A CONTRIBUTION TO THE KNOWLEDGE OF METAPSEUDIDS

Travaux du Muséum National d Histoire Naturelle «Grigore Antipa» Vol. LII pp. 101 125 Octobre 2009 A CONTRIBUTION TO THE KNOWLEDGE OF METAPSEUDIDS. DESCRIPTION OF A NEW GENUS AND THREE NEW SPECIES FROM THE CARIBBEAN SEA AND THE INDIAN OCEAN (CRUSTACEA: TANAIDACEA: APSEUDOMORPHA) MODEST GUÞU Abstract. Genus Vestigiramus n. g., from the Caribbean Sea, the waters of the Martinique Island, is described; it is characterized by the presence of a vestigial exopodite, uniarticulated and devoid of setae, at the cheliped level. The new genus (classified in the subfamily Chondropodinae Guþu, 2008) included two species: V. antillensis n. sp. and V. codreanui (Guþu, 1996), comb. nov. At the same time, it is recorded the presence of an unknown morphological component on the female chelipeds of the new genus and some amendments are mentioned for the description of the species V. codreanui, comb. nov. (from the Brazilian waters of the Atlantic Ocean) and for the diagnosis of the subfamily Chondropodinae. From Martinique (Caribbean Sea) and Réunion Islands (Indian Ocean), Apseudomorpha martinicana n. sp. and Pseudoapseudomorpha curtisetosa n. sp., respectively, are described, and the identification keys of the genera Apseudomorpha and Pseudoapseudomorpha species are presented. Also, from the waters of the Hispaniola Island (Dominican Republic), Pseudoapseudomorpha gomezi Guþu & Ortiz, 2009 is recorded. Résumé. On décrit, des eaux de la Mer des Caraïbes de l Île Martinique, le genre Vestigiramus n. g., caractérisé par la présence, au niveau des chélipèdes, d un exopodite vestigial, uniarticulé et dépourvu de sètes (genre classifié dans la sous-famille Chondropodinae Guþu, 2008), dans la composition du quel ont été incluses deux espèces, V. antillensis n. sp. et V. codreanui (Guþu, 1996), comb. nov. On signale, à cette occasion, pour la première fois, l existence sur les chélipèdes des femelles du nouveau genre d une formation inconnue et on apporte certains amendements à la description de l espèce V. codreanui comb. nov. (des eaux brésiliennes de l Océan Atlantique) ainsi qu à la diagnose de la sous-famille Chondropodinae. Des l îles Martinique (Mer des Caraïbes) et Réunion (Océan Indien) sont décrits Apseudomorpha martinicana n. sp. et, respectivement, Pseudoapseudomorpha curtisetosa n. sp. et on présente les clés d identification des espèces du genres Apseudomorpha et Pseudoapseudomorpha. On signale aussi, dans les eaux de l Île Hispaniola (Dominican Republique), l espèce Pseudoapseudomorpha gomezi Guþu & Ortiz, 2009. Key words: Tanaidacea, Metapseudidae, Vestigiramus antillensis n. g., n. sp., V. codreanui, comb. nov., Apseudomorpha martinicana n. sp., Pseudoapseudomorpha curtisetosa n. sp., P. gomezi. In the rich tanaidacean material from different marine areas of the World Ocean, sent by Dr. Hans-Georg Müller long time ago, I discovered a new species of the genus Apseudomorpha Miller, 1940 and another one, classified (together with Calozodion codreanui Guþu, 1996) in a new genus of the subfamily Chondropodinae Guþu, 2008. A special feature discovered in the new genus is, among others, an unknown morphological component on the chelipeds, only in adult females. Also, from the Indian Ocean (Réunion Island), I identified a new species belonging to the genus Pseudoapseudomorpha Guþu, 1991. Further on, I present the diagnosis of the new genus and the descriptions of the three new species, as well as the identification keys of the species of the genera Apseudomorpha Miller, 1940 and Pseudoapseudomorpha Guþu, 1991. Also, some amendments to the diagnosis of the

102 MODEST GUÞU subfamily Chondropodinae and the description of the species Calozodion codreanui, transferred in the new genus, are presented. Family Metapseudidae Lang, 1970 Subfamily Chondropodinae Guþu, 2008 In the diagnosis of the subfamily Chondropodinae, Guþu (2008: 74) mentioned: Chelipeds...with exopodite. By absence of some other specifications in Guþu s diagnosis, anybody interested in the study of apseudomorph tanaidaceans understands by exopodite, naturally, the presence of the classic exopodite, formed of three articles, the last one having some long plumose setae. The new genus (described further on, and classified in this subfamily) is characterized, among others, by the presence of a very small uniarticulated exopodite (difficult to detect), devoid of plumose setae. Not to create some confusions, I propose the subfamily diagnosis to be amended, the references on the cheliped being as follows: Chelipeds obvious dimorphic or not; exopodite present, with three articles, the last one ended in some long plumose setae, or, in very rare cases, with only one very small article, devoid of setae. Genus Vestigiramus n. g. Type-species: Vestigiramus antillensis n. sp. Diagnosis. Body elongate, of small size, more or less cylindrical. Carapace longer than broad; ocular lobes well defined; visual elements pigmented. Pereonites wider than long; at least, last four similar to a trapezium. Pleon, shorter than carapace length, with five free, short pleonites and a pleotelson; each pleonite with long circumplumose setae on sides. Antennule relatively long, biramous, with denticulated projections on the inner margin of the first peduncular article; inner flagellum short (one or two-articulated). Antenna squama small; first peduncular articles with distoinner margin ended in some acute processes; second and third article also with some spiniform projections. Cheliped exopodite very small, uniarticulated (hard visible), without terminal setae; male cheliped dimorphic, apparent similar to female, but with much shorter carpus and greater propodus. Pereopod II stronger than following ones, with normal exopodite; basis thick, with spiniform projections and long plumose setae on the tergal margin. Pereopods III, IV and VI propodus thin, curved and long (longer than merus or carpus), with a few sternal spines. Pleopods well developed, biramous, in five pairs. Uropod exopodite short, three-articulated; endopodite approximately as long as pleon length. Etymology. From the Latin vestigius, vestige, trace, and ramus, branch, alluding the reduced exopodite of chelipeds. Gender, masculine. Composition (2 species): Vestigiramus antillensis n. sp. and V. codreanui (Guþu, 1996), comb. nov. Geographical distribution. Eastern Caribbean Sea (Martinique Island) and Southwestern Atlantic Ocean (Coast of Brazil). Remarks. In diagnosis, I don t refer to mouthparts because I dissected only partially the single female I have in the type-material. As it results from the description of second species of genus (Vestigiramus codreanui, comb. nov., cf. Guþu, 1996: 98-101 and figs 30 A-E, 31 A, B), the main features (which are

A NEW GENUS AND THREE NEW SPECIES OF METAPSEUDIDS (CRUSTACEA: TANAIDACEA) 103 probable similar to those of the type-species of the genus) consist in mandible and maxillule palp (with three and two articles, respectively), and maxilliped setulation (with a few short simple and some long and thick circumplumose setae on the inner margin, and one midouter thick and long circumplumose seta on the second palp article). The main morphological feature by which Vestigiramus n. g. distinguishes the other six genera of the subfamily Chondropodinae consists in the morphology of the cheliped exopodite, formed of a single article, very small, devoid of plumose setae, in the new genus (Figs 2 A, B and 3), in contrast with three articles, with long plumose setae on the last article, in the other genera (Guþu, 2006: figs 395, 418-420). By the body and the appendages configuration (but especially by the cheliped aspect), Vestigiramus n. g. resembles better the genus Calozodion Gardiner, 1973 (Bamber & Sheader, 2005; Gardiner, 1973; Guþu, 1984, 2006, etc.), the main difference between them being the above-mentioned exopodite morphology. Another difference between the two genera, apparently less important, is the size of chelipedal carpus of females, longer in the new genus (Fig. 2 A, B) and shorter in Calozodion (Guþu: 1984, fig. 3 A; 1996: fig. 27 B). Checking of the type-material in the species Calozodion codreanui Guþu, 1996, from the waters of the Brazilian Atlantic (whose conclusions are detailed further on), made me to transfer it in the new genus. A special aspect, reported in the literature on tanaidaceans for the first time, is the presence of an unknown morphological component (or formation) in both species of the genus, present only on the sternal margin of merus and carpus of the adult female chelipeds, which I do not know how to approach. It is about two very similar formations (apparently as two compact aglutinated tufts of setae), of the same colour with the cheliped, covered with fine hairs (Figs 2 A, uc and 3, uc). In Vestigiramus codreanui, comb. nov., the two components are partially superposed and placed in the sternodistal angle of merus (Fig. 3, uc), and in V. antillensis n. sp. one of the formations is at the level of the merus-carpus joint, and the second one on carpus (Fig. 2 A, uc). The presence of these components in the same place of both chelipeds of each species made me think that they are a morphological element characteristic to both adult females of the new genus. At the same time, their presence in one of the species only on merus, and in the other one, on merus-carpus, can be considered a specific feature. Unfortunately, the presence of a single adult female (in the type-material belonging to both species) is a serious impediment for a doubtless conclusion. By the aspect of the cheliped exopodite, Vestigiramus n. g. resembles the monotypic genus Plectrocopus, described from the East-African waters (Mauritania coasts) of the Atlantic Ocean (Guþu 2006: 206 and figs 384-386). The difference between these last two genera consists in the body and appendages configuration, the genus Plectrocopus resembling Apseudomorpha Miller, 1940 (both of them classified in subfamily Metapseudinae) and, by no means, the genus Calozodion, as the new genus (included in subfamily Chondropodinae). Within this context, I also remind the presence of another genus with a reduced exopodite of the chelipeds (but formed of two articles) and devoid of plumose setae. It is about monotypic genus Ascumnella (family Parapseudidae), described from the waters of the Caribbean Sea of the Gran Cayman Island (Guþu & Heard, 2002: fig. 3 A-D).

104 MODEST GUÞU Vestigiramus antillensis n. sp. (Figs 1, 2) Material: 7 specimens (1 female with oostegites, 1 adult male, 1 juvenile and 4 manca), Caribbean Sea, Martinique Island, Madras, Baie de Tartane, in dead corals with seagrass beds, moderately exposed, 1-2 m depth, 14 April 1990; Station 14. Leg. Dr. Hans-Georg Müller. Holotype, female with oostegites (partially dissected), preserved in the collections of the Grigore Antipa National Museum of Natural History, Bucharest, No. 250427; Allotype, adult male, in the same collection, No. 250428; Paratypes, 5 specimens (1 juvenile and 4 manca) preserved as follows: - 2 specimens (manca) in the same collection, No. 250429; - 3 specimens (1 juvenile and 2 manca) in the collections of the Senckenberg Museum, Frankfurt-am-Main (Germany). Description of the female with oostegites Body (Fig. 1 A) more or less cylindrical, aproximately 5.5 times longer than the width of carapace; standard length, about 2.2 mm. Carapace, longer than broad, with some lateral circumplumose setae. Rostrum triangular, relatively short, with two obvious denticles in top and some very small tubercules, on its lateral margins. Ocular lobes well defined, with pigmented visual elements (Fig. 1 B). Epistome ended in a small spine. Pereon, two times longer than the carapace, with six relatively short but unequal pereonites. First pereonite, as long as the last one, approximately 2.5 times wider than long. Second and third pereonites, similar to a trapezium, a little longer than the first or the last ones, with a small tuberculiform prominence on the anterolateral margins, ended in a circumplumose seta. Fourth pereonite, relatively similar to the previous one, but longer than that. Fifth pereonite, slightly shorter than the fourth pereonite, but longer than others, with one anterolateral and two posterolateral circumplumose setae. Last pereonite with six or seven circumplumose setae on sides. Pleon, 2.8 times shorter than the pereon length, with five free pleonites and pleotelson. Each pleonite, short and very broad, with two circumplumose setae on sides (Fig. 1 C). Pleotelson, as long as all pleonites, with two circumplumose setae on the first half of sides, and the other two simple, very small, caudally. Antennule (Fig. 1 D) well developed, about as long as pereon. First peduncular article, shorter than the carapace length and five times longer than wide, with two denticles on the distal half of inner margin and another one in midouter side; some small circumplumose and broom setae are present on the inner and outer margin. Second article, about 2.5 times shorter than the first one, with one distal long simple seta; inner and outer margins with two circumplumose and some broom setae. Third peduncular article, shorter and narrower than the second one, with three distal simple setae. Inner flagellum two-articulated (without common article) ended in three unequal but long simple setae. Outer flagellum (without its terminal setae, shorter than the first peduncular article) with six articles, the last one being smaller than others; each article with about three unequal setae; last three articles with one strong aesthetasc. Antenna (Fig. 1 E) a little shorter than the half of antennule (both measured without distal setae). Squama, shorter than the third article, with one small and three

A NEW GENUS AND THREE NEW SPECIES OF METAPSEUDIDS (CRUSTACEA: TANAIDACEA) 105 D A B C 0.3 mm: B-E 1 mm: A E Fig. 1 Vestigiramus antillensis n. g., n. sp., female, holotype: A, body, dorsal; B, carapace, anterior part; C, pleon and left uropod; D, antennule; E, antenna. relatively long setae. Peduncle five-articulated. First peduncular article with three or four conspicuous denticles on the distoinner corner and a circumplumose seta on the inner margin. Second article thick, approximately two times longer than wide, with three distal strong spiniform projections; one and two short circumplumose setae are present on the outer and inner sides, respectively. Third peduncular article, shortest, with one and two acute denticles on the outer and inner corner; one long circumplumose seta is present in distoinner corner. Fourth article slightly longer than the third one, but a little narrower than that. Fifth peduncular article, longer than the previous one, with one long simple and three broom setae in distal end. Flagellum, about 2.2 times shorter than the peduncle, with five short and thin

106 MODEST GUÞU articles; excepting the last article which has five long but unequal setae, others have one to four short simple setae. Cheliped (Fig. 2 A) well developed. Exopodite very small, uniarticulated, devoid of setae (Fig. 2 A, e). Basis, 1.5 times longer than broad, with one midsternal spine and three distosternal long circumplumose setae; distotergally with two circumplumose setae. Merus narrow, with three sternal circumplumose setae. Carpus 1.5 times longer than wide, with ten tergal long circumplumose setae; sternal margin with three long circumplumose setae and two cone-shaped unknown morphological formations (similar to two tufts of compact aglutinated setae), imposible for me to understand their structure. Propodus large, about as wide as long (measured together with the fixed finger); sternal margin with six unequal simple setae and one median denticle; fixed finger, short and very thick at base, with two great and three small denticles, and some simple setae on the inner margin; claw robust. Dactylus thin and long, curved, with some very small spines on the inner margin, and three distotergal simple setae; claw stout, longer than the propodal one. Pereopod II (Fig. 2 C) with great exopodite, ended in four long plumose setae. Basis strong and thick, approximately two times longer than broad, with eight sternal long plumose setae and one distosternal spine; tergal margin with nine long plumose setae (first five much longer than the last four) and six denticles (alternating with the mentioned setae). Ischium very short. Merus, longer than carpus or propodus, with two unequal simple setae and one robust spine on the sternal margin; tergally with one median plumose seta and one distal spine. Carpus, as long as wide, with three sternal simple setae and one spine; tergally and distotergally with three long plumose setae, one spine and two simple setae. Propodus, as long as carpus but a little narrower than that, with four simple setae, which alternate with three unequal spines, on the sternal margin; tergally with one long plumose and two simple setae, and one distal spine. Dactylus thick, slightly shorter than propodus, with two sternal denticles and one tergal ciliate seta; claw curved, relatively small. Pereopod III (Fig. 2 D) slender. Basis, about three times longer than wide, with six and eight long plumose setae on the sternal and tergal margins, respectively. Ischium small. Merus thin, about as long as carpus, with one distosternal spine and one small seta; distotergally with one plumose seta. Carpus with two spines and two small setae on the sternal margin; distotergally with one plumose and two simple setae. Propodus thin and long, slightly curved, approximately as long as merus and carpus (measured together), with four spines and one seta, sternally, and two short setae, distotergally. Dactylus thin, about as long as carpus, with two sternal spinules; claw acute, curved, longer than the greatest propodal spine. Pereopod IV (Fig. 2 E) similar to pereopod III, excepting the plumose setae of basis, which are lost, and the spines on merus and carpus, present in a number of two and three, respectively. Pereopod V (Fig. 2 F) a little smaller than the previous pereopod. Basis, about 2.7 times longer than wide, with two proximotergal broom setae, and another one, very small, simple, distotergally; distosternally with one broom and two unequal simple setae. Ischium small, with three short setae. Merus, much shorter than carpus, with one distotergal long plumose seta; distosternally with two small spines and two simple setae. Carpus with two rows of three and four spines on the sternal margin, and two simple setae in distotergal corner. Propodus, as long as carpus but thinner than that, slightly curved, with two sternal small spines; midtergally with

A NEW GENUS AND THREE NEW SPECIES OF METAPSEUDIDS (CRUSTACEA: TANAIDACEA) 107 A B C e e uc 0.3 mm: A-H D E F G H 0.1 mm: I I Fig. 2 Vestigiramus antillensis n. g., n. sp., female, holotype, and male, allotype (B): A, B, chelipeds (e, exopodite; uc, unknown morphological component); C-H, pereopods II-VII; I, pleopod.

108 MODEST GUÞU one broom seta and distotergally with five short serrate and two long ciliate setae (detail fig. 2 F). Dactylus thin, approximately as long as merus, with two sternal spinules; claw short, acute and curved. Pereopod VI (Fig. 2 G) with three plumose setae on the tergal margin of basis. Ischium small. Merus, as long as carpus, with one long tergal plumose seta; sternally with two short setae and two small spines. Carpus with one plumose and one simple seta in distotergal corner, and two rows of two and three spines on the sternal side. Propodus, longer than merus or carpus, similar to the same of pereopods III or IV, with three sternal spines; distotergally with one broom and two simple setae. Dactylus and its claw similar to the same of pereopods III-V and VII. Pereopod VII (Fig. 2 H) with six or seven long plumose setae on each sides of basis. Ischium small. Merus, about 2.5 times shorter than basis, with three short simple and three long plumose setae on the sternal and tergal margins, respectively. Carpus, slightly shorter than merus, with one simple (very small) and one plumose seta, in the distosternal corner; tergally with two long plumose setae. Propodus, longer than merus or carpus, and three times longer than its width, with about 15 ciliate small setae disposed in distosternal and distal margins (detail fig. 2 H). Dactylus and its claw similar to those of pereopods III-VI. Pleopods (Fig. 2 I) biramous, in five pairs. Basal article well developed. Endopodite, approximately as long as exopodite, with nine long plumose setae, around, and four setulae in the first half of the outer margin. Exopodite with nine long plumose setae on the outer and distal sides, and six setulae on the distoinner margin. Uropod (Fig. 1 A, C) biramous. Protopodite relatively small. Endopodite, as long as pleon length, with ten slender articles, some of them having one-two short simple setae; terminally with four long but unequal setae. Exopodite very short, three-articulated, ended in three simple setae. Description of the male Male differs from female by the configuration of chelipeds, which have a shorter carpus (almost as long as wide) and two, one and three spiniform projections on the sternal margin of merus, carpus and propodus, respectively (Fig. 2 B), the female having an elongated carpus (1.5 times longer than wide) and one sternal spiniform prominence only on the propodus (Fig. 2 A). Also, the male cheliped have not the two unknown morphological components present only in adult females (Fig. 2 A, uc). Cheliped exopodite as in female (Fig. 2 A, B, e). Juveniles, similar to females, excepting the chelipedal unknown morphological formations (which are absent), as I already mentioned in the genus remarks and the description of female. Etymology. From the collecting area of species, Lesser Antilles. Remarks. As it results from the description of the species, I don t studied the mouthparts, the dissection made by me having in view only the chelipeds, pereopods and one pleopod (for protecting the single female I had in type-material). The main features by which Vestigiramus antillensis n. sp. differs from the V. codreanui (Guþu, 1996), comb. nov., consist in: (1) the number of articles of antenna flagellum (five in the new species, in contrast with three in V. codreanui), (2) the greater number in V. antillensis n. sp. of tergal denticles and sternal plumose setae of pereopod II basis, (3) the smaller number in V. codreanui of basal plumose setae of pereopods III and VII.

A NEW GENUS AND THREE NEW SPECIES OF METAPSEUDIDS (CRUSTACEA: TANAIDACEA) 109 Vestigiramus codreanui (Guþu, 1996), comb. nov. (Fig. 3) Guþu (1996: 101), in the description of the species Calozodion codreanui from the Brazilian waters of the Atlantic Ocean, makes the following specification on the chelipedal exopodite: I do not know if the exopodite is absent or it is very small and I could t observe it. The discovering of the uniarticulated chelipedal exopodite, devoid of terminal setae in Vestigiramus antillensis n. g., n. sp. from Martinique Island, made me to check attentively the type-material of C. codreanui, thinking that it might be possible that the exopodite of this species to be very small and hardly observed, as well as that one of the species from the Martinique Island. This way, I remarked both the similitude between the chelipedal exopodite of the two species (Figs 2 A, e and 3, e) and other common features, unoccurred in the species of the genus Calozodion, in which V. codreanui, comb. nov. was initially classified. It is about: (1) the rostrum configuration (with a small hollow, giving it a bifurcate aspect, more or less, fig. 1 B; Guþu, 1996: fig. 29 A), (2) the presence of the dimorphism, relatively minor, at the level of the chelipedal carpus (longer in females, fig. 2 A, B; Guþu, 1996: fig. 32 A, B), and (3) the different morphology of the female cheliped (by the presence of the two mentioned unknown formations, figs 2 A, uc and 3, uc). e 0.2 mm Fig. 3 Vestigiramus codreanui (Guþu, 1996), comb. nov., female cheliped: e, exopodite; uc, unknown morphological component. Subfamily Metapseudinae Lang, 1970 Genus Apseudomorpha Miller, 1940 Menzies (1953) described the genus Imitapseudes, very similar to Apseudomorpha Miller, 1940. The greatest difference between the two genera, as Menzies (op. cit.: 482, 483) mentions, is the number of pleopods: five pairs in the genus described by him and only one pair in the genus described by Miller (1940: 483). Lang (1968: 26) synonymyzed the two genera, specifying that A reexamination of the species of two genera has proved that all of them have five pairs of pleopods which, however, are difficult to detect.... In the end of his paper, Lang (op. cit.: 207) said: Apseudomorpha Miller has not one pair of pleopods but five pairs. Later, Guþu (2006, 2007), ignoring Lang s conclusion, classified two species (A. vestafricana and A. negoescuae, respectively, characterized by a reduced number of pleopods or their absence) in the genus discussed, and, at the same time, uc

110 MODEST GUÞU presenting a new diagnosis of the genus (Guþu, 2006). According to Menzies opinion (op. cit.), mentioned above, the two species should be classified in another genus. But, as it is known, also other genera have a variable number of pleopods (Pagurapseudes Whitelegge, 1901, Pseudoapseudomorpha Guþu, 1991, Pugiodactylus Guþu, 1995, etc.), this thing (at least for the time being) has to be admitted for the genus Apseudomorpha, too. Another disputable aspect is that one regarding the place in the apseudomorph systematics of the species Apseudes timaruvia Chilton, 1882 and Parapseudes hirsutus Stebbing, 1910 (now both classified in the genus Apseudomorpha). Referring to the place of some species in the systematics of family Apseudidae, Lang (1949: 5) expressed his opinion that the species A. timaruvia (from New Zeeland), belongs in all probability to the genus Apseudomorpha Miller, 1940. Sieg (1983: 136) classified the mentioned species in the same genus in incertae species, idea taken over by Guþu (1987, 2006) as a sure one. Reading again attentively Chilton s description (1882), I reached the conclusion that Apseudes timaruvia, although it resembles the species of the genus Apseudomorpha by several features, was wrongly included in this genus. For this opinion five arguments plead: (1) the large number of the articles of the inner and outer flagellum of the antennule (6 and, respectively, 14) and (2) of the antenna flagellum (formed of 8 articles), (3) the presence of two rows of spines on pereopod III propodus, (4) the configuration of cheliped propodus and (5) the setulation of pleopods. By the morphology of the pereopods III (and of the others, as it results from Chilton s description) the species resembles those of the genus Pseudoapseudomorpha, but with which it is in contradiction by the great length of the antennule and antenna flagella, the absence of the chelipedal expodite and the presence of the biramous pleopods with many plumose setae (Chilton, op cit.: 146-149, pl. XVIII, figs 7, 9 a, 11). In conclusion, the systematic position of Chilton s species has to be reconsidered, by all probabilities, it belonging to an unknown genus. The species Parapseudes hirsutus Stebbing, 1910, was reclassified by Sieg (op. cit.: 134) in the genus Apseudomorpha, an idea taken over by Guþu (1987, 2006) later. Analysing again Stebbing s description (1910) I reached the conclusion that the resemblances of the first three pereopods (including the large number of the sternal spines on carpus and propodus) corresponds only to the morphological features of the species of the genus Pseudoapseudomorpha. Also, the great length of the antennule and the large number al the articles of the flagella lead to the same conclusion. Unfortunately, in the absence of the description of the cheliped (from where it may results if the exopodite is present or not) any conclusion can lead to new mistakes. Considering the above remarks, the genus Apseudomorpha include 11 certains species: A. albida (Shiino, 1951), A. avicularia (Barnard, 1914), A. fontainei Guþu, 1987, A. glebosa (Menzies, 1953), A. magdalenensis (Menzies, 1953), A. martinicana n. sp., A. negoescuae Guþu, 2007, A. oahuensis Miller, 1940, A. ortizi Guþu, 2006, A. veleronis (Menzies, 1953) and A. vestafricana Guþu, 2006. The main differences between the species of the genus, as they result from descriptions (Barnard, 1914; Guþu, 1987, 2006, 2007; Menzies, op. cit.; Miller, 1940; Shiino, 1951), can be deduced from the identification key, which might include insignificant inaccuracies because of the numerous gaps of literature.

A NEW GENUS AND THREE NEW SPECIES OF METAPSEUDIDS (CRUSTACEA: TANAIDACEA) 111 Key to the species of the genus Apseudomorpha 1 - Pereopod II basis with a proximoanterior dentiform process...... A. negoescuae (&, %) - Pereopod II basis without a proximoanterior dentiform process... 2 2 - Last pleonite without a long seta on sides... A. avicularia (sex unknown) - At least the last pleonite with a long seta on sides... 3 3 - Pleopods absent... A. vestafricana (&; % unknown) - Pleopods present (uni- or biramous)... 4 4 - Only the last pleonite with a long seta on sides... 5 - Last pleonite and at least one more with a long seta on sides... 6 5 - First peduncular article of antennule with some dentiform processes on the proximal half of inner margin... A. ortizi (&; % unknown) - First peduncular article of antennule without dentiform processes on the proximal half of inner margin... A. albida (&, %) 6 - Last four pleonites with a long seta on sides... 7 - At most the last three pleonites with a long seta on sides... 8 7 - Pleopods biramous, with biarticulated exopodite... A. magdalenensis (&, %) - Pleopods biramous, with uniarticulated exopodite (in male), or uniramous (in female)... A. veleronis (&, %) 8 - Last three pleonites with a long seta on sides... A. oahuensis (&, %) - Only two pleonites (second and fifth) with a long seta on sides... 9 9 - Pleopods uniramous... A. fontainei (&, %) - Pleopods biramous... 10 10 - Pereopod II carpus and propodus with three and five sternal spines, respectively; one branch of pleopods with two long setae... A. glebosa (&, %) - Pereopod II carpus and propodus with two and four sternal spines, respectively; each branch of pleopods with one long setae...... A. martinicana n. sp. (&, %) Apseudomorpha martinicana n. sp. (Figs 4-6) Material: 234 specimens (27 adult and subadult males, 64 females with eggs or oostegites, 106 juveniles and 37 manca), Caribbean Sea, Martinique Island, collected as follows: - 177 specimens (21 adult and subadult males, 41 females with eggs or oostegites, 85 juveniles and 30 manca), La Trinité, bank reef of Ponte Rouge, Anse Riviere, exposed reef-flat, in dead corals, 0-2 m depth, 12 April 1990; Station 24. Leg. Dr. Hans-Georg Müller;

112 MODEST GUÞU - 57 specimens (6 adult males, 23 females with eggs or oostegites, 21 juveniles and 7 manca), Madras, Baie de Tartane, from deads corals in seagrass beds, moderately exposed, 1-2 m deep, 14 April 1990; Station 14. Leg. Dr. Hans- Georg Müller. Holotype, female with eggs, from the Station 24, preserved in the collections of the Grigore Antipa National Museum of Natural History, Bucharest, No. 250432; Allotype, adult male, from the Station 24, in the same collection, No. 250433; Paratypes, 72 specimens (20 adult and subadult males, 40 females with eggs or oostegites, 7 juveniles and 5 manca), also from the Station 24, preserved as follows: - 35 specimens (10 adult and subadult males, and 25 females with eggs or oostegites), in the collection of the Grigore Antipa National Museum of Natural History, Bucharest, No. 250434 and 250435, respectively; - 37 specimens (10 adult and subadult males, 15 females with eggs, 7 juveniles and 5 manca), in the collections of the Senckenberg Museum, Frankfurtam-Main (Germany). Remarks. Another material from the Station 24 (78 juveniles and 25 manca) preserved in the collections of the Grigore Antipa National Museum of Natural History, Bucharest, No. 250436 and 250437, respectively. The material from the Station 14 preserved in the collections of the Senckenberg Museum, Frankfurt-am-Main (Germany). Description of the female with eggs Body (Fig. 4 A) calcified, more or less cylindrical, approximately 4.5 times longer than the maximum width (at the level of the first pereonite); standard length, approximately 1.6 mm. Carapace (measured together with the rostrum) longer than wide. Frontal margin rounded, with some small denticles and a median short rostrum, three-forked in top (Fig. 4 B). Ocular lobes well defined; visual elements pigmented. Epistome with a relativerly small spine. Pereon, two times longer than carapace, with short but wide pereonites. First pereonite (about as long as last one but slightly shorter than other pereonites), rounded laterally, three times broader than long. Each of the following two pereonites, approximately as wide as first one, with one small anterolateral tubercle ended in a small seta. Last three pereonites, decreasing in width, as a trapezium; each of them with an evident anterolateral tuberculiform process ended in a small simple seta (Fig. 4 A, C). Pleon (Fig. 4 A, C), about as long as last two pereonites. Pleonites (measured together), shorter than the length of last pereonite; each of the five free pleonite about 8-10 times shorter than wide; lateral sides rounded in the first pleonite and acuted in the last four ones; second and fifth pleonites with one long lateral simple seta. Pleotelson, longer than the last pereonite, pointed caudally; anterolaterally with an obvious tuberculiform projection; antero- and posterodorsally with two long and two short simple setae. Antennule (Fig. 4 B) slightly longer than carapace. Peduncle three-articulated; first article stout, approximately four times longer than its thickness, with one distoexternal denticle, and two or three long simple and some broom setae on outer side; inner margin with one distal and three or four median spiniform expansions; second article, about 2.5 times shorter than the first one, with two simple and some

A NEW GENUS AND THREE NEW SPECIES OF METAPSEUDIDS (CRUSTACEA: TANAIDACEA) 113 A B D C 0.5 mm: A E F G 0.1 mm: E, F, H, I 0.1 mm: D 0.2 mm: B, C H I Fig. 4 Apseudomorpha martinicana n. sp., female, paratype: A, body, dorsal; B, carapace, anterior part, and antennule; C, last two pereonites, pleon and left uropod; E, right mandible; F, pars incisiva, lacinia mobilis and setiferous lobe of left mandible; G, labium (schematic); H, maxillule; I, maxilla. broom setae, distally; third article, much shorter and narrower than the previous one. Outer flagellum, as long as second peduncular article, with four short but thick articles; first three articles with one-two long simple setae, and the last one with four unequal but long simple setae; second and fourth article with one aesthetasc. Inner flagellum with one long but thin article (without common article), ended in two long simple setae. Antenna (Fig. 4 D) squama small, ended in two unequal simple setae. Peduncle five-articulated; first article strong, with an anteroinner expansion; second

114 MODEST GUÞU article short, a little thicker than long; third article, very short, with one stout spine in distoinner corner; fourth article, longer than the previous one, also with a robust spine in the distoinner corner; fifth peduncular article, a little longer than the second one, but approximately as long as two previous articles, with two long distoinner simple and some distal broom setae. Flagellum short, three-articulated, about as long as last peduncular article; second article much longer than the first or the last ones; first two articles with one and the last one with two long simple setae. Mandibles (Fig. 4 E, F) with three-articulated palp. First article of palp, as long as thick and about two times shorter than the second one, with one seta and one very small denticle (tuberculiform) on the distoinner corner; second article, approximately 2.2 times longer than wide, with three distoinner setae, the later smaller than the first two ones; third article, longer than the first one, but a little shorter than the second article, with seven ciliate setae in distoinner half, the distal ones being much greater than the first four. Left mandible with four and three denticles on pars incisiva and lacinia mobilis, respectively; setiferous lobe with four forked setae (Fig. 4 F). Right mandible with four distal denticles on pars incisiva; setiferous lobe with five forked setae, the first one much stronger than the others (Fig. 4 E). Pars molaris well developed, without special features. Labium (Fig. 4 G) with some distal denticles and some hairs on the outer and distoinner margins, respectively, of basal lobe. Palp ovate, about 1.5 times longer than wide, ended in two fine spines; lateral margins with fine setae. Maxillule (Fig. 4 H) with two-articulated palp, ended in three unequal setae, one of them very long. Outer endite with (?) ten terminal spines and two subterminal setae. Inner endite with five ciliate setae. Maxilla (Fig. 4 I) with some fine denticles or spines on both sides. Movable endite with seven long and five short setae on the outer and inner lobes, respectively. Fixed endite with eight setae (four simple and four forked in top) on the outer lobe, and eight setae (some of them ciliate) on the inner lobe. Maxilliped (Fig. 5 A, B). Coxa short and broad. Basis wider as long, with some denticles in the distoexternal corner, and a distoinner small seta. Palp fourarticulated; first article short with one long simple seta in the inner margin and a very small seta in the distoexternal angle; second article largest (about as long as basis, but narrower than that) with around 15 setae of different sizes on the inner margin, and one robust spine in the distoexternal corner; third article short and broad, with six thick and two fine setae on the inner margin; fourth article narrow, with one short and seven long distal setae. Endite (Fig. 5 B) with five fine and four thick setae on the rostral side; inner margin with six short plumose setae and two couplers. Cheliped (Fig. 5 C) slender, without exopodite. Basis relatively narrow, about two times longer than maximum broad, having one small spine and one short seta on the mid- and distosternal margin, respectively. Merus thin, two times shorter than carpus, with two small distosternal setae. Carpus, as long as basis, slender, slightly thinner proximally, four times longer than its median thickness. Propodus (measured together with fixed finger) a little shorter than basis or carpus, and, at the same time, narrower than basis; fixed finger well developed, relatively thick, with two and five short setae on the outer and inner margins, respectively; claw small. Dactylus curved, slightly thinner than the fixed finger; claw two-forked distally. Pereopod II (Fig. 6 A), longer than the cheliped or other perepods, without exopodite. Basis relatively thin, 3.5 times longer than broad, with one small spine

A NEW GENUS AND THREE NEW SPECIES OF METAPSEUDIDS (CRUSTACEA: TANAIDACEA) 115 A B C D E 0.1 mm: A, B 0.3 mm: C-E Fig. 5 Apseudomorpha martinicana n. sp., female, subadult (D) and adult males (E), paratypes: A, maxilliped; B, maxillipedal endite; C-E, chelipeds. and one simple seta in the distosternal corner. Ischium short, with two distosternal setae. Merus, 2.3 times shorter than basis, with one spine and four small setae on sternal side, and one spine and two setulae, distotergally. Carpus, slightly longer than wide, and much shorter than merus or propodus, with two stout spines and two unequal setae, sternally, and one spine and two setae, distotergally. Propodus, about as long as merus, with four spines and three small setae on sternal margin; distotergally with two unequal spines (one of them only a little shorter than the dactylus length, measured without claw), and one broom and one long simple seta; one short ciliate seta is present at base of dactylus joint. Dactylus, about two times shorter than propodus, with two sternal spinules; claw curved, much smaller than the greatest propodal spine. Pereopod III (Fig. 6 B) smaller than the previous one. Basis, four times longer than its thickness, with three and two setulae, tergally and sternally, respectively. Ischium with two unequal distosternal setae. Merus, longer than carpus, with two sternal spines and four short setae. Carpus, as long as wide and about two times shorter than propodus, with six spines and three setae, as in drawing (Fig. 6 B). Propodus, a little longer than merus, with three spines and one seta on the sternal margin, and two unequal spines and one long seta, distotergally; midtergally with one broom seta and near of dactylus joint with one small ciliate seta. Dactylus well developed, about as long as carpus, with two sternal spinules; claw relatively small. Pereopod IV (Fig. 6 C) similar to pereopod III, but slightly shorter than that. Pereopod V (Fig. 6 D) basis and ischium similar to those of pereopods III or IV. Merus, a little shorter than carpus, with two small spines and four unequal setae in distal end. Carpus, as long as propodus, with seven stout, sternal and distal, spines disposed in two rows, and four distal setae, two of them very long. Propodus,

116 MODEST GUÞU slightly thinner than carpus, with one sternal spine, and about ten serrate setae in distal end, around dactylus joint (detail fig. 6 D). Dactylus and its claw similar to the same of the previous pereopods. Pereopod VI (Fig. 6 E) basis and ischium relatively similar to those of previous pereopod. Merus, shorter than carpus, with one small spine and two long setae. Carpus, with two sternal rows of two spines, and two setae in the distotergal corner (one of them very long). Propodus, as long as carpus, with three spines and one short seta on the sternal margin, and two small substernal setae; one ciliate seta is present in sternodistal end, and one spine and one seta in distotergal corner. Dactylus and its claw similar to those of other pereopods. Pereopod VII (Fig. 6 F) basis and ischium no differ from those of the previous pereopod. Merus, shorter than the carpus, with one distotergal and three distosternal setae. Carpus, about 1.7 times longer than merus, with one spine and two unequal setae on the sternal margin, two distal and two distotergal setae, one of them very long. Propodus, as long as carpus, with two sternal spines; distotergally with one A B C D 0.2 mm: A-F 0.1 mm: G F E G Fig. 6 Apseudomorpha martinicana n. sp., female, paratype: A-F, pereopods II-VII; G, pleopod.

A NEW GENUS AND THREE NEW SPECIES OF METAPSEUDIDS (CRUSTACEA: TANAIDACEA) 117 broom and one simple seta, and one spine; distally, six unequal serrate setae are present (detail fig. 6 F). Dactylus, almost as long as merus, with one distosternal swelling; claw relatively short, curved. Pleopods (Fig. 6 G) biramous, in five pairs. Basal article long and narrow. Exo- and endopodite equal, uniarticulated, ended in a long plumose seta. Uropod (Fig. 4 C) biramous. Protopodite thick, well developed. Exopodite short (as long as pleonites, measured together), four-articulated, ended in three very long setae. Endopodite, as long as pleon or last two pereonites, with eight thin articles, some of them having one to three longer or shorter setae, excepting the latest one which has four long simple setae. Description of the male Males similar to females, excepting the chelipeds which are stronger. Chelipeds of adult males (Fig. 5 E) larger than those of subadult ones (Fig. 5 D). Basis broad, without exopodite; midsternally with one small spine. Merus narrow. Carpus very short and thick, with a dentiform sternal expansion. Propodus large, with a denticulated proximosternal swelling, and a relatively thick fixed finger; inner margin of fixed finger with two denticles and some setae; claw small, tuberculiform. Dactylus thin, curved, with four unequal denticles on the inner margin; claw greater than the same of fixed finger. Chelipeds of subadult males differ from the same of adult ones by the size and configuration of carpus and propodus, which are narrower and longer (Fig. 5 D). Etymology. From the collecting place (water of the Martinique Island). Remarks. Apseudomorpha martinicana n. sp. resembles very much A. ortizi Guþu, 2006, known also from the waters of the Caribbean Sea. The difference between the two species is represented by the presence of a long seta on the sides of the second pleonite in the new species (Fig. 4 C) and its absence in A. ortizi (Guþu, 2006: fig. 315), morphological feature also remarked in the specimens recently checked (Guþu & Ortiz, 2009). A. martinicana n. sp. and A. ortizi distinguished from Apseudomorpha sp. A (from Florida, recorded by Heard et al., 2004) by the rostrum configuration (Fig. 4 A, B; Guþu, 2006: fig. 314; Heard et al., op. cit.: fig. 70 c). The differences between A. martinicana n. sp. and the other species of the genus are mentioned in the identification key, presented above. Genus Pseudoapseudomorpha Guþu, 1991 Genus Pseudoapseudomorpha distinguishes Apseudomorpha by four well defined features, at least: (1) the greater length of the antennule and (2) the presence of the multiarticulated inner flagellum, (3) the similitude between the first three pereopods, and (4) the presence of the chelipedal exopodite. As I have already mentioned (in my comments on the genus Apseudomorpha) the species Apseudes timaruvia Chilton, 1882 and Parapseudes hirsutus Stebbing, 1910 (both reclassified in the genus Apseudomorpha, cf. Lang, 1949, Sieg, op. cit. and Guþu, 1987), are characterized by the first three mentioned features, present in the genus Pseudoapseudomorpha. Under these circumstances, till the knowledge of the fourth mentioned feature (regarding the presence or absence of chelipedal exopodite) the systematic status of the two species remain doubtful. Therefore, I consider that the genus Pseudoapseudomorpha contains 6 species: P. curtisetosa n. sp., P. gomezi Guþu & Ortiz, 2009, P. madagascariensis Guþu, 1991, P. ornata Guþu, 2006, P. tanzaniana Guþu, 2006 and P. wagait (Edgar, 1997).

118 MODEST GUÞU As it results from literature (Edgar, 1997; Guþu, 1991, 2006; Guþu & Ortiz, 2009), the main features by which the species of the genus Pseudoapseudomorpha can be distinguished, are resumed in the identification key presented further. Key to the species of the genus Pseudoapseudomorpha 1 - Pleopods in two pairs... 2 - Pleopods in three pairs or absent... 3 2 - Third and fourth pleonite with rounded sides, devoid of setae; antennule inner flagellum with two articles... P. madagascariensis (juvenile) - Third and fourth pleonite with pointed sides, ended in a short seta; antennule inner flagellum with three articles... P. tanzaniana (&, %) 3 - Second and third pleonites with short lateral setae...... P. curtisetosa n. sp. (&; % unknown) - At least second or third pleonite with a long lateral seta... 4 4 - Second and third pleonite with one long lateral seta... P. ornata (&, %) - Only second or third pleonite with one long lateral seta... 5 5 - Second pleonite with one long lateral seta... P. wagait (&, %) - Third pleonite with one long lateral seta... P. gomezi (&, %) Pseudoapseudomorpha curtisetosa n. sp. (Figs 7-9) Material: 2 males, Réunion Island, Southeastern Indian Ocean, reef-flat of La Saline-les-Bains, in dead corals covered with algae, 0.5-1 m depth; 18-20 January 1989. Leg. Dr. Hans-Georg Müller. Holotype, male, preserved in the collection of the Grigore Antipa National Museum of Natural History, Bucharest, No. 250430; Paratype, 1 male (dissected), in the same collection, No. 250431. Description of the male (female unknown) Body (Fig. 7 A) more or less cylindrical, approximately 5.3 times longer than the maximum width; standard length, 2 mm. Carapace (measured together with rostrum) about 1.5 times longer than broad, and two times shorter than the length of pereon. Rostrum well developed, thick at base and acute in top, with some lateral and dorsal tubercles (Fig. 7 B). Ocular lobes well defined; visual elements pigmented. Pereon, decreasing in width from the first to the last pereonite, approximately 3.7 times longer than the pleon. First pereonite, rounded laterally, about 2.5 times wider than long and slightly shorter than the second one. Second pereonite also with rounded sides. Third pereonite with two small, tuberculiform, prominences in the anterolateral margins, ended in a small seta. Fourth pereonite, 1.5 times longer than the shortest ones (the first and the last pereonites), with one tuberculiform and one lobate projections on the first lateral half, ended in a small seta. Last two pereonites, descreasing in length and width, similar to the fourth one.

A NEW GENUS AND THREE NEW SPECIES OF METAPSEUDIDS (CRUSTACEA: TANAIDACEA) 119 Pleon, shorter than the length of the last two pereonites but narrower than those, with five free, very short, pleonites and a pleotelson. Last four pleonites with a great lateral acute projection; second pleonite with two lateral short but unequal circumplumose setae, and third and fourth ones with only one small seta; first and last pleonites without lateral setae (Fig. 7 C). Pleotelson, pointed caudally, longer than pleonites (measured together), with a small dentiform prominence in the anterolateral margins; two long and two short circumplumose setae are present middorsally and distocaudally, respectively (Fig. 7 C). Antennule (Fig. 7 D) longer than carapace. First peduncular article, about four times longer than wide, with five dentiform processes on the inner margin (the last one, forward oriented, situated in the distoinner corner); one midouter spine and one great acute projection are present in distoexternal corner; on the sides and dorsal surface with 15-17 short circumplumose setae. Second peduncular article short, as long as width of the first article, with one great denticle on each distal corner, and some distal circumplumose setae. Third article smaller than the previous one. Inner flagellum, three-articulated (without common article), ended in three long simple setae. Outer flagellum, six-articulated, ended in four long setae; each article with two or three fine aesthetascs and one seta. Antenna (Fig. 7 E, F), measured without terminal setae, a little shorter than the antennule peduncle. First article with a large inner expansion (forward oriented) having some obvious tubercles on the outer margin. Second peduncular article, greatest, approximately as long as fourth and fifth ones (measured together), with three or four dentiform projections in the first half of the inner margin, and one strong distoexternal spine; squama small, a little longer than the adjacent article, ended in two unequal setae. Third peduncular article small, about four times shorter than the second one, with one long simple seta and one spine in the distoexternal corner. Fourth and fifth articles equal, the last one having one long and two short setae in outer corner, and another one in inner corner. Flagellum short, three-articulated, about as long as second peduncular article; first two articles with two short simple setae, and the last one with four ones, three of them longer than flagellum. Mandibles (Fig. 7 G) with thick three-articulated palp; first article, shorter than its thickness, with two short setae in the distoinner corner; second article, approximately 2.5 times longer than the first one, with about eight short simple and four long ciliate setae; third article as long as previous one, but thinner than that, with nine unequal ciliate setae. Pars incisiva of right mandible relatively small, apparently undenticulated. Setiferous lobe small, with one thick and four thin setae, forked distally. Pars molaris relatively small. Left mandible unstudied. Labium (Fig. 7 H) with outer margins of basal lobes smooth; rostral margin with some fine setae. Terminal lobes (palp) ovate, 2.2 times longer than broad, with many fine setae on both sides and two long spiniform setae in top. Maxillule (Fig. 7 I) with bi-articulated palp, ended in four unequal setae. Inner and outer endite with five ciliate setae and eleven strong spines, respectively. Maxilla (Fig. 7 J) without special features. Outer lobe of movable endite with six long setae and the inner one of same endite with five different short setae. Fixed endite with about nine setae (some of them serrate) on outer lobe; inner lobe with three strong, ciliate, and approximately 15 fine setae. Maxilliped (Fig. 8 A, B) basis with a distoventral great swelling, having a strong seta and some tubercles on the distoinner margin. Palp four-articulated; first article short and narrow; second article, wider distally (only slightly longer than its maximum