NEW DATA ON THE GENUS PARADOXAPSEUDES

Transcription

1 Travaux du Muséum National d Histoire Naturelle «Grigore Antipa» Vol. LI pp Novembre 2008 NEW DATA ON THE GENUS PARADOXAPSEUDES GUÞU, 1991, INCLUDING THE DESCRIPTION OF A NEW SPECIES. THE SYNONYMYSATION OF GOLLUMUDES BAMBER, 2000 WITH PARADOXAPSEUDES AND THE DESCRIPTION OF A NEW APSEUDID GENUS (CRUSTACEA: TANAIDACEA) MODEST GUÞU Abstract. As a result of a minute study made on some populations of the southern waters of Cuba (origin place of the species after which the description of the genus Paradoxapseudes Guþu, 1991 was made) the author reached the conclusion that the absence of the inner flagellum of the antennule in the type-species Paradaoxapseudes cubensis Guþu, 1991 (defining criterion for the above-mentioned monotypic genus) is the result of an anomaly. Also, it was established that the genus Paradoxapseudes is identical with Gollumudes Bamber, That is why the second one was synonymyzed with the first one. At the same time the Paradoxapseudes edgari n. sp., from the Cook Islands, is described. On this occasion, more or less shocking anomalies were observed (in a species of the genus Apseudes Leach, 1814 and in other two, also belonging to the genus Paradoxapseudes), which could led to a wrong description of a new taxa, as in the case of P. cubensis, above mentioned. In the final part of the paper, the new apseudid genus, Xanthapseudes, is described. Résumé. Suite à une étude détaillée de quelques populations provenant des eaux méridionales du Cuba (lieu de provenance de l espèce qui se trouve à la base de la description du genre Paradoxapseudes Guþu, 1991) on est arrivé à la conclusion que l absence du flagellum interne de l antennule chez l espèce-type Paradoxapseudes cubensis Guþu, 1991 (le critere qui définit ce genre monotypique) est la conséquence d une anomalie. On a aussi constaté que le genre Paradoxapseudes est identique avec Gollumudes Bamber, 2000, raison pour laquelle le second genre a été synonimisé avec le premier. En plus, on décrit une nouvelle espèce, Paradoxapseudes edgari, des Îles Cook. A cette occasion on a signalé plusieurs autres anomalies, plus ou moins choquantes (chez une espèce du genre Apseudes Leach, 1814 et chez deux autres, appartenant toujours au genre Paradoxapseudes) qui auraient pu conduire à la description erronée de nouveaux taxons, comme dans le cas de l espèce P. cubensis, évoquée ci-dessus. Dans la partie finale on décrit le nouveau genre d apseudides, Xanthapseudes. Key words: Xanthapseudes n. g., Paradoxapseudes Guþu, 1991, Gollumudes Bamber, 2000, Paradoxapseudes edgari, n. sp. In 1991, Guþu described genus Paradoxapseudes from the North of the Cariabbean Sea (Golfo de Ana Maria, South coast of Cuba), basing on an incomplete female (without an antennule and chelipeds). The main morphological characteristic in the genus diagnosis was the absence of the inner flagellum of the antennule. For the time being, there is another genus (also monotypic) with an uniflagellated antennule (Tanapseudes Bãcescu, 1978), after which Bãcescu (1978) described family Tanapseudidae. Under these circumstances, as it was normal, genus Paradoxapseudes was included in the mentioned family. Later, it was established that all the species belonging to the type-genus of the family Tanapseudidae had in fact a vestigial inner flagellum, represented by a very small tubercle (Bamber, 2000, fig. 4 B; Hansknecht et al., 2002, fig. 2 C; Guþu & Angsupanich, 2005, fig. 5 C), a feature which does not occur in Paradoxapseudes

2 18 MODEST GUÞU (Guþu, 1991, fig. 1 B). Besides, the presence of some morphological features completely different between the two genera led to their reclassification in different families. Therefore Tanapseudes (inclusively family Tanapseudidae, but with a subfamily rank) was initially reclassified (Bamber, 2000) in the family Sphyrapidae Guþu, Guþu (2001 a) and Hansknecht et al. (op. cit.) transferred subfamily Tanapseudinae in family Kalliapseudidae Lang, 1956, invoking the resemblances between many morphological features present in the two upper taxa. The genus Paradoxapseudes was considered to belong to family Apseudidae (Guþu, 2006), although it contravenes, by its uncommon morphology of the antennule, not only to the features of the species of this family but also to those of the entire suborder Apseudomorpha. The accidental discovery of an anomaly, represented by the tubercle-like configuration of the inner flagellum of one of the antennules (while the normal one has a flagellum with three articles) in a specimen of Apseudopsis caribbeanus Guþu, 2006, reminded me the genus Paradoxapseudes, suggesting me that it is also about an anomaly in the case of the inner flagellum absence of the antennule in the typespecies, Paradoxapseudes cubensis Guþu, My investigations made for removing the doubts on the real configuration of the inner flagellum of the antennule in the genus Paradoxapseudes confirmed the hypothesis on the presence of an anomaly at the antennule level in the type-species, P. cubensis, on the one hand, and on the other one, underlined the unquestionable resemblance between Gollumudes Bamber, 2000 and the discussed genus and the amending of Guþu s diagnosis (1991). In addition, the revision of the reclassified species in the genus Paradoxapseudes (as a result of its redefining) materialized in several comments regarding the validity of some of them, as well as by the identification of a new one from the waters of the South Pacific. In this paper, I reached these conclusions after the investigation made on a very large number of specimens originating both in the South of Cuba waters (as the type-species of the genus Paradoxapseudes) and in other geographical areas (including the South China Sea, the origin place of the type-species of the synonymyzed genus, Gollumudes). Material examined: a - 20 females and juveniles (from which only 9 with at least an intact antennule), Northern Caribbean Sea, South of Cuba, Golfo de Batabano (aproximately lat N long W), depth about 6 m; 30 May 1969; leg. Dr. Marian-Traian Gomoiu; b- 135 females and juveniles (30 with at least an intact antennule), Northern Caribbean Sea, South of Cuba, Golfo de Ana Maria (21 33 lat N long W), depth 3.5 m; 19 June 1973; leg. Dr. Modest Guþu; c - 37 females and juveniles (23 with at least an intact antennule), Northern Caribbean Sea, South of Cuba, Golfo de Guacanayabo (20 51 lat N long W), depth 5.5 m; 22 June 1973; leg. Dr. Modest Guþu; d - 25 females and juveniles (12 with at least an intact antennule), Northern Caribbean Sea, South of Cuba, Golfo de Guacanayabo (20 22 lat N long W), depth 10 m; 29 June 1973; leg. Dr. Modest Guþu; e - 60 specimens (females with eggs or oostegites, and juveniles), Northwestern Atlantic Ocean, Bermuda Islands, Walsingham Cave, 18 February 1982; leg. Dr. Thomas Iliffe; f - 29 specimens (12 females with eggs or oostegites and 17 juveniles), Northeastern Atlantic Ocean, Coast of Mauritania, February 1971 (without other collecting data); g - 93 specimens (66 females with eggs or oostegites, 6 males and 21 neutrium or juveniles), Western Indian Ocean, Coast of Tanzania; December 1973; leg. Romanian Expedition led by Dr. Mihai Bãcescu;

3 NEW DATA ON THE GENUS PARADOXAPSEUDES (CRUSTACEA: TANAIDACEA) 19 h - 12 specimens (9 females with eggs or oostegites and 3 juveniles), Southwestern Indian Ocean, La Réunion Island, Station Re-31, collected from lagoon near La Saline-les-Bains, after cyclon Firinga, mainly from dead corals; 30 January-4 February 1989; leg. Dr. Hans-Georg Müller; i specimens (54 females with eggs or oostegites and 47 juveniles), Northern Indian Ocean, (South of Sri Lanka), Ahangama, Station SL-20, collected in fringing reef, dead corals, partly covered with alge, moderately exposed reef-flat and reef channels, intertidal to shallow subtidal; 28 February, and 14 and 18 March 1993; leg. Dr. Hans-Georg Müller; j - 15 specimens (4 females with eggs or oostegites and 11 juveniles), Northeastern Indian Ocean, Strait of Malacca (Malaysia), Pulau Langkawi, Pantai Kok, Station MAS-35, collected from?agaricia coral, outer reef slope, depth 1 m; 1 April 1992; leg. Dr. Hans-Georg Müller; k specimens (178 females with eggs or oostegites, 28 males and 286 juveniles), Northeastern Indian Ocean, Strait of Malacca (Malaysia), Pulau Langkawi, Pantai Kok, Station MAS- 44, collected in outer reef margin, from dead corals, depth m; 3 and 5 March 1994; leg. Dr. Hans- Georg Müller; l - 51 specimens (15 females with eggs or oostegites and 36 juveniles), South China Sea, Malaysia, Tioman Archipelago, Pulau Babi Besar, Station MAS-3, medium part of reef-flat, collected in and under dead corals, intertidal, depth 1 m; 1 April 1991; leg. Dr. Hans-Georg Müller; m specimens (96 females with eggs or oostegites and 197 juveniles), South China Sea, Malaysia, Tioman Archipelago, Northwestern of the Babi Besar Pulau, Station MAS-6, collected in dead corals (Acropora sp., and Pocillopora damicornis) covered with alge, sponges, hydroids and ascidians, of the outer reef flat and reef margin, depth 1-2 m; 2-9 April 1991; leg. Dr. Hans-Georg Müller; n - 5 specimens (females with oostegites), South Pacific Ocean, East of Australia, Middle Banks, Moreton Bay; September-November 1976; o - 25 specimens (females with eggs or oostegites, and juveniles), Southern Pacific Ocean, Cook and Fiji Islands (more information are mentioned on the description of a new species) *10 30 *1 *2 *3 *4 *5 6* 7* *8 * * Fig. 1 Map of the studied areas (the letters between parantheses are the same with those from the list of examined material): 1, Northern Caribbean Sea, South of Cuba (a-d); 2, Atlantic Ocean, Bermuda Islands (e); 3, Atlantic Ocean, Coast of Mauritania (f); 4, Western Indian Ocean, Coast of Tanzania (g); 5, Southwestern Indian Ocean, Réunion Island (h); 6, Northern Indian Ocean, Sri Lanka Island (i); 7, Eastern Indian Ocean, Strait of Malacca, Malaysia (j, k); 8, South China Sea, Malaysia (l, m); Southern Pacific Ocean, East of Australia (n); 10, Southern Pacific Ocean, Cook and Fiji Islands (o).

4 20 MODEST GUÞU About the antennule inner flagellum of the genus Pardoxapseudes As I have already specified, the defining morphological feature of the monotypical genus Paradoxapseudes was the absence of the inner flagellum of the antennule. The observations made on 74 specimens with an entire antennule, at least (from a total of 217 specimens, partially incomplete, originating from South of Cuba, as the type-species Paradoxapseudes cubensis, but this time from Golfo de Batabano, Golfo de Ana Maria and Golfo de Guacanayabo, specified at the letters a- d in the list of the studied material), convinced me that the absence of the inner flagellum of the antennule in the mentioned species is not real, being the consequence of an anomaly. My conclusion bases on three main arguments: (1) indisputable resemblance (excepting the inner flagellum of the antennule) of the South of Cuba studied species and the type-species, P. cubensis, demonstrating that they belong to the same species, (2) the presence of the inner flagellum of the antennule in all studied specimens, and (3) discovery of an anomaly relatively similar (mentioned above) in another species of the suborder Apseudomorpha. All data regarding the size, configuration, spinulation and setulation of each part of the body and of each appendage support my conclusions from the items 1 and 2 (with little exceptions because of the variability phenomenon). The only differences observed at the body level are present in the configuration and apparent length of the pereonites four-six. Thus, the females with eggs or those with remains of marsupium (left after hatching) have the mentioned pereonites closer one another, and the anterolateral margins rounded (giving the impression of a more robust body) while the young specimens or those with oostegites have more spaced pereonites with small anterolateral prominences, more or less pointed (giving the impression of a more slender body). The differences at the appendages level are insignificant, too. These differences, influenced by variability, are represented, in general, by the number of some spines (among them also being those on the sternal margin of the pereopod II propod, rarely four instead of three) or of some setae (from the mandible, maxillule and maxilliped palp level, of the pereopods III-VII, etc., which vary between very small limits) of the number of outer flagellum articles of the antennule (usually 6 or 7, rarely 5 or 8, fig. 2 B, C) or that of the uropodal endopodite (which can have articles, fig. 2 F). A morphological feature, apparently inconstant, is the presence (or the absence) of the proximoinner denticulation from the first article of the antennule peduncle (difficult to be seen in many specimens, giving the impression that it is absent, fig. 2 A-C). At the same time, I remarked the constant presence of some features, apparently insignificant, but which can help a correct identification of some species. Especially, I take into consideration the displaying of the tergal setae on the basis of pereopods II and VII (Fig. 3 A, C) or on the tergal margin of pereopod II carpus (Fig. 3 A) and some appendages (as in the case of some setae on the inner margin of the first and second article of mandible palp, fig. 2 E). The mentioned resemblances and differences are present in all studied populations, not matter the collecting site was. As regards the inner flagellum of the antennule in the species P. cubensis, it usually has two articles (Fig. 2 B). Rarely it can have three (Fig. 2 C), or exceptionally, a single article (Fig. 2 A). Thus, from the total of 74 studied specimens with an entire antennule, provided from the Northern Caribbean Sea (9 from Golfo de Batabano, 30 from Golfo de Ana Maria and 35 from Golfo de Guacanayabo), 69 (representing around 93%) had the inner flagellum formed of two articles (8 from Golfo de Batabano, 27 from Golfo de Ana Maria and 34 from Golfo

5 NEW DATA ON THE GENUS PARADOXAPSEUDES (CRUSTACEA: TANAIDACEA) 21 A 0.1 mm: A 0.2 mm: B, C, D C D B E F 0.1 mm: E Fig. 2 Paradoxapseudes cubensis Guþu, 1991, females from Caribbean Sea, South of Cuba: A-C, antennules; D, antenna; E, mandible palp; F, last pleonite, pleotelson and uropods. de Guacanayabo), 4 (5.4%) of three articles (1 specimen from Golfo de Batabano and 3 from Golfo de Ana Maria), and 1 specimen (from Golfo de Guacanayabo, representing 1.35% from the total of the studied material) had the inner flagellum formed of a single article. Under these circumstances, I specify that I discovered another two specimens with the inner flagellum uniarticulated (from a number of 284 complete specimens, representing 0.7%), originating in the Strait of Malacca, Indian Ocean (mentioned at letter k from the studied material). Although the presence of the uniarticulated inner flagellum can be included in the anomaly category, because of its rarity, I consider it a very rare case of variability. Exceptionally, I considered an anomaly only the specimen from the Caribbean Sea, considering that both flagella of the antennule have an uncommon little number of articles (one and, respectively, four, fig. 2 A). The third mentioned argument refers to a specimen of Apseudopsis caribbeanus (preserved in the collections of Grigore Antipa National Museum of Natural History, Bucharest, with no ), originating in the Gulf of Batabano,

6 22 MODEST GUÞU A B C 0.2 mm: A A C Fig. 3 Paradoxapseudes cubensis Guþu, 1991, female from Caribbean Sea, South of Cuba: A, pereopod II; B, pereopod VI; C, pereopod VII. South of Cuba (Guþu, 2006), as the type-material. In this case, the inner flagellum of the left antennule is represented by a single tubercle (Fig. 9 A) while the right antennule is normal, three-articulated (Fig. 9 B). Admitting that, by accident, the specimens of A. caribbeanus had only the abnormal antennule (the normal one being lost during the material handling, as often happens), and the collecting geographical area had been another one than the origin place of the type-material, it would have surely been the base of the description of a new taxon, as in Paradoxapseudes. As a result of the above mentions, I can surely assert that the absence of the inner flagellum of the antennule in the specimens after which the monotypic genus Paradoxapseudes was described is because of an anomaly. One of the numerous causes of the appearance of this anomaly can be the excessive fragility of the antennules (as well as of the chelipeds, as a matter of fact), which rarely can lead to an incomplete regeneration, even aberrant. In this respect, I already mentioned that from the total of 217 specimens present in the mentioned samples (letters a-d from the studied material), only 74 (34%) had entire antennules, one or both, the other

7 NEW DATA ON THE GENUS PARADOXAPSEUDES (CRUSTACEA: TANAIDACEA) specimens (representing 66%) having both antennules broken (and lost) from the articles 2-3 level of the peduncle. On my opinion, this excessive fragility, known also in other apseudomorphs of shallow waters (Guþu, 1996, Bamber et al., 2003) does not manifest only during material collecting and handling but also under some influences of some natural physical factors, as the brutal turbulence of water (generated by the strong storms or by the tide), which, in its turn, draws coarse sediments from the bottom, creating serious injuries to the benthic microfauna. The synoymisation of the genus Gollumudes Bamber, 2000 with Paradoxapseudes Guþu, 1991 Correlation of the descriptions from literature with my own observations gathered during the study on the specimens collected from the three South of Cuba bays (assigned to the species Paradoxapseudes cubensis) but also on many others (originating in the waters of the South China Sea, Southern Pacific Ocean and in several areas of the Indian and Atlantic oceans, mentioned at the letters e-n in the list of the studied material) allowed me to establish that, at the level of the body and of the appendages, P. cubensis has numerous common morphological features with those of the species Apseudes bermudeus from the submarine caves of the Bermuda Islands (Bãcescu, 1980), A. garthi from the Gulf of California (Menzies, 1953), A. intermedius from the waters of the Cape Verde Islands (Hansen, 1895), from Morocco (Monod, 1925) and of the Mediterranean Sea (Larwood, 1940), A. larakia from Australia (Edgar, 1997; Guþu, 2006), A. littoralis from the Japanese waters (Shiino, 1952) and the Tanzanian ones (Guþu, 2007 b), A. mortoni from Southeastern Asia (Bamber, 1997) and A. tropicalis from Hawaii Islands (Miller, 1940), all of them transferred in genus Gollumudes (Bamber, 2000; Guþu, 2001 b, 2006, 2007 b). Moreover, I come to the conclusion that the species Apseudes heroe, described by Sieg (1986) from the Southwestern Atlantic Ocean (Coast of Argentina), is very similar to above mentioned ones, all together belonging to the same group. At the same time, the mentioned species have many common features with the other two species classified in the genus Gollumudes (G. botosaneanui, from the Curaçao Island, cf. Guþu, 2001 b, and G. basibidens, from the coast of Tanzania, cf. Guþu, 2007 b), this demonstrating that they belong to a single genus. Under these circumstances, basing on the priorities stipulated by the International Code of Zoological Nomenclature (1999), genus Gollumudes Bamber, 2000 becomes synonym with Paradoxapseudes Guþu, Genus Paradoxapseudes Guþu, 1991 New diagnosis (modified after Guþu, 1991, 2007 b and Bamber, 2000). Body of small size (2-3 mm length), dorsoventrally flattened. Carapace, longer than wide; rostrum prominent, acuted anteriorly; ocular lobes well defined, generally with pigmented visual elements. Pereon with last four pereonites similar and approximately equal. Pleon with five short pleonites having some lateral circumplumose setae. Antennule with a fine denticualtion on the proximal half of inner margin of the first peduncular article, generally hard visible; inner flagellum with two or three articles (accidentally, with only one). Antenna with fourth and fifth articles long and thin; squama with 3-6 setae. Mandibles with three-articulated palp; first article with 3-5 long simple setae; second article with second or third seta much longer than others. Labium palp ovate, slightly narrower distally, ended in three slender spines. Maxillule with two-articulated palp. Maxilliped with one long

8 24 MODEST GUÞU circumplumose seta in the distoinner corner of basis and the first article of palp; second palp article with one strong spine in distal half of outer margin. Epignath cup-shaped; spine well developed, slender, with long setae in the last half. Cheliped with exopodite; adult females with polymorph cheliped; males chelipeds unequal and different. Pereopod II fossorial, with exopodite; coxa with a small anterior prolongation, rounded in top, ended in two-three circumplumose setae; basis with three to five plumose setae (apparently simple in many cases) on the tergal (anterior) margin; propodus with three or four sternal spines. Pereopods VI and VII with relatively similar propodus, by the row of small ciliate setae (comb-like) disposed in the last sternal half. Pleopods biramous, in five pairs. Uropod exopodite with five articles; endopodite with at most 24 articles. Females with five pairs of oostegites. Type-species: Paradoxapseudes cubensis Guþu, Composition. By the synonymysation of the two genera and in conformity to the data from literature (Bãcescu, 1980; Bamber, 1997, 2000; Edgar, op. cit.; Guþu, 1991, 2001 b, 2006, 2007 b; Hansen, op. cit.; Menzies, op. cit.; Miller, op. cit.; Shiino, op. cit.), the genus Paradoxapseudes should have 12 species: P. basibidens (Guþu, 2007), comb. nov., P. bermudeus (Bãcescu, 1980), comb. nov., P. botosaneanui (Guþu, 2001), comb. nov., P. cubensis Guþu, 1991, P. garthi (Menzies, 1953), comb. nov., P. heroe (Sieg, 1986), comb. nov., P. intermedius (Hansen, 1895), comb. nov., P. larakia (Edgar, 1997), comb. nov., P. littoralis (Shiino, 1952), P. mortoni (Bamber, 1997), comb. nov., P. tropicalis (Miller, 1940) and P. edgari n. sp. In the above mentioned enumeration I haven t taken into consideration the synonymysation (wrong, on my opinion) mentioned by Sieg (1983: 59-60) between Apseudes garthi Menzies, 1953 and Apseudes intermedius Hansen, 1895 (reclassified in the genus Gollumudes, cf. Guþu, 2007 b, now the genus Paradoxapseudes), to which I shall refer further on. On the validity of some species. The analysis of the morphological features of the species Apseudes garthi Menzies, 1953, as they result from literature (Menzies, op. cit.), allowed me to establish that there is no significal difference between it and A. tropicalis Miller, 1940 (both reclassified in the genus Gollumudes, cf. Guþu, 2006, 2007 b). The different number of the articles of the inner flagellum of the antennule (three in A. tropicalis, cf. Miller, op. cit.: 307 and only two in A. garthi, cf. Menzies, op. cit.: 448) could suggest the presence of two different taxa. The difference from the number of the inner flagellum articles mentioned by Menzies (op. cit.) and Miller (op. cit.) is generated, on my opinion, by the way in which the two authors counted (with or without the common article of the two flagella, corresponding to the fourth article of the antennule). I maintain this idea because, as it is known, many previous authors (Bãcescu, 1961; Kudinova-Pasternak, 1970; Lang, 1956, 1968; Larwood, op. cit.; Norman and Stebbing, 1886, etc.) considered the common article of the two flagella of the antennule as belonging to the inner flagellum. From the correlation of the data of the text with those from the figures (in the species A. garthi) it results that Menzies (op. cit.: 448 and fig. 2 A) did not take into consideration the common article of the two flagella. Unfortunatelly, because of the unclear drawing presented by Miller (op. cit., fig. 4 c), I cannot make the same correlation in the species A. tropicalis. As it results from other Miller s descriptions (op. cit.) and clear figures, I think that he considered the common article of the two flagella of the antennule as belonging to the inner flagellum. My conclusion bases on the comparison between the data from the text with those from the drawing in the description of the species Apseudomorpha oahuensis (Miller, op. cit.: 315 and fig. 8

9 NEW DATA ON THE GENUS PARADOXAPSEUDES (CRUSTACEA: TANAIDACEA) 25 c). Even if my judgement would be wrong, the presence of the variability at the level of the antennule inner flagellum within the same population (of Paradoxapseudes cubensis, commented above) gives me the right to think that it could be about variability in the case of the two very similar species from the North Pacific, A. tropicalis (from Hawaii Islands) and A. garthi (from the Gulf of California). A D B E 0.2 mm: A, B, D F 0.1 mm: C F C Fig. 4 Paradoxapseudes littoralis (Shiino, 1952), comb. nov., female from Indian Ocean, Strait of Malacca: A, antennule; B, antenna; C, mandible palp; D, pereopod II; E, pereopod VI; F, pereopod VII. More than that, the correlation of the descriptions from literature of the Indo- South-Pacific species, reclassified in the genus Gollumudes (it is about A. mortoni Bamber, 1997, A. larakia Edgar, 1997, sensu Edgar, op. cit., and Guþu, 2006, and A. littoralis Shiino, 1952, sensu Shiino, op. cit., and Guþu, 2007 b) with the personal observations made on a rich material (letters g-n from the list of the studied material) originating in the waters of the Indian Ocean (the coasts of Tanzania, the Strait of Malacca, and of Sri Lanka and Réunion islands), the Southern Pacific (East of Australia) and those of the South China Sea (Tioman Archipelago), did not point out any major morphological difference between them and the North-Pacific species

10 26 MODEST GUÞU A C B 0.3 mm: A C Fig. 5 Paradoxapseudes littoralis (Shiino, 1952), comb. nov., from Indian Ocean, Strait of Malacca: A, B, left and right male cheliped, respectively; C, last pleonite, pleotelson and uropods (female). (A. garthi, A. tropicalis and A. littoralis). A simple comparison between the configuration of the body and of the main appendages (antennule, antenna, mandible palp and pereopods II, VI and VII, figs 4, 6) in the studied specimens and the mentioned species, as it results from the published data (Bamber, 1997, figs 1-3, 2000, fig. 1 E; Edgar, op. cit., figs 1-3; Guþu, 2006, figs , 2007 b, fig. 5; Menzies, op. cit., fig. 2; Miller, op. cit., fig. 4; Shiino, op. cit., figs 4, 5) absolve me of any comment. As it can be remarked, the only main difference is the different number of the articles of the antennule inner flagellum (which can have two, three or, rarely, a single article, as I have already mentioned). I consider that this inconsistency, observed even in the specimens of some populations which originate in the same collecting station (the case of those ones from the Strait of Malacca or South of Cuba, fig. 2 A-C, and the East of Australia, cf. Guþu, 2006, fig. 116) is due to the variability, and does not justify (in the absence of other differences) their declaration as bona species. As a matter of fact, the same variability can be also observed in the cheliped configuration (characterized by a strong polymorphism, cf. Bãcescu, 1980 and Guþu, 2007 b), in the number of the outer flagellum articles of the antennule (which can vary between 5 and 8), in the sternal spines on the pereopod II propodus (three, rarely four) or in that of the ciliate setules on the distosternal margin of the pereopod VI and VII propodus (Figs 4 E, F, 6 E, F). At the population level, the single observed differences are between the South of Cuba specimens and the South Asia ones. They are represented by the size of the proximoinner denticles

11 NEW DATA ON THE GENUS PARADOXAPSEUDES (CRUSTACEA: TANAIDACEA) 27 on the first article of the antennule peduncle (smaller and hard to be observed in the specimens from Cuba, fig. 2 A-C, but which are not mentioned or figured in the species described by Bamber, 1997, Edgar, op. cit., Miller, op. cit., and Shiino, op. cit.) and by the length of uropod endopodite and its article number: two times longer than the pleotelson, having articles, in South of Asia specimens (Fig. 5 C), and at least three times longer than the pleotelson, having articles, in the Northern Caribbean Sea ones (Fig. 2 F). In conclusion, in the absence of some significant morphological differences which allow the unquestionable identification of each of them, it is posible that at least some of the species Apseudes garthi Menzies, 1953 (from the Gulf of California), A. littoralis Shiino, 1952 (from the Japanese waters), A. mortoni Bamber, 1997 (from the South China Sea) and A. larakia Edgar, 1997 (from Australia) to be synonymous with Paradoxapseudes tropicalis (Miller, 1940), comb. nov. (from the Hawaii Islands), no matter they have the antennule inner flagellum, bi- or three-articulated. In above situation the identification main criterion would be the geographical one (without precise borders) to the prejudice of the morphological criteria, which should take priority, and in which absence taxonomy might become an empty word. Or, I don t say something new asserting that the two criteria, morphological and zoo-geographical, should identify each other. Coming back to the above example, it is not impossible to deal with a genus with cryptic species, but at the same time we can deal with a A B D E 0.2 mm: A, B, D F 0.1 mm: C F C Fig. 6 Paradoxapseudes littoralis (Shiino, 1952), comb. nov., from South China Sea: A, antennule; B, antenna; C, mandible palp; D, pereopod II; E, pereopod VI; F, pereopod VII.

12 28 MODEST GUÞU 0.2 mm: A, B, D F 0.1 mm: C A B D E F F C Fig. 7 Paradoxapseudes intermedius (Hansen, 1895), comb. nov., from Atlantic Ocean, Coast of Mauritania: A, antennule; B, antenna; C, mandible palp; D, pereopod II; E, pereopod VI; F, pereopod VII. cosmopolitan species, hypotheses which will be confirmed or annul only using some molecular studies. On the other hand, the investigations made on the specimens from the Bermuda islands and from the Mauritania waters (mentioned at the letters e and f, respectively, from the studied material) made me to consider the taxa Paradoxapseudes bermudeus (Bãcescu, 1980), comb. nov., and, respectively, P. intermedius (Hansen, 1895), comb. nov., as valid species. I reached the same conclusion for the species P. botosaneanui, comb. nov. Although all of them have the inner flagellum of the antennule constantly formed of three articles, giving the apparent impression that they belong to a single species, they distinguishes by some other features. Thus, P. intermedius distinguishes from the other species of the genus by the large number of the circumplumose setae at the level of the last two pereonites and of the five pleonites, the small length of the pleotelson, displaying and number of the setae on the second article of the mandible palp (Fig. 7 C), the presence of the five tergal setae and of four sternal spines on the basis and, respectively, on the pereopod II propodus (Fig. 7 D), and the large number of the circumplumose setae on the tergal margin of the pereopod VII basis (Fig. 7 F).

13 NEW DATA ON THE GENUS PARADOXAPSEUDES (CRUSTACEA: TANAIDACEA) 29 Within this context I underline that the morphological features of the species Apseudes garthi (=Paradoxapseudes garthi, comb. nov.) as it results from Menzies description, op. cit.) are different from those of A. intermedius (=P. intermedius, comb. nov.) and the synonymyzation of the first species with the second one, made by Sieg (op. cit.), becomes no longer valid. P. bermudeus characterises by the large number of the setae at the level of the last two articles of the mandible palp (Fig. 8 C), by the length/width ratio of the carpus and propodus of the pereopods VI and VII, but also by the much large number of the distosternal setules (comb-like aspect) on the propodus of the last two pereopods (Fig. 8 E, F). Although the species P. botosaneanui resembles P. intermedius, it distinguishes from the last one by the number of the setae of second article of mandible palp (greater in the second taxon), by the number of the sternal spines on the pereopod II propodus (3, instead of 4), the large number of the articles of the endopodit of the uropods (22-23, cf. Guþu, 2001 b, figs 1 A, D and 2 B, instead of only 17-18, etc.). By the article number of uropod endopodite, P. intermedius resembles P. cubensis, but these two species have some differences consisting in the A B D E 0.3 mm: A, B, D F C F 0.1 mm: C FigFig. 8 Paradoxapseudes bermudeus (Bãcescu, 1980), comb. nov., from Atlantic Ocean, Bermuda caves: A, antennule; B, antenna; C, mandible palp; D, pereopod II; E, pereopod VI; F, pereopod VII.

14 30 MODEST GUÞU number of the setae at the level of the second article of mandible palp and the basis pereopod VII (Figs 2 E, 3 C, 8 C, F), etc. By the smaller number of setae at the level of pereopod VII basis, P. heroe, comb. nov. (Sieg, 1986, fig. 5 P6) differs from P. botosaneanui (Guþu, 2001 b, fig. 2 G) and P. intermedius (Fig. 7 F), but by the greater number of the setae of mandible palp, the Sieg s species differs from all other ones, as it results from Bamber (1997, fig. 2 A), Edgar (op. cit., fig. 1 Mdr), Guþu, 1991 (Fig. 1 D), Guþu (2001 b, fig. 1 D), Guþu (2007, fig. 6 F), Menzies (op. cit., fig. 2 B), Miller (op. cit., fig. 4 d), Shiino (op. cit., fig. 5 E), Sieg (1986, fig. 3 Md), and the illustration of this paper (Figs 2 E, 4 C, 6 C, 7 C, 8 C). Some remarks on the males chelipeds. Guþu (2006, 2007 a), basing on literature (Bãcescu, 1961; Riggio, 1996) on the simultaneous hermaphrodism of the species Apseudes intermedius Hansen, 1895 (now Paradoxapseudes intermedius, comb. nov.), reclassified the mentioned taxon in the genus Muramurina Guþu, Also Guþu (2007 b) pointed out that the chelipeds, considered to belong to some males (Bamber, 1997, fig. 3 C; Edgar, op. cit., fig. 2) of the present genus Paradoxapseudes (it is about Gollumudes mortoni, and, respectively, Apseudes larakia, now both classified in the genus Paradoxapseudes) in fact belong to same females. On this occasion Guþu (2007 b) established that the chelipeds of the females of Gollumudes littoralis (= P. littoralis, comb. nov.) have a strong polymorphism (also remaked in P. bermudeus comb. nov., cf. Bãcescu, 1980, figs I, J, P), this explaining the confusions made by Bamber (1997) and Edgar (op. cit.). At the same time Guþu (2007 b) described, for the first time from the waters of Tanzania, the males of the species P. littoralis, comb. nov., characterized by unequal chelipeds, aspect confirmed today by the discovery of other 28 males with unequal chelipeds (very similar to Guþu s description, 2007 b, fig. 5 F, G), belonging to the same species (Fig. 5 A, B) provided from the waters of the Strait of Malacca (letter k from the studied material). Also, the males of another reclassified species in the genus Paradoxapseudes (respectively, P. basibidens Guþu, 2007, comb. nov.) have unequal chelipeds (Guþu, 2007 b, fig. 7 B, C). Unfortunately I haven t found specimens of P. bermudeus comb. nov. and P. intermedius, comb. nov., with unequal chelipeds which can certify the presence of the males, although I investigated large populations (mentioned at letters e and f from the studied material). Either they don t exist, because of the simultaneous hermaphrodism, as I have said that I was mentioned in literature (Bãcescu, 1961; Riggio, op. cit.), or they have a short (temporary) existence, or are extremely rare and were not discovered. The report of the males of Apseudes cf. bermudeus (=Paradoxapseudes cf. bermudeus, comb. nov.) by Suarez-Morales et al. (2004: 20, 24) in the Mexican waters of the Caribbean Sea is, on my opinion doubtful, in the absence of a minimum description of some specifications on dimorphism. I haven t included in these comments the species P. edgari n. sp., described futher on, after a relatively little number of specimens, which can explain the absence of the males. New anomalies recorded in the species of the genus Paradoxapseudes Verification of the morphological features of a large number of specimens from different geographical areas allowed me to discover other two morphological deviations (more or less serious), which can be add to that regarding the absence of the antennule inner flagellum in the type-species P. cubensis to which I refer in

15 NEW DATA ON THE GENUS PARADOXAPSEUDES (CRUSTACEA: TANAIDACEA) 31 the first part of the paper. In this deviation category the uniarticulated inner flagellum of the antennule, discovered in two specimens of P. littoralis, comb. nov., from the Strait of Malacca (letter k from the studied material) could have been included. As I have mentioned in the comments on the antennule variability, I included only the South of Cuba specimen (preserved in the collections of Grigore Antipa National Museum of Natural History, Bucharest, with no ) in the anomaly category because of the major morphological deviations present at the level of the both flagella of the antennule (inner flagellum having one article, and the outer one 4, instead of 2-3 and, respectively, 6-7, fig. 2 A). I discovered the second anomaly, much shocking by its importance, in a female of P. bermudeus, comb. nov., which could generate a major confusion in the apseudomorph systematics if it originated in another geographical area than the type-specimens, as in the case of the type-species P. cubensis (but also of the abovementioned species, Apseudopsis caribbeanus). The mentioned specimen (preserved in the collections of Grigore Antipa National Museum of Natural History, Bucharest, with no ) is from a sample with numerous specimens collected also from Walsingham Cave, Bermuda Islands (letter e, from the list of the studied 0.5 mm: A, C 0.2 mm: B A B C Fig. 9 Apseudopsis caribbeanus Guþu, 2006 (A, B) and Paradoxapseudes bermudeus (Bãcescu, 1980), comb. nov. (C): A, abnormal left antennule, with inner flagellum reduced to a tubercle; B, median part of right antennule, with normal inner flagellum; C, last two pereonites and an abnormal pleon, reduced to four pleonites.

16 32 MODEST GUÞU material) as the type-specimens of the species described by Bãcescu (1980). This time, the anomaly is represented in the diminishing the free pleonite number to four (Fig. 9 C), instead of five, as most of the apseudomorphs have (less the species of the subfamily Synapseudinae, family Metapseudidae). Also, the number of the pleopods is of four pairs (one on each pleonite) instead of five, occurred in P. bermudeus, comb. nov. From the analysis of the four pleonites, I remarked that the first segment is the longest (Fig. 9 C), which makes me to think that the anomaly is due to the intimate fusion between the first two pleonites and not to the wrong regeneration, as in antennules. The report of the three anomalies only in the species of a single genus, respectively Paradoxapseudes (out of which the complete absence of the antennule inner flagellum in P. cubensis and the diminishing of the pleonites and pleopods to four, instead of five, in P. bermudeus, comb. nov., I consider the serious anomalies), to which the wide variability of the article number of the two antennule flagella can be added, makes me to question, rhetorically: is this genus predisposed especially to such a phenomenon? My question is justified by the fact that, although I have made minutely observations on large populations of species belonging to other genera, I did not observe so many and serious morphological deviations which exceed the limits (restricted) of a normal variability. Paradoxapseudes edgari n. sp. (Figs 10-12) Material examined: 25 specimens provided as follows: 22 specimens (1 female with eggs, 6 females with remnants of marsupium, 6 females with ostegites, 5 juveniles and 4 manca) from the Mangaia Island (Cook Islands), 23 October 1987, and 3 specimens (1 female with eggs and 2 females with oostegites) from Fiji Island, 1988, both samples collected by Dr. Thomas Iliffe. Holotype, female with oostegites, from the Mangaia Island, preserved in the Collection of the Grigore Antipa National Museum of Natural History, Bucharest, No Paratipes, 15 specimens (1 female with eggs, 4 females with remnants of marsupium, 3 females with oostegites, 3 juveniles and 4 manca) from the Mangaia Island and 3 specimens from Fiji Island (1 female with eggs and 2 females with oostegites), in the same museum, No and , respectively, and 6 specimens (2 females with remnants of marsupium, 2 females with oostegites and 2 juveniles) from the Mangaia Island in the Collection of Smithsonian Institution, Washington. Description of the female with oostegites Body (Fig. 10 A) slender, flattened dorsoventrally, slightly decreasing distally, about 7.4 times longer than the carapace width; standard length, 2.7 mm. Carapace, longer than broad, with two small setae on each side in the first half. Rostrum well developed, acuted. Ocular lobes well defined, with pigmented visual elements. Pereon 2.8 times longer than the carapace. First pereonite shortest, approximately two times wider than long, with parallel sides; anterior and posterior corners rounded in dorsal view. Second pereonite, rounded laterally, a little longer than the first one but shorter than another ones, with four short simple setae on each

17 NEW DATA ON THE GENUS PARADOXAPSEUDES (CRUSTACEA: TANAIDACEA) 33 A 0.2 mm: C, D 0.1 mm: E, F, H D B C E 0.3 mm: B F G 1 mm: A H 0.05 mm: G Fig. 10 Paradoxapseudes edgari n. sp., female with oostegites, paratype: A, body, dorsally; B, right part of last two pereonites and pleon; C, antennule; D, antenna; E, right mandible, rostral part; F, pars incisiva, lacinia mobilis and setiferous lobe of left mandible; G, labium; H, maxillule.

18 34 MODEST GUÞU side. Last four pereonites similar, slightly shorter than wide, with some simple setae in anterolateral corners and laterally (Fig. 10 A, B). Pleon, as long as first four pereonites, with five short pleonites and a pleotelson. Each pleonite with four or five long circumplumose setae on lateral margins and other one, shorter, dorsolaterally. Pleotelson narrow, as long as last pereonite and first pleonite measured together, with some lateral long apparently simple setae (Fig. 10 A, B). Antennule (Fig. 10 C) much longer than the carapace. First peduncular article, 4.5 times longer than its maximum thickness, with some hard visible fine denticles in the proximoinner margin and some simple and broom setae on each side. Second peduncular article, about three times shorter than the first one, with some broom and simple setae, distolaterally. Third peduncular article, shorter and narrower than the second one, with two distolateral setae. Outer flagellum, seven-articulated, as long as first peduncular article (if we exclude the terminal flagellum setae); first six articles with one-two simple setae and the last one with three long setae; one aesthetasc is present on the fourth and sixth articles. Inner flagellum threearticulated (excluding the common article); first and second articles with two setae and the third one with three long simple setae. Antenna (Fig. 10 D) with the first peduncular article short. Second article, largest, as long as following two articles, with one median simple seta on each lateral margin and one dentiform process in distoinner corner. Third peduncular article very short with one distoinner seta. Third and fourth peduncular articles slender, each of them as long as the first two articles of flagellum. Squama well developed with five long simple setae. Flagellum five-articulated, as long as last three peduncular articles; first article with two long simple setae on the outer margin; second, third and fourth article with two simple, relatively short, setae, and the last article with three simple setae. Mandibles (Fig. 10 E, F) with acute tubercles on the outer margin (hard visible), near the palp insertion. Palp long, three-articulated; first article, approximately two times longer than broad and about 2.5 times shorter than the second article, with five long simple setae on the rostral margin; second article thin, about 5.5 times longer than its thickness, with eight ciliate setae on the rostral margin, the second one being much longer than the anothers; third article, about two times shorter than the preceding one, with nine unequal setae in the inner distal half and terminally. Pars incisiva of both mandibles four-denticulated. Setiferous lobes with four furcated and one simple seta. Lacinia mobilis of left mandible threedenticulated. Pars molaris without special features. Labium (Fig. 10 G) with some denticles on the outer side. Palp ovate, narrower distally, about two times longer than broad, with three setiform spines in top and long lateral hairs. Maxillule (Fig. 10 H) with biarticulated palp (ended in four subequal setae). Outer endite with eleven spines and two setae, distally and submarginally, respectively; both sides with numerous hairs. Inner endite with four ciliate setae. Maxilla unstudied. Maxilliped (Fig. 11 A) with short coxa. Basis, about as long as broad, with one long circumplumose seta in distoinner corner. First palp article short with one long circumplumose seta in distoinner corner and one setiform spine in outer angle. Second palp article, longer than broad, with one long simple and two long plumose setae, and at least 17 smaller and unequal simple setae on the inner margin; one long setiform

19 NEW DATA ON THE GENUS PARADOXAPSEUDES (CRUSTACEA: TANAIDACEA) 35 A B C 0.2 mm: A 0.4 mm: C E 0.2 mm: B, F D E F Fig. 11 Paradoxapseudes edgari n. sp., female with oostegites, paratype: A, maxilliped; B, epignath; C, cheliped; D, cheliped of another female; E, pereopod II; F, pleopod.

20 36 MODEST GUÞU spine is present in distoexternal margin. Third palp article with five long and three short simple setae in the second inner half. Fourth palp article with one very small and six long simple setae in the distal margin. Endite with some different setae on the rostral margin, and three couplers and some plumose setae on the inner margin. Epignath (Fig. 11 B) cup-shaped, approximately two times longer than broad, with one long spine having about 12 long setae in the last half; opposite to spine are present two small rounded lobes each of them with short hairs on the outer margin. Cheliped (Fig. 11 C) with exopodite. Basis well developed, at most two times longer than wide; tergal margin, relatively straight, with one median more or less triangular prominence and two distal small setae; sternal margin, rounded, with one proximal and two distal setae, and one median setiform spine. Merus with one simple seta in the distosternal rounded margin. Carpus, about as long as basis, much wider distally, with seven unequal simple setae on the sternal margin. Propodus very large, greater than the basis, with three small proximotergal and eight sternal setae; fixed finger thick, with two proximal long but unequal setae and other six smaller, distally, between which is present one great protuberance; claw small. Dactylus curved, thinner than the fixed finger, with three distal long simple setae; claw well developed. Pereopod II (Fig. 11 E) with exopodite. Coxa with an anterior small rounded prominence ended in two plumose setae. Basis, 3.3 times longer than broad; proximosternally with one long circumplumose seta, and distosternally with two unequal simple setae and one small spine; tergally with four long, apparently simple, setae. Ischium short with three distosternal unequal setae. Merus, two times shorter than the basis, with seven setae and one spine on the sternal margin; distotergally with four unequal setae and one fine spine. Carpus, about as long as merus, with four setae and two spines, sternally, and ten subequal setae (some of them very long) and one stout spine, tergally. Propodus, narrower and slightly shorter than the carpus; sternally with four setae which alternate with three spines, and one serrated seta at the basis of dactylus insertion; distotergally with two long setae and two spines. Dactylus slender, with two sternal spinules; midtergally and distosternally with one very small seta; claw relatively short. Pereopod III (Fig. 12 A) slender. Basis, four times longer than wide, with two long distosternal simple setae. Ischium small. Merus about 3.2 times shorter than the basis; sternally with three setae and one spine and distotergally with two long simple setae. Carpus, 1.5 times longer than the merus, with three sternal and five distotergal long simple setae, and two distal fine spines. Propodus, slightly narrower and longer than the carpus; sternally and distotergally with six slender spines and four long setae. Dactylus thin, a little longer than the adjacent spines; claw fine and long. Pereopod IV (Fig. 12 B) a little shorter and relatively similar to pereopod III. Pereopod V (Fig. 12 C) different from the other pereopods. Basis, four times longer than wide, with one midsternal broom seta and other one simple, distosternally. Ischium short with two unequal distosternal setae. Merus, only a little longer than the ischium; disosternally with one spine and one seta, and distotergally with one setiform spine. Carpus, two times longer than the merus; midsternally with two spines, and distosternally with one seta and two fine spines; distally and distotergally with three long setae and one spine. Propodus, as long as carpus, with one midsternal and one midtergal spine; distally with one very long ciliate seta and