THE REDESCRIPTION OF THE SPECIES LEPTOCHELIA BISPINOSA

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1 Travaux du Muséum National d Histoire Naturelle «Grigore Antipa» DOI: /v Vol. LIV (1) pp Juin 2011 THE REDESCRIPTION OF THE SPECIES LEPTOCHELIA BISPINOSA GUÞU AND L. CORSICA DOLLFUS, AND THE FIRST DESCRIPTION OF THE FEMALE OF L. TENUICULA MAKKAVEEVA (CRUSTACEA: TANAIDACEA) MODEST GUÞU Abstract. Leptochelia bispinosa Guþu, 2010 (from Réunion Island, Indian Ocean) and L. corsica Dollfus, 1898 (from Adriatic Sea) are redescribed and illustrated. The number of antennule articles in the specimens with female features belonging to the species L. bispinosa is commented. The female of L. tenuicula Makkaveeva, 1968 (from Martinique Island, Lesser Antilles) is recorded and described for the first time. At the same time, the diagnosis of the genus Leptochelia, as it is accepted now, is amended. Résumé. On a redécrit et illustré Leptochelia bispinosa Guþu, 2010 (de l Île Réunion, Océan Indien) et L. corsica Dollfus, 1898 (de la Mer Adriatique). On commente le nombre d articles de l antennule des exemplaires ayant des traits de femelle chez l espèce L. bispinosa. La femelle de L. tenuicula Makkaveeva, 1968 (de l Île Martinique, les Petites Antilles) est découverte et décrite pour la première fois. On ammende tout à la fois la diagnose du genre Leptochelia, tel qu il est accepté à présent. Key words: Tanaidacea, Tanaidomorpha, Leptochelia bispinosa, L. corsica, L. tenuicula. Investigating tanaidomorph tanaidaceans from different marine areas of the World Ocean, in the waters of the eastern coast of Italy (the Adriatic Sea) I rediscovered the species Leptochelia corsica, described by Dollfus (1898) from the Mediterranean Sea, considered by Lang (1973) synonymous (together with others) with L. dubia (Krøyer, 1842). On the other hand, Sieg (1983) reached the conclusion that all species synonymised by Lang (op. cit.) with L. dubia are, in fact, synonyms of L. savignyi (Krøyer, 1842). In a recent study, Bamber (2010) called in question the synonymisation of L. corsica with L. savignyi, made by Sieg, while Guþu (2010) considered Dollfus taxon a valid species. On the same occasion, I discovered a new species (L. bispinosa Guþu, 2010) from Réunion Island (Indian Ocean), very similar to L. corsica and to those with which the last mentioned species was synonymised by Lang (op. cit.) or Sieg (op. cit.), but also to L. tarda from the Andaman Sea (Larsen & Rayment, 2002), L. itoi from Japanese waters (Ishimaru, 1985), and L. daggi, L. dijonesae and L. opteros from Australian waters (Bamber, 2005, 2008), etc. Redescription and validation of the species L. corsica, as well as the description of the species L. bispinosa, formed the subject of the paper by Guþu (2010); however, this included only brief diagnoses (Guþu, 2010). In this report, the two species (L. bispinosa and L. corsica) are redescribed in detail, as well as an additional species, L. tenuicula Makkaveeva, 1968, from the shallow waters of Martinique Island (Lesser Antilles). This last species was described from the Atlantic Ocean (north-western coast of Cuba Island) on the basis of males only (Makkaveeva, 1968), females being recorded and described here for the first time. Some amendments are also proposed on Lang s

2 22 MODEST GUÞU diagnosis (op. cit.) of the genus Leptochelia, and a short comment on terminal female, pre-male protogynic neuter or final pre-male neuter (cf. Heard et al.: 100, 101), and pre-adult males (cf. Bamber, 2010: 293), is presented. Remarks on terminology. To avoid confusion, pereopods II-VII in this paper correspond to pereopods I-VI from the papers of most other authors. I also discuss here reference to specimens of Leptochelia that possess antennules of five articles, which were considered by Heard et al. (2004: 100, 101), to be terminal female, pre-male protogynic neuter or final pre-male neuter, and by Bamber (2010: 293), pre-adult males. Without going into detail, I note that the differences between males and females of Leptochelia are represented by numerous morphological features (which are very obvious at the antennule level, and with mouthparts, cheliped, pereopods, and the presence of the oostegites or eggs). To consider specimens with five-articulated antennules to be terminal female, pre-male protogynic neuter, final pre-male neuter or pre-adult males, ignores numerous other characteristic features of females (present in the discussed specimens), is not in concordance with reality. Females with eggs and five-articulated antennules are known in the species Leptochelia elongata, which was described by Larsen & Rayment (2002). Furthermore, in the species L. bispinosa (redescribed here), I discovered 6 females with oostegites and antennules of five articles. Therefore, to avoid confusions, I used the label specimens with female features for specimens with the antennules of four or five articles, but without oostegites, eggs or remains of a marsupium. I have stopped applying the recognised terms juvenil or neutrium, considering that these terms can generate confusion in the case of specimens with five-articulated antennules. Family Leptocheliidae Lang, 1973 Genus Leptochelia Dana, 1849 Lang (op. cit.: 224), in the diagnosis of the genus Leptochelia, mentioned that the female antenna has a tergal spine on the second and the third article each, and a sternal one on the second article (...zweites und drittes Glied beim Weibchen tergal mit einem Dorn, zweites Glied ausserdem sternal mit einem Dorn ). From this it follows that the third article of the antenna of the females of the genus Leptochelia has no sternal (ventral) spine. This assertion is inconsistent with features discovered in females of L. bispinosa Guþu, 2010 (a species belonging to Leptochelia-Gruppe 2, cf. Lang, op. cit., or to Leptochelia dubia-gruppe, cf. Sieg, op. cit.), whose antenna (Fig. 1 C) has a sternal spine on the third article, too (not on the second one, as Guþu, 2010: 46, incorrectly mentioned). Another feature which is contrary to the genus diagnosis refers to the number of setae on the female maxilliped basis, about which Lang (op. cit.: 224) specifies that there are three at least (...Basis mit wenigstens drei Borsten... ). Or, as regards the species L. tenuicula Makkaveeva, 1968 (included in Leptochelia minuta- Grouppe, cf. Sieg, op. cit., which corresponds to Leptochelia-Gruppe 1, cf. Lang, op. cit.), females have only two setae (Fig. 7 H), as in the genus Pseudoleptochelia Lang, 1973 (cf. Lang, op. cit.: 225). In addition, also in this species, the claw (unguis) of pereopod II is longer than the dactylus, and the uropod has the third article of the exopod obviously longer than the other articles, contrary to other species of the genus. Although these particularities could represent enough arguments for including the species L. tenuicula in a new genus, at least for the

3 ON THREE SPECIES OF LEPTOCHELIA (CRUSTACEA: TANAIDACEA) 23 present I think that the current classification should remain unchanged. Besides the confusions and contradictions commented by numerous specialists (out of which I mention only the recent ones, made by Larsen & Rayment, op. cit., and Bamber, 2008, 2010), the genus Leptochelia still hides too many enigmas, which could potentially emerge to contradict particular conclusions. Therefore, the morphological features commented above must be considered amendments to the diagnosis of the genus Leptochelia, as defined by Lang (op. cit.), as it is accepted at present. Leptochelia bispinosa Guþu, 2010 (Figs 1-4) Studied material: 434 specimens [78 females with eggs or remains of marsupium, 31 females with oostegites (6 having five-articulated antennule), 88 adult and subadult males and 237 specimens with female features (46 having fivearticulated antennule)], Southern Indian Ocean, Réunion Island, collected by the late Dr. Georg-Hans Müller, provided as follows: - 46 specimens [5 females with eggs, 3 females with oostegites (2 having fivearticulated antennule), 1 male and 37 specimens with female features (3 having five-articulated antennule)], Station Re-1, reef-flat of La Salineles-Bains, dead corals covered with algae, m depth, January, 1989; - 8 specimens (2 females with eggs, 1 female with oostegites, 1 male and 4 specimens with female features), Station Re-15, seagrass (Syringodium isoetifolium), near La Saline-les-Bains, m depth, 26 January, 1989; specimens [34 females with eggs or remains of marsupium, 12 females with oostegites (2 having five-articulated antennule), 41 males and 87 specimens with female features (21 having five-articulated antennule)], Station Re-31, lagoon near La Saline-les-Bains, after cyclone Firinga, mainly from dead corals, 30 January-4 February, 1989; specimens [37 females with eggs or remains of marsupium, 15 females with oostegites (2 having five-articulated antennule), 45 males and 109 specimens with female features (22 having five-articulated antennule)], Station Re-35, reef-flat of La Saline-les-Bains, after cyclone Firinga, from dead corals, m depth, 3-5 February, Type-material: Holotype, female with oostegites, from Station 31, preserved in the Collections of the Grigore Antipa National Museum of Natural History, Bucharest (Romania), No ; Allotype, adult male, from Station 31, in the same collection, No ; Paratypes (162 specimens) preserved as follows: - 80 specimens from the Station Re-31 (20 females with eggs or marsupium, with No ; 8 females with oostegites and four-articulated antennule, with No ; 2 females with oostegites and five-articulated antennule, with No ; 20 males, with No ; 20 specimens with female features, having four-articulated antennule, with No ; 10 specimens with female features, having five-articulated antennule, with No ) in the Collections of the Grigore Antipa National Museum of Natural History, Bucharest (Romania);

4 24 MODEST GUÞU - 82 specimens from the Station Re-35 (20 females with eggs or marsupium, 10 females with oostegites and four-articulated antennule, 2 females with oostegites and five-articulated antennule, 20 males, 20 specimens with female features, having four-articulated antennule, and 10 specimens with female features, having five-articulated antennule) in the Collections of the State Museum of Munich (Germany). Remarks. Specimens unmentioned in type-material are preserved in the Collections of the Grigore Antipa National Museum of Natural History, Bucharest (Romania). Description of the female with oostegites, paratype Body (Fig. 1 A) slightly dorsoventrally flattened, about 6.8 times longer than carapace breadth; standard length, 4.2 mm. Carapace, 1.2 times longer than wide, and as long as first two pereonites together, with three very short setae on sides. Rostrum very small. Ocular lobes well defined; visual elements pigmented. Pereon, 3.3 times longer than carapace. Each pereonite shorter than wide. First three pereonites and the sixth one, similar. Pereonites four and five similar, each of them slightly longer than any others. Pleon, slightly wider than pereon, with five very short pleonites and a pleotelson. First four pleonites with two and the last one with four small setae on sides. Pleotelson with four simple and two broom setae on the caudal margin, and another one, simple, on sides (Fig. 1 B). Antennule (Fig. 1 C), four-articulated, slightly shorter than carapace length. First article, longest, about 3.5 times longer than its thickness; setulation as in figure 1 C. Second article, three times shorter than the first one, and thinner than that, with two distal simple setae. Third article, as long as the second one, thinner distally, ended in two long simple setae and one aesthetasc. Fourth article (flagellum), very small, tuberculiform, with five long simple setae. Antenna (Fig. 1 C), a little shorter than antennule, six-articulated. First three articles short and thick, with similar dimensions; first article with one distosternal short seta; second and third article with one stout spine on distotergal and distosternal corners. Fourth article, 4.5 times narrower than long, and 2.5 times longer than the third article; midtergally with one seta, and distally with one broom and three simple setae (two of them much longer). Fifth article, as long as third article, ending with two long simple setae. Last article, tuberculiform, with five setae, four of them at least as long as last three articles. Mandibles (Fig. 1 D, E) with strong pars molaris. Pars incisiva of right mandible with two small denticles in top; outer margin slightly crenulated (Fig. 1 D). Pars incisiva of left mandible acute, with a very small denticle on outer margin; lacinia mobilis of the same mandible well developed, having four denticles on outer side (Fig. 1 E). Maxillule, unstudied. Maxilliped (Fig. 1 F) well developed. Basis, two times longer than broad, with six or seven long simple setae on distal side. Palp four-articulated; first article thick, as long as wide, naked; second article, equal with the first one, with one distoouter, and four disto-inner simple setae; inner margin with numerous setulae; third article, a little longer but slightly thinner than the previous one, with nine simple setae on inner margin; last article of palp, two times longer than wide and as long as the first article, with eight strong setae on distal and inner margin. Endite, as long as

5 ON THREE SPECIES OF LEPTOCHELIA (CRUSTACEA: TANAIDACEA) 25 A B C 0.4 mm: C 0.2 mm: D - F 0.5 mm: B D F E 1 mm: A Fig. 1 Leptochelia bispinosa Guþu, 2010, female, paratype: A, body, dorsal; B, right half of pleon with afferent uropod, dorsal; C, antennule and antenna; D, right mandible; E, pars incisiva and lacinia mobilis of left mandible; F, maxilliped. first palp article, with one simple seta and three unequal spiniform formations, rounded in top (one of them very short, tuberculiform), situated on the distal margin; disto-medially, with two very small setae. Cheliped (Fig. 2 A) as in other species of the genus. Basis, 1.4 times longer than maximum width, with one distal seta. Merus narrow, with one long and two short simple setae, sternally. Carpus thick, longer than basis, and 1.7 times narrower than its length, with three distosternal setae and other three ones, shorter, tergally. Propodus, only slightly shorter than basis, and 1.7 times narrower than its length (measured together with the fixed finger), with six setae on the inner surface, near the dactylus joint; fixed finger thick, with a median swollen and three simple setae

6 26 MODEST GUÞU A B 0.5 mm: A C D E H F G 0.3 mm: B - H Fig. 2 Leptochelia bispinosa Guþu, 2010, female, paratype: A, cheliped; B-G, pereopods II-VII, respectively; H, pleopod I. on the inner margin; one proximal seta is present on inner side, also, and another on the sternal margin; claw (unguis) small. Dactylus, as thick as fixed finger, with a median inner protuberance; proximotergally with a simple seta; claw (unguis) stout but small. Pereopod II (Fig. 2 B) slender, much longer than the following ones. Basis, 3.6 times longer than wide, with one simple and one broom seta, proximotergally. Ischium with one sternal seta. Merus, about 2.2 times shorter than basis, with one distotergal and two distosternal setae. Carpus, a little shorter than merus, with two distosternal and three distotergal setae. Propodus, 1.8 times longer than carpus and approximately five times longer than its median thickness, with four distal setae.

7 ON THREE SPECIES OF LEPTOCHELIA (CRUSTACEA: TANAIDACEA) 27 Dactylus as long as merus; together with the unguis as long as propodus; unguis 1.5 times shorter than dactylus. Pereopod III (Fig. 2 C) much shorter than the previous pereopod. Basis, 2.3 times longer than its median thickness, with one simple and two broom setae, proximotergally. Ischium with two sternal setae. Merus, 2.5 times shorter than basis; distosternally with one spine and one seta, and distotergally with one seta. Carpus, 1.3 times longer than wide, and slightly shorter than merus; distosternally with two small spines and one seta, and distotergally with two setae. Propodus, 3.2 times longer than wide, and longer but narrower than carpus; distally with one spine and three setae. Dactylus short but thick; unguis very small. Pereopod IV (Fig. 2 D) similar to the previous pereopod, except the distal setae of propodus, which are two in number. Pereopods V (Fig. 2 E) and VI (Fig. 2 F) similar. Basis, much thicker than those of the previous pereopods, about two times longer than wide. Ischium with two sternal setae. Merus, much narrower than basis, and about two times shorter than wide, with two distosternal small spines. Carpus, slightly shorter than merus, with two setae and three small spines, distally. Propodus, as long as carpus but narrower than that, with three ciliate spines and three setae, distally. Dactylus, joined with unguis, short and thick. Pereopod VII (Fig. 2 G) relatively similar to previous two pereopods, except some features. Propodus, distally with two spines and eight setae (six of them shorter). Pleopods (Fig. 2 H) well developed, biramous, in five pairs. Basal article thick, with an inner plumose seta. Endopod, two times longer than wide, with one midinner plumose seta; outer margin with 20 unequal plumose setae, the apical one being stronger and shorter than the following ones. Exopod, as long as endopod but slightly narrower, with 30 unequal plumose setae on the outer margin. Uropod (Fig. 1 A, B) biramous. Peduncle shorter than first two endopod articles. Exopod, uniarticulated, as long as first endopod article, having a short midlateral seta and other two distally, much longer. Endopod with six articles, each of them longer than wide; first five articles with two relatively short setae; last article with five setae, two of them longer, as long as last four articles of endopod. Description of adult male, paratype Body (Fig. 3 A) approximately four times longer than the maximum width, at the level of the first pereonite; standard length, 4.5 mm. Carapace, about as long as broad, a little longer than first three pereonites; eyes large, pigmented. Pereon, two times longer than wide, each pereonite being wider than long; first pereonite shortest and the fourth one longest; pereonites two and three equal, each of them being shorter than any of following pereonites; each pereonite with two short setae on sides. Pleon as long as carapace and first pereonite, combined; first two pleonites with one lateral small seta; pleonites three and four with two setae on sides, and the last one with four lateral setae; pleotelson, longer than any pleonite, with one lateral and four terminal short setae. Antennule (Fig. 3 B) approximately half body length. Peduncle, threearticulated, as long as carapace and first pereonite (together); first article, a little shorter than carapace length; second article two times shorter than the first one; third

8 28 MODEST GUÞU A B 0.3 mm: B C D E 1 mm: A 0.5 mm: C 0.2 mm: D 0.5 mm: E Fig. 3 Leptochelia bispinosa Guþu, 2010, adult and subadult (D) males, paratypes: A, body, dorsal; B, antennule and antenna; C, D, chelipeds; E, uropod. article 2.2 times shorter than the second; each peduncular article with three distal setae; flagellum, shorter than peduncle, with 9-10 articles, the last very short (tuberculiform), with three long terminal setae; each article with four relatively short aesthetascs. Antenna (Fig. 3 B), six-articulated, as long as first two articles of antennule peduncle. First three articles short and thick; second article with one distotergal and one distosternal seta, and the third with one distotergal spine and one distosternal seta. Fourth article thin and very long (about six times longer than wide), with one median and three distal setae. Fifth article, as long as previous article, with two

9 ON THREE SPECIES OF LEPTOCHELIA (CRUSTACEA: TANAIDACEA) 29 A B C D E F 0.3 mm: G 0.5 mm: A - F G Fig. 4 Leptochelia bispinosa Guþu, 2010, adult male, paratype: A-F, pereopods II-VII, respectively; G, pleopod I.

10 30 MODEST GUÞU distal long setae. Sixth article very small (tuberculiform), ending in one short and three very long simple setae (only a little shorter than last three articles). Mouth parts atrophied. Cheliped (Fig. 3 C) long and strong. Basis, 1.5 times longer than the maximum width. Merus thick and short, with three sternal setae. Carpus, about as long as basis and merus (measured together) and 2.4 times longer than wide, with two midsternal setae. Propodus (with its fixed finger), much longer than carpus; palm, shorter than fixed finger, with a row of setae on the inner surface; fixed finger thin and long, having two well developed denticles on the inner margin; 11 simple setae are disposed as in drawing; unguis small. Dactylus thin and long, curved, with numerous spinules on the inner margin; unguis small. Pereopods II-VII as in figure 4 A-F. Pleopods (Fig. 4 G) biramous, in five pairs. Basal article thick and short. Endopod, about 2.5 times longer than wide, with one midinner plumose seta and another 24 on outer margin. Exopod, about as long as endopod but slightly narrower, with 33 unequal plumose setae on the outer margin. Uropod (Fig. 3 E) biramous. Basal article small, as long as first two endopod articles, with some short distal setae. Exopod very small, uniarticulated, having one short midouter seta and two longer distal setae. Endopod filiform, five-articulated (as long as first three pleonites); each article with four or five setae. Subadult male, relatively similar to the adult, except the cheliped (Fig. 3 D) which is smaller, the main differences consisting of shape of basis (as long as carpus) and of propodus fixed finger (shorter than that in adult stage). Etymology. From the Latin bispinosus, with two spines, referring to the number of the spines present on the antenna third article. Type locality: Réunion Island (South-western of Indian Ocean). Remarks. Returning to the assertions made in the first part of paper regarding females with oostegites and antennules of five articles (representing 19.35% from all females with oostegites), I think that this phenomenon is due to polymorphism. An argument which supports this idea is that at the level of the specimens with female features (with four- or five-articulated antennule, but without eggs or oostegites) I discovered the same percentage (19.41%, from all specimens with fivearticulated antennule). The main morphological feature that distinguishes Leptochelia bispinosa n. sp. from other species of the genus is the presence of a distosternal spine on the third article of the female antenna (Fig. 1 C), a fine seta on the same article in males (Fig. 3 B). In order to remove doubt, I emphasize that I found this spine (or seta in the case of the males) in all 60 specimens (20 females with eggs or oostegites, 20 specimens with female feature and 20 males) randomly verified. In addition to the antenna spines, the new species differs from others by the combination of the following features: (1) female body seven times longer than carapace width (only 4.7 times in male); (2) maxilliped basis with six or seven long setae; (3) uropod exopod as long as endopod first article; (4) uropod endopod with five slender articles. Leptochelia corsica Dollfus, 1898 (Figs 5, 6) Studied material: 2 females, syntypes (preserved in the Collections of the Grigore Antipa National Museum of Natural History from Bucharest, with No , cf. Guþu, 1996: 151), Mediterranean Sea, Strait of Bonifacio, South of

11 ON THREE SPECIES OF LEPTOCHELIA (CRUSTACEA: TANAIDACEA) 31 Corsica Island (Campagne Melita ), depth 10 m, 21 July, 1890; 1 female, Adriatic Sea, Bari, Coast of Italia (Expedition Calypso ), 9 September 1977 (without other collecting data). Description of the female with oostegites, syntype Body (Fig. 5 A) slender, slightly dorsoventrally flattened, about 8.7 times longer than the carapace width; standard length, 8 mm. Carapace (Fig. 5 A, B), as long as first two pereonites, 1.4 times longer than its width, with three short setae on sides. Rostrum very small. Ocular lobes well developed; visual elements pigmented. Pereon (Fig. 5 A), four times longer than carapace. First pereonite, 1.8 times wider than long, shortest, having a small seta in anterolateral corners. Second pereonite, 1.7 times longer than the first, as long as wide. Third pereonite slightly longer than the preceding pereonite. Fourth pereonite longest, about two times longer than the first. Fifth pereonite slightly shorter than the previous. Last pereonite similar to the second. Pereonites two to six with one short seta on rear lateral margins. Pleon (Fig. 5 A), longer than the carapace but shorter than any two pereonites measured together (excluding from this calculation the first one), with five very short pleonites and a pleotelson. Pleonites with two short setae on sides, except the fifth which has three setae. Pleotelson, a little longer than any pleonite, with one small seta on sides and four on caudal margin, the median longest (Fig. 5 C). Antennule (Fig. 5 B, D) four-articulated, a little shorter than carapace length. First article, about 4.3 times longer than its thickness, with one median and one distal simple seta. Second and third articles similar in length; measured together shorter than the first article. Fourth article, very small, tuberculiform, with terminal setae. Antenna (Fig. 5 B, D), six-articulated. Second and third articles thick and short, each of them having one distodorsal stout spine; one distoventral robust spine is also present on the second article. Fourth article, longer than two previous together and about five times longer than its thickness, with one dorsomedian and three distoventral simple setae. Fifth article, slender, longer than any of first three articles, but approximately two times shorter than the fourth, with two long distal simple setae. Last article, tuberculiform, ending in five very long but unequal simple setae. Mandibles, maxillule and labium unstudied. Maxilliped (Fig. 5 E) with three long distoinner simple setae on basis. First article of palp short, naked. Second article with one disto-outer and two disto-inner setae. Third article, about two times longer than wide, with eleven simple setae on inner margin. Last article of palp, shorter than the preceding article and wider distally, with 12 simple setae. Cheliped (Fig. 5 F) relatively similar to that of other species of the genus. Basis about 1.8 times longer than maximum width. Merus, a little longer than half of carpus, with three ventral simple setae. Carpus, a little shorter than basis, and 2.3 times longer than wide, with one ventral and two dorsal short setae. Propodus well developed, with eight setae near the joint of dactylus (detail fig. 5 F); fixed finger thick, with two and three simple setae on the outer and inner sides, respectively; claw small. Dactylus curved, a little shorter than fixed finger; claw stout but small. Pereopod II (Fig. 6 A) much longer than following pereopods. Basis slender, about 4.8 times longer than wide, with one proximodorsal seta. Ischium short, with

12 32 MODEST GUÞU A B 0.3 mm: C C 0.3 mm: B 0.2 mm: D D 0.2 mm: E E 2 mm: A F 0.3 mm: F Fig. 5 Leptochelia corsica Dollfus, 1898, female with oostegites, syntype: A, body, dorsal; B, first pereonite, carapace, cheliped, antennule and antenna, lateral; C, last part of pleotelson and uropods; D, antennule and antenna, lateral; E, maxilliped; F, cheliped. a ventral seta. Merus, 2.5 times shorter than basis, naked. Carpus approximately as long as merus, with four distal setae. Propodus, slender, a little shorter than previous two articles together, with four distal setae. Dactylus, slender and long, with one proximodorsal seta; together with unguis much longer than propodus; unguis, shorter than dactylus. Pereopod III (Fig. 6 B). Basis, thicker than that in the previous pereopod, about 2.5 times longer than maximum width. Ischium short, with two ventral setae. Merus, a little thinner than basis and 2.5 times shorter, with one spine and one seta on distoventral corner. Carpus, a little shorter than merus, with two distodorsal setae, and one seta and one small spine, distoventrally. Propodus a little thinner than carpus but longer, with three distodorsal setae and one distoventral spine. Dacylus shorter than carpus; unguis small. Pereopod IV (Fig. 6 C, G) differs from the previous pereopod only by number of spines or setae, as shown in figure. Pereopod V (Fig. 6 D, H) slightly longer than previous pereopod. Basis, much wider than in pereopods II-IV, two times longer than maximum thickness. Ischium

13 ON THREE SPECIES OF LEPTOCHELIA (CRUSTACEA: TANAIDACEA) 33 A B C D E F 0.1 mm: G - J 0.3 mm: A - F, K G H I J K Fig. 6 Leptochelia corsica Dollfus, 1898, female with oostegites, syntype: A-F, pereopods II-VII, respectively; G-J, distal end of pereopods III-VII, respectively; K, pleopod I. small, with two ventral setae. Merus, much thinner than basis and three times shorter, with two distoventral small spines. Carpus, approximately as long as merus, with two spines and one seta, distally. Propodus, a little longer than carpus but thinner than that, with three distodorsal setae and two distoventral spines. Dactylus thick and relatively short; unguis small, apparently joined with dactylus (Fig. 6 H). Pereopod VI (Fig. 6 E, I) with minor differences in comparison with previous pereopod, as illustrated. Pereopod VII (Fig. 6 F, J) similar to two preceding pereopods, except the configuration of distal part of propodus, characterized by the presence of two spines and nine setae, seven of them short and obvious ciliate. Pleopods (Fig. 6 K) well developed, biramous, in five pairs. Basal article short and thick. Endopod, 2.8 times longer than wide, with one midinner long plumose seta; outer margin with 15 unequal plumose setae; apical seta much thicker

14 34 MODEST GUÞU than others. Exopod, as long as endopod but slightly narrower, with 33 unequal plumose setae on the outer margin. Uropod (Fig. 5 A, C) biramous. Peduncle a little shorter than first two endopod articles but thicker than these. Exopod uniarticulated, with one midlateral and two distal, relatively short, setae. Endopod filiform, as long as first three pleonites, with six slender articles, each of them with at least two setae. Remarks. The female of Leptochelia corsica differs from others of the genus by the combination of the following features: (1) the big length of body (6.2-8 mm); (2) the length of the pereonites four and five (each of them longer than broad); (3) the pleon length (longer than first two pereonites, but shorter than the combination of any two adjoining pereonites, measured together); (4) the maxilliped basis with three long setae; (5) the pereopod II propodus shorter than dactylus (together with unguis); (6) the pereopod VII propodus with nine setae; (7) the uropod endopod with six thin articles. Males, as indicated by Dollfus description (op. cit.: 43, 44), are similar to males belonging to the Leptochelia-Gruppe 2 (cf. Lang, op. cit.: 211) or to Leptochelia dubia-grouppe (cf. Sieg, op. cit.). Leptochelia tenuicula Makkaveeva, 1968 (Figs 7-9) Studied material: 29 specimens (3 females with eggs, 6 females with oostegites, 6 adult males and 14 specimens with female features, having fourarticulated antennule), Martinique Island, Lesser Antilles, Caribbean Sea, Madras, Baie de Tartane, Station Maq-14, collected in dead corals, in seagrass beds, 1-2 m deep; 14 April, Leg. Dr. Georg-Hans Müller. Description of the female with oostegites Body (Fig. 7 A), dorsoventrally flattened, 7.7 times longer than carapace width; standard length, approximately 1.5 mm. Carapace, as long as first two pereonites, and about 1.5 times longer than broad, with one median short seta on sides. Rostrum very small. Eye lobes well defined; visual elements pigmented (Fig. 7 B, D). Pereon, 3.2 times longer than carapace. First pereonite 1.5 times wider than long; second and third pereonites a little longer than the first one, 1.3 times shorter than broad; fourth pereonite longest, as long as wide; fifth pereonite shorter than the fourth one, but longer than other pereonites; sixth pereonite as the first one. Each pereonite with a small seta in anterolateral corners. Pleon, as long as carapace, with five short but wide pleonites and a pleotelson. Each pleonite with two short setae on sides. Pleotelson, a little longer than any pleonites, with one small seta on sides; caudal margin with six setae, the two median being longer (Fig. 7 C). Antennule (Fig. 7 D), a little shorter than carapace, with four articles, the last very small (tuberculiform). First article, as long as following ones (measured together), about 2.5 times longer than its thickness. Second and third articles, decreasing in thickness, 3.2 and 2.2 times shorter than the first one, respectively. The setulation as in drawings (Fig. 7 D). Antenna (Fig. 7 D) six-articulated. Second article with two spines, tergally and sternally. Third article with only one tergal spine. Fourth article, as long as

15 ON THREE SPECIES OF LEPTOCHELIA (CRUSTACEA: TANAIDACEA) 35 B A C 0.2 mm: B, C 0.1 mm: D, I D E F H I G 0.5 mm: A 0.1 mm: E - H Fig. 7 Leptochelia tenuicula Makkaveeva, 1968, female with oostegites: A, body, dorsal; B, anterior part of carapace, dorsal; C, last pleonite, pleotelson and uropods, dorsal; D, antennule and antenna, lateral; E, left mandible, distal half; F, pars incisiva of right mandible; G, maxillule; H, maxilliped; I, cheliped. previous two together, but thiner, with one short and two long setae, distally. Fifth article, thiner distally, 1.7 times shorter than fourth, with three long distal setae. Sixth article very small (tuberculiform), ending in three very long, but unequal, setae. Mandibles (Fig. 7 E, F) with pars molaris ending in some denticles. Pars incisiva ending in two small denticles in both mandibles. Left mandible with fivedenticulated lacinia mobilis. Maxillule (Fig. 7 G) with ten distal spines and setose distoexternal margin. Palp with two unequal simple setae. Maxilliped (Fig. 7 H) with a large basis, with two unequal, simple distal setae. First palp article short, naked. Second article with four setae in distoinner corner.

16 36 MODEST GUÞU Third article, longest, with six setae on inner margin. Fourth palp article with one midouter seta, and another seven situated on distal and inner sides. Endite, as in other species of the genus, with one distoexternal seta and three unequal tuberculiform spines. Cheliped (Fig. 7 I) with a well developed basis (two times longer than median width), having a conspicuous distotergal swelling and one simple seta. Merus small, with two sternal setae, one of them much longer than the other. Carpus, about 2.5 times longer than wide, with three short tergal and three long sternal setae. Propodus, a little narrower than carpus, with eight small setae near the dactylus joint, on the inner surface; fixed finger stout, with crenulate inner margin, and three tergal and two sternal setae; claw very small. Dactylus a little thinner than the fixed finger, with one proximal seta; claw small. A B C D E F 0.1 mm: A - G G Fig. 8 Leptochelia tenuicula Makkaveeva, 1968, female with oostegites; A-F, pereopods II-VII, respectively; G, pleopod IV.

17 ON THREE SPECIES OF LEPTOCHELIA (CRUSTACEA: TANAIDACEA) 37 Pereopod II (Fig. 8 A) longer than following pereopods. Basis, approximately 3.4 times longer than median thickness, with one short proximotergal simple seta. Ischium short, with a simple seta. Merus naked, 2.5 times shorter than basis. Carpus a little longer than merus, with one distotergal simple seta. Propodus, 1.5 times longer than carpus, with two distotergal setae (one of them very short) and one diststernal simple seta. Dactylus and unguis (together), as long as propodus; unguis about 1.5 times longer than dactylus. Pereopod III (Fig. 8 B) with short coxa. Basis, about 2.5 times longer than maximum thickness, naked. Ischium small. Merus, aproximately 2.6 times shorter than basis, with one distoventral small spine. Carpus, as long as merus, with one very short seta and one small spine in the distosternal corner, and one distotergal seta. Propodus, 1.5 times longer than carpus, with one distosternal spine and two distotergal setae, one of them very short. Dactylus, thicker proximally, a little shorter than carpus; unguis small. Pereopod IV (Fig. 8 C) a little shorter than the preceding pereopod, but relatively similar, except spines of merus and carpus (which are two in number). Pereopod V (Fig. 8 D) without coxa. Basis swollen, 1.9 times longer than thickness. Ischium short, with a sternal seta. Merus, 2.3 times shorter than basis, with two small distosternal spines. Carpus, shorter than merus, with two distosternal spines. Propodus, about two times longer than carpus, with one distosternal spine, and one very small spine and two setae, distotergally. Dactylus, joined with the unguis, curved, as long as merus. Pereopod VI (Fig. 8 E), relatively similar to previous pereopod; propodus with two distosternal spines, and two setae and one spine in distotergal corner. Pereopod VII (Fig. 8 F) as long as preceding pereopod. Basis, 2.2 times longer than thick. Merus with one distosternal spine. Carpus, as long as merus, with two distal spines. Propodus, 2.2 times longer than wide, and 1.5 times longer than carpus, with one distosternal spine; distotergally with one very small spine and four simple setae. Dactylus as in previous two pereopods. Pleopods (Fig. 8 G) biramous, in five pairs. Basal article about two times longer than wide. Exo- and endopod approximately equal, about two times longer than broad. Endopod with one inner plumose seta; outer margin with eight plumose setae, seven of them situated in distal half. Exopod with 12 unequal plumose setae along outer margin. Uropod (Fig. 7 A, C) biramous. Peduncle thick, relatively short. Exopod very small, uniarticulated, as long as first endopod article, with three setae, one very short, midlaterally, and two longer, terminally. Endopod filiform, five-articulated, about as long as last pleonite and pleotelson, combined; third article longest (as long as first two ones); third and fourth articles with one and two setae, respectively, and the fifth with one broom and four simple setae. Description of adult male Body (Fig. 9 A) slender, 7.3 times as long as carapace width; standard length, approximately 1.5 mm. Carapace, slightly longer than the first two pereonites. Eye lobes (Fig. 9 B) only a little larger than that in female. Pereon approximately three times longer than carapace; first three pereonites subequal, each 1.6 times shorter than wide; fourth and fifth pereonites longest, as long as broad; last pereonite shortest, length 1.8 times width.

18 38 MODEST GUÞU Pleon, as long as two longest pereonites; each pleonite with two short setae on sides (Fig. 9 C); pleotelson with six setae on caudal side. Antennule (Fig. 9 A, D) about half body length. Peduncle three-articulated; first article, approximately seven times longer than wide, as long as carapace; second article 2.6 times shorter than the first; third peduncular article short, as long as any flagellum article. Flagellum, as long as first peduncular article, with eight articles, each having 4-6 very long aesthetascs (as long as three articles of flagellum); penultimate and ultimate articles with one and three long simple setae, respectively. Antenna (Fig. 9 D), much shorter than the first peduncular article of antennule, with seven articles; first three articles short and about equal in length; second and third articles with one short tergal seta, only the second article with one sternal seta; each of the following three articles approximately times longer than the third article; last article very small, tuberculiform, ending in three setae. Cheliped (Fig. 9 D) slender, approximately as long as half of the body length; basis about two times longer than width, and 1.9 times shorter than the carapace; merus short, with two sternal setae; carpus, 4.9 times longer than thickness, with two sternal and two tergal setae; propodus (together with fixed finger), as long as carpus, with a row of setae on the inner surface, near the dactylus joint, as in drawing; fixed finger slender, half length of propodus palm, curved distally; inner side with seven small denticles in the proximal half, and an obvious swelling and three setae, distally; distoexternal side with one seta; claw very small. Dactylus, as long as fixed finger, curved distally, with a proximotergal seta and about six denticles on the inner margin; claw short. Pereopod II (Fig. 9 E) slender. Basis, five times longer than thickness, with one proximotergal seta. Ischium short, with one sternal seta. Merus, one-third length of basis, with one distosternal seta. Carpus, a little longer than merus, with four distal setae. Propodus, about as long as preceding two aticles together, with one distosternal spine and one distotergal seta. Dactylus (together with its unguis) shorter than propodus; unguis longer than dactylus. Pereopod III (Fig. 9 F) a little shorter than the previous pereopod. Basis slender, 3.6 times longer than wide. Ischium small, with one sternal short seta. Merus thin, about 4.5 times shorter than basis, having a small sternal seta. Carpus, slightly longer than merus; distally with one spine and two setulae. Propodus, longer than the length of merus and carpus (combined), with one distosternal spine and one distotergal seta. Dactylus (together with its unguis) a little longer than carpus. Unguis stronger than propodal spine. Pereopod IV relatively similar to pereopod III. Pereopod V (Fig. 9 G) with two distosternal spines on merus; carpus with four distal spines and one seta; propodus with two distosternal and two distotergal setae. Dactylus as in pereopods III or IV; unguis, apparently joined. Pereopod VI (Fig. 9 H) relatively similar to pereopod V; carpus with two distal spines, and propodus with three distotergal setae. Pereopod VII (Fig. 9 I) similar to the previous pereopod, except the size of propodus spines, which are slightly longer. Pleopods biramous, in five pairs, relatively similar to females ones. Uropod (Fig. 9 C) biramous. Basal article thick and short, a little longer than the first two endopod articles. Exopod uniarticulated and thin, as long as first two endopod articles combined, with one midouter and two distal long setae. Endopod

19 ON THREE SPECIES OF LEPTOCHELIA (CRUSTACEA: TANAIDACEA) mm: D D A 0.5 mm: A 0.1 mm: E - I B C E F G 0.2 mm: B, C H I Fig. 9 Leptochelia tenuicula Makkaveeva, 1968, adult male: A, body, dorsal; B, anterior part of carapace, dorsal; C, last part of pleon and uropods, dorsal; D, anterior part of body, lateral; E-H, pereopods II, III, V, and VI, respectively; I, distal part of pereopod VII.

20 40 MODEST GUÞU five-articulated, decreasing in thickness from the first to the last article; third article as long as any two other articles (measured together); setulation as in drawing. Remarks. The females of L. tenuicula distinguish the species from others in the genus, by the combination of the following morphological features: (1) the maxillipedal basis (Fig. 7 H) with only two simple setae (other species have at least three ones); (2) the pereopod II unguis longer than dactylus (Fig. 8 A); (3) the smaller number of the setae of pereopods II-VII propodus (Fig. 8 A-F); (4) the small number of plumose setae present on the outer margin of the pleopods rami (Fig. 8 G); (5) the long third article of uropod endopod (Fig. 7 C). Males are distinguished from others in the genus by: (1) the great length of the antennule aesthetascs (Fig. 9 D); (2) the number of antenna articles (seven, fig. 9 D); (3) the long sixth article of antenna (Fig. 9 D); (4) the configuration of cheliped (Fig. 9 D), and by some features mentioned for females (points 2-5). ACKNOWLEDGEMENTS I express my sincere thanks to my colleagues, Mrs Mihaela Achim-Barcan, who translated this paper, and Dr. Gabriel Chiºamera, who help me in the scanning of the figures, both from the Grigore Antipa National Museum of Natural History. Also I wish to thank to Prof. Dr. Richard Heard, of the Gulf Coast Research Laboratory of the University of Southern Mississippi and to anonymous reviewer for their valuable comments on the manuscript. REDESCRIEREA SPECIILOR LEPTOCHELIA BISPINOSA GUÞU ªI L. CORSICA DOLLFUS ªI PRIMA DESCRIERE A FEMELEI DE L. TENUICULA MAKKAVEEVA (CRUSTACEA: TANAIDACEA) REZUMAT Sunt redescrise speciile Leptochelia bispinosa, din apele marine ale Insulei Réunion (pentru care Guþu, 2010, a prezentat doar o succintã diagnozã) ºi L. corsica, din Marea Adriaticã (descrisã foarte sumar de Dollfus, 1898, ceea ce a condus la sinonimizarea ei eronatã cu L. dubia sau L. savignyi de cãtre Lang, 1973 ºi, respectiv, Sieg, 1983). L. tenuicula (descrisã incomplet de Makkaveeva, 1968, de pe coasta nord-vesticã a Insulei Cuba doar în baza a doi masculi) a fost redescoperitã în Insula Martinica, femela fiind descrisã pentru prima datã. Cu aceastã ocazie, într-un scurt comentariu, este contrazisã opinia unor specialiºti (Heard et al., 2004; Bamber, 2010), conform cãreia exemplarele de Leptochelia cu trãsãturi morfologice caracteristice femelelor, dar cu antenula formatã din cinci articule, ar fi femele terminale, pre-masculi neutri în stadiu final sau masculi pre-adulþi. De asemenea, au fost aduse unele amendamente la diagnoza genului Leptochelia. LITERATURE CITED BAMBER, R. N., 2005 The Tanaidaceans (Arthropoda: Crustacea: Peracarida: Tanaidacea) of Esperance, Western Australia, Australia. Pp In: F. E. Wells, D. I. Walker and G. A. Kendrick (eds), The Marine Flora and Fauna of Esperance, Western Australia, Western Australian Museum, Perth. BAMBER, R. N., 2008 Tanaidaceans (Crustacea: Peracarida: Tanaidacea) from Moreton Bay, Queensland. Memoirs of the Queensland Museum-Nature, 54 (1): BAMBER, R. N., 2010 In the footsteps of Henrik Nikolaj Krøyer: the rediscovery and redescription of Leptochelia savignyi (Kroyer, 1842) sensu strictu. Proceedings of the Biological Society of Washington, 123 (4): DOLLFUS, A., 1898 Campagnes de la Melita. Tanaidae récoltés par M. Ed. Chevreux dans l Atlantique et dans la Mediterranée. Mémoires de la Société Zoologique de France, 11: GUÞU, M., 1996 The tanaidacean types (Crustacea) present in the collections of the Oceanographic Museum from Monaco. Travaux du Muséum National d Histoire Naturelle Grigore Antipa, 36:

21 ON THREE SPECIES OF LEPTOCHELIA (CRUSTACEA: TANAIDACEA) 41 GUÞU, M., 2010 The genus Leptochelia Lang, 1973 (sic): systematic novelties (Crustacea: Tanaidacea). Pp In: D. Murariu, C. Adam, G. Chiºamera, E. Iorgu, L. O. Popa, O. P. Popa (eds), Annual Zoological Congress of Grigore Antipa Museum - Book of abstracts. Grigore Antipa National Museum of Natural History, Bucharest, Romania. HEARD, R. W., T. HANSKNECHT, K. LARSEN, 2004 An Illustraten Identification Guide to Florida Tanaidacea (Crustacea: Peracarida) Occuring in Depths of Less Than 200 m. State of Florida, Department of Environmental Protection, Tallahassee. ISHIMARU, S., 1985 A new species of Leptochelia (Crustacea, Tanaidacea) from Japan, with a Redescription of L. savignyi (Krøyer, 1842). Publication of the Seto Marine Biological Laboratory, 30 (4/6): LANG, K., 1973 Taxonomische und phylogenetische Untersuchungen über die Tanaidaceen (Crustacea). 8. Die Gattungen Leptochelia Dana, Paratanais Dana, Heterotanais G. O. Sars und Nototanais Richardson, Dazu einige Bemerkungen über die Monokonophora und ein Nachtrag. Zoologica Scripta, 2: LARSEN, K., H. RAYMENT, 2002 New species of Leptochelia (Crustacea: Tanaidacea) from the Andaman Sea, North-Eastern Indian Ocean. Phuket Marine Biological Center, Special Publication, 23 (1): MAKKAVEEVA, E. B., 1968 Vidovoi sostav i raspredelenie tanaidovyh i ravnonoghi rakov v pribrejnom raione o. Kuba. In: Anonimus, Issledovanija Centralno Amerikanskih Morei, Naukova Dumka, Kiev, 2: SIEG, J., 1983 Tanaidacea, Part 6. Pp In: H. E. Gruner and L. B. Holthuis (eds), Crustaceorum Catalogus, Hague. Received: March 14, 2011 Accepted: May 31, 2011 Muzeul Naþional de Istorie Naturalã Grigore Antipa ªos. Kiseleff 1, Bucureºti 2, România mgutu@antipa.ro