal seta ventrally; unguis 1.5 times as long as dactylus; unguis and dactylus somewhat shorter than propodus. Pereopod-2 (Fig. 70C): Of walking type; basis just as long as merus, carpus and propodus combined, with three short seta along article; ischium with one seta; merus as long as carpus, with two simple and spiniform setae distally; carpus with two spiniform setae distally; propodus with spiniform seta ventrally and with two setae dorsally and one more robust than the other; dactylus shorter than unguis, combined shorter than propodus. Pereopod-3 (Fig. 70D): Similar to pereopod-2, but basis as long as rest of articles combined. Pereopod-4 (Fig. 70E): Of clinging type; basis with two pinnate setae ventrally; ischium with two setae; merus almost as long as carpus, with two subdistal spiniform setae; carpus with hooks distally and with large (over half as long as article) prickly tubercles ventrally; propodus twice as long as carpus, with two spiniform setae ventrally, one distal seta reaching over half of dactylus; dactylus tipped by simple unguis; dactylus and unguis as long as propodus. Pereopod-5 (Fig. 70F): Similar to pereopod-4. Pereopod-6 (Fig. 70G): Similar to pereopod-5, but propodus with three terminal setae and carpus with two dorso-distal setae. Pleopods 1–5 (Fig. 70H): All pleopods similar; exopod with eleven plumose setae on outer margin and with one plumose seta one inner margin; endopod with twelve plumose setae on outer margin; both rami with clear gap between proximal outer seta and others. Uropod (Fig. 68D): Basal article shorter than endopod proximal article; exopod and endopod two-articled; exopod articles subequal, combined shorter that endopod proximal article; distal article tipped by one long and one short seta; endopod proximal article with six spiniform setae on inner margin; endopod distal article with one subdistal and four distal setae. Distribution: known from Arctic Davis Strait (Hansen 1913) and Iceland-Rockall sector of North Atlantic (Bird personal comm.) Remarks: T. spinicauda differs from T. squamiger n. sp. in having smaller spiniform setae on the merus, carpus and propodus of pereopods 2 and 3, and much smaller spiniform setae ventrally on the propodus of pereopods 4–6. These spiniform setae are especially robust in T. squamiger. Also, the scale-like cuticular structures covering the antennules, chelae and pereopods are more distinct in T. squamiger than in T. spinicauda. Those scale-like structures occur also on the cheliped bases and in specific light conditions can be visible as ‘teeth’ when seen laterally.



FIGURE 70. Typhlotanais spinicauda, female paratype. A) Cheliped; B) Pereopod-1; C) Pereopod-2; D) Pereopod-3; E) Pereopod-4; F) Pereopod-5; G) Pereopod-6; H) Pleopod. Scale = 0.1 mm.

Typhlotanais squamiger n. sp. (Figs 71–73) Material examined: Holotype: non-ovigerous female (K 41418), ANT XXII/3, PS 67/88-8 E, 68°3.66'–68° 3.61'S, 20°27.90'–20°27.52' W, depth 4929–4931 m, epibenthic sledge, 27 Feb 2005. A REVISION OF FAMILY TYPHLOTANAIDAE SIEG 1984


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Paratype: non-ovigerous females (dissected on slides), (K 41437) the same locality. Diagnosis: Strong spiniform setae on pereopods 2 and 3 merus, carpus and propodus and pereopods 4–6 propodus; pereopods 2 and 3 propodus with one regular and one strong setae dorso-distal; antennule, cheliped and pereopods 1–3 with scales, cheliped merus strongly calcified. Etymology: squamiger (Lat.) – bearing a scale. The name reflects the presence of scale-like structures on the chelipeds and pereopods. Description: Non-ovigerous female. Body length 2.8 mm (Fig. 71A,B), long, 8.3 times as long as wide; carapace smooth, margins parallel tapering proximally, 1.3 times as long as wide; rostrum weakly pointed; pereonites smooth, rectangular, with parallel lateral margins; pereonite-1 shortest, less than twice as wide as long; pereonites 2 and 6 0.8 times as long as wide; pereonite-3 square; pereonites 4 and 5 subequal, slightly shorter than wide. Pleon as long as carapace and pereonite-1 combined; pleonites 1–5 similar in size; pleotelson rounded; caudal projection with two strong spines.

FIGURE 71. Typhlotanais squamiger n. sp. A) Holotype, female, dorsal view; B) Holotype, female, lateral view. Scale = 1 mm.



Antennule (Fig. 72A): Article-1 about three times as long as wide, and twice as long as article-3, with two small simple setae and two pinnate spines distally, numerous scale-like structures along article; article-2 as long as wide, with two small setae distally; article-3 over three times as long as article-2.

FIGURE 72. Typhlotanais squamiger n. sp., female paratype. A) Antennule; B) Antenna; C) Labrum; D) Left mandible; E) Right mandible; E’) Molar process; F) Maxillule; G) Maxilla; H) Labium; I) Maxilliped. Scale: A, B = 0.1 mm; C–I = 0.01 mm.



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Antenna (Fig. 72B): Article-1 broken; article-2 twice as long as article-3, with one simple seta; article-3 with numerous minute scales distally; article-4 twice as long as article-5, with one long, one short and one pinnate setae distally; article-5 with simple seta; article-6 with four simple subterminal and terminal setae. Mouth parts: Labrum (Fig. 72C) hood-shaped, covered by numerous, relatively long setae. Mandible (Figs 72D,E) stout; molar process well-developed, with strongly crenulated edges; lacinia mobilis well-developed, crenulated. Maxillule (Fig. 72F) endite with eight terminal setae, additional (ninth) seta visible centrally, three short simple setae on outer margin; palp lost during dissection. Maxilla (Fig. 72G) oval. Maxilliped (Fig. 72I) bases nearly twice as long as wide, without simple setae distally; each endite with two setae and two large tubercles on distal margin; palp article-1 with microtrichiae; article-2 wedge-shaped, with two simple and one serrated setae on inner margin and one short simple seta on outer margin; article-3 trapezoidal, with four setae on inner margin and scale structures on outer margin; article-4 with one simple seta on outer margin and five strong terminal setae. Labium (Fig. 72H) with bunch of setae on outer corner of inner lobe; outer lobe with small setae. Cheliped (Fig. 73A): Basis twice as long as wide, naked; merus with simple seta ventrally; carpus twice as long as wide, covered by numerous scales, with two long and one short setae ventrally and two short setae dorsally, carpal shield moderately developed; chela almost as long as carpus, twice as long as wide; fixed finger with three setae on inner margin and with two simple setae ventrally; dactylus slightly curved, with one short seta proximally. Pereopod-1 (Fig. 73B): Of walking type; coxa naked, with small spur; basis just as long as merus and carpus combined, with two setae dorsally and three setae ventrally; ischium short with one simple seta; merus almost as long as carpus but shorter than propodus, with two short setae distally; carpus with three short setae and one spiniform seta distally; propodus with two serrated subdistal setae dorsally; unguis twice as long as dactylus; unguis and dactylus slightly shorter than propodus. Pereopod-2 (Fig. 73C): Of walking type; basis as long as merus, carpus and half of propodus combined, with two simple setae ventrally; ischium with one seta; merus as long as carpus, with two simple and one spiniform setae distally; carpus with few scales ventrally and with three short setae and one spiniform seta distally; propodus with few scales ventrally, strong spiniform setae ventrally and with one thick and one thin seta dorsally; dactylus shorter than unguis, combined little shorter than propodus. Pereopod-3 (Fig. 73D): Similar to pereopod-2, but basis as long as rest of articles combined. Pereopod-4 (Fig. 73E): Of clinging type; basis naked; ischium with two setae; merus almost as long as carpus, with two subdistal, spiniform setae; carpus with hooks distally, two sensory setae dorsally, and large (over half as long as article) prickly tubercle ventrally; propodus a little longer than carpus, with two strong spiniform setae ventrally, and one distal seta reaching half of dactylus; dactylus tipped by simple unguis; dactylus and unguis as long as propodus. Pereopod-5 (Fig. 73F): Similar to pereopod-4, but propodus with medial, pinnate seta on propodus dorsal margin. Pereopod-6 (Fig. 73G): Similar to pereopod-5, but propodus with three terminal setae reaching half of unguis. Pleopods 1-5 (Fig. 73H): All pleopods similar; exopod outer margin with twelve plumose setae, andwith one plumose seta one inner margin; endopod arranged with sixteen plumose setae on outer margin; both rami with clear gap between proximal outer seta and others. Uropod (Fig. 73I): Basal article shorter than endopod proximal article; exopod and endopod two-articled; exopod about 0.4 times length of endopod; proximal article 0.6 times as long as distal article; distal article tipped by two simple setae; endopod proximal article 1.5 times as long as distal article, with six spiniform setae on inner margin; distal article with one seta at middle and four terminal setae. Distribution: Antarctic, Weddell Sea.



FIGURE 73. Typhlotanais squamiger n. sp., female paratype. A) Cheliped; B) Pereopod-1; C) Pereopod-2; D) Pereopod-3; E) Pereopod-4; F) Pereopod-5; G) Pereopod-6; H) Pleopod; Uropod. Scale = 0.1 mm.



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‘cornutus’ group Diagnosis: Body less than or about five times as long as wide; all pereonites wider than long. Antennule as long as carapace, article-1 with some simple setae. Mandible molar with some blunt, regular teeth. Maxilliped basis with long seta; endites with two seta and two tubercles. Chelae compact (carpus twice as long as wide). Pereopod-1 with simple setae only. Pereopods 2 and 3 with spiniform setae on carpus and propodus; merus and carpus with setae about as long as half of following article. Pereopods 4 and 5 propodus with seta longer than bifurcated unguis. Pleopod proximal seta separated by a gap from the others. Uropod rami two-articled. Species included: Typhlotanais cornutus G.O. Sars, 1879; ?T. adipatus sensu Tzareva, 1982; ? T. crassus Dojiri & Sieg, 1997, T. andeepae n. sp. Remarks: The short body (less than five times as long as wide), compact chelae (carpus only twice as long as wide), pereopods 4–6 with distal propodus seta longer than bifurcated unguis and two-articled rami of uropods distinguish members of the ‘cornutus’ group from the other typhlotanaids. In the figure by Tzareva (1982, page 53, Fig. 9) Typhlotanais adipatus has a simple unguis on pereopod-4 and uni-articled rami of uropods. The material of Tzareva (1982) is not available for study so it cannot be confirmed whether this character was correctly interpreted. However, if those specimens do have a bifurcated unguis on pereopods 4–6 then the species fits comfortably in the ‘cornutus’ group. A stout body (less than five as long as wide) is characteristic also for typhlotanaids such as T. penicillatus G. O. Sars, 1882, T. spinipes Kudinova-Pasternak, 1982, and Typhlotanais sp. A (present paper) and T. eximius Hansen, 1913, but those species have antennules twice as long as the carapace and the distal propodal seta of pereopods 4 and 5 is shorter than the dactylus.

Typhlotanais andeepae n. sp. (Figs 74–76) Material examined: Holotype: non-ovigerous female, (K 41336), ANT XXII/3, EBS 133-2-S, 62°46.49'– 62°46.38'S, 53°03.50'–53°03.98'W, depth 1594–1579 m, epibenthic sledge, 15 Mar 2005; Paratypes: eight non-ovigerous females (one dissected on slides), the same locality; one non-ovigerous female, (K 41339), ANT XXII/3, PS 67/81-9, 70°32.94'–70° 33.15'S, 14°34.40'–14° 32,74’W, depth 4390–4392 m, Agassiz trawl, 24 Feb 2005; two females, body length 1.8 mm and 1.1 mm, (K 41341), ANT XXII/3, PS 67/74-6, 71°18.35'–71°18.28'S, 13°57.71'–13°57.31'W, depth 1030–1040 m, epibenthic sledge, 20 Feb 2005; one nonovigerous female, (K 41340), ANT XXII/3, PS 151-7-E, 61°45.46'–61°45.34'S, 47°07.57'–47°07.78’W, depth 1181–1188 m, epibenthic sledge, 20 Mar 2005; one female, (K 41338), ANT XXII/3, PS 80-9, 70°39.07'– 70°39.22'S, 14°43.36'–14°43.39'W, depth 3103–3102 m, epibenthic sledge, 23 Feb 2005; two females, (K 41337), ANT XXII/3 PS 88-8-E, 68°03.66'-68°03.61'S, 20°27.90'–20°27.52'W, depth 4929–4931 m, epibenthic sledge, 27 Feb 2005. Etymology: The species is named to acknowledge the ANDEEP Expedition during which the species was collected. Diagnosis: Body 5.3 times as long as wide. Carapace slightly tapering proximally, rounded laterally. All pereonites wider than long, clearly rounded laterally (width: length ratio 3.3, 2.3, 2.0, 1.4, 1.6, and 1.6 respectively). Pleotelson gently rounded. Antennule as long as carapace; article-1 with about five simple setae along margin; antenna article-2 and 3 without spines; cheliped compact; basis reaches the anterior edge of pereonite1, carpus with moderately developed carpal shield, twice as long as wide, slightly longer than chela. Pereopods 1–3 coxa without acute projection, merus without spiniform seta; pereopod-1 propodus with long subdistal seta on ventral margin (over ten times as long as wide); pereopods 2–3 carpus with spiniform seta, merus and carpus with long setae (reaching half of following article); pereopods 4–6 carpus with large round prickly tubercles (at least as long as half of article), unguis tip bifid, propodus distal setae reaching over unguis. Both



pleopod rami with proximal seta separated from others by gap. Uropods rami two-articled; endopod proximal article twice as long as distal. Description: Non-ovigerous female. Body length 2.1 mm (Figs 74A, B), short, 5.3 times as long as wide; carapace smooth, rounded laterally, 1.2 times as long as wide; rostrum weakly pointed; pereonites smooth, wider than long, with rounded lateral margins; pereonite-1 short, 3.3 times as wide as long; pereonite-2 and 3 slightly longer; pereonites 4–6 the longest, 1.5 as wide as long. Pleonites 1–5 similar in size; pleon as long as pereonites 4 and 5 together; pleotelson rounded; caudal projection poorly developed. Antennule (Fig. 75A): Article-1 more than twice as long as wide; 1.5 times as long as articles 2 and 3 combined, with five simple setae and five pinnate setae along article and distally; article-2 half as long as article-3, with two simple setae distally; article-3 with six simple terminal setae. Antenna (Fig. 75B): Article-1 lost during dissection; article-2 twice as long as article-3, with one simple seta; article-3 with one simple seta; article-4 is 1.5 times as long as article-5, with three simple and four pinnate setae distally; article-5 with simple seta; article-6 with five simple subterminal and terminal setae. Mouth parts: Labrum (Fig. 75C) hood-shaped, covered by numerous minute setae. Mandible (Figs 75D, E) stout; molar process well-developed, with strongly crenulated edges; lacinia mobilis well-developed, crenulated. Maxillule (Fig. 75F) endite with seven terminal setae (two bifurcated); palp lost during dissection. Maxilla lost during dissection. Maxilliped (Fig. 75H) bases little longer than wide; each with simple setae reaching over endite; each endite armed with two plumose setae and two tubercles on distal margin; palp article-1 naked; article-2 wedge-shaped, with three setae on inner margin and one short, simple seta on outer margin; article-3 trapezoidal, with four setae on inner margin; article-4 with one simple seta on outer margin and five terminal setae. Labium (Fig, 75 G) with group of setae on outer corner of inner lobe; outer lobe with small setae. Cheliped (Fig. 75I): Basis 1.5 times as long as wide, with one simple seta on outer margin, posterior margin rounded; merus wedge-shaped, with seta on ventral margin; carpus twice as long as wide (length:width ratio 2.0), with two simple setae and one minute seta ventrally and two short setae dorsally; carpal shield moderately developed; chela almost as long as carpus, twice as long as wide; fixed finger with three setae on inner margin and with two simple setae ventrally; dactylus slightly curved, with two spiniform setae on inner margin and with one short seta proximally. Pereopod-1 (Fig. 76A): Of walking type; coxa with one simple setae; basis as long as merus, carpus and propodus combined, with three proximal setae dorsally; ischium short with one simple seta; merus almost as long as carpus but shorter than propodus, with one short and one long setae distally; carpus with two long and one short setae distally; propodus with three subdistal setae dorsally and one long subdistal simple setae ventrally; unguis twice as long as dactylus; unguis and dactylus little shorter than propodus. Pereopod-2 (Fig. 76B): Of walking type; basis slightly shorter than rest of articles combined, with three proximal setae dorsally; ischium with one seta; merus almost as long as carpus with three long setae distally (reaching half of carpus); carpus with two long, one short and small spiniform seta distally; propodus with two subdistal setae dorsally and one spiniform seta ventrally; dactylus shorter than unguis. Pereopod-3 (Fig. 76C): Walking type; basis a little shorter than rest of articles combined, with three proximal setae dorsally; ischium with one seta; merus with one long seta distally; carpus with two long and two short setae and one spiniform seta distally and with a few combs of spines ventrally; propodus with one subdistal setae reaching over end of dactylus and one spiniform seta on ventrally; dactylus shorter than unguis. Pereopod-4 (Fig. 76D): Of clinging type; basis wide, with three simple setae; ischium with two setae; merus almost as long as carpus, with two subdistal spines and numerous microtrichae on ventral margin; carpus with hooks distally, one sensory seta dorsally and rounded, large (at least half as long as article) prickly tubercles ventrally; propodus 1.7 times as long as carpus, with one pinnate seta dorsally, two spiniform setae ventrally and one serrated distal seta reaching over unguis; dactylus and unguis half as long as propodus, unguis with bifid tip.



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Pereopod-5 (Fig. 76E): Similar to pereopod-4. Pereopod-6 (Fig. 76F): Similar to pereopod-5, but propodus with three distal setae.

FIGURE 74. Typhlotanais andeepae n. sp., holotype, female. A) lateral view; B) dorsal view. Scale = 0.1 mm



FIGURE 75. Typhlotanais andeepae n. sp., female paratype. A) Antennule; B) Antenna; C) Labrum; D) Left mandible; E) Right mandible; F) Maxillule; G) Labium; H) Maxilliped; I) Cheliped. Scale: A, B, I = 0.1 mm; C–H = 0.01 mm.

Pleopods 1–5 (Fig. 76G): All pleopods similar; exopod outer margin with ten plumose setae, inner margin with one plumose seta; endopod with thirteen plumose setae on outer margin; both exopod and endopod with small gap between most proximal seta and remind setae. Uropod (Fig. 76H): Basal article shorter than endopod proximal article; exopod and endopod two-articled; exopod little shorter than endopod; exopod proximal article almost half as long as distal article, with one simA REVISION OF FAMILY TYPHLOTANAIDAE SIEG 1984


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ple seta distally; article-2 tipped by two setae (short and long); endopod proximal article twice as long as distal one, with two simple setae distally; distal article with five terminal simple setae.

FIGURE 76. Typhlotanais andeepae n. sp., female paratype. A) Pereopod-1; B) Pereopod-2; C) Pereopod-3; D) Pereopod-4; E) Pereopod-5; F) Pereopod-6; G) Pleopod; H) Uropod. Scale = 0.1 mm.



Distribution: Antarctic: Eastern Weddell Sea, east of Elephant Island and the Orkney Islands at depths of 1030 to 4931 m. Remarks: Typhlotanais andeepae n. sp. has two bifurcated terminal spiniform setae on the maxillule. This character has been observed before in a few abyssal tanaidacean species such as Collettea minima Hansen, 1913, Arthrura andriashevi Kudinova-Pasternak, 1966, Curtichelia expressa Kudinova-Pasternak, 1987, Parafilitanais mexicana Larsen, 2002, Cetiopyge mira Larsen & Heard, 2002, Isopodidus janum, Larsen & Heard, 2002, and Collettea lilliputa, Błażewicz-Paszkowycz & Larsen 2005, representing various Paratanoidea families and emphasizing the polyphyletic character of this feature (Larsen, 2002). Typhlotanais andeepae is most similar to T. cornutus from the North Atlantic, sharing a cluster of characters listed in the diagnosis of the group. The new species can be distinguished from T. cornutus by having a carapace longer than wide (length:width ratio 1.2) and pereonite-4 only 1.5 times as wide as long. The carapace of T. cornutus is clearly shorter than wide (width: length ratio 0.83) while pereonite-4 is over twice as wide as long. A third species which shares the characters listed above (with exception of one-articled uropod rami) with T. cornutus and T. andeepae is Typhlotanais (=Peraeospinosus) adipatus sensu Tzareva, from shallows off the Adelie Islands. Because the type material of T. adipatus sensu Tzareva is no longer available any conclusions about its similarity to T. cornutus and T. andeepae can only be based on Tzareva’s drawings—Fig. 9, page 53. However, it is almost certain that Typhlotanais adipatus sensu Tzareva and sensu Sieg, 1982 represent two different taxa. Tzareva’s species is compact (about five times as long as wide), in contrast to Sieg’s species that is almost seven times as long as wide, has short distal setae on propodus of pereopods 4–5, and dense minute setation in antenna articles 2–4 that is lacking in T. cornutus and T. andeepae.

‘plicatus’ group Diagnosis: Pereonites 1–3 with minute transversal ridges. Species included: Typhlotanais longimanus, Dollfus, 1897; Typhlotanais plicatus Kudinova-Pasternak, 1993, ?T. variabilis Hansen 1913 Remarks: Typhlotanais longimanus and T. plicatus share the unique character of a corrugated cuticle on the first three pereonites. The third species, T. variabilis, has this character weakly developed (G. Bird. pers. obs.). The material obtained for this study was the poorly preserved type material of T. longimanus and T. variabilis, and one manca of T. plicatus. This slender material precluded dissecting any specimen and establishing a reliable set of characters for this morpho-group. Nevertheless, the corrugated pereonites 1–3 is a feature specific enough for provisionally distinguishing the ‘plicatus’ morpho-group.

Typhlotanais plicatus Kudinova-Pasternak, 1993 (Fig. 77) Typhlotanais plicatus: Kudinova-Pasternak (1993) 127: 142–143.

Material examined: one manca (K 41419), body length 1.5 mm, ANT XXII/3, PS 67/102-13, 65°34.32'– 65°34.40'S, 36°31.32'–36°31'W, depth 4805–4803 m, epibenthic sledge, 6 Mar 2005. Diagnosis: Pereonites 1–3 corrugated. Carapace oval, 1.3 times as long as wide. Antennule article-3 3.5 times as long as article-2. Distribution: The specimen collected during ANDEEP is the second record of the species. The first record was by Kudinova Pasternak (1993) in the Weddell Sea (62°36.4'–62°36.4'S, 15°28.1'–15°32.7'W).



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Remarks: T. plicatus and T. longimanus share a peculiar character – a corrugated pereonites 1–3. The species however can be distinguished by comparing the carapace, which is oval and 1.4 times as long as wide in T. plicatus, while it is rectangular and almost as long as wide in T. longimanus. More conspicuous differences are seen in article-1 of the antenna which is six times as long as wide in T. longimanus but only 3.5 times as long as wide in T. plicatus.

FIGURE 77. Typhlotanais plicatus Kudinova-Pasternak, 1993, manca, lateral view.

‘eximius’ group Diagnosis: Body short, less than five times as long as wide. All pereonites wider than long. Antennule twice as long as carapace. Cheliped elongated (carpus four times as long as wide). Species included: Typhlotanais eximius Hansen, 1913; T. spinipes Kudinova-Pasternak 1982; Typhlotanais sp. A. Remarks: The species included in this group share characters listed in the diagnosis. Because of the scarcity or unavailability of the material, further analysis to justify the erection of this (‘eximus’) group is not possible at the moment. Such quandaries explain the question marks ahead of the names above.

Typhlotanais sp. A (Fig. 78) Material examined: one non-ovigerous female, body length 1.5 mm ANT XXII/3, PS 81-9, 70°32.94'– 70°33.15'S, 14°34.40’–14°32,74'W, depth 4390–4392 m, Agassiz trawl, 24 Feb 2005. Diagnosis: Body compact, short, 4.5 times as long as wide; carapace 1.3 times as wide as long; cheliped basis reaches pereonite-1 ventrally; pereonite-1 short (almost four times as wide as long), slightly wider than carapace in dorsal view; pleon 25% of total body length; antennule twice as long as carapace. Description: Non-ovigerous female (Fig. 78) body length up to 1.5 mm, 4.5 times as long as wide. Carapace smooth, rounded laterally, 15% of overall length, 1.3 times as wide as long; rostrum pointed; pereonites smooth rounded laterally; cheliped basis reaches pereonite-1 ventrally; pereonite-1 four times as wide as long, little wider than carapace; pereonite-2 longer than pereonite-1, about three times as wide as long; pereonite-3 longer than pereonite-2, about twice as wide as long; pereonite-4 twice as wide as long but slightly narrower than pereonite-3; pereonite-5 a little longer than pereonite-4, 1.7 times as wide as long; pereonite-6 three



times as wide as long. Pleon 26% of body length, somewhat wider than pereonites 4–6, pleonites 1–5 similar in size.

FIGURE 78. Typhlotanais sp. A. female, A) dorsal view; B) lateral view; C) carapace ventral side. Scale = 0.1 mm.



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Distribution: Eastern Weddell Sea, at depths of 4390–4392 m. Remarks: This species was represented by only one specimen and it is not formally described here owing to the lack of material. The similar Typhlotanais penicillatus described by Sars (1882) from shallow waters of North Atlantic differs from Typhlotanais sp. A in having a carapace about as long as wide and a shorter antennule, 1.3 times as long as the carapace. A similar compact body and long antennule are observed in T. eximius (Fig. 79), from the abyssal of the South-West Iceland, North Atlantic, but that species has long cheliped which is almost four times as long as wide, while it is only 2.5 times as long as wide in both Typhlotanais sp. A and T. penicillatus.

FIGURE 79. Typhlotanais eximius Hansen, 1913, holotype, female. A) lateral view; B) dorsal view. Scale = 0.1 mm.



A short body habitus is also characteristic for T. solidus (Fig. 80), although its relatively short antennule (little longer than carapace) suggests it may constitute a valid morpho-group. Because the species is known from the holotype specimen only that is covered by crystals, it is difficult to indicate its similarity to the others short-body morpho-groups at the moment.

FIGURE 80. Typhlotanais solidus Hansen, 1913, female holotype. Scale = 0.1 mm.



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TABLE 2. List of the species determined to have invalid world-wide distribution Species

Reference

Chaulipleona armata (Hansen, 1913)

Guerrero-Kommritz, 2004

Meromonakantha macrocephala (Hansen, 1913)

present study

Paraleptognathia gracilis (Kryer, 1842)


Paraleptognathia multiserrata (Hansen,1913)


Peraeospinosus kergulenenesis (Beddard, 1886)

Błażewicz-Paszkowycz, 2005

Peraeospinosus peculiaris (Lang, 1968)

Błażewicz-Paszkowycz, 2005

Tube building ability of nototanaids and typhlotanaids The clinging type of pereopods are the most evident feature that distinguishes typhlotanaids from nototanaids. Those appendages have strongly widened basis (length: width ratio ca.2–3) and have reduced setation that is limited only to one distal seta on the propodus of pereopods 4–5 and three setae on pereopods 6. In place of setation, the clinging appendages are supported by various hooks, tubercles, thorns and spines (clinging apparatus) on the merus and carpus, while the shortened unguis is bifurcated (e.g. T. aequiremis, Torquella) or trifurcated (Hamatipeda). These appendages are held bent upward, rather close to the body, and in a position inside the tube against the tube-wall. This clinging apparatus undoubtedly enables the animal to hold the inside the tube firmly, but it is probably also an adaptation for efficient spinning of the tube. That may explain why we know more about the tubes of typhlotanaids than those of other genera (Hassack & Holdich, 1987). Primarily tanaidomorphs were considered all tube-dwellers (Sieg, 1983a). Later, because for many species tubes has never been recorded, numerous tanaidomorphs were considered free-living forms (Greve, 1967, Hassack & Holdich, 1987). Observations made on samples taken by SCUBA in shallow muddy postglacial coves (Admiralty Bay, South Shetland Islands) revealed aggregations of long and tangled corridors made of soft sediment particles inhabited by Nototanais antarcticus (Hodgson 1902). Although the corridors had a regular lining inside, they were flimsy structures which easily disintegrated while washing or sieving material. Dwelling in rigid corridors within sediments is an alternative behavior to tube-construction that may explain why tubes has been never found for tanaids with spinning glands (Larsen, 2005).

Remarks on world-wide Tanaidacea distribution in deep-water The hypothesis of a high local endemism in contrast to reports of wide distribution of tanaidaceans has been discussed by Larsen (2005). The mechanism of dispersal by tube-dwelling tanaidomorphs having demersal and non-dispersive larva is not fully understand. The world distribution of tanaidaceans suggests more dispersal capacity of deep-sea species than shallow-water species, despite the limited influence of rafting via human invention in the abyssal environment (Larsen, 2005), although historical aspect of the distribution should be considered (Bamber & Sheader, 2003). Attempts at explaining the wide deep-sea distribution of tanaidaceans involve numerous biological (inbreeding, food supply, predation and mortality) or abiotic factors (topography, currents, temperature, salinity, hydrostatic pressure, upwelling and benthic storms). Far more likely than all of those factors is misidentification of species (Larsen, 2005). The poor recognition of the tanaidaceans, the reduction of setation (as a consequence of the tubicolous life style), the presence of cryptic or sibling species that can be sympatric (Larsen, 2005) has made misidentification a key problem in the apparent wide distribution of tanaidaceans. Morphological analysis of the material seems to identify misidentification of some tanaidomorphs as the cause for the apparent wide distribution of the species (Table 2). Deep-sea



tanaidacean species (mostly surface-living apseudomorphs and neotanaidomorphs) can be widely distributed by near-bottom currents (Reidenauer & Thistle, 1985) although mortality of relatively fragile tanaidaceans during such rafting is likely to be high (Larsen 2005). Nevertheless, the main oceanic currents could potentially explain a wide distribution of species within one basin (e.g. North/South Atlantic or Circumantarctic distribution), but still a putative distribution of species in both North Pacific and West Antarctic is questionable.

Acknowledgements I am grateful to Prof. Angelika Brandt from the Zoologisches Institut und Museum der Universität Hamburg (Germany) for access to the material collected during the ANDEEP Expeditions. Material for this study was also loaned from the Zoological Museum University Copenhagen (Denmark), the Zoological Museum, University of Lomonosow (Moscow, Russia), the National Museum of Natural History, the Smithsonian Institute (Washington DC), Icelandic Museum of Natural History (Reykjavic) and the Museum of Oceanography (Monaco). I am especially grateful to Dr Roger Bamber (Natural History Museum, London) for improoving English through the manuscript and to Dr Graham Bird for critical notes and productive discussion of the subject. Financial support was provided by the Ministry of Science and Higher Education (Grant 2PO4C 089 29/ 2005) and by the EU SYNTHESYS Project.

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