Invertebr . Taxon., 1994. 8. 683-747 Review of the Family Janiridae (Crustacea :Isopoda :Asellota) George D . F. WilsonA and Johann- Wolfgang Wagele A Australian Museum. P O Box A285. Sydney South. NSW 2000. Australia Universitat Bielefeld. PO Box 8640. D-4800 Bielefeld 1. Germany . Contents . Page Abstract ................................................................................................................684 Introduction ........................................................................................................... 684 Literature Sources and Methods .................................................................................. 685 Systematics ................................................................................................................ 685 Family Janiridae G . 0. Sars. 1897a ......................................................................... 685 History of the Janiridae ..........................................................................................685 Problems in the Classification of the Janiridae ............................................................ 686 Composition of the Janiridae ...............................................................................687 Adjustments to the composition of the Janiridae .........................................................688 Key to janiroidean genera once classified as 'janirids' ........................................................689 Genera of the Janiridae ............................................................................................. 692 692 Austrofilius Hodgson. 1910 ...................................................................................... Caecianiropsis Menzies & Petit. 1956 .........................................................................693 Caecijaera Menzies. 195 l a ....................................................................................... 694 Carpias Richardson. 1902 .....................................................................................695 Ectias Richardson. 1906 ..................................................................................... 697 Hawaianira Miller. 1967 .........................................................................................698 Heterias Richardson. 1904a ...................................................................................... 699 Iais Bovallius. 1886 ................................................................................................ 700 Ianiropsis Sars. 1897a ............................................................................................. 702 Iathrippn BovaIlius. 1886 .......................................................................................704 Jaera Leach. 18 14 .................................................................................................. 706 Janaira Moreira & Pires. 1977b ................................................................................708 709 Janira Leach. 1814 ................................................................................................. Janiralata Menzies. 195 1b ........................................................................................ 711 712 Remarks on the 'Janiralata' Complex ................................................................... Janthura Wolff. 1962 ..........................................................................................714 Mackinia Matsumoto. 1956 ....................................................................................715 Microjaera Bocquet & LCvi. 1955 .............................................................................. 716 Microjanira Schiecke & Fresi. 1970 ........................................................................... 717 Neojaera Nordenstam. 1933 ................................................................................ 718 Protocharon Chappuis. Delamare-Deboutteville & Paulian. 1956 .....................................719 Family Microparasellidae Karaman. 1934 .......................................................................720 Provisional Diagnosis .............................................................................................. 720 History and Composition of the Microparasellidae ...................................................... 720 Genera of the Microparasellidae ................................................................................... 722 Angeliera Chappuis & Delamare. 1952 ....................................................................... 722 Microcharon Karaman. 1934 ...................................................................................723 Microparasellus Karaman, 1933 ................................................................................724 Paracharon Coineau 1968 .................................................................................725 . 684 G. D. F. Wilson and J.-W. Wagele Contents (continued) Janiroidea incertae sedis ............................................................................................... 726 726 Tole Ortmann, 1901 ............................................................................................... 728 Jaerella Richardson, 1911b ....................................................................................... 728 Rhacura Richardson, 1908 ....................................................................................... 729 Trichopleon Beddard, 1886a .................................................................................... Xostylus Menzies, 1962a .......................................................................................... 730 731 Acknowledgments ....................................................................................................... References ................................................................................................................. 731 Abstract The family Janiridae is reviewed, primarily using the published literature, as a step for revising the systematics of the Asellota. The history of janirid taxonomy is reviewed, and all taxa that have been attributed to the family are listed and discussed. A new composition for the family is provided with synonymies and references to all genera. The genera included in the Janiridae are: Austrofilius Hodgson, Caecianiropsis Menzies & Petit, Caecijaera Menzies, Carpias Richardson, Ectias Richardson, Hawaianira Miller, Heterias Richardson, Zais Bovallius, Zaniropsis Sars, Zathrippa Bovallius, Jaera Leach, Janaira Moreira & Pires, Janira Leach, Janiralata Menzies, Janthura Wolff, Mackinia Matsumoto, Microjaera Bocquet & LCvi, Microjanira Schiecke & Fresi, Neojaera Nordenstam, Protocharon Chappuis, DelamareDeboutteville & Paulian. A key is provided for identification of genera belonging to the Janiridae and related families. Generic compositions are revised where necessary, and diagnoses, lists of species and most references are provided for each genus discussed. The composition of the Microparasellidae is discussed and a provisional diagnosis is given. The Microparasellidae is limited to the following genera: Angeliera Chappuis & Delamare, Microcharon Karaman, Microparasellus Karaman, Paracharon Coineau. Some genera are excluded from the Janiridae and temporarily left as family incertae sedis: Tole Ortmann (=Zolella Richardson), Jaerella Richardson, Rhacura Richardson, Trichopleon Beddard, Xostylus Menzies. Introduction Isopods of the superfamily Janiroidea Sars, 1897a (suborder Asellota) are ecologically important and biogeographically interesting. Systematic knowledge on most janiroidean families, however, is extremely limited and often superficial. The Asellota have been the subject of several major monographs and numerous smaller papers (papers prior to 1960 reviewed by Wolff 1962). In the last three decades, great interest in the Janiroidea has been generated by the discovery of its high diversity of both species and morphological types in the deep sea (Menzies 1962a; Wolff 1962; Birstein 1963a; Hessler and Sanders 1967; Hessler 1970; Hessler et al. 1979). This interest has resulted in the rapid accumulation of new species and genera described from deep-sea samples taken since the early 1960s. Continued sampling, especially in the tropics, has yielded a steady supply of new shallowwater taxa. Despite this new information, few major reorganisations of the suprageneric taxa have been attempted since the monographs of Wolff (1962) and Menzies ( 1 9 6 2 ~ '1962b). The family-level groups have become poorly defined as they have been forced to include a broad variety of taxa. Unusual taxa have been placed in monotypic families often because sufficient comparative information was unavailable. The family Janiridae is especially in need of revision. This family occupies a central position in the understanding of the approximately 22 families in the Janiroidea, because the deep-sea forms are probably derived from a janirid-like ancestor (Kussakin 1973, 19796; Hessler et al. 1979). The Janiridae, however, are not the earliest-derived janiroideans (Wilson 1987). Therefore, a systematic revision of this family is urgent, because it is not possible to define the sister group of the Janiridae o r to understand the origin of the 'higher Janiroidea' sensu Wilson (1987). This paper reviews the genera that have been included in the Janiridae by Wolff (1962) and later authors, with the ultimate goal of revising the family's composition and definition. Unfortunately, available information cannot allow a complete revision of the family. Review of the Janiridae 685 Although we offer suggestions for reclassifying the janirid genera and related families, more study of individual taxa is necessary before this important assemblage of shallow-water Janiroidea can be better understood. Therefore, our goal in this paper is to provide a thorough listing of the genera, species and literature for future revisions. For this reason, we have not provided illustrations of each genus. Most genera would benefit from reillustration of their type species, but this is well beyond the scope of a single paper because so many taxa are involved. A phylogenetic analysis of janirid genera is presented in Wilson (1994) t o demonstrate those difficulties that remain. Literature Sources and Methods A systematic review such as this would not be possible without access to a great deal of literature. Literature information prior to 1975 was obtained from Torben Wolff's taxonomic catalogue. Morphological information on janirid genera was derived from photocopy catalogues maintained separately by both authors. Literature since 1978 was checked by computerised searches of Zoological Record, BIOSIS, and Current Contents databases. Specimens from the Los Angeles County Museum and the Australian Museum were used to verify the morphology of many genera, especially Janiralata, Janira, Zais and lathrippa. The generic diagnoses are not strictly limited to diagnostic characters, but also include the data used in the phylogenetic analysis (Wilson 1994). The generic synonymies include only the first instance of a synonymous name, and a separate section, 'Additional References', provides additional later instances of both synonymous names as well as other generic names of species now of a particular genus. The generic synonymies and associated additional references provide most taxonomic and related references on the species of each genus, although in some cases, especially Jaera, not all references are included. In listings of species in each genus, we often have raised subspecies to the species rank using a phylogenetic species definition. The subspecies rank was not used unless some evidence of introgression between subspecies or other genetic information was available (Jaera is best known in this regard). Although this paper is taxonomic in content, we have included discussion of biology or biogeography in the Remarks section of genera where interesting topics have arisen recently. Systematics Superfamily JANIROIDEA Sars, 1897a Family JANIRIDAE Sars, 1897a Ianiridae Sars, 1897a: 98 (Janiridae misspelled).-Hurley, 1961: 262; Menzies, 1962a: 181; 19626: 69; Jang and Kwon, 1990: 193-4. Janiridae. -Ohlin, 1901: 30; Richardson, 1905b: 448; Stebbing, 1905: 48; Nierstrasz and Stekhoven, 1930: 116; Gurjanova, 1932: 20; 1933: 397; 1936b: 39; Hatch, 1947: 171; Wolff, 1962: 32, 33-40, 215, 220, 282, 283, 295, 300; Gruner, 1965: 121; Miller, 1967: 187; Schultz, 1969: 250; Kensley and Schotte, 1989: 81-82. Parasellidae (pars) Hansen, 1905: 315-16. Jaeridae Stebbing, 1910b: 224.-Monod, 1925: 238. Ianirini Hansen, 1916: 12.-Monod, 1926: 13; Nordenstam, 1933: 172; Nierstrasz, 1941: 282; Menzies, 1951b: 123; Hurley, 1957: 17. Microjaniridae Bocquet and Lkvi, 1955: 117. - Birstein, 1961 : 137. History of the Janiridae Sars (1897a: 98) defined the family Ianiridae for the genus Ianira (original spelling Janira) and included the genera Ianthe (junior synonym of Tole), Ianiropsis and Zaera (correctly spelled Jaera). Sars and later authors adopted the unfortunate tendency of exchanging 'J' with 'I' or vice versa in naming the genera and the family (Wolff 1965), despite their correct original spellings. Because Janira Leach, 1814 is the correct spelling of the type of the family, its name must be Janiridae, although the authorship still resides with Sars, 1897a. The period between 1886 to 1962 saw the addition of 34 genera (see list below) and many new species t o the Janiridae; some like Santia (previously Antias: Sivertsen and Holthuis 1980) and Joeropsis (correct spelling for Jaeropsis: ibid.) were removed to other families before 1962. During this time, the genus Janira received a great burden of different forms. 686 G. D. F. Wilson and J.-W. Wkigele Hansen (1916) subsumed all of Sars' families into the Parasellidae (a family-level nomen nudum because Parasellus is not an available name) and reduced the Janiridae to the tribe 'Ianirini'. This was followed by Nordenstam (1933), who added new family-group names such as the Jaeropsini (correctly spelled Joeropsidini: see Sivertsen and Holthuis 1980). Menzies (1956, 1962b) recognised the distinctness of the Janiridae, and created several new families, Janirellidae and Acanthaspidiidae, to contain genera previously classified as janirids. Taxonomists working with the interstitial isopods created new families or subfamilies to contain various dwarfish janirid-like isopods: the Microparasellidae Karaman, 1934, and the Microjanirinae Bocquet & LCvi (1955) of the family Janiridae, later elevated to family status by Birstein (1961). The first comprehensive review of the Janiridae and these related families was Wolff (1962), where Janira was pared down from over 30 species to 8 species. Wolff's definition of the Janiridae (see below) included a great deal of diversity, and subsumed several wholly or primarily deep-sea families. Later workers (Menzies and Schultz 1968; Kussakin 1973; Hessler and Thistle 1975; Bowman and Abele 1982) have not used his composition of the family, although no one has provided a useful definition of the Janiridae. Miller (1967) followed Wolff's definition in adding Hawaianira to the family. Recently, a definition of the Janiridae even broader than Wolff's has been proposed (Kussakin 1988) that includes the Thambematidae Birstein, 1963a and the Acanthaspidiidae. Because of the broad definition for the Janiridae, unusual genera such as Katianira Hansen, 1916, Xostylus Menzies, 1962a, or Dactylostylis Richardson, 1911a are still considered part of the family or have been removed only recently (e.g. Katianira to the Katianiridae Svavarsson, 1987). After 1962, 18 genera either have been brought out of synonymy or were newly created. Some are junior synonyms of older taxa, and others belong in other families. Problems in the Classification of the Janiridae Several problems that recur in janirid systematics are evident in Wolff's (1962) diagnosis of the family, reproduced here with italics added to the variable parts. 'Body oblong, depressed. Head and pereonites free, with the lateral parts lamellarly expanded in most genera and pereonites subequal in length. Pleon usually consists of two segments; when present, the first is narrow and inconspicuous, the second large and shieldshaped. Eyes often present, situated on the upper surface. Antennae almost always longer than antennulae, with six joints in peduncle and with squama on joint 3 generally well developed. Mandibles with palp, and normally with well developed molar process. Palp of maxillipeds usually with joint 3 in particular, expanded; its width almost always equal to that of the endite or broader. First pair of pereopods usually more or less prehensile; the remainder are walking legs of moderate and almost equal length. Dactylus almost always with two or three claws. Uropods always on the lateral or the terminal margin, with peduncle, and biramous or, occasionally, uniramous.' This diagnosis defines a generalised janiroidean or asellote isopod (e.g. Hessler et al. 1979) using largely plesiomorphic characters. All defined character states, with their built-in variation, can be found in most families and therefore are phylogenetically uninformative. These characters are repeatedly used to define other janiroidean families by stating any derived character states and filling out the definition from the characters above. Our analysis of the Janiridae was obstructed by the absence of descriptions of useful characters in many species (e.g. the fourth and fifth pleopods: Menzies and Frankenburg 1968). Some taxa have been traditionally illustrated with a few limb parts to describe new species or genera, especially in the interstitial isopod literature. Such descriptions, while minimally sufficient for parochial taxonomic purposes, are nearly useless for higher-level considerations, and are difficult to use for field workers. Specialists should illustrate aN limbs and should show the whole organism in several aspects. Missing from practically all illustrations of Janiridae are lateral and ventral views of various parts of the body, especially the cephalon and pleotelson. Review of the Janiridae 687 Other, possibly useful, characters have been only recently discovered. The spermathecal duct ( =cuticular organ), a female copulatory organ currently known only from the Asellota (Veuille 1978b; Wilson 1986b, 1987, 1991), is a prime example. For example, the spermathecal duct reported for Haploniscidae (Wolff 1962) differs from that in the Dendrotiidae (Lincoln and Boxshall 1983). Research on the swimming family Munnopsididae (Wilson and Hessler 1980, 1981; Wilson 1985, 1989) shows that the complex morphology of the head may also provide many useful characters. Head morphology is largely ignored in most descriptions of Asellota. Composition of the Janiridae The following list contains the names of all genera that have been included into the Janiridae under the broad definitions of Wolff (1962) or Kussakin (1988). Senior synonyms and taxa moved to other families are indicated in parentheses ('='indicates a senior synonym, and ':' indicates the reference establishing the synonymy or transferring the genus to another family). Abyssijaera Menzies, 1962a ( = Katianira: Katianiridae Svavarsson, 1987) Acanthaspidia Stebbing, 1893 (Acanthaspidiidae: Menzies and Schultz 1968) Angeliera Chappuis & Delamare-Deboutteville, 1954 (Microparasellidae: Coineau 1986) Austrofilius Hodgson, 1910 Austroniscoides Birstein, 1963b ( = Janthura: Menzies and George 1972) Bagatus Nobili, 1906 ( = Carpias: Bowman and Morris 1979) Caecianiropsis Menzies & Pettit, 1956 Caecijaera Menzies, 195l a Carpias Richardson, 1902 Dactylostylis Richardson, 191l a (Janirellidae: as Spinianirella Menzies, 1962a) Ectias Richardson, 1906 Fritzianira De Castro & Lima, 1977 (subgenus of Heterias: Bowman et al. 1987) Hawaianira Miller, 1967 Heterias Richardson, 1904a Zais Bovallius, 1886 Zaniroides Kensley, 1976 (=Ectias: Kensley 1978) Zaniropsis Sars, 1897a Zanisera Kensley, 1976 ( =Neojaera: Kensley 1984c and herein) Zanthopsis Beddard, 1886b (Acanthaspidiidae: Menzies and Schultz 1968) Zathrippa Bovallius, 1886 Zolella Richardson, 1905b ( = Tole, family incertae sedis: herein) Jaera Leach, 1814 Jaerella Richardson, 1911b (family incertae sedis: herein) Janaira Moreira & Pires, 1977b Janatus Carvacho, 1983 (=Carpias: herein) Janira Leach, 1814 Janiralata Menzies, 1951b Janirella Bonnier, 1896 (Janirellidae Menzies, 1956) Janthura Wolff, 1962 Janthurella Kussakin, 1982 ( =Katianira: Katianiridae Svavarsson, 1987) Jehaia Wagner, 1990 ( = Protocharon: herein) Katianira Hansen, 1916 (Katianiridae Svavarsson, 1987) Mackinia Matsumoto, 1956 Microcharon Karaman, 1934 (Microparasellidae Karaman, 1934) Microjaera Bocquet & LCvi, 1955 Microjanira Schiecke & Fresi, 1970 Microparasellus Karaman, 1933 (Microparasellidae Karaman, 1934) Microprotus Richardson, 1909 (Munnopsididae: Wilson et al. 1989) Microthambema Birstein, 1961 (Thambematidae: Harrison 1987) Natalianira Kensley, 1984b (Katianiridae: Svavarsson 1987) 688 G. D. F. Wilson and J.-W. Wagele Neoectias Winkler, 1992 ( = Zais: herein) Neojaera Nordenstam, 1933 Paracharon Coineau, 1968 (Microparasellidae: Coineau 1986) Protocharon Delamare-Deboutteville & Chappuis, 1956 (Microparasellidae: Coineau 1986) Protojanira Barnard, 1927 (Protojaniridae: Sket 1982) Pseudasellus Chappuis, 1951 ( = He{erias: Williams 1968) Pseudojanira Barnard, 1925 (Pseudojaniridae Wilson, 1987) Rhacura Richardson, 1908 (family incertae sedis: herein) Spinianirella Menzies, 1962a (=Dactylostylis: Hessler 1968) Tole Ortmann, 1901 Thambema Stebbing, 1912 (Thambematidae: Harrison 1987) Trichopleon Beddard, 1 8 8 6 ~(family incertae sedis: herein) Vermectias Sivertsen & Holthuis, 1980 (Vermectiadidae Just and Poore, 1992) Xostylus Menzies, 1962a (family incertae sedis: herein) Adjustments to the Composition of the Janiridae We here limit the Janiridae to a smaller group of genera. The following paragraphs indicate the current placement of genera removed from the janirids. These adjustments are based on reasons external to the definition of the Janiridae, such as the revision of a previously included family. Microprotus, despite its complete lack of swimming pereopods, is a derived member of the Munnopsididae sensu lato (Wilson 1989; Wilson et al. 1989). Its closest relative in the Munnopsididae seems to be Storthyngura Vanhoffen, 1914 (ibid.). The genera Abyssijaera, Janthurella, Katianira and Natalianira have been removed to the new family Katianiridae by Svavarsson (1987), and are reduced to two genera. Katianira now contains the species of Janthurella and Abyssijaera. Natalianira is retained as a valid genus of the Katianiridae. Protojanira and Pseudojanira have been removed from the Janiroidea. Protojanira is placed in its own family with the genera Enckella Fresi, Idato & Scipione, 1980 and Anneckella Chappuis & Delamare-Deboutteville, 1957. The Protojaniridae is considered to belong to either the superfamily Protojaniroidea (see Sket 1982; Wagele 1983) or the Gnathostenetroidoidea (see Wilson 1987). Pseudojanira was placed in its own monotypic family and superfamily (Wilson 1986b, 1987). Poore and Just (1990) confirm the concepts of the Pseudojaniridae and the Pseudojaniroidea with the description of abundant material of a new species, P. investigatoris from Bass Strait. The family Microparasellidae Karaman, 1934, has continued to be recognised (Birstein and Ljovushkin 1965a, 1965b; Coineau 1968, 1969, 1986), despite its elimination by Wolff (1962). We discuss this family below. The genera of Microparasellidae are limited to Microparasellus, Angeliera, Microcharon and Paracharon (but see Coineau 1969, 1986). Janirella and Dactylostylis (senior synonym of Spinianirella; see Hessler 1968) belong to the Janirellidae Menzies, 1956, following the composition of family of Menzies (1962a, 1962b). We, however, exclude the genus Rhacura from the Janirellidae until this genus can be reillustrated and revised. These genera have synapomorphies that clearly separate them from the Janiridae (see also outgroup discussion in Wilson 1989), so their classification in this family by Kussakin (1988) is not used here. The family Acanthaspidiidae Menzies, 1962a is currently recognised (Bowman and Abele 1982; Brandt 1991). Menzies and Schultz (1968), who added several new genera to it, offered no arguments rebutting Wolff's (1962) removal of the family, although Brandt's (1991) treatment places the family on firmer ground. Therefore, we do not follow Kussakin (1988) who included Acanthaspidia into the Janiridae. Acanthaspidiids can be defined as janiroideans that have enlarged pereonal lappets, narrow or finger-like mandibular molars, broad maxillipedal endites with narrow palps, third pleopods with many plumose setae on both rami, and elongate uropodal sympods. Most species of this family also have dorsal spines. The family now contains Acanthaspidia and Zanthopsis (see Brandt 1991), although the latter heterogeneous taxon would benefit from a thorough revision. Zolanthe Beddard, 1886a, Paracanthaspidia Menzies & Schultz, 1968, and Exacanthaspidia Menzies & Schultz, 1968 were synonymised with Acanthaspidia by Brandt (1991). Review of the Janiridae 689 The Thambematidae Stebbing, 1912, including Thambema and Microthambema, is a welldefined family (Harrison 1987). Consequently, we do not follow Kussakin (1988) in including this family into the Janiridae. Jaerella, Rhacura and Tole (senior synonym of Zolella) may have characters in common but are assigned nevertheless to incertae sedis owing to their poor descriptions. Trichopleon and Xostylus have no place in the Janiridae: their dactylar claw form, anal position and third pleopod exopod are all characteristic of more-derived deep-sea genera. These two genera need revision before their exact affinities can be resolved, so they are temporarily assigned to incertae sedis. Vermectias has been recently placed into its own family, Vermectiadidae by Just and Poore (1992), because this genus has been found to have non-janiroidean pleotelson, pleopods and penes. The male copulatory morphology of Vermectias might show some relationship to genera of the Microparasellidae, specifically Angeliera (see Just and Poore 1992). Key to Janiroidean Genera once classified as 'Janirids' To enter this key, a specimen must be clearly a member of the Janiroidea, based on an inspection of the first pereopod and the pleopods 1-11 (see diagnosis in Wilson 1994). The specimen also should lack specific apomorphies of other families. The family keys of Wolff (1962) or Kussakin (1988), which recognise a broad definition to the Janiridae, may be used first to identify many janiroidean family-level taxa, especially deep-sea taxa. All genera or families that have been placed in the 'janirid' group in the past are included in this key (see list in Introduction). This key references the following families as single entries and the component genera are not keyed out: Acanthaspidiidae, Janirellidae, Joeropsididae, Katianiridae, Munnopsididae and Thambematidae. The genera of the other 'janirid' family, the Microparasellidae, are keyed out separately. Rhacura Richardson, 1908 was left out of this key. The key was constructed so as to require minimal dissection, and with some recognition that incomplete specimens might be at hand. The key is polyotomous, at least at the beginning using the form of the uropod. la. lb. lc. Id. le. If. 2a. 2b. 2c. 3a. 3b. 4a. 4b. 5a. 5b. 6a. 6b. Uropod large with normal rami, total length near that of pleotelson, uropodal endopod longer than protopod ............................................................................................ 24 Uropod short, distinctly shorter than pleotelson but not squat, endopod longer than protopod ................................................................................................................. 2 Uropod with squat rami and sympod, rami shorter than sympod ............................ 8 Uropod with thin, subequal sympod and rami; pereopods 11-VII with only 1 or 2 dactylar claws ........................................................................................................ 13 Uropod with elongate, robust protopod that is longer than rami .............................19 Uropods broken off but large socket present .......................................................31 Frontal area with broad and quadrate rostrum ....................................................... ..........................................Austrofilius Hodgson, 1910 (Antarctic shallow marine) Frontal area with pointed and elongate rostrum .... Jaerella Richardson, 1911b (deep sea) Frontal area broad and/or rounded and sloping, never quadrate or pointed .............. 3 Pereopods 11-VII with 3 dactylar claws or 2 dactylar claws and accessory seta; head often with a few ocelli .........................................................................................4 Pereopods 11-VII with only 1 or 2 dactylar claws; blind ........................................ 5 Pereopod I gnathopod, carpus enlarged with robust sensillate setae .........................................................Ectias Richardson, 1906 (free-living marine) Pereopod I leg-like, carpus slender with few setae ..................................................... .... Iais Bovallius, 1886 (free-living in fresh to brackish water to commensal on marine Sphaeromatid isopods) 6 Pleotelson tapering posteriorly to distal spine ..................................................... Pleotelson distally rounded ............................................................................ 7 Body broad; anterior pereonites with anteriorly projecting points; antennula elongate, flagellum with around 16 articles .................. Trichopleon Beddard, 1886a (deep-sea) Body narrow; anterior pereonites laterally rounded; antennula short, flagellum with only 2 articles .........................................Microjaera Bocquet & Lkvi, 1955 (interstitial) .............................. G. D. F. Wilson and J.-W. Wagele 10a. lob. 1la. llb. Body extremely thin and attenuated, pereonites much longer than wide; pleopods 1-11 may not be opercular (pleopod I11 exposed) ............................................................... ....... Thambematidae (Thambema Stebbing, 1912, and Microthambema Birstein, 1961) (deep-sea) Body narrow, but some pereonites wider than long; pleopods 1-11 always opercular (pleopod 111 not exposed) ............................................. Xostylus Menzies, 1962b (deep-sea) Pereopods 11-VII with 3 dactylar claws, or 2 dactylar claws and subdistal accessory seta ................................................................................................................. 9 Pereopods 11-VII with only 1 or 2 dactylar claws, no accessory seta ........................ 10 Uropodal protopod lacking medial curved spine; antenna normal; frontal area lacking rostral projection ................ Jaera Leach, 1814 (shallow marine to brackish and commensal with sphaeromatid isopods) Uropodal protopod with medial curved spine; antenna strongly geniculate between enlarged article 5 and article 6, proximal flagellar articles conjoint and swollen; frontal area with separate rostra1 projection ................................................................................. ................... Joeropsididae (Joeropsis Koehler, 1885) (shallow marine, cosmopolitan) Uropods uniramous ................Microparasellus Karaman, 1933 (interstitial, fresh water) Uropods biramous ......................................................................................... 11 Pereopods 11-VII with ventral dactylar claw subequal to dorsal claw; antennal flagellum proximal articles wider than long, conjoint, near width of article 6 .......................... ............................................................Hawaianira Miller, 1967 (shallow marine) Pereopods 11-VII with ventral dactylar claw distinctly smaller than dorsal claw; antennal flagellum proximal articles separate, not conjoint, longer than wide ....................12 Antenna1 articles 5-6 near length of articles 1-4; maxillipedal palp wider than endite ............................... Caecijaera Menzies, 1951a (commensal with limnoriid isopods) Antennal articles 5-6 distinctly longer than articles 1-4; maxillipedal palp distinctly narrower than endite .......................................Neojaera Nordenstam, 1933 (shallow marine) 14 Body with dorsal and/or lateral denticles or spines ............................................... Body without denticles or spines ........................................................................ 15 Uropods biramous; pereonites 5-7 and pleotelson fused together .................................. ................. Munnopsididae, in part (Microprotus Richardson, 1909) (deep sea, blind) Uropods short and uniramous; pereonites 5-7 and pleotelson free ................................ ..........................................Janirellidae (Janirella Bonnier, 1896) (deep sea, blind)) Maxillipedal palp narrower than endite, articles subequal; antenna articles 5-6 length subequal to articles 2-3 ................................................................................................. ......... Protocharon Delamare-Deboutteville & Chappuis, 1956 (fresh water on islands) Maxillipedal palp wider than or subequal to endite, articles subequal; antenna articles 5-6 distinctly longer than articles 2-3 .............................................................16 Frontal area broad, rounded, projecting between antennulae ..................................... ............................................. Microjanira Schiecke & Fresi, 1970 (shallow marine) Frontal area narrow, antennulae not widely separated 17 Male pleopod I1 stylet shorter than protopod, thin, with distinct angle in distal part ....... ...........................................Mackinia Matsumoto, 1956 (subterranean fresh water) Male pleopod I1 stylet much longer than protopod, distally coiled ........................... 18 Antennal scale absent; some pereopodal coxae visible in dorsal view ............................ ............ Heterias (Fritzianira) De Castro & Lima, 1977 (South American fresh water) Antennal scale present; no pereopodal coxae visible in dorsal view ............................... Heterias (Heterias) Richardson, 1904a (Australian and Tasmanian fresh water) Uropods without rami, made of single segment; pereopod I chelate with large seta forming fixed finger on propodus ........................ Katianiridae (Katianira Hansen, 1916, and Natalianira Kensley, 1984~)(deep sea, blind) Uropods with at least one ramus or biarticulate; pereopod I leg-like or subchelate ..... 20 Body thin, worm-like; head longer than broad; pereopods inserting dorsolaterally 21 Body broad, often with dorsal or lateral spines; head broader than long; pereopods inserting 23 ventrally .................................................................................................... Antennal scale absent; maxillipedal palp thin, much narrower than endite, with only 4 articles, distal article pointed; pereopods 11-VII with 3 dactylar claws ...................Angeliera Chappuis & Delamare-Deboutteville, 1954 (interstitial marine) Antennal scale present; maxillipedal palp with 5 articles; pereopods 11-VII with only 2 dactylar claws .......................................................................................................22 ...... .......................................... .......... ...... ................................. Review of the Janiridae 69 1 Pereopodal coxae visible in dorsal view ................................................................... ............................... Microcharon Karaman, 1934 (interstitial marine to fresh water) Pereopodal come not visible in dorsal view, hidden by small tergal plate ...................... .......................... Paracharon Coineau, 1968 (interstitial freshwater, tropical islands) Maxillipedal endite much broader than palp; pleopod I11 exopod broad with multiple plumose setae ............................ Acanthaspidiidae [Acanthaspidia Stebbing, 1893 (blind) and Ianthopsis Beddard, 1886bl (shallow marine to deep sea) Maxillipedal width near that of palp; pleopod 111 exopod narrow without plumose setae .............................. Tole Ortmann, 1901 (senior synonym of Iolella) (Arctic marine) Uropodal exopod minute or absent ..................... Caecianiropsis Menzies & Pettit, 1956 (excluding C. ectiformis) (shallow marine interstitial) Uropodal exopod near length of protopod, not minute or absent 25 Pereopods 11-VII with only 1 large dactylar claw, second claw seta-like, third claw absent ...............................................................Janthura Wolff, 1962 (deep-sea, blind) Pereopods 11-VII with 3 dactylar claws or 2 dactylar claws and subdistal accessory seta ..................................... ......................................................................26 Body vermiform; pleotelson tiny, much smaller than pereonites; 3 anterior pleonites free ventrally ............................Vermectiadidae (Vermectias Sivertsen & Holthuis, 1980) (shallow marine, subantarctic) Body normal; pleotelson longer than pereonites; no more than 2 anterior pleonites visible ventrally .................................................................................................... 27 Pereopod I propodus with proximoventral denticles; pereonites often with large lateral lappets or spines ................. Janiralata Menzies, 19511, (shallow marine to deep sea) Pereopod I propodus lacking distinct proximoventral denticles; pereonites without spines and with only small lappets ..........................................................................28 Male pereopod I propodus and carpus enlarged, often swollen .................................... ...............................................Carpias Richardson, 1902 (tropical shallow marine) Male pereopod I carpus elongate, not swollen; male propodus never enlarged, always slender ............................................................... Ianiropsis Sars, 18976 (shallow marine) Male pereopod I carpus and propodus not elongate, near length of pereopods 11-VII ... 29 Head with small dorsally curved rostrum; eyes large, often bulging laterally; uropodal rami flattened ................. Iathrippa Bovallius, 1886 (southern hemisphere shallow marine) Head generally lacking small rostrum; eyes compact, set on head without bulging distinctly laterally; uropodal rami round in cross-section ................................................. 30 Male pleopod I without protruding lateral lobes; maxillipedal endite much narrower than broad palp .....................Janaira Moreira & Pires, 1977b (Brazilian shallow marine) Male pleopod I with distinct lateral lobes separate from medial lobes; maxillipedal endite width subequal to palp ..................................Janira Leach, 1814 (shallow marine) Pereopods 11-VII with 3 dactylar claws or 2 dactylar claws and subdistal accessory seta ............................................................................................................32 Pereopods 11-VII with only 1 or 2 dactylar claws, no accessory seta ........................ 33 Maxillipedal endite width near that of palp; pleopod 111 exopod narrow without plumose setae .........................................................................................................27 Maxillipedal endite much broader than palp; pleopod I11 exopod broad with multiple plumose Acanthaspidiidae (see 23a) setae ........................................................................ Pereopod I chelate with large seta forming fixed finger on propodus ............................ ...................................................................................Katianiridae (see 19a) Pereopod I leg-like, not chelate ......................................................................... 34 Caecianiropsis Menzies & Pettit, 1956 (interstitial marine) Body narrow, worm-like Body broad, often with spines ...........................................................................35 Pereonites 5-7 and pleotelson fused together ............................................................ .................Munnopsididae, in part (Microprotus Richardson, 1909) (deep sea, blind) Pereonites 5-7 and pleotelson free ..................................................................... 36 Head without anteriorly projecting rostrum Janthura Wolff, 1962 (deep sea, blind) Head with anteriorly projecting rostrum .............................................................. 23 ............................ .. ......... ........ 692 G . D. F. Wilson and J.-W. Wagele Genera of the Janiridae Austrofilus Hodgson, 1910 Austrofilius Hodgson, 1910: 51. Type species: Austrofilius furcatus Hodgson, 1910. Species Included A. furcatus Hodgson, 1910: 51; A. serratus Vanhoffen, 1914: 554. (Not included: A. arnaudi Kussakin & Vasina, 1980; 'Austrofilius furcatus' Branch, Griffiths, Kensley & Sieg, 1991.) Other References Austrofilius. -Vanhoffen, 1914: 554; Nordenstam, 1933: 252; Barnard, 1940: 434; Schultz, 1976: 24, 27; Kussakin and Vasina, 1980: 362; 1984:12; Kussakin, 1982: 82. Neojaera.-Menzies, 19626: 74; Wolff, 1962: 252, 254, 259, 290; Amar and Roman, 1973: 571; Siebenaller and Hessler, 1977: 20; Kensley, 1978: 139. Diagnosis Body broad and flattened. Cephalon with broad quadrate rostrum and dorsal eyes; lateral margins concave, sometimes with denticles and pointed anteriorly. Lateral margins of pereonites smoothly concave, coxae not visible in dorsal view. Pleotelson shield-like, with rounded lateral margins and generally with lateral denticles. Antennula short, with only 5-6 articles; first article broad, flagellum with 2-3 articles. Antenna with conspicuous scale (exopod); proximal articles of the flagellum conjoint and swollen. Mandibular molar process truncate with broad grinding surface, palp with 3 articles. Maxillipedal palp articles 2-3 expanded, distinctly broader than articles 4-5 and endite. Pereopod I ambulatory, dactylus short, with 2 claws; propodus and carpus with row of setae on ventral margin. Pereopods 11-VII dactyli with two distal claws, ventral claw smaller than dorsal claw. Male pleopod I lateral lobes curving, projecting posteriorly, medial lobes broad and rounded, also distinctly projecting; proximal region narrow with straight or concave lateral margins. Sympod of male pleopod I1 acutely pointed or rounded, setiferous; endopod curved, sometimes elongate and coiled, much longer than sympod, with proximal article sometimes enlarged. Female operculum broad ovate, distally rounded. Pleopod I11 endopod with 3 plumose setae having distinct gap between medial seta and two lateral setae; exopod narrower and longer than endopod; exopod with 2 segments, lacking distal plumose setae. Uropods biramous, branches of subsimilar length; sympod squat, distinctly shorter than rami. Remarks Austrofilius has had a varied history since it was first described by Hodgson (1910). The genus was recognised by authors who studied Antarctic isopods, including Nordenstam (1933), until recently. Menzies (1962b), however, dubiously created a synonymy with Neojaera, and later authors followed this opinion (Wolff 1962; Siebenaller and Hessler 1977). Schultz (1976) reinstated Austrofilius, addressing the broad rostrum that Neojaera species lack. Kussakin (1982) and Kussakin and Vasina (1980) recognised Austrofilius, but the former paper included a new species of Austrofilius that lacks a rostrum and thus belongs in Neojaera. The enlarged endopodal proximal article of male pleopod I1 with the elongate coiled stylet, characteristic of Neojaera, is also found in A. serratus but not in the type species, A. furcatus. The pleopods 1-11 of the males in both species indicates that they are related (similar form, distribution of setae on protopod of pleopod 11). On the other hand, a fully mature male of A. furcatus may not have been illustrated to date. This feature is left variable in the diagnosis for this reason. The diminutive, Iais-like Austrofilius arnaudi Kussakin & Vasina, 1980 departs rather substantially from A. furcatus. The cephalon and pereonites do not extend laterally into broad plates, the pereopodal coxae are visible in dorsal view, and the rostral area of the head is rounded rather than quadrate. A. arnaudi, however, is not a member of Iais, owing to its broad rostral area and the biunguiculate pereopods. The body form is reminiscent of Review of the Janiridae 693 Protocharon, but the rostrum and normal shape of the maxilliped prevents this species from being placed here. For the moment, the species arnaudi must be classified as janirid genus incertae sedis. The species 'Austrofilius furcatus' Branch, Griffiths, Kensley & Sieg, 1991 (not Hodgson, 1910) appears to be similar to arnaudi and also should be excluded from Austrofilius. Caecianiropsis Menzies & Pettit, 1956 Caecianiropsis Menzies and Pettit, 1956: 442. Species Included C. psammophila Menzies & Pettit, 1956: 442; ?Austroniscus ectiformis Vanhoffen, 1914: 553; C. birsteini Kussakin, 1979a: 107. Other References Austroniscus. -Branch et al., 1991: 28. Caecianiropsis.-Birstein, 1961: 137; Wolff, 1962: 36, 38, 236, 276, 251, 291; Kussakin, 1967: 339 (English translation 1968: 341); Schultz, 1969: 256-7; Kensley, 1976: 295; Coineau, 1986: 468; Kussakin, 1988: 160. Diagnosis (based primarily on C. psammophila) Body elongate and thin, more than 5 x longer than wide. Cephalon anterior margin with rounded, nonprojecting rostrum; eyes absent; lateral margins of head rounded, never denticulate, lacking anterior points. Pereonites laterally rounded lacking lappets, coxae visible in dorsal view. Pleonite 1 near width of pereonite 7. Pleotelson oval, posteriorly rounded, approximately same width as pereonites, lateral margin with tiny denticles anterior to insertion of uropods. Antennula first article longer than broad, flagellum with 2-3 articles. Antenna article 3 distinctly longer than articles 1, 2, and 4, with tiny rudimentary scale; flagellum with 15 or more distinct articles, each longer than wide, proximal articles not conjoint; articles 5-6 distinctly longer than basal articles. Mandibular molar process truncate but tapering distally; palp functional with large setose distal article. Maxillipedal endite longer than broad; palp articles 2-3 expanded, broader than endite; palp articles 4-5 distinctly narrower than proximal articles. Pereopod I resembling ambulatory legs, carpus elongate, dactylus short with 2 claws; propodus and carpus with few spine-like setae. Pereopods 11-VII dactyli with two distal claws, ventral claw near same size as dorsal claw. Male pleopod I medial lobes distally rounded; lateral lobes angular, projecting laterally. Male pleopod I1 with exopod inserting subdistally on sympod; endopod with long coiled slender stylet and expanded proximal article; protopod narrow, more than 2.5 x longer than wide. Pleopod I11 endopod with 3 plumose setae having distinct gap between medial seta and two lateral setae; endopod broader than and around same length as exopod; exopod, with 2 segments, lacking plumose setae. Uropods elongate, biramous; protopod distinctly longer than wide, shorter than endopod; exopod inserting apically, shorter than endopod. Remarks Caecianiropsis is a marine interstitial genus with 3 known species. The genus has an apparent bipolar distribution with two morphologically distinct groups. The northern group, containing C. psammophila from Northern California (Menzies and Pettit 1956) and C. birsteini from the Bering and Okhotsk Seas (Kussakin 1979a), has elongate antenna1 flagellae, a rostrum only slightly wider than the basal antennular article, and a pleotelson that is broadest just anterior to the uropods. The southern group includes only C. ectiformis from Kerguelen (Kussakin 1974) and St Paul Islands (Kensley 1976) in the South Indian Ocean and possibly from Marion and Prince Edward Islands in the South Atlantic (Branch et al. 1991). This species has a compact antennular flagellum, its rostrum is much broader than antennular article 1, and its pleotelson either narrows posteriorly or is at least parallelsided laterally. The male pleopods are poorly known in C. ectiformis, although the pleopod I differs from that of the northern group in lacking the angular lateral extensions. Menzies and Pettit (1956) uncertainly assigned C. ectiformis to this genus, and Kensley (1976) 694 G . D. F. Wilson and J.-W. Wagele maintained this classification. Kensley in a recent paper (Branch et al. 1991) erroneously placed the species in Austroniscus, Vanhoffen's (1914) original placement. C. ectiformis may not belong in this genus owing to the differences between the northern and southern groups, but a redescription is necessary to demonstrate this. In C. psammophila, the male second pleopods are similar to those seen in Neojaera, especially the elongate coiled stylet. This would argue for a close relationship between the two genera, although the generality of this observation is unknown. Caecijaera Menzies, 195l a Caecijaera Menzies, 1951a: 3. Type species: Caecijaera horvathi Menzies, 1951a. Species Included Caecijaera horvathi Menzies, 1951a: 3; C. mirabilis Kussakin, 1962: 53; C. borealis Kussakin, 1962: 59; C. derjugini Kussakin, 1962: 56. Other References Caecijaera.-Menzies, 19626: 70; Wolff, 1962: 39,236,251,254,276,281,291; Kussakin, 1988: 168; Schultz, 1969: 257; Cooke, 1977: 105; Svavarsson, 1982: 224. Diagnosis Body broad, setose, lateral margins broadening posteriorly. Cephalon broad, lateral margin rounded or linear, with rounded or pointed anterior projection; eyes lacking; rostrum absent, vertex projecting anteriorly. Tergites of pereonites 2-4 with single anterolateral lappets, posterolaterally concave; pereopodal coxae visible in dorsal view on pereonites 1-4; coxae on pereonites 5-7 not visible in dorsal view. Pleonite 1 short, much narrower than pereonites or pleotelson. Pleotelson as broad or broader than pereonite 7; lateral margins setose without denticles. Antennula article 1 wider than long, with distinct setose medial lobe; flagellum with 2-3 articles. Antenna basal articles 1-4 length subequal; article 3 with large setose scale; articles 5 and 6 short and stout, near length of article 3; flagellum with 8-11 articles, each separate and longer than wide, but not basally conjoint. Mandible thin, elongate with reduced and tapering molar process (broader in subgenus Caecijaerella); palp distal article with few setae, not strongly curved. Maxilliped endite longer than wide; palp broader than endite with distal 2 articles distinctly narrower than basal articles. Pereopod I similar to walking legs; propodus narrower than carpus with no ventral setae; carpus subrectangular with few ventral setae. Pereopods 11-VII dactylus with only 2 distal claws, with ventral claw tiny, much smaller than dorsal claw. Male pleopod I distal tip lacking large laterally projecting lobes. Male pleopod I1 proximal article of endopod near maximum width of stylet or somewhat thicker than stylet; stylet varying from shorter than protopod (C. horvathz) to longer than sympod with distal part elongate and curving but not distally coiled. Pleopod I11 endopod with many plumose setae, sometimes with gap between medial seta and lateral setae; exopod with 2 free articles, lacking plumose setae, narrower but longer than broad endopod. Uropods with squat rami and sympod; exopod apical; sympod longer than rami with distinct setose lateral lobe. Remarks Caecijaera has a body form reminiscent of Heterias and Jaera because it broadens posteriorly to a broad and rounded pleotelson; this form is best expressed in the three boreal species described by Kussakin (1962) in his subgenus Caecijaerella: C. mirabilis, C. borealis, and C. derjugini. Caecijaera apomorphies include a broad antennular article 1 with a distinctive medial lobe, short and squat antennae with a large scale, and many plumose setae on the broad pleopod I11 endopod. An elongate male pleopodal stylet forming only a single coil seems to be a synapomorphy of subgenus Caecijaerella, while the stylet of C. Caecijaera horvathi is more normal, being shorter than the protopod. The molar process also varies Review of the Janiridae 695 considerably among the four species: from more or less normal in C. borealis t o greatly reduced in the type species, C. horvathi. The entire mandible is rather attenuated i n this latter species. Caecijaera species are small commensals o f the isopod wood-borer, Limnoria: they live in Limnoria's burrows and may feed o n the fungi and bacteria living on the burrow walls (Svavarsson 1982; Bowman 1988). C. horvathi described from Hawaii (Cooke 1977) may be a distinct species because its male pleopods have distinctly elongate lateral lobes, while the individuals described b y Menzies ( 1 9 5 1 ~have ) lateral lobes that only slightly exceed the length o f the medial lobes. Carpias Richardson, 1902 Carpias Richardson, 1902: 294.-Bowman and Morris, 1979: 650. Bagatus Nobili, 1906: 268.-Nobili, 1907: 42; Pires, 1980: 96. Janatus Carvacho, 1983: 289. Type species: C. berrnudensis Richardson, 1902. Species Included Carpias bermudensis Richardson, 1902: 294; C. floridensis Menzies & Kruczynski, 1983: 92; C. harrietae Pires, 1981a: 206; Bagatus stylodactylus Nobili, 1906; Janira algicola Miller, 1941: 317; B. brachydactylus Pires, 1982: 239; B. brucei Monod, 1974: 1123; Janira crosslandi Stebbing, 1910a: 108; B. galloprovincialis Amar, 1950: 36; B. ichthyoxenos Monod, 1961: 68; B. longidactylus Nordenstam, 1946: 14; B. longimanus Pillai, 1954: 19; Janira minuta Richardson, 1902: 297; Janira? nanus Stebbing, 1905: 50; B. nereus Pires, 1982: 254; Janiropsis parva Omer-Cooper, 1921: 1; B. punctatus Kensley, 1984a: 61; B. serricaudus Menzies & Glynn, 1968: 78; B. stebbingi Monod, 1933: 169; B. triton Pires, 1982: 25 1 ; Janatus villalobosi Carvacho, 1983: 289. Other References Carpias.-Richardson, 1905b: 452; Omer-Cooper, 1921: 79; Nierstrasz, 1941: 287; Menzies, 1962b: 70; W o l f f , 1962: 39, 236, 276, 281, 290; Miller, 1968: 23; Schultz, 1969: 260; Pires, 1980: 97; Menzies and Kruczynski, 1983: 92; Kensley and Snelgrove, 1987: 196; Kensley, 1988: 41; Kensley and Schotte, 1989: 82; Schotte et al., 1991: 253; Muller, 1992: 331, 337. Bagatus. -Nierstrasz, 1941: 284; Amar, 1952: 355; W o l f f , 1962: 39, 235, 276, 281, 292, 300; Fresi, 1968: 271; Bellan-Santini, 1969: 30; Geldiay and Kocatas, 1972: 26; Pires, 1981b: 32; Ortiz, 1983: 9; Kensley, 1984b: 33; Muller, 1989: 203-7; Muller, 1992: 331-2. Janira?. -Vanhoffen, 1914: 528; Brian and Dartevelle, 1949: 93. Janira. -Richardson, 1905b: 471; V a n h o f f e n ,1914: 528; Monod, 1931c: 408; Nierstrasz, 1941: 283; Nordenstam, 1946: 15; Barnard, 1962: 244; Kensley, 1978: 139; Menzies and Glynn, 1968: 79; Schultz, 1969: 258. Diagnosis Body moderately broad, pereonite 4 broadest, tergites o f pereonites 2-4 laterally concave; coxae visible in dorsal view o n most pereonites. Cephalon broad, laterally rounded, with dorsolateral eyes, vertex straight or concave, lacking rostrum. Pleotelson broad, laterally rounded, lacking denticles on posterolateral margins. Pleonite 1 free, short, distinctly narrower than pleotelson or pereonite 7 . Pleotelson same width as posterior pereonites; lateral margins o f pleotelson lacking denticles. Antennula basal article only slightly broader than second article, length decidedly greater than width; flagellum multiarticulate, approximately 7-11 articles in males. Antenna articles 1-4 subequal with large scale o n article 3; antenna1 articles 5 and 6 longer than basal articles 1-4; flagellum multiarticulate, with more than 20 articles, proximal articles o f flagellum not conjoint, separate and longer than wide. Mandibles normal with truncate, nontapering molar process; palp elongate, distal article curved and setose. Maxillipedal endite longer than wide; palp broader than endite, distal articles 4-5 thin, straight-sided, much narrower than broad palp articles 1-3. In female, 696 G . D . F. Wilson and J.-W. Wagele pereopod I carpus enlarged, oval, with two rows of spine-like sensory setae. In male, pereopod I enlarged, without large spine-like setae on carpus, but with strong spines on subchelate margin of carpus; carpus subchelate (distoventral) margin variously inflated or rounded; propodus elongate, curved, often distally inflated. Pereopod I1 of male generally similar to pereopods 111-VII. Pereopods 11-VII dactylus with 2 large subequal distal claws on dactylus with third large subdistal accessory claw. Male pleopod I distal tip 'V' shaped, with merging medial and lateral lobes; lateral lobes projecting posteriorly. Male pleopod I1 stylet never coiled, often near length of sympod, proximally; basal article of endopod not enlarged, only slightly wider than proximal part of basis. Pleopod I11 endopod either with 3 plumose setae, either equally spaced or with gap between medial seta and 2 lateral setae; exopod of both sexes narrower than endopod and same length as or longer than endopod; exopod uniarticulate, distally rounded, lacking plumose setae. Uropods large, near length of pleotelson or longer, with narrow elongate rami and sympod; exopod apical; sympod shorter than rami. Remarks Carpias Richardson is a common tropical and subtropical genus. Specimens of this genus are often encountered in shallow (less than 10 m) marine habitats around coral reefs. Carpias belongs to the shallow-water group of three-clawed janirids that includes Ianiropsis, Janaira and Janira. Species of Carpias and its junior synonym Bagatus display similar sexual dimorphism in pereopod I: the male has an enlarged and carpochelate pereopod I, with Carpias perhaps being more extreme than Bagatus. Bagatus was synonymised into Carpias Richardson by Bowman and Morris (1979), but Pires (1980) recognised and maintained their distinctness. The genus Carpias Richardson, 1902, was also synonymised (incorrectly) by Menzies and Glynn (1968) into Bagatus Nobili, 1906. According to Pires (1980), Bagatus and Carpias are without doubt closely related, but are not identical. She gives the following 'key characters' of Carpias (ibid., p. 96). 'Pereonite I nearly 1.5 times wider than head. Pereopod I: ischium and merus long, subequal in length; basal third of carpus narrow, the remaining portion very enlarged with lateral outer margin globularly swollen; propodus narrow, with the apex roundish but never dilated.' In the discussion, she also pointed out that 'the copulatory organ of pleopod I1 does not reach the distal margin of the sympod . . .' In her concept, Bagatus is similar but has long copulatory organs on pleopod 11, the pereopod I carpus is never swollen, the pereopod I ischium and merus are short, and pereonite 1 is subequal to the width of the head. Significantly, Pires (1980) does not include any of these key characters in the diagnosis for Carpias, except for the shape of the distal articles of the male pereopod I. Pires (1981~) added a species, Carpias harrietae, that conflicts with her key features for the genus, except for the swollen carpus of pereopod I. The stylet of the male pleopod I1 of C. harrietae is long and coiled, although Pires rationalises its length by the statement that it does not extend beyond the distal margin of the sympod (Pires 1981~).Elongate stylets, however, are typically coiled in most janirids. The length of the stylet should be measured on the medial margin from its proximal articulation with the first endopodal article to the distal tip (as done in Wilson and Hessler 1980). The definition of Carpias was not adjusted in Pires (1980) to take account of the similarities of C. harrietae to species of Bagatus. A third species, C. floridensis Menzies & Kruczynski (1983), is also near the definition of Bagatus given by Pires (1982) except that the male first pereopod I has a swollen carpus and propodus. Therefore, the only significant character in Pires' (1982) concept that separates the two genera is the form of the male pereopod I carpus. Because this character is within the possible structural variation of these two genera (e.g. see Pires 1982: B. stylodactylus and B. platydactylus, her figs 43 and 45 from Nobili 1907), Carpias as defined by Pires (1980) does not appear to be a monophyletic group. Notably, the published illustrations of the distal tip of male pleopod I for species of Carpias are more similar to species of Bagatus than they are to each other. Review of the Janiridae 697 More recently, Miiller (1992) claimed that the two genera can be distinguished by the presence of one or more large teeth on the proximal ventral margin of the mature male propodus. Using this feature, he proposes the transfer of the species punctatus, brachydactylus, brucei and minutus from Bagatus to Carpias. The presence or absence of denticles on the male propodus I is not a convincing character because Pires (1982, her figs 7, 8) clearly shows that males of minutus change from a Bagatus form to a Carpias form during maturation. Moreover, the male pereopod of minutus is little different from other species in Bagatus sensu Pires (1982). Because the two genera cannot be clearly defined, the synonymy of Carpias and Bagatus is retained, with the senior synonym being Carpias Richardson (see Bowman and Morris 1979). Carvacho (1983) created a new genus, Janatus, for the species villalobosi, because this species seemed intermediate between Janira and Bagatus. Janatus was erroneously justified further by its intermediate geographic position between Janira and Bagatus. Janatus is well within the known morphological variation of Carpias (especially in the male pleopod I: see Pires 1980, 1981a, 1982), and it has an enlarged male pereopod I, an autapomorphy of Carpias. Therefore, Janatus is a junior synonym of Carpias. C. villalobosi (Carvacho 1983) may belong to the minutus group of Pires (1982). C. galloprovincialis (Amar 1950) is recognised as a full species rather than a subspecies of C. stebbingi (Monod). Ectias Richardson, 1906 Ectias Richardson, 1906: 14. Zaniroides Kensley, 1976: 295 (created for Janira angusta). Type species: Ectias turqueti Richardson, 1906. Species Included Ectias turqueti Richardson, 1906: 14; Janira angusta Barnard, 1920: 404. Other References Ectias. -Richardson, 1913: 18; Tattersall, 1921: 202; Nordenstam, 1933: 186; Hale, 1937: 31; Nierstrasz, 1941: 288; Stephensen, 1947: 8; Wolff, 1962: 36, 38, 39, 236, 276, 252, 289, 291; Kussakin, 1967: 339; 1968: 341; Arnaud, 1974: 646; Schultz, 1976: 30; Kensley, 1978: 135; Kussakin, 1982: 82. Diagnosis Body narrow, with all pereonites approximately same width; length approximately 6 x width. Cephalon lateral margin rounded, with no spines; vertex convex or obtusely angular but lacking rostrum; small eyes on dorsal surface near lateral margin. Tergites of pereonites laterally rounded, with coxae visible only as narrow crescents on anterior edges of pereonites 1-4 and posterior edges of pereonites 5-7. Pleonite 1 short, distinctly narrower than pleotelson or pereonite 7. Lateral margins of pleotelson rounded, lacking any denticles. Antennula short but longer than cephalon, antennular article 1 straight-sided, length decidedly greater than width; flagellum with 5-6 articles. Antenna1 basal articles (1-4) all same length with distinct scale on article 3; articles 5 and 6 elongate, much longer than basal articles 1-4; flagellum multiarticulate, numbering 15 or more articles; subproximal flagellar articles separate, wider than long but not conjoint and not wider than antenna1 article 6; most proximal flagellar article longer than wide. Mandibular molar process truncate but distally tapering; palp elongate with curved and setose distal tip. Maxillipedal endite longer than wide; only slightly narrower than palp; palp distal articles 4-5 narrow and straight-sided, narrower than broad basal palp articles 1-3. Pereopod I carpus subchelate, enlarged and subtriangular, with 2 rows of few spine-like setae on margin opposing propodus; propodus with no spine-like setae. Pereopod 11-VII dactylus with ventral claw near same size as dorsal claw, and accessory seta enlarged into third claw. Male pleopod I narrow, distally 'V'-shaped, with medial and lateral lobes merged into elongate posterolateral projections. Male pleopod I1 proximal article of endopod enlarged, decidedly rounded, and 698 G . D. F. Wilson and J.-W. Wagele thicker than stylet; stylet much longer than sympod; elongate and curving, but not distally thin and tightly coiled. Pleopod I11 endopod with 3 plumose setae, with distinct gap between medial seta and outer 2 setae; exopod more slender but longer than endopod, with many plumose setae, with 2 free articles with interior suture strongly slanting to body axis. Uropods reduced, distinctly shorter than pleotelson; endopod flattened with rounded margins; exopod inserting apically; sympod subrectangular, shorter than rami. Remarks Kensley (1976) erected the genus Ianiroides for Janira angusta Barnard, 1920, but later placed this species in Ectias (Kensley 1978). Apomorphies of Ectias include a carposubchelate pereopod I with an enlarged carpus, elongate and widely separated lateral lobes on the male pleopod, a long curling stylet on the male pleopod I1 (but not distally coiling), and plumose setae on the exopod tip of pleopod 111. Hawaianira Miller, 1967 Hawaianira Miller, 1967: 187. Type species: H Q W Q ~ peleae U ~ ~ ~Miller, Q 1967. Species Included Hawaianira peleae Miller, 1967: 187; Janira caudata Richardson, 1910: 33. Other References Ianthopsis.-Wolff, 1962: 41, 254. Janira. -Nierstrasz, 1941: 282. Diagnosis Body compact, flattened, pereonite widths subequal. Cephalon broad, lateral margins with acute anterior projections, lacking denticles; vertex lacking rostrum, slightly convex; eyes dorsal. Pereonites laterally truncate, without notches or lappets, coxae not visible in dorsal view. Pleonite 1 fused to pleotelson. Pleotelson shield-like, same width as anterior pereonites; lateral margins indented at posterolateral insertions of uropods, with small posterolateral denticles. Antennular article 1 enlarged, width near length or wider, with distinct medial lobe; flagellum with 3 articles. Antenna long; basal articles (1-4) subequal length, with small scale on article 3; articles 5 and 6 much longer than basal articles 1-4; flagellum with 40-45 articles; proximal flagellar articles wider than long, conjoint, not swollen, basal articles near width of article 6. Mandibular molar normal, elongate, straight-sided, distally truncate; palp large with elongate first article and setose straight distal article. Maxillipedal endite longer than wide, broader than palp; palp articles 4-5 straight-sided, elongate, narrower than palp articles 1-3. Pereopod I carpus leg-like; propodus with 3 spine-like setae and numerous ctenate scales; carpus with only fine setae in 2 rows. Pereopod 11-VII dactyli with only 2 distal claws, ventral claw near same size as dorsal claw. Male pleopod I with short but distinct medial and lateral lobes; medial lobe setose and rounded; lateral lobe short and roughly triangular with slight terminal notch for stylet guide. Male pleopod I1 not modified; proximal article of endopod near maximum width of stylet; stylet shorter than protopod, only slightly curved. Pleopod I11 endopod truncate with 6-8 plumose setae; exopod distally rounded, finely setose, narrower and shorter than endopod, with 2 free articles, interior suture at angle to body axis. Uropods reduced, distinctly shorter than pleotelson; with squat rami and protopod; rami shorter than protopod; uropod exopod apical. Remarks The affinities of Hawaianira are somewhat unclear because this genus has characters of an unspecialised Joeropsis or Caecijaera (see Miller 1967). However, Hawaianira shares several apomorphies with Caecijaera: short broad uropods and antennular first articles Review of the Janiridae 699 (seen also in Janthura and Neojaera), a broad flattened body, and a broad pleopod I11 endopod with many plumose setae. Janira caudata Richardson, 1910 appears to belong in this genus, not in Ianthopsis as suggested by Wolff (1962). H.-G. Muller (in litt.), reports discovering a new species of Hawaianira from RCunion Island. Heterias Richardson, 1904a Janirella Sayce, 1900: 124 Heterias Richardson, 1904a: 6. Pseudasellus Chappuis, 1951: 7. Fritzianira Castro and Lima, 1977: 1. Type species: Heterias pusilla (Sayce, 1900). Species Included Janirella pusilla Sayce, 1900: 124; Janira exul Muller, 1892: 209; Pseudasellus nichollsi Chappuis, 1951: 7; Heterias petrensis Roberts, 1975: 21. Other References Heterias. -Barnard, 1927: 161; Nicholls, 1929: 290, 295; Wolff, 1962: 39, 41, 236, 276, 281, 290; Williams, 1968: 145; 1974: 88; 1980: 155-8; Bowman et al., 1987: 275; Horwitz and Knott, 1991: 29-32. Fritzianira.-Castro and Lima, 1977: 1; Ringuelet, 1981: 88; Taberner, 1982: 19; Bowman et al., 1987: 278. Janiropsis. -Van Name, 1942: 323; Roberts, 1975: 30. Pseudasellus. -Wolff, 1962: 39, 207, 250, 276, 281, 290; Williams, 1974: 88; 1981: 1116. Parasellidae sp. - Bachman, 1964: 491. Diagnosis Body broad, lateral margin broadening posteriorly to broad pleotelson, with many fine setae. Cephalon lateral margin rounded, lacking spines or denticles; eyes dorsal if present; vertex straight or rounded, lacking rostrum. Pereonites laterally rounded or truncate, coxae not visible in dorsal view (in subgenus Heterias), or anterior pereonites laterally concave, coxae slightly visible in dorsal view on pereonites 2-7 (in subgenus Fritzianira) . Pleonite 1 extremely short and narrower than pleotelson or pereonite 7. Pleotelson broader than anterior pereonites; lateral margins rounded, lacking denticles or incision for uropods. Antennula short but not compact; article 1 short and broad, length near width; flagellum with 3-4 articles. Antenna elongate; basal articles 1-4 subequal, article 3 with (subgenus Heterias) or without (subgenus Fritzianira) scale; articles 5-6 distinctly longer than basal articles 1-4; flagellum with more than 20 articles, proximal articles separate, longer than wide, not conjoint. Mandible normal; molar process distally truncate, only slightly tapering; palp large and functional, near length of mandible body, proximal article elongate, distal article slightly curved and setose. Maxillipedal endite longer than wide, narrower than palp; palp articles 4-5 narrower than palp articles 1-3. Pereopod I leg-like, carpus not enlarged, with few or no spine-like setae; propodus with only fine setae. Pereopod 11-VII dactylus with only 2 distal claws, with ventral claw shorter than dorsal claw but approximately of same proximal width. Male pleopod I with well-differentiated medial and lateral lobes; lateral lobes point laterally; medial lobes broad, near width of pleopod, distal flattened and dorsally curved. Male pleopod I1 proximal article of endopod enlarged, decidedly rounded, and thicker than stylet; stylet much longer than sympod, distal part thin and coiling. Pleopod I11 endopod with no plumose setae; exopod longer and broader than endopod, with no plumose setae and 2 free articles sutured at angle to body axis. Uropods slender, overall length subequal to pleotelson length; exopod inserting apically; protopod shorter than rami. G. D. F. Wilson and J.-W. Wagele 700 Remarks The unique copulatory apparatus on the male pleopods is a distinctive apomorphy of Heterias. This feature has been unevenly described in most species and should be carefully restudied (Bowman et al. 1987 provide scanning electron micrographs). Other features of Heterias include a Jaera-like body form (broadening posteriorly), 2 claws on the walking legs, long slender uropods, a short antennula with only three articles in the flagellum, a long antenna with more than 20 articles, and a pleopod I11 with no plumose setae on the endopod. Wolff (1962) assigned Janira em1 to Heterias, but De Castro and Lima (1977) apparently were unaware of this assignment as well as of Robert's (1975) work when they created Fritzianira for this species. Fritzianira currently is a junior synonym of Heterias, but Bowman et al. (1987) retained this name for a subgenus because H. em1 lacks the antennal scale seen in the Australian and Tasmanian species. Williams (1968, 1974) suggested that Pseudasellus nichollsi Chappuis, 1951 belongs in Heterias, owing to the similarity of the male pleopods. H. petrensis Roberts retains eyes and an antennal scale, suggesting that it may be the least-derived species of the genus. Both Chappuis (1951, his fig. 21) and Roberts (1975, his fig. 6b) mistook a juvenile male as an intersex of the species. In both papers, buds of male pleopods are shown developing inside the female operculum, typical of most janiroideans (e.g. Wilson 1981, his fig. 10C). The distribution of Heterias in fresh waters of South Australia, Tasmania and South America suggests that this genus is an ancient Gondwanaland relict (Bowman et al. 1987), much like the protojanirid and phreatoicidean isopods. On the other hand, marine janirid taxa are being found in increasing numbers, so the presence of a marine Heterias-like form should not be discounted. Indeed, a Heterias-like isopod has been reported from the Galapagos Islands (Brusca 1987), although these undescribed specimens may not fit comfortably into Heterias. Bowman et al. (1987) mention that some of their specimens came from tidally influenced streams, possibly indicating a close association with marine waters. Moreover, Bowman et al. (1987) did not attempt to explain the disjunct distribution of Heterias exul between Brazil and Chile. If this 'species' is derived from marine ancestors, its distribution would indicate multiple invasions of freshwater. Another possibility is that Heterias may not be monophyletic. The distinctive male copulatory morphology, under such a hypothesis, would be homoplastic and uninformative about relationships. Further morphological study of this genus is therefore warranted to distinguish among these possibilities. Iais Bovallius, 1886 Iais Bovallius, 1886: 50-2. Jais Cltret, 1973: 32. Neoectias Winkler, 1992: 174. Type species: Jaera pubescens Dana, 1852. Species Included Jaera pubescens Dana, 1852: 744 (Atlas, 1855, plate 49, fig. 9a-d) (senior synonym of Iais hargeri Bovallius, 1886: 50, originally designated type); I. aquilei Coineau, 1977: 424; I. elongata Sivertsen & Holthuis, 1980: 103; I. solangeae Coineau, 1985: (ref. in Coineau, 1986: 468); Janiropsis californica Richardson, 1904~:223 (senior synonym of I. pubescens longistylis Chilton, 1912: 132); Iais singaporensis Menzies & Barnard, 1951: 136; Neoectias chilense Winkler, 1992: 174; Protocharon antarctica Chappuis, 1958: 16. Other References Iais. -Dollfus, 1891: 70; Stebbing, 1900: 549; Chilton, 1909: 649; Ortmann, 1911: 645-6; Barnard, 1914: 436; Vanhoffen, 1914: 530; Giambiagi, 1925: 17; Monod, 1926: 13, 1931a: 1; 1931b: 13; Stephensen, 1927: 356; Nordenstam, 1933: 177; Hale, 1937: 31; Kesselyak, 1938: 221; Nierstrasz, 1941: 287; Menzies, 1952: 135; Pillai, 1954: 19; Hurley, 1956: 715, Review of the Janiridae 701 1961: 262; Holdgate, 1961: 14, 19; Menzies, 1962b: 74; Wolff, 1962: 39, 236, 251, 276, 293; Barnard, 1965: 199; Kussakin, 1967: 308; Schultz, 1969: 269; Coineau, 1971: 134-5; Rotramel, 1972: 194; 1975: 21, 247; Iverson, 1974: 165; Miller, 1975: 149; Kensley, 1978: 135-6; Kussakin and Vasina, 1980: 361; Marsden, 1982: 233; Coineau, 1986: 468, 472; Kussakin, 1988: 135; Branch et al., 1991: 28; Miiller and Brusca, 1992: 73. Jaera.-Smith, 1876: 63; Beddard, 1886b: 19-20. Janiropsis. -Richardson, 1904b: 666; 1905b: 455. Protocharon. -Wolff, 1962: 38, 250. Diagnosis Diminutive isopods with pereonites often broadening posteriorly. Cephalon lateral margin rounded, not elaborate or spined; vertex convex but not rostrate; eyes dorsal, reduced to few separate ocelli. Tergites of all pereonites with slight rounded anterolateral projections on pereonites 1-4; all coxae visible in dorsal view emerging from posterior margin of each pereonite. Pleonite 1 dorsally separate, short, distinctly narrower than pereonite 7 and near same width (more than approximately 75%) of pleotelson. Pleotelson ovoid, small, narrower than pereonite 7, lacking deep incisions for uropods or denticles. Antennula article 1 much broader than article 2 but length greater than width, distally rounded; flagellum short, with 3 articles. Antenna basal articles (1-4) subequal, article 3 lacking scale; articles 5 and 6 somewhat longer than basal articles 1-4 but shorter than length of head; flagellum long, with 20-25 articles, proximal articles separate and longer than wide, not conjoint. Mandible slender but normal; molar process distally truncate, straight-sided or slightly tapering distally; palp large, near length of mandible, proximal article long and near length of distal article, distal article slightly curved and setose. Maxillipedal endite length near width, either wider or near width of palp article 2; palp articles 4-5 narrow and straight-sided, palp article 3 only slightly wider than palp articles 4-5 and distinctly narrower than broad palp articles 1-2. Pereopod I leg-like, with carpus and propodus straight-sided with few fine setae and no spine-like setae. Pereopods 11-VII stout, but ambulatory, dactyli with 3 claws, ventral claw near length and basal width of dorsal claw, accessory seta enlarged into third claw near length of ventral claw. Male pleopod I medial and lateral lobes short, distinctly separated and posteriorly projecting; lateral lobes distally pointed and subtriangular. Male pleopod I1 endopod proximal article near maximum width of stylet; stylet shorter than length of protopod, only slightly curving. Pleopod I11 endopod and exopod entirely lacking plumose setae; exopod thin, distally pointed, longer and narrower than endopod, biarticulate with suture at angle to body axis. Uropods short, distinctly shorter than pleotelson; protopod rectangular, near length of rami; exopod shorter than endopod, inserting apically. Remarks Like Jaera, the body of Iais tends to expand posteriorly, although the pleotelson is narrower, not wider, than the last pereonite. In the non-ectocommensal species, the pereonal expansion is much less pronounced. Several behavioural similarities (discussed below) with a species of Jaera suggest perhaps a close relationship between the two genera (see also cladogram in Vieulle 1979). Protocharon shares a general body form with Iais, although Protocharon (among other differences) has only two claws on the walking legs rather than three as in Iais. The nominal species, I. pubescens, is morphologically variable and broadly distributed in the subantarctic (many publications with Iais in the synonymy above contain illustrations of I. pubescens). This species may be a complex of species, requiring a detailed revision. If the Falkland Island species I. hargeri were found to be distinct from I. pubescens from Tierra del Fuego, the type of Iais would be I. hargeri, as indicated by Bovallius (1886). A study of Chappuis' (1958) description of Protocharon antarctica led us to the inescapable conclusion that this species is a member of Iais (see discussion of Protocharon). Coineau (1986: 472, footnote), in describing I. aquilei, agrees with this observation. A similar species, I. chilense, was originally presented as a new genus, Neoectias, but Winkler (1992) disregarded similarities of this species with I. aquilei. G. D. F. Wilson and J.-W. Wagele 702 Iais singaporensis Menzies & Barnard, 1951 was synonymised to I. californica (see Hurley 1956; Rotramel 1972) in recognition that the widespread sphaeromatid host, Sphaeroma quoyanum Milne-Edwards, 1840 might have carried its commensal worldwide on wooden ships. More recently, I. singaporensis has been redescribed and recognised as a distinct species (Miiller and Brusca 1992). Because I. pubescens longistylis Chilton, 1912 inhabits a variety of host sphaeromatids in New Zealand (Hurley 1956), the synonymy of this species with I. californica also may be in doubt. The species of Iais that live ectocommensally on large sphaeromatid isopods are found in most oceans. Interestingly, Jaera hopeana shares this habit. Exceptions exist: I. aquilei, I. antarctica and I. chilense were found interstitially at or near sea level (Chappuis 1958; Coineau 1977; Winkler 1992); specimens of 'I. pubescens' were found free-living in freshwater streams and pools above sea level (Chilton 1909; Holdgate 1961; Barnard 1965); I. elongata was found free-living in 'a pool of brackish water 2 m above the beach' (Sivertsen and Holthuis 1980:106). Australian Museum collections from Macquarie Island have three species of Iais that occur in all three environments mentioned above (G.D.F.W., unpublished observations). Mature males of I. aquilei were observed by Coineau (1977) to carry manca stage individuals with their specialised fourth pereopods. I. cf. pubescens from Macquarie Island also has this habit. Miiller and Brusca (1992) suggest that the reduced male pereopod IV may be diagnostic of the genus, although J. hopeana has a similarly specialised limb (Franke 1993). This behaviour may be an extended form of precopula (Wilson 1991). Females of small janiroidean species are functionally capable of receiving sperm via their spermathecal duct ('cuticular organ', see Vieulle 1977), prior to preparatory stage (Franke 1993; unpublished observations on Pleurocope and other genera). Sperm are held by the juvenile female until she enters the brooding stage. Therefore, an adult male's fitness would benefit if he were to hold a female prior to her becoming sexually receptive. Precopula is widely observed in isopods (Ridley 1983), and this behaviour is advanced to an earlier stage owing to the uniqueness of the janiroidean genital system (Wilson 1991). The manca-holding behaviour by males is precopula rather than offspring-holding as in the brooding females (Franke 1993). Manca-holding behaviour has been observed in other species of Iais (B. Kensley, personal communication), in Munna (R. Hessler, personal communication), and in Jaera hopeana (Veuille 1979; Franke 1993). This behaviour may be more common in janiroideans than previously realised. Ianiropsis Sars, 1897a Ianiropsis Sars, 1897a: 102. Janiropsis Richardson, 1904c: 221. Type species: Janira breviremis Sars, 1883. Species Included I. analoga Menzies, 1952: 141; Janira breviremis Sars, 1883: 64; I. chiliensis Menzies, 19626: 80; 1. epilittoralis Menzies, 1952: 149; I. kincaidi Richardson, 1904c: 221; Janiropsis derjugini Gurjanova, 1933: 82; I. koreaensis Jang & Kwon, 1990: 195; 1. kussakini Carvacho, 1982: 200; I. longiantennata Thielemann, 1910: 70; I. longipes Sivertsen & Holthuis, 1980: 109; I. magnocula Menzies, 1952: 145; I. minuta Menzies, 1952: 155; I. montereyensis Menzies, 1952: 152; Janira neglecta Chilton, 1909: 648; I. notoensis Nunomura, 1985a: 130; I. pallidocula Kussakin, 1962: 44; I. palpalis Barnard, 1914: 222; I. perplexus Menzies, 1962b: 78; I. picta Kussakin & Mezhov, 1979: 151; I. punctulata Kussakin & Mezhov, 1979: 153; Janiropsis serricaudis Gurjanova, 1936a: 251; I. setifera Gurjanova, 1950: 282; I. tridens Menzies, 1952: 156; I. sp. Nunomura, 19856: 109. Other References Ianiropsis.-Barnard, 1955: 75; Spasski, 1961: 296; Wolff, 1962: 39, 234, 235, 251, 254, 276, 287, 292, 300; Kussakin, 1967: 339 (English translation 1968: 341); George and Review of the Janiridae 703 Stromberg, 1968: 235; Miller, 1968: 24; Schultz, 1969: 261; Kussakin, 1972: 164, 1974: 258; Monod, 1974: 1133; Kussakin, 1975: 67; Kensley, 1976: 305-7; Kussakin, 1988: 88. Ianthopsis. -Wolff, 1962: 41, 251; Iverson, 1974: 165. Janira.-Stephensen, 1927: 355; Nierstrasz, 1941: 283; Hatch, 1947: 171; Hurley, 1961: 262; Kussakin, 1967: 339; 1968: 341. Janiropsis.-Richardson, 1905b: 456; Gurjanova, 1935: 30; 19368: 44; 1938: 231; Nierstrasz, 1941: 285; Hatch, 1947: 172; Gurjanova, 1950: 281; Kussakin, 1956: 114; Hauksson, 1980: 32; Nunomura, 1981: 44; Kensley, 1989: 148. Diagnosis Medium-sized, somewhat elongate janiroideans. Cephalon broad, laterally rounded or quadrate, lacking large spines; vertex smoothly convex, lacking rostrum; eyes dorsal. Body moderately broad, pereonites all near same length or widest medially; tergites of pereonites 2 and 3 laterally concave with reduced lappets; all coxae visible in dorsal view. Pleotelson broad, shield-shaped, laterally rounded, sometimes with small denticles on lateral margins anterior to shallow insertions of uropods. Antennula basal article only slightly broader than second article; flagellum multiarticulate, with around 5-25 articles (median 10 articles). Antenna elongate; basal articles 1-4 subequal, article 3 with distinct scale; articles 5-6 much longer than articles 1-4; flagellum multiarticulate, generally more that 20 articles; proximal flagellar articles narrower than antennular article 4, not conjoint, wider than long except most proximal article (generally longer than wide). Mandibles normal with truncate molar process and elongate palp; palp article 3 curved and setose. Maxillipedal endite longer than broad, narrower than broad palp; palp articles 4-5 thin, straight-sided, narrower than palp articles 1-3, in males of some species longer than antennulae; palp article 3 much wider than long, medial margin tapering from broadest point distally. Pereopod I of male extremely long; carpus elongate, with 2 rows of spine-like sensory setae, basis and ischium also much longer than in posterior legs; propodus with row of spine-like sensory setae. Pereopod I in female similar to more posterior walking legs. Pereopods 11-VII dactylus with 2 large subequal distal claws and third large subdistal accessory claw. Male pleopod I distal tip with medial lobes straight or only slightly rounded, merging smoothly into pointed lateral lobes with no separating notch; lateral lobes not curling medially. Stylet of male pleopod I1 not coiled and not longer than sympod. Pleopod I11 endopod with distinct gap between medial plumose seta and 2 lateral plumose setae; exopod of both sexes narrower and longer than endopod; exopod with 2 articles, distally rounded or pointed and lacking plumose setae, suture perpendicular to ramal axis. Uropods large, often near length of pleotelson (but short and squat in some species); rami and sympod narrow; exopod near length of endopod, inserting apically; protopod shorter than endopod. Remarks Ianiropsis is a broadly distributed shallow-water genus. Its features are similar to those of Janira and Carpias, including three claws on the walking legs, coxae visible in dorsal view, and small pereonal lappets in some species. Its most distinctive apomorphy is an elongate male pereopod I, although the carpus is not broad and spiny as in Carpias; the elongation is especially apparent in the basis and ischium. Moreover, the distal tip of the first pleopod of Ianiropsis males is very similar in most species: a broad nonprojecting medial lobe curving smoothly into the laterally pointed lateral lobes. In view of the variability of this feature in most other janirids, the constancy of the male pleopod I in Ianiropsis is somewhat surprising, although very useful. Sars (1897a) mentions that the antennulae of I. breviremis are different from those of Janira maculosa (they are shorter), but the simple length of antennulae and antennae cannot be trusted as a generic character. As in several other janirid genera, Ianiropsis can only be definitively identified from the males (see remarks under Janira). Ianiropsis bisbidens Barnard, 1955, placed in Janira by Carvacho (1981), has broad, leaf-like uropods, large third pleopods, and first male pleopods with long medial lobes and short but protruding lateral lobes. All these features indicate that this species is a member of the genus lathrippa. G. D. F. Wilson and J.-W. Wagele 704 The species Janira neglecta Chilton, 1909 was placed by Wolff (1962) into Zanthopsis, although the male first pereopod clearly has an elongated but not expanded carpus, an apomorphy of Zaniropsis. In addition, the figured pleotelson is more typical of Zaniropsis and its allies than of Ianthopsis, which typically has tubercles and thick spines on the dorsal and lateral surfaces. The species neglecta should be assigned provisionally to Zaniropsis. Menzies (1952) considered I. kincaidi Richardson and I. derjugini Gurjanova as subspecies of I. kincaidi. Separate morphological diagnoses, illustrations, and positions in a key were provided for each presumed subspecies, and Menzies even reported that each form lives in a separate part of the intertidal zone. He gave no evidence for substantial genetic introgression between the two taxa. Therefore, we consider these two taxa separate and distinct species. The sympatry of I. kincaidi and I. derjugini and the relationship between Monterey County (California) and Komandorski Islands (Bering Sea) populations deserves more study. Kussakin (1982) and Jang and Kwon (1990) also consider I. kincaidi and I. derjugini to be separate species but do not mention the sympatry problem of Menzies (1952). Several cryptic species may be classified as I. kincaidi. Two distinct groups are apparent within Zaniropsis. The palpalis-group has enhanced sexual dimorphism in the form of an elongate male maxillipedal palp; this group includes I. palpalis Barnard, 1914, 1. serricaudis Gurjanova, 1936, I. notoensis Nunomura, 1985a, and I. epilittoralis Menzies, 1952. Males of the palpalis-group have maxillipedal palps that protrude well beyond the basal articles of the antenna in dorsal view. The elongation is greatest in the fifth and fourth articles, although the third article is also much longer than wide. In most other Zaniropsis species, the maxillipedal third article is shorter than wide, on the basis of literature descriptions. I. epilittoralis was not described as having this enlarged male palp, although an inspection of specimens from San Diego, California shows that it, too, belongs to thepalpalis-group. If the enlarged male palp was omitted from other descriptions, this character may have a more widespread distribution among species of Ianiropsis. The second group, thepicta-group, includes two species by Kussakin and Mezhov (1979): 1. picta and I. punctulata. This group has a robust maxillipedal palp with short stocky distal articles. Maxilliped article 5 is shorter than wide, while descriptions of all other Ianiropsis species show article 5 being longer than wide. Zathrippa Bovallius, 1886 Zathrippa Bovallius, 1886: 3 1. Notasellus Pfeffer, 1887: 85. Jorina Nierstrasz, 1918: 134. Zantra (Zathrippa) Nordenstam, 1933: 278 (lapsus calami). Type species: Janira longicauda Chilton, 1884. Species Included Ianiropsis bisbidens Barnard, 1955: 75; Janira capensis Barnard, 1914: 220; Jorina chiliensis Nierstrasz, 1918: 134; Janira hirsuta Carvacho, 1981: 135; Stenetrium inerme Haswell, 1881: 479; Janira longicauda Chilton, 1884: 250; I. menziesi Sivertsen & Holthuis, 1980: 99 ( = I . chiliensis Menzies, 1962a: 70; see Sivertsen and Holthuis 1980); 1. multidens Menzies, 1962b: 72; Notasellus sarsi Pfeffer , 1887: 85; Notasellus trilobatus Richardson, 1910: 649; Janira tristani Beddard, 1886a: 105. Other References Iathrippa. -Hutton, 1904: 264; Nierstrasz, 1941: 286; Hurley, 1957: 17; Wolff, 1962: 39, 41, 234, 235, 277, 294, 300; Kussakin, 1967: 339; Schultz, 1976: 17; Kussakin and Vasina, 1980: 363. Zanira (Iathrippa). - Nordenstam, 1933: 172; Hurley, 1961: 262. Janira.-Beddard, 1886b: 6; Thomson and Chilton, 1886: 157; Thomson, 1888: 265; Stebbing, 1905: 49; Nierstrasz, 1918: 134; Tattersall, 1921: 200; Stephensen, 1947: 7; Barnard, 1955: 6; Wolff, 1962: 42, 251; Kensley, 1976: 303; 1989: 148. Review of the Janiridae 705 Notasellus. -Hodgson, 1902: 251; Richardson, 1906: 13; Hodgson, 1910: 49; Richardson, 1913: 17; Vanhoffen, 1914: 532; Tattersall, 1921: 202; Giambiagi, 1925: 16; Hale, 1937: 31; Schultz, 1976: 17; Kensley, 1978: 139; Kussakin and Vasina, 1980: 364; Kussakin, 1982: 89; Wilson, 1986a: 297; 1987: 264; Branch et al., 1991: 29. Stenetrium.-Haswell, 1882: 309; Whitelegge, 1889: 60; Bovallius, 1886: 21. Diagnosis Moderately broad janiroideans with setose dorsal surfaces. Cephalon broad, lateral margins rounded, without spines; vertex with small to elongate rostrum protruding from frons; eyes lateral, sometimes bulbous on distinct projections. Pereonites laterally concave with all coxae visible in dorsal view, small but distinct lappets present on pereonites 2-4. Pleonite 1 short, distinctly narrower than pereonite 7 and pleotelson. Pleotelson broad, laterally rounded and smooth, sometimes with small angular projections lateral to insertions of uropods; anal part of pleotelson demarcated by small rounded posterior projection. Antennula basal article only slightly broader than second article, with multiarticulate flagellum (more than 15 articles). Antenna with elongate, multiarticulate flagellum; basal articles 1-4 subequal, article 3 with large scale; articles 5-6 elongate, much longer than articles 1-4; proximal flagellar articles longer than wide, not conjoint. Mandibles normal with truncate molar process and elongate palp; distal article elongate and setose. Maxillipedal endite longer than wide; palp basally broader than endite; palp articles 4-5 elongate, straight-sided, narrower than palp articles 1-3. Pereopod I leg-like, enlarged i n male; carpus robust, setose, with 2 rows of spine-like sensory setae. Pereopod 11-VII dactylus with 2 large subequal distal claws and third large subdistal accessory claw. Male pleopods I and I1 not opercular, distinctly smaller than pleopod I11 exopod. Male pleopod I distal tip with distinctly separate medial and lateral lobes; lateral lobes pointing laterally and posteriorly; medial lobes distinctly longer than lateral lobes. Male pleopod I1 endopod proximal article somewhat larger than maximum width of stylet, not enlarged or inflated; stylet subequal to or longer than sympod. Opercular female pleopod I1 with distal indentation. Pleopod I11 endopod with distinct gap between medial seta (sometimes 2 medial setae) and 2 lateral setae; exopod of both sexes with 2 articles, distally rounded, lacking plumose setae; exopod of male broader and longer than endopod, opercular; exopod of female narrower and longer than endopod. Uropods large, with broad flattened rami; rami much longer than sympod; exopod shorter than endopod, inserting apically. Remarks Zathrippa, like some species of Jaera, retains the asellotan plesiomorphic structure of the pleopods in the adult male: pleopods I and I1 are much smaller than pleopod I11 and the latter is opercular (Wilson 1987). Other distinctive characters of the genus are the large uropods with flattened rami, a pointed or rounded rostrum, and usually large protruding eyes. Of these characters, the enlarged, generally leaf-like uropods are the most useful synapomorphy of Zathrippa species. Zathrippa has been subject to some confusion because a junior synonym, Notasellus, was named one year after the first species was described, and later workers tended to mix the classifications of the New Zealand and South American/Antarctic species. Nordenstam (1933) established the synonymy, although his concept is almost certainly too broad. The most recent, full discussion of the genus was by Schultz (1976), in which the species in the two genera were separated only by the number of claws on the pereopods: two claws in the former and three in the latter. Wilson (1986a, 1987) followed Schultz's (1976) classification in referring to I. sarsi as Notasellus sarsi. Nevertheless, the species assigned to these two genera are so similar that they should be classified together. The species reported to have two claws should be restudied; if they indeed do lack the third claw, then this must be considered a secondary reduction within the genus. Indeed, subantarctic specimens attributed to I. longicauda have been reported (but see below) to have 3 claws (Kussakin and Vasina 1980). I. inerme Haswell, 1881 has three claws, as an inspection of material in the Australian Museum collection has shown. The third claw in I. inerme is present as a large subterminal accessory seta and therefore might be not classified as a third claw by 706 G. D. F. Wilson and J.-W. Wagele previous workers; this may also be the case in I. longicauda. Sivertsen and Holthuis (1980) did not use Schultz's division of the species, and maintained the junior synonymy of Notasellus. These latter authors also created a new species name, I. menziesi, for the junior homonym I. chilensis Menzies, 1962a, not I. chilensis (Nierstrasz, 1918). Janira hirsuta Carvacho, 1981 belongs in Iathrippa, although the species has been drawn coarsely without a rostrum. In addition to its broad uropodal rami, which help define the species as Iathrippa, the male pleopod I11 is much larger than pleopod 11. As mentioned above, Ianiropsis bisbidens Barnard, 1955 belongs in Iathrippa. Kensley (1978) assigned Janira capensis Barnard, 1914 to Notasellus, which is here considered a junior synonym of Iathrippa. The species Iathrippa longicauda is subject to further confusion, in addition to the problem of the number of dactylar claws. This species, originally described from New Zealand, has been attributed to specimens collected from numerous subantarctic islands, as well as South America (see synonymy and distribution in Kussakin and Vasina 1980). This broad distribution and variety of habitats listed in the records suggests that I. longicauda is a complex of species. Jaera Leach, 1814 Iaira Meinert, 1877: 80. Joera Koehler, 1885: 1-2. Iaera Sars, 1897a: 103. Jaera Leach, 1814: 434. Jaeridina Milne-Edwards, 1840: 150. Metajaera Verhoeff, 1943: 283. Type species: Jaera albifrons Leach, 1814. Species Included Jaera albifrons Leach, 1814: 434; Janira nordmanni Rathke, 1837: 388; J. bocqueti Veuille & Kocatas, 1979: 597; J. caspica Kesselybk, 1938: 240; J. forsmani Bocquet, 1953: 225; J. hopeana Costa, 1853: 1; J. ischiosetosa Forsman, 1949: 455; J. istri Veuille, 1979: 208; J. italica Kesselybk, 1938: 238; Janira nordmanni Rathke, 1837 ( = Jaera nordmanni nordmanni Lemercier, 1960);J. nordmanni guernei Dollfus, 1889: 133; J. nordmanni illyrica Veuille, 1979; J. nordmanni massiliensis Lemercier, 1958b: 3688; J. nordmanni nordica insulana Veuille, 1976b; J. nordmanni occidentalis Veuille, 1979: 206; J. nordmanni illyrica Veuille, 1979: 206; J. nordmanni balaerica Jaume & Garcia, 1990: 85; J. nordmanni brevicaudata Jaume & Garcia, 1990: 81; J. nordica Lemercier, 1958b: 3688; J. petiti Schulz, 1953: 59; J. posthirsuta Forsman, 1949: 457; J. praehirsuta Forsman, 1949: 458; J. sarsi Valkanov, 1936: 3 14; J. schellenbergi Kesselyak, 1938: 232; J. sorrentina Verhoeff, 1943: 284; J. syei Bocquet, 1950: 1; ?Joera wakishiana Bate, 1865: 668. Other References Jaera.-Delage, 1881: 155; Sye, 1887: 1; Zirwas, 1910: 94; Hansen, 1916: 12; Monod, 1925: 238; Gurjanova, 1932: 21; Arcangeli, 1934: 280; Valkanov, 1938: 57; Strouhal, 1942: 150; 1954: 15; Forsman, 1944: 1; Verhoeff, 1949: 422; Birstein, 1951: 136; Margalef, 1952: 209; Karaman, 1953: 102; Margalef, 1953: 106; Staiger and Bocquet, 1956: 10; Renault, 1958: 63; Lemercier, 1958a: 86; 1960: 23; Wolff, 1962: 21, 40, 235, 276, 291, 297; LCcher, 1962: 561; Prunus, 1963: 2043; Bocquet and Prunus, 1963: 343; Gruner, 1965: 127; Haahtela, 1965: 309; Haahtela and Naylor, 1965: 367; Naylor and Haahtela, 1966: 209; 1967: 19; George and Menzies, 1968: 381; Schultz, 1969: 269; Kussakin, 1969: 427; 1988: 137; Steele and Steele, 1972: 205; Naylor, 1972: 52; Geldiay and Kocatas, 1972: 25; Schiecke, 1973: 154; Jones, 1974: 727; Bubel and Jones, 1974: 737; Monod, 1975: 240; Veuille, 1976a: 1235; 1976b: 1334-7; Pantoustier and Prunus, 1977: 361; Cariou, 1977: 6; 1985: 281; Veuille, 1978a: 300; 1978b: 391; Radu, 1979: 177; Ingolfsson, 1980: 97; Solignac, 1981: 387; Klochenko, 1986: 85; Sket, 1990: 44; Reboreda and Otero, 1990: 73; Franke, 1993: 65. Asellus. -Bate and Westwood, 1868: 320. Review of the Janiridae laera.-Sars, 18976: 298; Hansen, 1916: 12; Nierstrasz, 1941: 286. Joera. -Bate, 1866: 282. Oniscus. -Fabricius, 1780: 252. Metajaera.-Verhoeff, 1949: 419; Birstein, 1951: 139; Kussakin, 1988: 154. Diagnosis Body moderately broad, tergites subequal or becoming more broad posteriorly (in males). Cephalon lateral margin rounded, lacking spines; anterior margin with rounded vertex and rounded frons protruding between antennae; eyes dorsal, generally well developed. Pereonites broad, laterally rounded, coxae not visible in dorsal view. Pleonite 1 dorsally separate, short, distinctly narrower than pereonite 7 or pleotelson. Pleotelson same width as anterior pereonites (in males) or narrower; lateral margins of pleotelson without denticles; incisions for uropods merged into single posterior notch. Antennular article 1 flattened, squat but longer than wide; flagellum with 2-3 articles. Antenna basal articles (1-4) subequal, article 3 without scale; articles 5 and 6 longer than basal articles 1-4; flagellum with more than 20 articles; proximal articles separate and longer than wide, not conjoint. Mandibular molar normal distally truncate, parallel-sided or slightly tapering posteriorly; palp large, distal article curved and setose. Maxillipedal endite longer than wide; palp narrower or broader than endite; palp articles 4-5 straight-sided, narrower than broad palp articles 1-2; palp article 3 distally narrowing to width of palp articles 4-5, distinctly narrower than palp article 2. Pereopod I leg-like, carpus not enlarged, with no spine-like setae. Pereopod 11-VII dactylus with 3 claws, ventral claw near same size as dorsal claw, accessory seta enlarged into third claw. Male pleopod I broad, lateral lobes with projecting copulatory horns (laterally or ventrally); medial lobes various, but always distinct from lateral lobes. Male pleopod I1 proximal article of endopod width greater than width of stylet, but not enlarged and rounded; stylet elongate and curving, but not distally thin and coiled. Pleopod I11 endopod with no plumose setae; in males endopod narrower and shorter than exopod (especially when opercular); exopod without plumose setae, with 2 free articles at distinct angle to body axis. Uropods inserting adjacent to one another, reduced, distinctly shorter than pleotelson, with squat rami and sympod; exopod apex distinctly shorter than endopod; sympod longer than rami. Remarks Jaera has been known for nearly two centuries and, of all Janiroidea, probably has the best known behaviour, sexual biology and genetics. Living in shallow intertidal, brackish or freshwater habitats makes species of Jaera readily accessible to researchers. Some species are hardy and are easily cultured (Solignac 1981). The exact number of valid species is difficult to establish, as some species are polymorphic (i.e. the 'albifrons' complex or superspecies: Solignac 1981). Jaera nordmanni has several local races adapted to brackish or fresh water and Jazdzewski (1969) shows that some species can interbreed (e.g. J. syei with J. ischiosetosa). Veuille (1979) provides a useful review of some species, and their historical zoogeography around the Mediterranean. Although the variation observed in Jaera has prompted many authors to recognise numerous subspecies in larger superspecies, Veuille (1979) regards many subspecific forms as species. Veuille, however, also recognises several additional subspecies, such as the subspecific race insulana in J. nordmanni nordica (Veuille 1976b), which is listed here in the species list. Despite polymorphism at the species level, the genus Jaera is easily recognised. The body is broad, sometimes becoming more so posteriorly. The cephalon has lateral projections and the eyes are sessile on the dorsal surface. The broad rounded pleotelson has a posterior indentation that bears abbreviated uropods. The antennula is short with only a few articles, while the antenna can be as long as the pereon. Some species have the primitive configuration of the male pleopods as seen in Iathrippa: the pleopods I and I1 are smaller than pleopod 111, and the latter is more or less opercular (e.g. illustrations of J. nordmanni in Karaman 1953). This janiroidean plesiomorphy cannot be used as evidence for a close relationship between Jaera and Iathrippa. 708 G . D. F. Wilson and J.-W. Wagele Jaera has two recognised subgenera (Kussakin 1988) with all species except one in J. Jaera, and Jaera hopeana in subgenus Metajaera Verhoeff, 1943. J. Metajaera can be separated from J. Jaera by an indistinct incision in the pleotelson for the uropods. Veuille (1979: 197-8) additionally recognises four separate groups and provides characters for their recognition: 'groupe Atlantique', including the J. albifrons superspecies (J. albifrons, J. ischiosetosa, J. posthirsuta, J. praehirsuta, J. forsmani); 'groupe mCditerranCen', including J. italica, J. schellenbergi, J. nordmanni, J. nordica, J. massiliensis, J. bocqueti; 'groupe ponto-caspien', including J. sarsi, J. caspica, J. istri; and 'quatrikme groupe' of J. hopeana. Veuille's grouping implicitly recognises the existence of the subgenus Metajaera, as well as further plausible divisions of the genus. Veuille (1979) also suggests that Zais and Jaera share a close relationship. The phylogenetic analysis of the janirid genera (Wilson 1994) corroborates this hypothesis. Moreover, Jaera and Zais share, at least in Jaera hopeana, the habit of holding mancas with a specialised pereopod IV, a possibly useful synapomorphy. Franke (1993) provides a detailed description of this manca precopula and general mating behaviour for J. hopeana. Jaera, Zais and possibly others could be grouped under the available family name Jaeridae Stebbing (1905), once a phylogenetically based division of the Janiridae is determined. Janaira Moreira & Pires, 1977b Janaira Moreira and Pires, 1977b: 23. Type species: Janaira gracilis Moreira & Pires, 1977b. Monotypic. Other References Janaira. -Moreira and Pires, 1977a: 181; Pires, 1981b: 31; Miiller, 1989: 207. Diagnosis Body not especially broad, with of all tergites subequal. Cephalon broad, laterally quadrate; vertex anteriorly rounded, lacking rostrum; eyes dorsal, well developed. Pereonal tergites laterally rounded, coxae visible in dorsal view. Pleonite 1 short, narrower than pereon or pleon. Pleotelson somewhat narrower than pereonite 7, laterally rounded, lacking spines or denticles, with only slight notch at insertions of uropods. Antennula basal article longer than broad, only slightly broader than second article; flagellum with 7 articles. Antenna basal 4 articles subequal, article 3 with tiny scale; articles 5-6 elongate, much longer than basal articles 1-4; flagellum elongate, multiarticulate, having more than 30 articles, basal articles narrower than article 6, wider than long but not conjoint. Mandibles normal with truncate molar process and elongate palp, distal article curved and setose. Maxillipedal endite longer than broad, narrower than broad palp; palp articles 4-5 thin, straight-sided, narrower than palp articles 1-3; palp article 3 much wider than long, subrectangular. Pereopod I of male carpus enlarged, oval, with 2 rows of spine-like sensory setae; pereopod I in female similar to posterior walking legs; propodus of both sexes with row of spine-like sensory setae. Pereopods 11-VII dactylus with 2 large subequal distal claws and third large subdistal accessory claw. Male pleopod I distal tip with medial lobes straight, angling smoothly into posteriorly pointed lateral lobes; lateral lobes curling medially only slightly. Male pleopod I1 endopod basal article around same width as maximum width of stylet; stylet of male pleopod I1 much shorter than sympod; not elongate or coiled. Female pleopod I1 with distinct distal notch. Pleopod I11 endopod with 3 equally spaced plumose setae; exopod of both sexes narrower than and same length as endopod; exopod uniarticulate, distally rounded, lacking plumose setae. Uropods large, longer than pleotelson, with narrow rami longer than propodus; exopod inserting apically; protopod much longer than wide. Remarks Janaira is very similar to Janira, Carpias and Janiropsis. It differs from Janira by having broad maxillipedal palp articles 2-3, and a male first pleopod that is not distally bifurcate. Review of the Janiridae 709 Janaira has n o laterally protruding lateral lobes on the male pleopod I and the male pereopod I is not elongate as in Zaniropsis or Carpias. Despite the claims o f Moreira and Pires (1977b) t o the contrary, Janira does have three claws o n the walking legs as does Janaira, and the fourth pleopods o f both genera are very similar. As the membership o f some species in Janira is in doubt (especially Janira operculata W o l f f ,1962), the type o f the genus, Janira maculosa Leach, should be used for comparison with possibly new groups. T h e differences that remain between Janaira and Janira are subtle, and may not be sufficiently significant for a genus-level division. Nevertheless, Janaira should be retained until Janira is completely revised. Janira Leach, 1814 Janira Leach, 1814: 434. Oniscodes Berthold, 1828: 282. Oniscoda Latreille, 1829: 140. Henopomus Krdyer, 1849: 366. Asellodes Stimpson, 1853: 41. Ianira Meinert, 1877: 78. Type species: Janira maculosa Leach, 1814. Species Included Janira maculosa Leach, 1814: 434 (possible senior synonym o f Zanira hanseni Menzies, 1962a: 181); Asellodes alta Stimpson, 1853: 41; J. japonica Richardson, 1909: 114; J. denticulata Gourret, 1891: 34. Other References Janira.-Leach, 1815: 373; Rathke, 1843: 24; Dana, 1852: 716; Bate and Westwood, 1868: 338; Harger, 1879: 158; 1880: 321; Sars, 1883: 16, 64; Beddard, 1886a: 105; 1886b: 6 ; Norman, 1894: 280; Richardson, 1900: 300; 1901: 556; 1905a: 468; Rathbun, 1905: 44; Zirwas, 1910: 92; Dahl, 1916: 30; Monod, 1925: 241; Fee, 1927: 21; Nierstrasz and Stekhoven, 1930: 117; Gurjanova, 1932: 23; 1933: 399; Dons, 1935: 43; Hult, 1941: 40; Stephensen, 1943: 37; Hatch, 1947: 171; Stephensen, 1948: 74; Holthuis, 1949: 178; 1950: 13; 1952: 75; 1956: 107; Staiger and Bocquet, 1956: 4; Menzies, 1962b: 70; Barnard, 1962: 244; W o l f f , 1962: 39-43, 233-5, 277, 281, 289, 293, 300; Gruner, 1965: 122; Miller, 1968: 23; Schultz, 1969: 259; Carvacho, 1981: 131; Kussakin, 1988: 116. Asellodes.-Verrill, 1873: 437; Verrill, 1874a: 411, 502; 1874b: 350. Bagatus. -Monad, 1974: 1123. Henopomus. -Delage, 1881: 155. Zanira.-Bovallius, 1886: 24; Hansen, 1888: 190; 1916: 13; Sars, 1897a: 98; Nierstrasz, 1941: 282; Hurley, 1961: 262. Oniscoda. -Milne-Edwards, 1840: 15 1. Diagnosis Body broadest at pereonites 4-5. Cephalon broad, with rounded lateral margins lacking large spines; eyes dorsal; vertex anteriorly rounded or with small rostrum. Pereonites 2-4 laterally concave, coxae visible in dorsal view between reduced lappets. Pleonite 1 short, narrower than pereonite 7 or pleon. Pleotelson broad, near same width as pereonite 7 , laterally rounded, with small row o f denticles o n posterolateral margins; slight notch for insertions o f uropods. Antennula basal article longer than broad, only slightly broader than second article; flagellum with more than 20 articles. Antenna basal articles 1-4 subequal, article 3 with distinct scale; articles 5-6 elongate, longer than basal articles 1-4; flagellum multiarticulate, with more than 30 articles, basal articles length subequal t o or greater than width, articles not swollen or conjoint. Mandibles normal with truncate molar process; palp elongate, distal article slightly curved and setose. Maxillipedal endite longer than wide, width subequal t o palp; palp articles 4-5 narrow, straight-sided, distinctly narrower than broad articles 1-3, palp article 3 distally broad, setose, not tapering, near width o f article 2. 710 G . D. F. Wilson and J.-W. Wagele Pereopod I carpus of male enlarged, oval, with 2 rows of spine-like sensory setae; propodus with fine denticles and spine-like sensory setae on ventral margin; pereopod I of female leg-like. Pereopod I1 of male elongated, distinctly longer than pereopods I or 111, and distinctly longer than pereopod I1 of female. Pereopods 11-VII dactylus with 2 large subequal distal claws and third large subdistal accessory claw. Male pleopod I distal tip with distinctly separated medial and lateral lobes; lateral lobes curling medially. Male pleopod I1 endopodal basal article not enlarged, approximately same width as widest part of stylet; stylet not coiled, shorter than sympod. Female pleopod I1 rounded with slight notch on distal margin. Pleopod I11 endopod either with 3 equally spaced plumose setae or with distinct gap between distal seta and 2 lateral setae; exopod of both sexes narrower than endopod and same length as or longer than endopod; exopod uniarticulate, distally pointed, lacking plumose setae. Uropods large, with narrow rami and propodus; endopod inserting apically; protopod subequal to endopod. Remarks The diagnosis of Janira is based on the type species, Janira maculosa Leach, and includes features that separate it from other similar genera such as Ianiropsis or Carpias. Females of J. maculosa are essentially similar to those of Ianiropsis and Carpias. In these genera, the definitive genus-level characters are found on the adult males. Sexual dimorphism also is present in at least J. maculosa. Although the male first pereopod is only slightly larger than that of the female, the second pereopod is decidedly larger than pereopod 111. In females, both pereopods are approximately the same size. Assuming this difference holds for other species of Janira, the enlarged male pereopod I1 is a useful synapomorphy of this genus. Field workers should be aware that fully mature males are required to apply a genus name to a species collection of typical janirids. As can be seen from the synonymies of other janirids, many species were originally included in Janira. Wolff (1962: 41-2) began the process of removing many species to other genera, whose changes we recognise here. Carvacho's (1981) review, which generally followed that of Wolff (1962), included several species in Janira that belong elsewhere. Janira hirsuta Carvacho, 1981, J. capensis Barnard, 1914, J. tristani Beddard, 1886a and Ianiropsis bisbidens Barnard, 1955 all belong in Zathrippa (see above). J. tricornis (Krdyer, 1849) belongs in Janiralata. J. biunguicula Hooker, 1985, a species with an atypically broad rostrum, was described only briefly, although a comparison of this species with Microjanira dentifrons Schiecke & Fresi, 1970 shows that they are congeneric. J. biunguicula Hooker, therefore, is moved to Microjanira. The only known specimen of a deep-sea species from the Gulf of Panama, J. operculata Wolff, 1962, lacks the cephalon and the male pleopods; therefore, its generic position cannot be established at present. J. abyssicola Beddard, 1886a is an unfigured dubious species (Wolff 1962); this species should be genus incertae sedis until the types are redescribed. Wolff's (1962) synonymy of J. maculosa and J. hanseni (Menzies, 1962a) is provisionally accepted, although the study of variation in this species should be extended beyond the inspection of coloration and the male pleopods. Vanhoffen (1914) included a Janira sp. with little description; this species lacks dorsally visible coxal epimera, making its assignment to Janira unlikely. Nierstrasz (1941) mentions J. monodi, erroneously attributed to Nordenstam, although this species must be a nomen nudum because a careful literature search for this species has found no source (in agreement with Wolff 1962). In addition to the type species, we recognise the following species as belonging to Janira: J. alta (Stimpson, 1853), J. japonica Richardson, 1909, J. denticulata Gourret, 1891. J. alta has a rostrum of possibly varying lengths (Hansen 1916; Kussakin 1988), unlike J. maculosa (see Wolff 1962). J. japonica Richardson, 1909 has not been reviewed since being described with only two simple drawings. Monod (1925) suggested that J. denticulata Gourret, 1891 from the Gulf of Marseille may be identical with the Nordic species J. maculosa. J. denticulata has been overlooked by later authors. These actions limit the definition of Janira to those species with 3 claws on the walking legs, although the species remaining in the genus will need careful revision to establish the above assertions with certainty. It is ironic that the type genus of the Janiridae is so poorly known. Review of the Janiridae Janiralata Menzies, 1951b Janiralata Menzies, 1951b: 124. Janirata Nunomura, 1991 : 37-8. Type species: Janiralata davisi Menzies, 1951b. Species Included J. abberrantis Kussakin & Mezhov, 1979: 163; J. bifurcata Mezhov, 1981: 11; J. bilobata Kussakin & Mezhov, 1979: 158; J. bisinuata Kussakin, 1972: 156; Iolella chuni Thielemann, 1910: 72; J. davisi Menzies, 1951b: 124; Ianthe erostrata Richardson, 1899: 858; J. gurjanovae Kussakin, 1962: 25; J. hexadentata Birstein, 1970: 294; Tole holmesi Richardson, 1905a: 216; J. intermedia Mezhov, 1981: 43; J. koreaensis Jang, 1991: 64; J. kurilensis Kussakin, 1962: 31; J. microphthalma Kussakin, 1972: 159; J. modesta Mezhov, 1981: 16; J. obliterata Kussakin, 1972: 162; Janira occidentalis Walker, 1898: 280; J. ochotensis Kussakin, 1962: 28; J. pilosa Kussakin, 1962: 34; J. problematica Kussakin & Mezhov, 1979: 166; Janira pulchra Hansen, 1916: 19; J. rajata Menzies, 1951b: 128; J. rhacuraeformis Birstein, 1963a: 17; Iolella sarsi Richardson, 1905a: 467; J. serrata Birstein, 1963: 12; J. shiinoi Kussakin, 1962: 28; Janira solasteri Hatch, 1947: 172; Janira soldatovi Gurjanova, 1933: 81; Ianthe triangulata Richardson, 1899: 857; Henopomus tricornis Krdyer, 1849: 372; J. vitjaZi Kussakin, 1962: 32. Other References Janiralata.- Menzies, 1952: 119; Menzies and Barnard, 1959; 11; Kussakin, 1988: 25; Wolff, 1962: 41, 234-5, 254, 277, 291, 297; Miller, 1968: 24; Schultz, 1969: 265; Brandt, 1993: 130. Zanira. - Hansen, 1916: 17. Ianthe. -Richardson, 1899: 857; 1900: 299. Zanthopsis. - Wolff, 1962: 41; Brandt, 1991: 224. Jolanthe.-Ortmann, 1900: 39-40 (nomen nudum). Iolella. -Richardson, 1905b: 462; Nierstrasz, 1941: 283. Janira. -Richardson, 1905b: 472; Stafford, 1913: 183; Boone, 1920: 7; Stephensen, 1943: 37; Gurjanova, 1950: 281; Wolff, 1962: 41, 254; Schultz, 1969: 259-60. Janthe. - Stephensen, 1913: 70. Jolella. -Oldevig, 1917: 42. Tole. -Ortmann, 1901: 157. Diagnosis Body broad and flattened. Cephalon with dorsal eyes; lateral margins straight, with anterolateral points or spines; vertex obtusely pointed to rostrate. Lateral margins of pereonites 2-3 divided into 2 lobes by a deep cleft or a broad niche, coxae visible on dorsal view; pereonite 1 with coxal lappet dorsally visible in anterolateral border; coxae visible dorsally on posterior margins of pereonites 4-7, if at all. Pleonite 1 very short, only barely visible, much narrower than pereonite 7 or pleotelson. Pleotelson of varying outline, generally broad, near width of pereonite 7; usually with notch at insertions of uropod and denticles on lateral margins. Antennulae multiarticulate, with approximately 12-20 flagellar articles; antennal article 1 longer than wide. Antenna article 3 with conspicuous scale (exopod); basal articles 1-4 subequal or article 3 somewhat more robust than other basal articles; articles 5-6 long, distinctly longer than basal articles 1-4; flagellum multiarticulate, with more than 20 articles; basal articles of flagellum wider than long but not conjoint. Mandibular molar process truncate; palp long, article 3 elongate, curved and setose. Maxillipedal endite longer than wide, width subequal to palp; palp articles 4-5 narrow, straight-sided, distinctly narrower than broad articles 1-3, palp article 3 distally broad, setose, not tapering, near width of article 2. Pereopod I carposubchelate, dactylus short, with 2 claws; propodus on proximal inferior border serrated, carpus with 2 rows of short spine-like setae on ventral margin. Pereopod 11-VII dactylus with 2 subequal claws, with subdistal accessory seta, sometimes enlarged into third claw. Male pleopod I distal tips 712 G. D. F. Wilson and J.-W. Wagele laterally expanded, with projecting subtriangular lateral lobes, and broad setose medial lobes merging smoothly into lateral lobes. Male pleopod 11 distal tip of protopod blunt, setiferous; basal article of endopod only somewhat wider than maximum stylet width, but not inflated; stylet elongate, curved, longer than sympod. Female pleopod I1 broad, ovate, distally with median concavity. Pleopod I11 endopod with 3 plumose setae having distinct gap between medial seta and 2 lateral setae; exopod narrower and longer than endopod; exopod with 2 segments, with distal plumose setae. Uropods biramous, rami subequal; exopod inserting apically; sympod about as long as rami. Remarks on the 'Janiralata' Complex Janiralata Menzies, 1951b, is the most heterogeneous genus of the 'Janiridae'. A majority of its species have been described from the northern Pacific Ocean, primarily owing to the taxonomic efforts of Kussakin and his coworkers (see Kussakin 1988). According to Menzies (1952), the characteristic feature is the serrated proximoventral margin of the propodus of the first pereopod. This structure is unique to Janiralata and is its primary defining synapomorphy of the various forms included below. In Janira, serrations occur at the same place, but they are finer and extend over the whole propodus. Besides the single feature used by Menzies, some further common structures are present: the carpus of the first pereopod is elongate and oval, bearing an irregular row of sensory spine-like setae near the edge of the ventral margin on both sides. This type of carpus is also found in Janira, Ianiropsis and Iathrippa. The first coxa has an anterolaterally directed lobe visible in dorsal view and often shaped as the tergal lobe of the same segment. The sympod of the second male pleopod is broad and rectangular. The species of Janiralata partly resemble Iathrippa, Ianiropsis or Tole (senior synonym of Iolella). Janiralata may not be a monophyletic unit a* some species-groups may be related to Tole or the Acanthaspidiidae. This genus requires a careful revision to define genera and possibly families. The similarities of some species with Iathrippa deserve some comment: Iathrippa has, as do many species of Janiralata, a protruding rostrum, coxae that are visible in dorsal view, a first pereopod with two rows of sensory spine-like setae on the carpus. The serrations on the propodus are not present in Iathrippa. In comparison with Janiralata, Iathrippa has a more plesiomorphic, broad third pleopod and astonishingly small first male pleopods, thus clearly differing from Janiralata. In the following, Janiralata is limited to groups of species. All species-groups have pairs of lateral tergal lappets on the pereonites 2 and 3, which enclose the coxae (see Wagele, 1989: his fig. 40). As already mentioned, the lobe of the first coxa is similar in shape to the tergal lobe. All coxae have a ventrolateral notch, which in some species deepens to two separate coxal lobes on pereopods 11-IV. This morphology can be derived from the structures already present in Janira, Iathrippa, or Ianiropsis by enlargement of the tergal and coxal lobes. Only in the rajata-group (see below) are the coxae similar to those of Janira. Whether this situation is plesiomorphic or a secondary acquisition by reduction cannot be decided. (A) The rajata-group has a dorsal outline similar to that of Zaniropsis; i.e. compared with Iathrippa, the rostrum is reduced. Pereopod I and the male pleopod I1 show the typical features of Janiralata. Lateral coxal and tergal margins are formed as in Iathrippa. The pleotelson is distally rounded. The following species belong to this group: J. koreaensis Jang, 1991 J. microphthalma Kussakin, 1972 J. modesta Mezhov, 1981 J. obliterata Kussakin, 1972 J. rajata Menzies, 1951b (B) The tricornis-group. Here the rostrum is present (very acute in J. tricornis), the pleotelson is broader than in the foregoing group and the distal margin has shallow concavities above the insertion of the uropods. The species are: Review of the Janiridae J. erostrata (Richardson, 1899) J. kurilensis Kussakin, 1962 J. pulchra (Hansen, 1916) J. soldatovi (Gurjanova, 1933) J. tricornis (Krdyer, 1849) J. vitjazi Kussakin, 1962 Iolella alascensis Richardson, l9O5b and Tole libbeyi Ortmann, 1901 are junior synonyms o f J. tricornis (see Kussakin 1988). (C) The following species have a broad pleotelson with posterolateral points, a concave distal telsonic margin medial to the points, and a small convex medial apex. The uropods insert in the concave areas. A short rostrum can be present or reduced. (C.l) In this subgroup (solasteri-group) the rostrum is present, but it does not surpass the anterolateral point o f the cephalon. The lateral tergal lappets are well developed (exception: J. intermedia). The species are: J. bisinuata Kussakin, 1972 J. intermedia Mezhov, 1981 J. occidentalis (Walker, 1898) J. pilosa Kussakin, 1962 J. solasteri (Hatch, 1947) (?) J. triangulata Richardson, 1899 A species described by Richardson (1905b) as 'Iolella sarsi', which is not identical with Iathrippa sarsi (Pfeffer,1887), may also belong to this group. (C.2) The bifurcata-group. The tergal lappets are clearly longer than wide, the coxae o f pereopods XI-IV also have pairs o f long lateral lobes. The group includes those species having a cephalon with a blunt anterolateral horn-like spine in the species J. holmesi (Richardson, 1905b) J. bifurcata Mezhov, 1981 and those with two anterolateral horns in the species J. bilobata Kussakin & Mezhov, 1979 J. problematica Kussakin & Mezhov, 1979 J. problematica has the same shape as found in the genus Rhacura Richardson, 1908. In Rhacura, only the outline o f the body has been illustrated, and Richardson's (1908) description lacks the information that would allow a taxonomic assessment o f their relationships (see Rhacura section below). ( D ) The following species bear plumose setae on the exopod o f pleopod 111, and the pair o f lateral coxal lobes on pereopods XI-IV have an anterior lobe that is twice as long as the posterior. (D.l) The gurjanovae-group. Without rostrum, pleotelson distally much narrower than proximally, with two apical notches above the insertion o f the uropods. The species are: J. gurjanovae Kussakin, 1962 J. ochotensis Kussakin, 1962 J. serrata Birstein, 1963a (D.2) The davisi-group. Rostrum small or absent, coxae not visible in dorsal view, lateral tergal lappets broadened, pleotelson wider than long, distally rounded. Much o f the diagnosis above was derived using J. davisi (type species o f the genus!). The species are: J. davisi Menzies, 195 1 b J. chuni Thielemann, 1910 The genus Janthura W o l f f , 1962 has the same body form as does Janiralata davisi. The genus contains deep-sea species [Janthura abyssicola W o l f f , 1962, J. bougainvillei (Birstein, 1963a)l which are more specialised than Janiralata davisi: eyes lacking, the molar 714 G . D. F. Wilson and J.-W. Wagele process acute, the exopod of pleopod 3 with no setae (plesiomorphic state?). Nevertheless, Janthura could be closely related to J. davisi, because the broad tergal lappets are present on pereonites 2 and 3. (D.3) The hexadentata-group. The following species have three pairs of tergal lappets instead of two as in the former groups, the third pair is present on the fourth pereonite. The rostrum is longer than in J. tricornis, the margin of the pleotelson has four lateral points and, between these, shallow concavities. In dorsal view only one coxal lobe is visible. The species are: J. rhacuraeformis Birstein, 1963a J. hexadentata Birstein, 1970 J. aberrantis Kussakin & Mezhov, 1979. The group D.3 is very similar to Tole Ortmann. The latter only lacks the lateral points on the pleotelson, and the outline of the pleotelson is more similar to the species of group D.1. Furthermore, Tole bears dorsal spines on the tergites and has no plumose setae on the exopod of pleopod 111. Tole shares the typical feature of group D, namely three pairs of tergal lappets and coxae that insert more ventrally than in other species of Janiralata; the coxae have small or reduced caudal lateral lobes. Group D could as well be the sister group of the Acanthaspidiidae, which have three pairs of lappets and plumose setae on the exopod of pleopod 111. The unusual polymorphism of the genus Janiralata can be explained partly with the parallel migration into deep water observed in some of the species-groups. Shallow-water species such as J. rajata (group A), J. solasteri (C.l), J. gurjanovae (D.l) or J. davisi (D.2) have a smooth outline of t h e body and larger eyes. Species from deep water can be recognised by their longer tergal and coxal lobes, with variation in the telson shape (especially in the groups D.l and D.3). Janiralata species possibly have a common origin within the more primitive 'Janiridae', possibly providing a link to deep-sea families that have a morphology similar to Tole. Janthura Wolff, 1962 Janthura Wolff, 1962: 45. Austroniscoides Birstein, 1963b: 828. Type species: Janthura abyssicola Wolff, 1962. Species Included J. abyssicola Wolff, 1962: 46; Austroniscoides bougainvillei Birstein, 19636: 828. Other References Janthura.-Wolff, 1962: 34, 40, 236, 279, 281, 290; Menzies and George, 1972: 95; Siebenaller and Hessler, 1977: 18. Diagnosis Body broad and flattened, all pereonites width subequal. Cephalon lateral margins nearly straight, tapering anteriorly, setose, lacking spines; vertex broadly concave, not rostrate; eyes absent. Lateral margins of pereonites 2-3 divided into 2 lobes by cleft; lateral margin of pereonite 1 bilobed, consisting of coxal and pereonal lappet; coxae not visible in dorsal view on other pereonites. Pleonite 1 short, much narrower than pereonites or pleon. Pleotelson wider than long, narrower than pereonites; margins sinuate above insertions of uropods, lacking denticles or spines. Antennula short, with few articles; article 1 more or less tubular, length decidedly greater than width, tapering distally; flagellum with 3 articles. Antenna1 basal articles 1-4 subequal, article 3 bearing scale; articles 5 and 6 longer than basal articles 1-4; flagellum with greater than 20 articles, proximal articles separate and longer than wide, not swollen or conjoint. Mandibular molar process reduced, tapering distally with few setae; incisor process slender; palp longer than body of mandible, distal Review of the Janiridae 715 article elongate, slightly curved, setose. Maxillipedal endite narrow, narrower than second joint of palp; palp articles 4-5 broad, near width of endite, marginally curved; palp article 3 short, distally tapering, narrower than broad articles 1-2. Pereopod I leg-like; carpus rectangular, lacking spine-like sensory setae; propodus slender, lacking spine-like sensory setae; dactylus short. Pereopod 11-VII dactylus with only 2 distal claws, ventral claw distinctly smaller than dorsal claw; dactylus lacking accessory seta. Male pleopod I lateral lobe pointing laterally; medial lobe expanded, rounded. Male pleopod I1 sympod broad; endopod elongate, longer than protopod, curved, styliform; proximal article of endopod inflated, distinctly broader than stylet. Female pleopod I1 rounded with small rounded protrusion distally. Pleopod 111 endopod with 3 plumose setae having distinct gap between medial seta and 2 lateral setae; exopod width subequal to endopod, length greater than endopod, distally rounded, with 2 segments, lacking distal plumose setae, articulation between segments of exopod laterally indented, proximal article of exopod with distinct posterolateral lobe. Uropods elongate, longer than pleotelson, with thick sympods; exopod inserting apically, length near endopod length; sympod distinctly shorter than rami. Uropods biramous, branches elongate, subsimilar. Remarks Janthura is similar in body form to some species of Janiralata (see discussion of the Janiralata davisi group above). Species of Janthura, however, have specialised mouth parts and an unusually elongate stylet on the male pleopod 11. The last feature is seen also in Neojaera, Ectias and Heterias. Menzies and George (1972) assigned Austroniscus bougainvillei Birstein, 1963b to Janthura; Siebenaller and Hessler (1977), who revised the Nannoniscidae, agreed with this synonymy. This genus is known only from the deep sea with a depth range of 2250-9034 m. Mackinia Matsumoto, 1956 Mackinia Matsumoto, 1956: 1219. Type Species: Mackinia japonica Matsumoto, 1956. Species Included M. birsteini Henry & Magniez, 1991: 223; M. continentalis Birstein & Ljovushkin, 1965b: 1006; M. coreana Matsumoto, 1967; M. japonica dilatata Matsumoto, 1967; M. japonica japonica Matsumoto, 1956: 1219; M. troglodytes Matsumoto, 1967. Other References Mackinia.-Birstein, 1961: 137; Wolff, 1962: 36, 38, 236, 250, 276, 281, 290; Coineau, 1986: 468. Diagnosis Body elongate, flattened, more than 4x longer than broad, pereonite widths subequal or slightly broadening posteriorly. Cephalon lateral margins rounded, lacking spines; vertex concave, lacking rostrum; eyes absent. Lateral margins of pereonites rounded, without lappets, coxae not visible in dorsal view. Pleonite 1 short, narrower than pereonite 7 or pleon. Pleotelson ovate, narrower than pereonite 7 but width subequal to pereonite 1 width; margins rounded, not denticulate, lacking notches near uropods. Antennula short, flagellum with 4 articles; article 1 more or less tubular, length decidedly greater than width. Antenna1 basal articles 1-4 length subequal, article 3 with scale; articles 5 and 6 longer than basal articles 1-4; flagellum with more than 20 articles; proximal articles separate and longer than wide, not conjoint or swollen. Mandibular molar process thin, parallel-sided, distally truncate, with broad grinding surface; palp large, longer than body of mandible, distal article curved and setose. Maxillipedal endite longer than wide, narrower than palp; palp articles 4-5 narrow, straight-sided; palp article 3 distally tapering, basally near width of palp article 2. Pereopod I leg-like, as slender as pereopods 2-7, carpus elongate, with few G. D. F. Wilson and J.-W. Wagele 716 spine-like setae; propodus with spine-like setae; dactylus short. Pereopod 11-VII dactylus with only 2 distal claws and no accessory seta, ventral claw distinctly smaller than dorsal claw. Male pleopod I lateral lobe reduced, stopping at lateral margin of medial lobes; medial lobes produced posteriorly, distally semicircular. Male pleopod I1 sympod distally produced, rounded; proximal article of endopod broader than maximum width of stylet but not inflated or rounded; stylet shorter than protopod, thin, with distinct angle in distal part. Female pleopod I1 rounded with slightly concave distal tip. Pleopod I11 endopod lacking plumose setae; exopod broader and longer than endopod; exopod broadly rounded distally, with 2 segments, lacking plumose setae. Uropods biramous, longer than or subequal to pleotelson; exopod shorter than endopod, inserting proximal to apex; sympod elongate, subequal to endopod. Remarks This dwarfish janiroidean has been excluded from the Microparasellidae because it lacks an enlarged pleonite 1 (see below) and the mandible still conserves a grinding surface on the molar process. It was once classified in the now disused Microjanirinae Birstein, 1961. Most differences in Mackinia species from more typical janirids are simply reductions for the interstitial habit. Mackinia might be related to Heterias: their body forms are somewhat similar and their pleopods lack plumose setae. The phylogenetic analysis of the janirid group (Wilson 1994) supports some relationship between these two genera. This genus is currently known only from fresh ground-waters of Japan, Korea and the far east of Russia. Microjaera Bocquet & Ltvi, 1955 Microjaera Bocquet and LCvi, 1955: 118. Type species: Microjaera anisopoda Bocquet & Levi, 1955. Monotypic. Other References Microjaera. - Spooner, 1959a: 1696; Birstein, 1961: 137; Wolff, 1962: 35, 38, 277, 281, 290; Naylor, 1972: 60; Schiecke, 1973: 152; Coineau, 1986: 468; Kussakin, 1988: 183. Diagnosis Dwarfish isopods; body about 6 x longer than wide, width of all pereonites subequal. Cephalon quadrate, lateral margin straight-sided, lacking spines; eyes absent; bases of antennulae separated by short broadly rounded rostra1 process. Pereonites laterally rounded or straight-sided, lacking spines or lappets; coxae not visible in dorsal view. Pleonite 1 short, only somewhat narrower than pereonite 7 or pleotelson. Pleotelson ovate, narrower than pereonite 7, with a terminal spine-like process, margins lacking denticles or notches near uropods. Antennula short, flagellum with 2 articles; article 1 more or less tubular, length greater than width, shorter than article 2. Antenna1 basal articles (1-4) all subequal, article 3 lacking scale; articles 5 and 6 longer than basal article; flagellum with 12 short articles, proximal articles wider than long, near width of article 6 but not conjoint and swollen. Mandibular molar process without grinding surface, distally tapering to rounded setose tip; lacinia mobilis on left mandible conspicuous; palp shorter than body of mandible but still functional; distal article of palp curved and setose. Maxillipedal endite longer than wide, narrower than article 2 of palp; palp articles 1-3 enlarged; article 4 long, narrow, straight-sided; article 5 much shorter than long article 4; article 3 broad, not distally tapering. Pereopod I stouter than pereopods 11-VII but still leg-like; carpus with few spinelike setae; dactylus short, with 2 unequal claws, dorsal claw as long as dactylus. Pereopod 11-VII dactylus with only 2 distal claws, ventral claw near same size as dorsal claw. Male pleopod I lateral lobes elongated into copulatory horns; medial lobes distally flattened, merging directly into lateral lobes. Male pleopod I1 sympod oval; stylet very slender, long, curved, much longer than sympod (basal article of endopod not described). Female operculum broad, distal margin broad, slightly concave. Pleopod 111 endopod with only single medial plumose seta; exopod narrower than endopod, length subequal to endopod Review of the Janiridae 717 length, uniarticulate. Uropods biramous, endopod 3 x longer than exopod; exopod inserting apically; syrnpod shorter than exopod, not visible in dorsal view. Remarks A terminal spine on the pleotelson is a useful autapomorphy of Microjaera anisopoda. Otherwise this species is similar to the other diminutive interstitial janirids that lack the more extreme characteristics of the Microparasellidae. The broadly rounded rostrum on the head appears in other taxa, such as Xostylus or Caecianiropsis, although the details of its form are uncertain. Microjaera is most similar to Caecianiropsis in antenna1 form, body form, and possibly in the male second pleopods, although the last character is poorly illustrated in this species. Microjanira Schiecke & Fresi, 1970 Microjanira Schiecke and Fresi, 1970: 243. Type species: Microjanira dentifrons Schiecke & Fresi, 1970. Species Included M. dentlfrons Schiecke & Fresi, 1970: 243; Janira biunguicula Hooker, 1985: 268. Other References Microjanira.-Schiecke, 1973: 149; Coineau, 1986: 468. Diagnosis Dwarfish isopods, body flattened, roughly elliptical, somewhat less than 3 x longer than wide. Cephalon laterally rounded with broadly pointed anterolateral margin lacking denticles; apex with long and broad, apically truncate rostra1 projection separating bases of antennulae; eyes absent or dorsolateral with few ocelli. Pereonites 2-4 laterally rounded or concave; coxae on all pereonites visible in dorsal view. Pleonite 1 very short, narrower than pereonite 7 or pleotelson. Pleotelson ovate, distal margin rounded, margin lacking denticles or notch near uropods. Antennula short; article 1 enlarged, length near width; flagellum with 3-4 articles. Antenna1 basal articles (1-4) lengths subequal; article 3 with scale; articles 5 and 6 longer than basal articles 1-4; flagellum probably multiarticulate, proximal articles separate, longer than wide. Mandibular molar process distally tapering, truncate; incisor process and body of mandible slender; palp large, near length of mandibular body, distal article strongly curved and setose. Maxillipedal endite longer than wide, narrower than palp; palp article 2 of maxillipedal palp expanded, much broader than articles 1 or 3; articles 4-5 thin, straight-sided; article 3 tapering distally, distinctly narrower than article 2. Pereopod I leg-like, with thin, elongate carpus lacking spine-like sensory setae, dactylus short. Pereopod 11-VII dactylus with only 2 distal claws, ventral claw smaller than dorsal claw; accessory seta absent. Male pleopod I distally truncate, not laterally expanded; medial lobes setose and rounded; lateral lobes merged into medial lobes. Male pleopod I1 sympod distally tapering; proximal article of endopod near maximum width of stylet; stylet shorter than protopod, distal tip narrows abruptly to thin point. Pleopod 111 endopod broader than exopod, length subequal to exopod, distal tip rounded with 3 plumose setae, with gap between medial seta and lateral 2 plumose setae. Uropods thin, shorter than length of pleotelson; sympod slender, nearly as long as exopod; exopod shorter than endopod, inserting apically on sympod. Remarks The cephalon of Microjanira is Austrofilius-like, although the thin, somewhat elongate uropods of this genus set the two taxa apart. As discussed above, Janira biunguicula Hooker, 1985 also belongs in Microjanira. The family Microjaniridae Birstein, 1961 is a nomen nudurn because Microjanira was not described until 1970, and the proposed family included only the genera Microjaera, Caecianiropsis, Protocharon and Mackinia. If this 718 G . D. F. Wilson and J.-W. Wagele genus is moved to a new family grouping not included in one of the recognised families, the reviser will have the choice of using some name other than 'Microjaniridae' if desired (recommended). Neojaera Nordenstam, 1933 Neojaera Nordenstam, 1933: 188. Ianisera Kensley, 1976: 299. Type species: Jaera antarctica Pfeffer, 1887. Species Included Jaera antarctica Pfeffer, 1887: 94; Neojaera caeca Kussakin & Vasina, 1984: 12; N. elongata Menzies, 1962b: 74; Ianisera expansa Kensley, 1984a: 275; N. hirsuta Sivertsen & Holthuis, 1980: 107; Jaera octodentata Vanhoffen, 1914: 529; ?Austrofilius pallidus Kussakin, 1982: 82; Jaera pusilla Barnard, 1925: 404; Jaera serrata Barnard, 1914: 433; Ianisera trepidus Kensley, 1976: 299. Other References Neojaera. -Kesselyhk, 1938: 221, 247; Nierstrasz, 1941: 287; Wolff, 1962: 234, 235, 277, 294, 300; Kussakin, 1967: 306, 340; Monod, 1974: 1133; Arnaud, 1974: 562, 646; Schultz, 1976: 24; Siebenaller and Hessler, 1977: 20. Zanisera.-Branch et al., 1991: 28. Jaera. -Nordenstam, 1930: 550. Diagnosis Body flattened, elongate, at least 3.5 x longer than wide. Cephalon anterior margin rounded or sinuate, lacking rostrum so that bases of antennulae not separate medially; eyes small and dorsal, sometimes absent; lateral margins often denticulate with anterior points. Pleonite 1 narrower than pleotelson or anterior pereonites. Pleotelson narrower than anterior pereonites, lateral margin often with denticles; posterior margin slightly sinuate above insertion of uropods. Antennula article 1 broad, wider than long, with anteromedial lobe, flagellum with 2-3 articles. Antenna with scale (exopod), flagellum with 20 or more articles, proximal articles longer than broad, not conjoint. Mandibular molar process truncate with broad grinding surface; palp near length of mandibular body, distal article curved and setose. Maxillipedal endite length subequal to width, width subequal or broader than palp, palp articles 1-3 expanded, about as broad as endite; article 3 not tapering; palp article 4 straight-sided; palp article 5 shorter than article 4, rounded. Pereopod I leg-like; carpus elongated, rectangular, with no spine-like setae; propodus slender, with spine-like setae; dactylus short with 2 claws. Pereopods 11-VII dactyli with 2 distal claws, ventral claw smaller than dorsal claw; accessory seta absent. Male pleopod I medial lobes rounded; lateral lobes sometimes prolonged; proximal region of pleopod I extremely broad and rounded, not straight. Male pleopod I1 with exopod inserting subapically on sympod; endopod with long, often coiled slender stylet and greatly expanded proximal article. Female pleopod I1 outline circular. Pleopod I11 endopod with 3 plumose setae having distinct gap between medial seta and 2 lateral setae; exopod narrower than and around same length as endopod; exopod with 2 segments, lacking plumose setae. Uropods short, biramous, often emerging from notches in pleotelson; protopod short, squat, shorter than rami; exopod inserting apically. Remarks See Austrofilius (above) for part of the remarks for this genus. We regard Neojaera to be fairly heterogeneous because it contains both broad and thin forms. All species seem to have distinctive male pleopods: pleopod I expanded and rounded laterally; and pleopod I1 with a narrow protopod, an inflated proximal article of the protopod and an extremely long, coiled stylet. Some characters also are found in Austrofilius and Caecianiropsis, but the Review of the Janiridae 719 absence of a rostrum makes Neojaera distinct. Ianisera Kensley, 1976 was later suggested by Kensley (1984~)to be congeneric with Neojaera although no name changes were made. A comparison of published descriptions of both taxa shows that this synonymy is indeed valid. In fact, Ianisera trepidus Kensley is similar to N. antarctica (Pfeffer) as described by Vanhoffen (1914). More recently, Kensley (in Branch et al. 1991) continued to use the name Ianisera. Further study of the type specimens is probably needed. Kussakin (1982) described Austrofilius pallidus as lacking a rostrum; this species therefore is a Neojaera. N. caeca Kussakin & Vasina (1984) has decidedly Janiralata-like characteristics, although the crude drawing of the male pleopods indicates that this species should be classified in Neojaera. Protocharon Chappuis, Delamare-Deboutteville & Paulian, 1956 Protocharon Chappuis, Delamare-Deboutteville and Paulian, 1956: 58. Jehaia Wagner, 1990: 187. Type species: Protocharon arenicola Chappuis, Delamare-Deboutteville & Paulian, 1956. Species Included Protocharon arenicola Chappuis, Delamare-Deboutteville & Paulian, 1956: 58; Jehaia stocki Wagner, 1990: 187. Other References Protocharon.-Chappuis and Delamare-Deboutteville, 1960: 354; Birstein, 1961: 137; Wolff, 1962: 35, 38, 250, 276, 281, 290; Schiecke and Fresi, 1970: 249; Monod, 1975: 1010; Coineau, 1986: 472. Diagnosis Tiny isopods about 4 x longer than wide, all pereonites subequal width. Cephalon rounded, lacking rostrum, vertex straight; eyes reduced to tiny dorsal spots. Pleonite 1 short or not visible. Pleotelson ovate, lateral margins denticulate, distal margin truncate, width subequal to pereonites width; uropods inserting directly on posterior margin. Antennula first article broad, longer than wide; flagellum with 3-4 articles. Antenna1 [known only in P. stocki] basal article 1 fused or absent; articles 2-3 more than twice as broad as distal articles; article 3 with short antenna1 scale; articles 5-6 shorter than article 3, flagelliform; flagellum with 8 articles. Mandibular molar process thin, distally rounded and setose, without grinding surface; mandibular body slender; palp elongate, distal article slightly curved with few distal setae. Maxilliped endite much broader than palp, distally quadrate with row of large rounded setulose setae (modified fan setae); palp slender, all segments longer than wide, widths subequal. All pereopods similarly slender; dactyli with 2 claws, ventral claw near same size as dorsal claw. Male pleopod I tapering posteriorly; medial lobe on distal tip projecting and rounded, paucisetose; lateral lobe rounded, protruding only slightly from margin of pleopod. Sympod of male pleopod I1 broad, laterally oval, tapering distally, stylet much shorter than sympod. Female operculum broad, subcircular or oval. Pleopod I11 narrow, distally rounded, endopod lacking plumose setae; exopod narrower and longer than endopod; exopod uniarticulate, distally rounded, lacking plumose setae. Pleopod VI rudimentary and pleopod V absent [in P. stocki]. Uropods near length of pleotelson, or shorter; sympod near length of endopod; exopod shorter, inserting subapically on sympod. Remarks This tiny freshwater janirid genus has previously been placed either in the Janiridae (Wolff 1962) or the Microjaniridae (Birstein 1961); this is discussed further with respect to the Microparasellidae below. Protocharon is possibly related to Mackinia. The concept of Protocharon is here restricted to P. arenicola and P. stocki because Protocharon antarctica Chappuis, 1958 has different mouth parts, and should not be in the same genus. Wagner (1990) erected the new genus Jehaia because he believed that the differences in chaetotaxy 720 G . D. F. Wilson and J.-W. Wagele of the antennae, the free pleonite 1 and outline of the uropods are substantially different from Protocharon. The synapomorphic similarities of the antennae, the mouthparts and the male pleopod I in both taxa argues for their monophyly. Consequently, we here subsume Jehaia under Protocharon. Taking a cue from Coineau (1986, footnote on p. 472), we re-inspected the original description of Protocharon antarctica. Typical of many interstitial workers' papers, Chappuis did not illustrate the whole animal, providing only a scattered selection of rather poor limb illustrations. His illustrations, however, reveal mouth parts that are clearly similar to those of Iais aquilei and I. chilense, and unlike those of P. arenicola. Although Chappuis (1958) states that the pereopods have only two claws, his illustration of pereopod VII in the species antarctica appears to have an over-large terminal seta, unlike any found on most janirids; this could be the tip of a subterminal third accessory seta or claw. Furthermore, he notes that the pereopod IV of the male displays some secondary sexual differences (Chappuis, 1958: 18). Iais aquilei also has dimorphic pereopods IV: in the male, it is shorter than the other pereopods, and it lacks the subdistal third claw (Chappuis et al. 1956); the fourth pereopod of the females resembles its neighbours. This dimorphism may be a synapomorphy of the Iais group. Therefore, we remove Protocharon antarctica to Iais, pending a thorough revision of Protocharon and all species of Zais. The species Austrofilius arnaudi (genus incertae sedis) may be related to Protocharon or Zais, as suggested in the Austrofilius discussion, but some illustrated features of A. arnaudi conflict with the published figures of P. arenicola. Protocharon has enlarged fan setae on the distal tip of the maxillipedal endite, which also apparently occurs in Iais (e.g. I. aquilei Coineau, 1977). These maxillipedal setae cannot be used as a diagnostic character because the endite is poorly illustrated in most groups. The similarity of body form between Iais and Protocharon is at least suggestive of a relationship, although the two genera have differing numbers of claws on the pereopods and rather different mouthparts overall. Family MICROPARASELLIDAE Karaman, 1934 Provisional Diagnosis Tiny janiroideans with slender body, pleonite 1 free and width subsimilar to pereonites; all somites approximately same width. Eyes and chromatophores absent. Flagellum of antennula with maximally 4 articles. Mandibular molar process distally pointed, without grinding surface, with several terminal setae. First pereopod similar to walking legs, without subchela. Pereopods II-VII with 3 dactylar claws, accessory seta or claw 3 often reduced or absent. Pleopod I11 endopod with 3 plumose setae (often reduced or absent), exopod slender. Exopods of uropods, when present, inserting subapically on sympod. Anus terminal, not covered by pleopods. History and Composition of the Microparasellidae The family Microparasellidae was erected by Karaman (1934) to include the genera Microparasellus Karaman, 1933 and Microcharon Karaman, 1934. The family is characterised by free first pleonites, eyes absent, a short first antenna, a second antenna shorter than the body, furnished with a scale on article 3, and pereopods with two claws. Wolff (1962) compared features of this family with those of other slender janiroideans, namely the Thambematidae Stebbing, 1912 and the Microjanirinae Bocquet & L h i , 1955. The Thambematidae are a specialised group of deep-sea isopods with only one large claw on pereopods II-VII and with no closer affinities to the Janiridae. The status of the Thambematidae as a separate family is generally accepted (see also Harrison 1987). The Microjanirinae have two claws, a subcylindrical molar process, a slightly differentiated or undifferentiated first pereopod and a male pleopod I which is narrower at its base. Birstein (1961) accepted this division with the Microjanirinae as a separate family containing the genera Microjaera, Caecianiropsis,Protocharon and Mackinia. Wolff (1962), on the other hand, believes that the Microjanirinae and Microparasellidae cannot be Review of the Janiridae 72 1 separated from the Janiridae, arguing that their defining features are reductions and adaptations to the interstitial habitat that might evolve independently several times in different taxa. The common features, reduction of the eyes and the slender body, are not enough to define the families. Kussakin (1988) follows Wolff (1962) in not recognising the Microparasellidae. This criticism is in conformity with the requirements necessary to obtain a phylogenetic system, but the Janiridae cannot be defined as a monophyletic family (Wilson 1994). Therefore, monophyletic groups of genera within the 'Janiridae' should be sought. The Microparasellidae possibly are such a group. The genera to be discussed in this context are Angeliera Chappuis & Delamare-Deboutteville, 1952, Caecianiropsis Menzies & Pettit, 1956, Mackinia Matsumoto, 1956, Microcharon Karaman, 1934, Microjaera Bocquet & LCvi, 1955, Microparasellus Karaman, 1933, Paracharon Coineau, 1968, and Protocharon Chappuis et al., 1956. Since Wolff's (1962) comments, the discussion of the validity of the family Microparasellidae has been controversial (see Coineau 1986). Of the above-mentioned genera only Angeliera, Microcharon, Microparasellus, Paracharon and Protocharon are usually kept as microparasellids (Coineau and Schmidt 1979; Coineau 1986). With the exception of Protocharon, a freshwater genus with broader posterior pereonites that should not be included in the Microparasellidae, these animals are more slender than any 'janirid', the free pleonite being relatively long in comparison with the pereonites, all somites having the same width. A useful apomorphy is the subapical insertion of the exopod on the sympod of the uropods. Additional apomorphies also occur in other 'Janiridae': the antennular flagellum has fewer than four articles, the third pleopod exopod is slender, and the molar process is pointed and has no grinding surface. The eyes and chromatophores are reduced, although these are almost certainly of little phylogenetic value because of the almost universal reduction of these features in interstitial and hypogean animals. Symplesiomorphies are the short dactylus with three claws (third claw only retained in Angeliera), anus terminal, not covered by pleopods. The body outline of Microparasellus varies with respect to other microparasellids: the lateral borders of the tergites are rounded and slightly protruding, bearing numerous small cuticular scales, and cephalon and pleotelson are shorter than in the remaining microparasellids. Furthermore, the uropods of Microparasellus are reduced and the exopod is absent. The family concept of the Microparasellidae may be open to challenge because Microparasellus is distinct from the other three genera in these autapomorphies. Angeliera, Paracharon and Microcharon all have their pereopodal insertions rotated dorsally -unlike Microparasellus. Characteristic autapomorphies of the other genera are: the maxillipedal palp of Angeliera has only four articles, the sympod of uropods is expanded in Microcharon. Moreover, Just and Poore (1992) note Angeliera with male penes attached to the coxae and the male pleopods being at variance with what is expected for janiroideans. Microcharon also has coxal penes (see generic section below). Although we continue to recognise a reduced concept of the Microparasellidae, the composition of this family will require further study. Zoogeographic data (summary in Coineau 1986) do not help to understand the origin of the Microparasellidae because this family has a world-wide distribution, especially around the area of the ancient Tethys. A freshwater origin is not probable, and the dwarfish freshwater genus Protocharon should be excluded as a model for the ancestor of the microparasellids. Stock (1977) suggested that the freshwater microparasellids were 'stranded' during the Miocene sea-level regressions, and subsequently became adapted to hyposaline conditions. Species of Angeliera live mesopsammally in sandy beaches and sublittoral interstitial biotas, Microcharon is known from sandy beaches, some freshwater species are stygobiontic in Mediterranean karst areas. Microparasellus is known from perimediterranean ground water, and Paracharon is known only from beaches of New Caledonia. Wagele (1990) discusses the distribution of the microparasellids further. With the above-mentioned apomorphies to define the family, the hypothetical ancestor of the group belongs to those janiroideans that had a short dactylus with three terminal claws on the pereopods 11-VII, and the exopod of pleopod I11 with three plumose setae. 722 G. D. F. Wilson and J.-W. Wagele Protocharon and Mackinia do not have these plumose setae, similar to Angeliera. Caecianiropsis and Microjaera, which also show adaptations to the interstitial habitat, have only two claws, the body is broader and the free pleonite much shorter than in most microparasellids, the antennae are separated by a broad rostrum, the propodus of pereopod I bears sensory spine-like setae, and in Caecianiropsis the molar process is not acute. Therefore, the following genera may form a monophyletic group, for which the name Microparasellidae is available: Angeliera, Microcharon, Microparasellus and Paracharon. This hypothesis will need further evaluation as the evolutionary relationships of the Asellota become better known. Genera of the Microparasellidae Angeliera Chappuis & Delamare-Deboutteville, 1952 Angeliera Chappuis and Delamare-Deboutteville, 1952: 2014. Brevipleonida Gnanamuthu, 1954: 258. Type species: Angeliera phreaticola Chappuis & Delamare-Deboutteville, 1952. Species Included A. cosettae Coineau & Rao, 1972; A. dubitans Stock, 1977: 69; Brevipleonida gracilis Gnanamuthu, 1954: 258; A. ischiensis Schulz, 1954: 253; A . phreaticola Chappuis & Delamare-Deboutteville, 1952: 2014; A. psamathusKensley, 1984a: 64; A . racovitzai Coineau & Botosaneanu, 1973: 196; A. rivularis Stock, 1985: 92; A. xarifae Siewing, 1959: 365. Other References Angeliera.- Chappuis and Delamare-Deboutteville, 1954: 123; Chappuis and DelamareDeboutteville, 1960: 321; Birstein, 1961: 137; Wolff, 1962: 35, 38, 236, 250, 276, 281, 289, 291, 300; Coineau and Renaud-Mornant, 1977: 346; Renaud-Mornant and Coineau, 1978: 1249; Coineau, 1981: 20; Coineau, 1986: 468-9; Kensley and Schotte, 1989: 91. Diagnosis Dwarfish slender isopods, body 7-10 x longer than broad. Cephalon anteriorly broader than posteriorly; lateral margin straight, lacking lateral spines or plate-like extensions; vertex straight, without rostrum, frontal margin medially compressed; eyes absent. Pereonal tergites dorsally reduced; pereopods inserting dorsolaterally. Pleonite 1 enlarged, width subequal to pleotelson width. Pleotelson longer than broad, nearly rectangular in dorsal view, lateral margins lacking denticles. Antennula short; article 1 more or less tubular, longer than wide; flagellum with 4 articles. Antenna longer than antennula; article 3 enlarged compared with articles 1-2 and 4, without scale; articles 5 and 6 length subequal to article 3; flagellum usually with 8 articles, proximal articles separate and longer than wide. Mandible molar process without grinding surface, distally pointed; palp with strongly curved distal article. Maxillipedal endite longer than wide, narrow at insertion of palp and broadening distally; palp much narrower than endite, with only 4 articles, distal article pointed, proximal article longest. Maxillipedal palp slender, with 4 articles, distal article with sickle-shaped tip; endite broad, distally rectangular. All pereopods similarly slender, with 3 claws. Pereopod I leg-like; carpus slender, margins slightly curving, with few spine-like setae; propodus without spine-like setae. Pereopods 11-VII with 3 distal claws on dactyli, ventral distal claw smaller than dorsal claw. Pereopod V of male carposubchelate. Male pleopod I with distally rounded medial lobes, sometimes laterally expanded; lateral lobes reduced, confluent with lateral margin. Male pleopod I1 sympod not broad, subrectangular; proximal article of endopod near maximum width of stylet; stylet distally very slender, shorter than syrnpod. Pleopod I11 if present lacking exopod; endopod small, rounded, lacking setae. Female operculum distally bilobed, with median incision. Uropods inserting terminally, large sympods broader than rami; exopod shorter than endopod, inserting subterminally on sympod. Review of the Janiridae 723 Remarks Angeliera has the morphology of a typical mesopsamrnal isopod, with a slender body, dorsolateral insertions of the pereopods, reduced eyes. The uropodal sympod is enlarged, the exopod inserts subapically. Three dactylar claws are present, and the first pereopod bears no subchela. A peculiar feature is the outline of the fourth (last) palp article of the maxilliped, which ends with a sickle-shaped apex. The species are known from sandy marine beaches. Just and Poore (1992) report that a species of Angeliera from Australia retains the primitive isopod configuration of the male penes attached directly to the coxae. Specimens of Microcharon in the U.S. National Museum of Natural History were also found to have coxal penes. The presence of coxal penes has important implications for the eventual placement of the Microparasellidae among the Asellota, and may require a new rearrangement of the families of the Asellota that is at odds with the current superfamily scheme. An alternative hypothesis, as suggested in the character analysis of janirid taxa (Wilson 1994), is that the coxal position of the penes might be a reversion owing to the repositioning of the coxae. The homology of this feature in Angeliera and Microcharon obviously requires further consideration. Microcharon Karaman, 1934 Microparasellus Karaman, 1933: 20. Microcharon Karaman, 1934: 44. Duslenia Lhi, 1950: 42. Type species: MicroparaseNus stygius Karaman, 1933. Species Included (see 'Other References' for page numbers) M. acherontis Chappuis, 1942; M. anatolicus Pesce & Galassi, 1990; M. angelicae Pesce & Galassi, 1988b;M. angelieri Coineau, 1963a;M. antonellae Galassi, 1991; M. apolloniacus Cvetkov, 1964; M. arganoi Pesce & Tett, 1981 [cited in Coineau, 19861; M. boui Coineau, 1968; M. bureschi Cvetkov, 1976; M. coineauae Galhano, 1970; M. comasi Coineau, 1985 [nomen nudum cited in Coineau, 1986: 4691; M. doueti Coineau, 1968; M. eurydices Cvetkov, 1965; M. galapagoensis Coineau & Schmidt, 1979; M. halophilus Birstein & Ljovushkin, 1965b;M. harrisi Spooner, 1959b; M. heimi Coineau, 1968; M. hercegovinensis Karaman, 1959;M. herrerai Stock, 1977;M. hispanicus Pesce & Galassi, 1988a;M. juberthiei juberthiei Coineau, 1968; M. juberthiei ramosus Coineau, 1968;M. karamani Pesce & TetB, 1978b; M. kirghisicus Jankowskaja, 1964;M. latus latus Karaman, 1933; M. latusprespensis Karaman, 1954; M. letiziae Pesce & Galassi, 1988b; M. longistylus Pesce & Galassi, 1988b; M. luciae Sket, 1990 ; M. lydicus Pesce & Galassi, 1990; M. major Kararnan, 1954; M. margalefi Sabater & de Manuel, 1988; M. marinus Chappuis & Delamare-Deboutteville, 1954; M. cf. marinus Pesce & Galassi, 1988a; M. meijersae Pesce & Galassi, 1988a; M. monnioti Bocquet, 1970; M. mooreanensis Coineau [nomen nudum cited in Coineau, 1986: 4721; M. motasi Serban, 1964; M. notenboomi Pesce & Galassi, 1988a; M. nuragicus Pesce & Galassi, 1988b; M. oltenicus Serban, 1964; M. orghidani Serban, 1964; M. orphei Cvetkov, 1977; M. othrys Argano & Pesce, 1979; M. phlegetonis Cvetkov, 1967; M. phreaticus Coineau & Botosaneanu, 1973; M. profundalis beranensis Karaman, 1940; M. profundalis kosovensis Karaman, 1940; M. profundalis kumanovensis Karaman, 1940; M. profundalis profundalis Karaman, 1940; M. raffaellae Pesce, 1979; M. reginae Dole & Coineau, 1987; M. rouchi Coineau, 1968;M. sabulum Kensley, 1984a; M. salvati Coineau, 1968; M. silverii Pesce & Galassi, 1988b; M. sisiphus Chappuis & Delamare-Deboutteville, 1954; Microparasellus stygius Karaman, 1933; M. stygius hellenae Chappuis & DelamareDeboutteville, 1954; M. tantalus Birstein & Ljovushkin, 1965b; M. teissieri LBvi, 1950; M. thracicus Cvetkov, 1965; M. ullae Pesce, 1981; M. zibani Pesce & Tett, 1978a; M. sp. Pesce & Galassi, 1988a; M. sp. Schotte, Heard & Kensley, 1991. G. D. F. Wilson and J.-W. Wagele 724 Other References Microcharon.-Karaman, 1940: 33; Chappuis, 1942: 120; 1944: 6; Birstein, 1951: 129; Levi, 1950: 42-7; Birstein, 1952: 227; Chappuis and Delamare-Deboutteville, 1954: 111; Karaman, 1954: 108; 1958: 11; 1959: 333; Spooner, 1959a: 1696; 1959b: 58; DelamareDeboutteville, 1960: 295; Chappuis and Delamare-Deboutteville, 1960: 301; Birstein, 1961: 137; Wolff, 1962: 35, 38-9, 276, 281, 290; Coineau, 1963a: 711; 1963b: 197; Serban, 1964: 342; Coineau, 1968: 146; Wells, 1963: 85; Cvetkov, 1964: 201; Jankovskaja, 1964: 975; Birstein and Ljovushkin, 1965b: 1001; Cvetkov, 1965: 312; Galhano, 1966: 11; Cvetkov, 1968: 108; Galhano, 1970: 83; Bocquet, 1970: 85; Coineau and Botosaneanu, 1973: 191; Cvetkov, 1976: 81; 1977: 70; Stock, 1977: 70; Pesce and Tett, 1978a: 992; 1978b: 116; Coineau and Schmidt, 1979: 145; Pesce and Argano, 1979: 129; Argano and Pesce, 1979: 178; Argano, 1979: 36; Coineau, 1981: 20; Pesce, 1979: 237; 1981: 58; Kensley, 19846: 66; Coineau, 1986: 469-71; Dole and Coineau, 1987: 200; Kussakin, 1988: 179; Kensley and Schotte, 1989: 91; Pesce and Galassi, 1988a: 308; 1988b: 250; 1990: 173; Sket, 1990: 41; Galassi et al., 1990: 265; Galassi, 1991: 201; Schotte et al., 1991: 254; Coineau, 1992: 101. Diagnosis Dwarfish, slender isopods; body more than 7 x longer than broad, all somites of subequal width. Cephalon anteriorly broader than posteriorly; lateral margin straight, lacking lateral spines or plate-like extensions; vertex straight, without rostrum, frontal margin medially compressed; eyes absent. Pereonal tergites dorsally reduced; pereopods inserting dorsolaterally. Pleonite 1 enlarged, width subequal to pleotelson width. Pleotelson longer than broad, longer than any pereonite, liearly rectangular in dorsal view, lateral margins lacking denticles. Antennula short; article 1 more or less tubular, longer than wide; flagellum with 2-3 articles. Antenna longer than antennula; article 3 enlarged compared with articles 1-2 and 4, with scale; length of articles 5 and 6 subequal to article 3; flagellum with 8-12 articles, proximal articles separate and longer than wide. Mandibular molar process without grinding surface, distally pointed; palp with strongly curved distal article. Maxillipedal palp broader than endite, articles 2-3 slightly expanded. Pereopod I leg-like; carpus slender, margins straight, with few or no spine-like setae; propodus without spine-like setae. Pereopods 11-VII with 2 distal claws on dactyli, accessory seta absent. Penes inserting on coxa of pereopod VII. Male pleopod I with distally rounded medial lobes, never expanded; lateral lobes reduced, confluent with lateral margin; sperm channel open groove, not closed tube. Male pleopod I1 sympod broadly oval; proximal article of endopod near maximum width of stylet; stylet distally very slender, shorter than sympod. Female pleopod I1 operculum oval or rectangular, distal margin sometimes with median concavity. Pleopod I11 endopod plumose setae if present with distinct gap between medial seta and 2 lateral setae; exopod narrower than and around same length as endopod; exopod pointed, with 2 segments, lacking plumose setae. Uropods with conspicuously long and broad sympods, inserting terminally on pleotelson; both rami short and slender, exopod inserting subapically. Remarks Species of Microcharon have a body form similar to Angeliera, but the maxillipedal palp is not modified and has five articles. The sympod of the uropods is usually larger and longer than in Angeliera. Only two dactylar claws are present. The species are found in sandy beaches, at low depth in marine sand and in phreatic water (Coineau 1986). Like Angeliera, Microcharon also has penes on the coxae of pereopod VII (observations of M. acherontis Chappuis, 1942, U.S. National Museum of Natural History, cat. no. 855509), a significant fact for future revisions of the Microparasellidae (see above under Angeliera). Microparasellus Karaman, 1933 Microparasellus Karaman, 1933 : 17. Type species: MicroparaseNus puteanus Karaman, 1933. Review of the Janiridae 725 Species Included M. aloufi Coineau, 1978 [cited in Coineau 1986: 1491; M. hellenicus Argano & Pesce, 1979: 173; M. libanicus Chappuis & Delamare-Deboutteville, 1954: 108; M. puteanus Karaman, 1933: 18. Other References Microparasellus. - Karaman, 1940: 39; 1954: 107; Chappuis and Delamare-Deboutteville, 1960: 295; Birstein, 1961: 137; Wolff, 1962: 35, 38, 250, 276, 281, 290; Coineau, 1968: 149; Coineau, 1986: 471-2. Diagnosis Dwarfish isopods, body 4-6 x longer than wide. Cephalon lacking eyes; lateral margins rounded; vertex broadly angular but lacking rostrum. Tergites laterally protruding, margins straight or rounded with denticles; coxae not visible in dorsal view. Pleonite 1 enlarged, width subequal to pleotelson width. Pleotelson of varying outline, often oval, longer than wide, longer than any pereonite, lateral margins with denticles. Antennula short; article 1 subrectangular, longer than wide, with denticles; flagellum with 3-4 articles. Antenna short; article 3 enlarged compared with articles 1-2 and 4, lacking scale; articles 5-6 length subequal to article 3 length; flagellum with 7-8 articles, proximal articles separate and longer than wide. Mandibular molar process without grinding surface, pointed; palp short, near length of mandibular body, with truncate and slightly curved distal article. Maxilliped normal; endite longer than wide; palp with slightly expanded articles 2-3; endite about as broad as palp; palp article 3 subrectangular; palp articles 4-5 narrow, straight-sided. Pereopod I carpus long, cylindrical with 1 or no spine-like sensory setae; propodus lacking spine-like sensory setae; dactylus shorter than dorsal claw. Pereopods 11-VII similarly slender, dactyli with only 2 distal claws, ventral claw distinctly smaller than dorsal claw, accessory seta absent. Male pleopod I with distally rounded medial lobes, never expanded; lateral lobes reduced, confluent with lateral margin; sperm channel ventrally closed, not open groove. Male pleopod I1 sympod broad, distally tapering; proximal article of endopod near maximum width of stylet; stylet distally very slender, shorter than sympod. Female pleopod I1 operculum broadly oval. Pleopod 111 endopod lacking plumose setae; exopod narrower and longer than endopod; exopod slender, with 2 segments, lacking plumose setae. Uropods short, squat, uniramous, inserting terminally on pleotelson; single ramus shorter than sympod. Remarks Microparasellus puteanus, the first discovered microparasellid (Karaman 1933), lives in phreatic water, as do all of the hitherto known species of the perimediterranean genus. In the same publication, Karaman (1933) described M. stygius, for which he later (1934) erected the genus Microcharon. The species of Microparasellus, whose morphology is not well known, have a broader body than other microparasellid genera, with laterally protruding tergites (see Coineau 1968, her fig. 1) bearing denticles. The uropods are very short and uniramous, the latter condition being found in no other 'janirid' or microparasellid genus. Paracharon Coineau, 1968 Paracharon Coineau, 1968: 153. Type species: P. renaudae Coineau, 1968. Monotypic. Other References Paracharon.-Coineau, 1970: 373; 1971: 29, 107; 1986: 472. G. D. F. Wilson and J.-W. Wiigele 726 Diagnosis Dwarfish isopods, body about 5-6 x longer than wide. Cephalon and pleotelson longer than any of the pereonites. Cephalon lacking eyes; nearly rectangular in dorsal view; lateral margins rounded; vertex straight-sided, lacking rostrum or projection. Pereonites with coxal insertions dorsolateral, but with small tergal projection dorsally; coxae visible in dorsal view. Pleonite 1 short, narrower than pleotelson or pereonites. Pleotelson with rounded tip and shallow marginal grooves above insertion of uropods; lateral margins lacking denticles. Antennula short; article 1 more or less tubular, length greater than width; flagellum with 2 articles. Antenna short; article 3 enlarged compared with articles 1-2 and 4, with distinct scale (exopod); articles 5 and 6 short, subequal to articles 3; flagellum with 6 articles, proximal articles separate and longer than wide. Mandibular molar process without grinding surface; palp longer than body of mandible, with large slightly curved distal article. Maxilliped endite longer than wide, endite about as broad as palp; palp articles 2-3 slightly expanded, article 3 not tapering, palp articles 4-5 slender. Pereopod I leg-like; carpus slender, oval, lacking spine-like sensory setae; propodus lacking spine-like sensory setae; dactylus short, claws somewhat shorter than dactylus, enlarged and oval. Pereopods 11-VII dactyli with only 2 distal claws, accessory seta absent, ventral claw subequal to dorsal claw. Penes on posteromedial margin of sternite, not on coxae of pereopods VII. Male pleopod I medial lobes rounded; lateral lobes short and hook-like. Male pleopod I1 sympod slender, laterally rounded, tapering distally; endopod proximal article near maximum width of stylet; stylet shorter than sympod. Female pleopod I1 operculum nearly quadrangular with rounded corners. Pleopod I11 endopod with 3 plumose setae having distinct gap between medial seta and 2 lateral setae; exopod narrower and longer than endopod; exopod distally pointed, with 2 segments, lacking plumose setae. Uropods long, inserting terminally on pleotelson; syrnpod about as long as endopod, exopod shorter, inserting subterminally on sympod. Remarks This monotypic genus is known only from New Caledonia. Paracharon is similar to Microcharon, but the coxae of pereopods are not fully visible in dorsal view, their insertion being more ventral than in Microcharon. The uropods are less broad than in Microcharon, and the pereonites less slender. Janiroidea incertae sedis Tole Ortmann, 1901 Ianthe Bovallius, 1881: 5. Tole Ortmann, 1901: 157 (available misspelling of Iole nom. praeocc.). Janthe Ortmann, 1901: 158. Iolella Richardson, 19056: 457. Jolella Vanhoffen, 1914: 536 (not Richardson, 1905b). Lanira Boone, 1920: 7. Type species: Janira spinosa Harger, 1879. Species Included Iolella glabra Richardson, 1908: 71; Janira laciniata Sars, 1872: 92; Janira spinosa Harger 1879: 158 (senior synonym of Ianthe speciosa Bovallius, 1886: 34); Ianthe spinosissima Stephensen, 1936: 7; Ianthe vilhelminae Stephensen, 1913: 94. Other References Io1ella.-Thielemann, 1910: 72; Nierstrasz, 1941: 283; Menzies, 19626: 83; Wolff, 1962: 33, 39-40, 234-5, 277, 293, 300; Schultz, 1969: 251; Kussakin, 1988: 125; Wagele, 1989: 76. 1anthe.-Richardson, 1900: 299; 1901: 555; Sars, 1897a: 101. Janira. -Harger, 1880: 323; Stephensen, 1913: 68; Barnard, 1914: 436; Gurjanova, 1932: 23; Barnard, 1955: 6 . 1anira.-Hansen, 1888: 191; 1916: 20. Review of the Janiridae 727 Diagnosis Cephalon with a prominent acute rostra1 process nearly as long as remaining cephalon; pair of acute anterolateral projections directed anteriorly; eyes conspicuous dorsally. Body flattened; tergites with acute lateral projections: one on pereonite 1 directed anteriorly, pair on each side of pereonites 2-4, single caudally curved projections on each tergite 5-7. Pereonites often with sharp dorsal spines. Pleotelson with characteristic pair of posterolateral, pointed lappets; uropods inserting between these lappets. Antennula short, but with multiarticulated flagellum. Antenna with scale (exopod), peduncular articles 5-6 elongated; flagellum long, multiarticulated. Mandibular molar process with grinding surface of varying size. Maxillipedal palp articles 1-3 broader than 4-5, endite about as broad as palp. Pereopod I somewhat stouter than pereopods 11-VII, carpus long-oval, with 1 row of acute sensillate setae on palm; dactylus very short, with 2 terminal claws. Pereopods 11-VII slender, with 2 terminal claws; coxa, merus and dactylus short, other articles long-cylindrical. Tips of male pleopod I apically truncate. Sympod of male pleopod I1 distally narrower than proximally, endopod barely surpassing sympod. Pleopod I11 endopod with 3 plumose setae having distinct gap between medial seta and 2 lateral setae; exopod narrower and longer than endopod; exopod having 2 segments, lacking plumose setae. Female operculum nearly quadrangular, with rounded corners, distal margin with a shallow medial concavity. Uropods slender, elongated, sympod longer than rami. Remarks The type species of Tole, Janira spinosa Harger, 1879 (type locality: north of Cape Cod), was assigned to Ianthe (with Ianthe speciosa) by Bovallius, 1881 (type locality: Baffin Bay). Tole Ortmann, 1901 was proposed to replace the preoccupied name Ianthe Bovallius, 1881, which Ortmann (1901: 158) misspelled as Janthe in his list of species in his new genus Tole. In a letter to Richardson, Ortmann explained that Iole was the intended name in his 1901 publication, and that Tole was an error. Thus, Richardson (1905b: 457 footnote) created the genus Iolella to replace the preoccupied name Iole. Nevertheless, Tole is an available name (Neave 1939) and is a senior synonym of Iolella. Tole could be suppressed under the plenary powers of the Commission using the 50-year rule (ICZN article 79), but a prima facie case that stability is threatened cannot be made. Iolella has been used in only 8 different publications, some prior to 1943 (50 years preceding this writing). Moreover, corrections of commonly used generic names to little-used forms has been the rule in isopod taxonomy (e.g. Jaeropsis to Joeropsis Koehler in Sivertsen and Holthuis 1980), so we recommend that priority apply in this case. Tole replaces Iolella as the name of the taxon whose type species is Ianthe speciosa Bovallius. T. speciosa is a junior synonym of T. spinosa (synonymy proposed by Hansen 1916). Tole was placed in the family Janiridae (as Iolella) by Wolff (1962), and in the Janirellidae by Menzies (1962b, spelled 'Ianirellidae'). Tole has some features in common with the Janiralata species-group D.3 (see above), namely the acute rostrum and the outline of the tergal lappets, but it differs from Janiralata in the lack of plumose setae on the exopod of pleopod 3. Furthermore, the lateral coxal lobes are shorter and barely visible in dorsal view. Tole also cannot be included in any of the lappet-bearing families as they are presently defined (Mictosomatidae, Mesosignidae, Acanthaspidiidae, Janirellidae, Katianiridae) because the species of Tole have three claws on the pereopods 11-VII (in contrast to e.g. Janirella). Therefore, Tole should be considered incertae sedis. The species Janira exstans Barnard, 1914 was included in Iolella by Wolff (1962: misspelled J. extans). An inspection of the original description, however, shows that this species is undoubtedly a santiid, perhaps in the genus Santia. The following species are assigned to the genus Janiralata: Iolella alascensis Richardson, 1905b [junior synonym of J. tricornis (Krdyer, 1849)l; Ianthe erostrata Richardson, 1899 (placed in Janiralata by Kussakin 1962); Tole holmesi Richardson, 1905a; Iolella chuni Thielemann, 1910; Tole libbeyi Ortmann, 1901 (junior synonym of J. tricornis); Iolella sarsi Richardson, 1905b (placed in Janiralata by Menzies 1951b); Ianthe triangulata Richardson, 1899. Iolella sp. G. D. F. Wilson and J.-W. Wagele 728 (Vanhoffen, 1914) might be a member of Ianthopsis. Kussakin (1988) provides thorough synonymies for most species in Tole and Janiralata, including many taxonomic decisions listed here. Jaerella Richardson, 1911b Jaerella Richardson, 191 1b: 634. Zaerella Nierstrasz, 1941: 285. Type species: Jaerella arrnata Richardson, 191 1 b. Monotypic. Other References Jaerel1a.-Wolff, 1anira.-Hansen, 1962: 34, 40, 236, 258, 278, 281, 291; Kussakin, 1988: 187. 1916: 13. Diagnosis Body broad, pereonite widths subequal. Cephalon with elongate anterolateral spines and lateral spines; eyes large, rounded, dorsally situated; vertex with elongate rostrum extending to tip of antennulae; pair of elongate dorsal spines between eyes. Pereonite 1 with single elongate lateral marginal spines and one pair of dorsal spines. Pereonites 2-7 lateral margins with 2 elongate lappet-like spines; each tergite with 2 dorsal spines except for pereonite 4 which has 4 dorsal spines. Pleonite 1 fused to pleotelson. Pleotelson broad, nearly as broad as pereonites, somewhat flattened, dorsally lobed; margins with single long posterior projecting lateral spines and pair of long terminal spines; uropods emerging from under concave posterior margin. Antennular flagellum with 10-1 1 articles. Antenna1 basal articles (1-4) all subequal; antenna1 articles 5 and 6 longer than basal articles 1-4; flagellum probably exceeding 12 articles, proximal articles separate and longer than wide, not conjoint or swollen. [Mouthparts not described or illustrated.] Pereopod I prehensile; pereopods 11-VII ambulatory [claws not described]. [Pleopods not described.] Uropods not modified, reduced, distinctly shorter than pleotelson, with short protopod and slightly unequal rami; exopod inserting apically. Remarks Jaerella may be closely related to either Ianthopsis or Rhacura, but is too poorly described at the moment to allow an appropriate assignment. Unfortunately, the only type of the genus is a dried male specimen residing at the US National Museum of Natural History. Rhacura Richardson, 1908 Rhacura Richardson, 1908: 73. Type species: Rhacura pulchra Richardson, 1908. Monotypic. Other References Rhacura. -Menzies, 1962a: 171; Wolff, 1962: 34, 39, 236, 263, 279, 281, 290; Schultz, 1969: 253; Kussakin, 1988: 72. 1anira.-Hansen, 1916: 13. Diagnosis Body broad, pereonites broadening posteriorly, all body margins with minute acute spinules. Cephalon lateral margin with 2 large subtriangular lateral spines; minute eyes situated dorsally near posterior margin of cephalon; rostrum absent, although vertex broadly triangular. Pereonites with broad triangular spines on lateral margins: 1 spine on pereonites 1 and 4; 2 spines on pereonites 2-3 and 5-6; 3 spines on pereonite 7. Pleonite 1 very short, narrower than pleotelson or pereonite 7. Pleotelson near width of anterior pereonites; lateral margins with 4 large broad subtriangular spines; posterior margin convex. Review of the Janiridae 729 Antennular article 1 more or less tubular, length greater than width; flagellum with around 18 articles. Antenna1 basal articles 1-4 all subequal, article 3 with large scale [articles 5-6 and flagellum unknown]. [Mandible not described.] Maxillipedal endite longer than wide, width subequal to basal articles of palp; palp article 3 broad, quadrate, not tapering; palp articles 4-5 narrow, straight-sided. Pereopod I prehensile [not described further]. Pereopods 11-VII ambulatory, dactyli with 2 claws [not described further]. [Pleopods unknown.] [Uropoda unknown, probably large.] Remarks Rhacura is another genus that would benefit from a complete revision. The type species, R. pulchra, was described as having minute eyes and only two claws on the pereopods, as well as multiple pointed lateral projections on the pleotelson and pereon. This genus seems to fall within the range of variation of Ianthopsis, which would indicate that Rhacura may be near the Acanthaspidiidae in affinities. Bowman (in litt: Wolff, 1962: 34) reports that Rhacura has coxal plates similar to that of Janiralata. In addition, J. problematica Kussakin & Mezhov, 1979 of the Janiralata bifurcata group (see above) has a dorsal outline nearly identical to that of R. pulchra. If these two species should be in the same genus, the concept of Janiralata should be revised to distinguish the two genera. Nevertheless, the uropods, pleopods, and mandible of the genus are unknown, and the dactylar claw count needs to be checked again. No further action should be taken until the genus is revised, perhaps after more specimens are found (only a single damaged specimen has been described). Rhacura was included tentatively by Menzies (1962a) in the Janirellidae, a conclusion not followed here. Trichopleon Beddard, 1886a Trichopleon Beddard, 1886a: 106. Type species: Trichopleon ramosurn Beddard, 1886. Monotypic. Other References Trichop1eon.-Beddard, 1886b: 21; Wolff, 1962: 40, 236, 259, 274, 278, 281, 291. Diagnosis Body broad anteriorly, pereonites narrowing posteriorly; all body surfaces devoid of denticles or tubercles. Cephalon lateral margin without denticles or spines, slightly curved, tapering posteriorly; eyes completely absent; rostra1 area broad, rounded, sloping into frons. Pereonite tergites 1-4 laterally truncate, with anteriorly directed spine; pereonite 5-7 posteriorly directed, pereonite 5 laterally truncate, pereonites 6-7 laterally rounded. Pleonite 1 fused to pleotelson. Pleotelson narrower than pereonites, widest anteriorly, tapering posteriorly to distal spine; lateral margins smoothly curving, fringed with slender setae. Antennulae long; antennular article 1 broad, length greater than width, with small rounded lobe medially; flagellum with 16 articles. Antenna longer than body; article 3 slightly enlarged compared with basal articles 1-2 and 4, with distinct scale; articles 5 and 6 much longer than basal articles 1-4; flagellum with more than 30 articles; flagellar proximal articles separate and longer than wide, not conjoint or swollen. [Mandibles not described.] Maxillipedal endite wider than long, broader than palp; palp articles 1-3 broad; palp article 3 quadrate, not tapering; palp articles 4-5 elongate, straight-sided, much narrower than basal articles. Pereopod I leg-like, with elongate setae [but apparently no sensillate setae]. Pereopod 11-VII dactyli with only 1 large distal claw, ventral claw [if present] much smaller than dorsal claw. [Pleopods unknown.] Uropods unmodified, shorter than pleotelson; exopod shorter than endopod, inserting apically; sympod elongate, subequal to rami. G . D. F. Wilson and J.-W. Wagele 730 Remarks The only specimen of this blind, deep-water isopod genus (a preparatory female mounted in Canada balsam) was collected south of the Philippine Islands at 500 fathoms. More specimens and species will be needed for a complete redescription. In the descriptions of Trichopleon, Beddard (1886a, 18866) reports features found in the janirids (large uropods, maxillipedal palp articles 4 and 5 narrow and elongate), as well in some deep-sea taxa (broad rostral area of cephalon, single-clawed dactyli). In addition, he illustrates some odd 'sculpturing' of the cuticle that may indicate that this animal has a thick opalescent cuticle like that of Xostylus and other deep-sea taxa. An inspection of the type specimen from the British Museum indicates that Trichopleon has a cephalon similar to Xostylus. As for Xostylus (see below), a proper family assignment for Trichopleon cannot be made at this time. Xostylus Menzies, l962a Xostylus Menzies, 1962a: 179. Type species: Xostylus parallelus Menzies, 1962a. Species Included X. parallelus Menzies, 1962a; X. longiflagellus Birstein, 1970: 297. Other References Xostylus. -Wolff, 1962: 63, 236, 263, 279, 281, 290; Kussakin, 1988: 165. Diagnosis (based on X. parallelus) Body narrow, straight-sided, all pereonites subequal. Cephalon lateral margin without denticles or spines, slightly curved, tapering posteriorly; eyes completely absent; rostral area broad, rounded, sloping into frons. Pereonites laterally rounded, coxae visible in dorsal view only on posterolateral margins of pereonites 5-7. Pleonite 1 short, only slightly narrower than pleotelson and pereonite 7. Pleotelson only slightly narrower than anterior pereonites; lateral margins smooth, lacking denticles or spines; uropods emerging posteriorly from slight notches in posterolateral margin; anus separated from pleopodal chamber. Antennular article 1 more or less tubular, length greater than width; flagellum with 5 articles [8 articles in X. longij7agellatus]. Antenna1 basal articles (1-4) subequal, article 3 bearing small rounded scale; articles 5 and 6 longer than basal articles 1-4; [flagellum unknown in X. parallelus]. Mandibular molar process slender, distally truncate; palp shorter than body of mandible, distal article short, curved, setose. Maxillipedal endite length subequal to width, width subequal to palp; palp articles 1-3 broad, article 3 quadrate, not tapering; palp articles 4-5 straight-sided, much narrower than basal articles 1-3. [Pereopod I unknown in X. parallelus.] Pereopod 11-VII dactyli with only 2 distal claws, with ventral distinctly smaller than and closely adpressed to dorsal claw. [Pleopods and uropods unknown in X. parallelus.] Remarks Xostylus was originally classified by Menzies (1962a) in the Abyssianiridae (now synonyrnised with Paramunnidae, see Just 1990), although Wolff (1962) felt that it was more similar to the janirids Ectias Richardson or Caecianiropsis Menzies & Pettit. Neither grouping seems correct, as Xostylus has several characters that would ally it with several deep-sea families: an exposed anus completely separate from the branchial chamber and placed ventrally as in the Haploniscidae, Nannoniscidae, Desmosomatidae or Ischnomesidae (i.e. not an abyssianirid); a ventral dactylar claw distinctly smaller than the anterior claw (not a typical 'janirid' feature); and a pleopod I11 exopod that is short, laterally rounded, Review of the Janiridae 73 1 fused into a single segment, and distally pointed, i.e. similar to that of the Nannoniscidae or the Haploniscidae and dissimilar to the 'janirid' condition. In addition, the cuticle of Xostylus spp. is heavily calcified as in many of the deep-sea forms. In many respects, this genus seems somewhat generalised; for example, an ischnomesid conceivably could be derived from a Xostylus-like ancestor. The species X. longiflagellus Birstein, 1970 has features that are reminiscent of the coarse drawings of Trichopleon, especially in its long antennae and antennulae, serially similar pereopods, and anteriorly pointed pereonites 1-4. Moreover, X. longiflagellus also has several characters that are different from X. parallelus: its dactylar claws are more equal in size and separate, it has large anterolateral setae on the pereonites, the antennula is longer, and the mandibular molar process is distally bifurcate rather than truncate. For this reason, the diagnosis above was constructed without using X. longiflagellus. If Trichopleon were better known, possibly X. longiflagellus could be put in that genus. Xostylus and Trichopleon may belong to the same family-level taxon, perhaps related to one of the deep-sea families. Acknowledgments Torben Wolff (Zoologisk Museum, Copenhagen) loaned one of us (G.D.F.W.) his taxonomic catalogue of all Janiroidea until approximately 1975. His generous sharing of information added considerably to the detailed literature information presented here. We are grateful for financial support of the National Science Foundation, grants BSR8604573 and BSR88-18448 to G.D.F.W., Australian Research Council Grant No. A1913 1795 to G.D.F.W., and Deutsche Forschungsgemeinschaft (the German Science Foundation) grant Wa-530/2 to J.-W.W. J.-W.W. is grateful for help, advice and laboratory facilities provided during the stay at the Scripps Institution of Oceanography by the first author, R. R. Hessler and W. A. Newman. Useful reviews of the manuscript were provided by G. C. B. Poore, and by two anonymous reviewers of an earlier version of this paper. E. L. Ho and P. 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