AÇOREANA, Suplemento 6, Setembro 2009: 183-200

THE TANAIDACEANS (ARTHROPODA: : ) OF SÃO MIGUEL, AZORES, WITH DESCRIPTION OF TWO NEW SPECIES, AND A NEW RECORD FROM TENERIFE

Roger N. Bamber1 & Ana Cristina Costa2

1The Natural History Museum, Cromwell Road, London SW7 5BD, U.K. e-mail: [email protected] 2 CIBIO-Pólo Açores, Department of Biology, University of the Azores, 9501-801 Ponta Delgada, São Miguel, Azores, Portugal

ABSTRACT During the Third International Workshop of Malacology and Marine Biology in São Miguel, Azores, in July 2006, sampling of the littoral and sublittoral benthos was under- taken in order to characterize the smaller marine fauna of this region, including tanaidaceans. In the event, 338 tanaidacean specimens representing three species were collected; two of these species, Leptochelia caldera and Paratanais martinsi, are new to sci- ence. In addition, previous tanaidacean material collected around São Miguel in 1996 and 1997 was analyzed, from which a further 272 specimens (three species, one not found in 2006) were identified. No apseudomorph tanaidaceans were found. All of the material is described, and an attempt is made to investigate the provenance of the Azorean tanaidacean fauna, although the general lack of data from the North-east Atlantic pre- cludes any reasonable interpretation other than some possible links with the Mediterranean. However, recent material of Zeuxo exsargasso collected from Tenerife, in the Canary Islands, does suggest Macaronesian links with the West Atlantic.

RESUMO Durante o 3º Workshop Internacional de Malacologia e Biologia Marinha em São Miguel, Açores, em Julho de 2006, fizeram-se amostragens do bentos litoral e sublitoral de modo a caracterizar a fauna de pequenos artrópodes marinhos dessa região, incluindo os tanaidáceos. Assim, recolheream-se 338 espécimens de tanaidáceos representando três espécies; duas dessas espécies, Leptochelia caldera e Paratanais martinsi, são novas para a ciência. Para além disso, analisou-se material recolhido previamente à volta de São Miguel em 1996 e 1997, do qual foram identificados 272 especimens ( três espécies, uma das quais não encontrada em 2006). Não se encontraram tanaidáceos apseudomorfos. Todo o material foi descrito, e abordou-se a questão da proveniência da fauna tanaidácea Açoreana, embora a ausência generalizada de dados do Nordeste Atlântico impeça qualquer interpretação razoável para além de algumas possíveis ligações com o Mediterrâneo. No entanto, material recente de Zeuxo exsargasso recolhido em Tenerife, Ilhas Canárias, sugere ligações Macaronésicas com o Atlântico Oeste.

INTRODUCTION waters from two directions: the Azores Drift, a diffuse southerly arm of the Gulf he Azores are a group of islands Stream breaking off from the North Tsomewhat isolated in the north-east Atlantic Drift supplies water from the Atlantic, lying adjacent to the Mid- Americas, while the somewhat less-sig- Atlantic Ridge some 1300 km west of nificant western eddies of the Canary Portugal and 1730 km southeast of Current bring waters from Spain and Newfoundland. The main surface water North Africa; below these, the midwater currents reaching the archipelago bring current brings warm, hyperhaline water 184 AÇOREANA 2009, Sup. 6: 183-200 from the Mediterranean outflow (Gofas, Finally, the record of “Paratanais atlanti- 1990; Morton et al., 1998). This hydrogra- cus” of Dollfus (1897) is incertae sedis, but phy clearly has implications for the colo- not attributable to Paratanais (see below). nization of the islands by benthic marine The nine confirmed species recorded species. from around the Azores are: Tanaidaceans are a group of the with minimal dispersive abil- Suborder Apseudomorpha ity; the larvae are not planktonic, and Leviapseudes leptodactylus (Beddard, there are only limited examples of adults 1886), Azores 1830 m. swimming (Bamber, 1998). Some species are known to have spread in fouling com- Suborder Tanaidomorpha munities on ship’s hulls (Bamber, 1977), Superfamily Tanaoidea and others are known to live in floating grimaldii Dollfus, 1897, Azores, algae (Sieg, 1980) or ectoparasitically on littoral, 5-6 m. turtle tests and on manatees (see Morales- Vela et al., 2008). It is therefore of some Superfamily Paratanaoidea interest to determine the suite of species Siphonolabrum mirabile Lang, 1872, which has colonized the Azores archipel- Azores 3500-4165 m. ago, and their provenance. Agathotanais hanseni Lang, 1971, E. Previous records of tanaidaceans from Pacific & Azores, 2861-4165 m the Azores are very sparse, and largely Leptognathia abyssorum (Dollfus, 1897), incidental, although Dollfus (1897) Azores, 1287 m. reported on the tanaidacean material col- Paratyphlotanais richardi (Dollfus, 1897), lected around the Azores during the W Ireland, Azores, 699-1287 m. cruise of the Hirondelle in 1887 and 1888. Typhlotanais spiniventris Dollfus, 1897, All but two of the species recorded in litt. Azores, 130-1287 m. are from deep-water (>100 m; mostly Mesotanais dubius Dollfus, 1897, >500 m). Dollfus (1897) reported Azores, 1287 m. Leptochelia savignyi Krøyer, 1842 from Leptochelia savignyi Krøyer, 1842, Horta; despite the controversy regarding Azores, 5 to 6 m. earlier records of sibling species of this taxon, Dollfus’ records appear to be sub- During the Third International stantiated (see below); L. savignyi was Workshop of Malacology and Marine described originally from Madeira, so its Biology at Vila Franca do Campo, São occurrence in the Azores is not surprising. Miguel, in July 2006, sampling of the lit- Dollfus (1897) also recorded Tanais toral and sublittoral benthos was under- dulongii (Audouin, 1826) (as Tanais taken in order to characterize the smaller cavolinii Milne-Edwards, 1828) from Baïe marine arthropod fauna of this region, de Fayal, and described as new including tanaidaceans. In the event, 338 T. grimaldii from Horta, distinguishing the specimens representing three species two on the shape of the cephalon (his new were collected; two of these species are species having a shorter cephalon) and new to science. In addition, a previous the number of uropod articles; there is collection from around the island was doubt whether his T. dulongii specimens made available, from which a further 272 were in fact of that species (see below); specimens (three species, one additional Morton et al. (1998) mention the occur- to the above three) were identified. No rence of Tanais in littoral algal mats. apseudomorph tanaidaceans were found. BAMBER & COSTA: TANAIDACEANS FROM SÃO MIGUEL 185

In addition, recently collected materi- Quente, Faial da Terra, Nordeste, Porto al from Tenerife, in the Canary Islands, Formoso, Ladeira de Velha, Ribeirinha, kindly supplied to us by Brian Morton, Lactoaçoreana, Cofaco and São Vicente. revealed a new record of a tanaidacean, These sites variously represented exposed relevant to the origins of the and sheltered shores, undisturbed, natu- Macaronesian fauna. All of the material is rally disturbed (near shallow-water vent described below. sites or stream mouths) and polluted shores. MATERIAL AND METHODS Finally, a collection of littoral algal turf was made from a rocky intertidal The present Azores material comes platform at Playa de Fanabe, Costa Adeje, from two sources. During the Workshop Tenerife, Canary Islands, in June 2007, at Vila Franca do Campo in July 2006, a from which a further species not found in number of littoral and infralittoral habi- the São Miguel material was extracted. tats on the island of São Miguel were sam- Terminology used herein recognizes pled for tanaidaceans, including crevice the first pair of antennae as antennules, habitats, macroalgae and soft sediments. the second pair as the antennae. The first The principal sampling areas were the lit- maxilla is termed the maxillule. The first toral sediments, rocks and algae below pair of pereopods (of six) is the pair the Clube Naval building (the old Vila immediately posterior to the chelipeds. Franca do Campo abattoir); the sediments Serially repetitive body-parts, such as the and algae within the caldera of the Ilhéu pereonites and subdivisions of antennal de Vila Franca; and the soft sediments off flagella are segments, others (such as the Vila Franca do Campo (ca N37º 43’ W25º parts of the limb) are articles. Total length 25’), from 12 to 200 m depth. Offshore is measured axially from the tip of the sediments were sampled using a 0.025 m2 rostrum to the posterior edge of the pleo- van Veen grab and various dredges. All telson; measurements were made dorsal- samples were washed through a 0.5 mm ly on the body and antennules, and later- mesh sieve, and specimens sorted alive. ally on the pereopods and antennae. The Some of these specimens were fixed in term ‘spines’ is used in the traditional absolute ethanol to allow DNA analysis. sense to distinguish between rigid ‘thorn- Extensive material collected in 1996 like’ structures and the more flexible and 1997, from 11 (1996) and 20 (1997) ‘hair-like’ setae (in keeping with their ety- infralittoral rocky-substratum sites mology and all historic literature); non- around São Miguel, was also analysed in articulating spine-shaped extensions of detail. Samples of algae (Stypocaulon sco- the cuticle are considered to be apophy- paria, Halopteris filicina and Zonaria tourne- ses; comb–rows of fine setules, occasion- fortii) were collected by SCUBA diving at ally present on maxillae and pereopod depths from between 5 and 16 m. Details articles, inter alia, are referred to as of the sampling and protocols are given microtrichia. by Costa & Ávila (2001). The sampling Voucher and type-material has been sites, anti-clockwise around the Island lodged in the collections of The Natural from the north-west, were Mosteiros, History Museum, London (NHM). The Ponta da Ferraria, Santa Clara, Pesqueiro, higher is based on Guţu & Sieg Emissário, Sinaga, Atalhada, Ribeira da (1999). Praia, Caloura, Porto de Vila Franca do Campo, Ilhéu de Vila Franca, Ribeira 186 AÇOREANA 2009, Sup. 6: 183-200

SYSTEMATICS November 1996; 1 manca, Ribiera Quente, 18 June 1997; 1 manca, Nordeste, 21 SÃO MIGUEL MATERIAL November 1996; 1 male, 1 brooding female, 1 neuter, 2 mancae. Nordeste, 2 Suborder TANAIDOMORPHA Sieg, 1980 May 1997; 1 female, Porto Formoso, 30 Superfamily Tanaoidea Dana, 1849 October 1996; 4 males, 1 female with oost- Family Tanaidae Dana, 1849 egites, 3 neuters, 2 mancae, Porto Subfamily Tanainae Dana, 1849 Formoso, 19 June 1997; 2 males, 3 mancae, Genus Tanais Latreille, 1831 Ladeira da Velha, 19 June 1997; 4 females Tanais grimaldii Dollfus, 1897 (1 brooding, 2 with oostegites), 1 male, Figure 1A, B 4neuters, 1 manca, Ribeirinha, 8 October 1996; 2 females with oostegites, 7 neuters, Sieg, 1980, pp. 84-91; Figure 31, 22. 5 mancae, Ribeirinha, 25 June 1997; 4 females (1 brooding), 8 neuters, 9 mancae, Material: 1 female with oostegites, 1 Lactoaçoreana, 25 June 1997; 1 male, 1 female with empty brood pouch, 2 female, Cofaco, 7 October 1996; 4 females subadult females, 2 neuters, 1 manca (1brooding), 7 males, 6 neuters, 1 manca, (Registration Nos NHM.2007.764-771), Isl. São Vicente, 7 October 1996; 3 females, 3 24.2, drift algae within the flooded crater neuters, 6 mancae, São Vicente, 6 May of the Ilhéu de Vila Franca, 24 July 2006, 1997. All coll. A.C.C. & João Brum. coll. A. Salvador, R. Robbins & R.B.; 12 females with oostegites, 4 brooding Remarks: Tanais grimaldii is one of the only females, 6 males, 148 neuters, 24 mancae two littoral tanaidacean species recorded (NHM.2007.772-781), Isl. 24.4, attached previously from the Azores: the type- low-littoral algae, south wall of the flood- locality is Horta (Faial), at 5 to 6 m depth ed crater of the Ilhéu de Vila Franca, 24 (Dollfus, 1897). T. grimaldii is distin- July 2006, coll. A. Salvador, R. Robbins & guished from its only northeast Atlantic R.B.; 4 females with oostegites, 1 brood- congener, T. dulongii (Audouin, 1826) by ing female, 4 males, 31 subadult females, the conformation of the laciniae mobili of 42 neuters, 28 mancae, Isl. 24.5, mid- the mandibles (see Sieg, 1980, Figure 31), lagoon algae, within the flooded crater of but also adults of the present species have the Ilhéu de Vila Franca, 24 July 2006, coll. one more article in the uropod (basis plus A. Salvador, R. Robbins & R.B.; 2 neuters, three-segmented endopod: Figure 1B) 2 mancae (NHM.2007.782-785), WVF011, than does T. dulongii (basis plus two-seg- northeastern side of Ilhéu de Vila Franca mented endopod). Dollfus (1897) also do Campo, 16 m, scuba dive collection by recorded (Audouin, 1826) Gonçalo Calado, José Pedro Borges, Joana (as Tanais cavolinii Milne-Edwards, 1828) Xavier, Paola Rachello, Patrícia Madeira, from Baie de Fayal, distinguishing it and 20 July 2006. T. grimaldii on the shape of the cephalon 1 brooding female, 1 neuter, 1 manca (his new species having a shorter II, Mosteiros, 17 June 1997; 1 neuter, 1 cephalon) and the number of uropod arti- manca II, Ponta da Ferraria, 17 June 1997; cles; it is unclear whether his specimens 3 females (1 brooding), 2 males, 1 manca, of putative T. dulongii were fully mature, Emissário, 2 November 1996; 1 manca, as only adult T. grimaldii have four seg- Atalhada, 30 May 1997; 1 female, 4 ments to the uropod endopod (T. dulongii neuters, 15 mancae, Caloura, 26 June can be distinguished from immature 1997; 5 neuters, 1 manca, Ribeira Quente, T. grimaldii with three-segmented BAMBER & COSTA: TANAIDACEANS FROM SÃO MIGUEL 187

FIGURE 1. A, B, Tanais grimaldii Dollfus, 1897: A, dorsal; B, uropod; C to E, Leptognathia breviremis (Lilljeborg, 1864): C, dorsal; D, uropod; E, cheliped. (A redrawn after Sars, 1886; B, Azores speci- men; C to E modified from Sars, 1899).

endopods, as the penultimate segment is records from the western Mediterranean twice as long as the ultimate segment in and Casablanca (Sieg, 1980). the former, only slightly longer in the lat- All the material reported above was ter). Dollfus did not examine the collected from algae at less than 17 m mandibular structure, and indeed would depth. Brooding females and mancae have been unaware of its significance. were present in all the months sampled It is likely that all previous records of (May, June, July, October and November). Tanais dulongii from the Azores in fact refer to T. grimaldii. The present species is Superfamily Paratanaoidea Lang, 1949 also recorded from the Italian Family Anarthruridae Lang, 1971 Mediterranean (Gulf of Naples, Ischia Subfamily Leptognathiinae Sieg, 1976 and Linosa); there are unconfirmed Genus Leptognathia Sars, 1882 188 AÇOREANA 2009, Sup. 6: 183-200

cf. Leptognathia breviremis (Lilljeborg, Description of female: body (Figure 2A) 1864) slender, holotype 3.6 mm long, 7.3 times Figure 1C to E as long as wide. Cephalothorax subrec- tangular, 1.6 times as long as wide, as Material: 1 headless specimen, Ponta da long as pereonites 2 and 3 together, with Ferraria, 17 June 1997, coll. A.C.C. & João slight rostrum, eyelobes rounded, eyes Brum. present and black, single setae at posteri- This damaged specimen is almost or of eyelobes and posterolaterally. Six unidentifiable; the cheliped and uropods are free pereonites; pereonite 1 shortest, pere- appropriate to L. breviremis. This species is onites 2 and 3 subequal, 1.3 times as long known from the eastern North Atlantic, but as pereonite 1; pereonites 4 and 5 sube- has been recorded (dubiously) from the qual (pereonite 4 longest) and 1.25 times North Pacific. A figure of L. breviremis (mod- as long as pereonite 2; pereonite 6 just ified from Sars, 1899) is given to aid possible longer than pereonite 1 (all pereonites recognition of this taxon in the Azores in respectively 1.9, 1.4, 1.3, 1.1, 1.0 and 1.4 future (Figure 1, C to E). times as wide as long). Pleon of five free subequal pleonites bearing pleopods; Family Leptocheliidae Lang, 1973 each pleonite about 3.5 times as wide as Genus Leptochelia Dana, 1849 long, with paired lateral setae. Pleotelson Leptochelia caldera sp. nov. (Figure 2M) semicircular, 0.16 times as Figures 2, 3 long as pleon, 2.8 times as wide as long, with paired lateral setae, paired postero- Material: 1 female with oostegites, holo- lateral setae on each side and two distal type (NHM.2007.424), 1 male, allotype setae. (NHM.2007.425), 13 females, paratypes Antennule (Figure 2D) of four taper- (NHM.2007.426-435), Isl. 24.4, attached ing articles, proximal article 3.6 times as low-littoral algae, south wall of the flood- long as wide, 1.5 times as long as distal ed crater of the Ilhéu de Vila Franca, 24 three articles together, with two long July 2006, coll. A. Salvador, R. Robbins & outer and single short dorsal and inner R.B.; 2 females, paratypes setae; second article twice as long as wide, (NHM.2007.436-437), Isl. 24.5, mid- distal outer seta shorter than article; third lagoon algae, within the flooded crater of article 1.3 times as long as second and the Ilhéu de Vila Franca, 24 July 2006, coll. with one aesthetasc; fourth article minute, A. Salvador, R. Robbins & R.B.. eccentric, with five distal setae. 2 females, Pesqueiro, 7 November Antenna (Figure 2G) of six articles, 1997; 3 males, 32 females (3 brooding), 5 proximal article compact, naked; second neuters, Emissário, 2 November 1996; 3 article as long as wide, with single ven- females, 1 male, Emissário, 12 June 1997; 1 trodistal and dorsodistal slender spines; manca, Sinaga, 30 May 1997; 4 males, 40 third article 1.3 times as long as wide, females (2 brooding), 9 neuters, 2 mancae, with dorsodistal slender spine; fourth Atalhada, 25 October 1996; 1 male, 14 article longest, three times as long as females, 7 neuters, Atalhada, 30 May wide; fifth article half as long as fourth; 1997; 1 female, Caloura, 26 June 1997; 1 sixth article minute. neuter, Faial da Terra, 12 June 1997; 1 Labrum (Figure 2H) rounded, setose, female, 1 neuter, Nordeste, 12 May 1997; 1 typical of genus. Left mandible (Figure male, 1 neuter, Ribeirinha, 8 October 1996; 2I) with crenulate lacinia mobilis wider all coll. A.C.C. & João Brum. than pars incisiva, proximal crenulation BAMBER & COSTA: TANAIDACEANS FROM SÃO MIGUEL 189

FIGURE 2. Leptochelia caldera sp. nov., A, holotype female, dorsal; B, allotype male, dorsal; C, allo- type male, lateral; D, female antennule; E, male antennule; F, male antenna; G, female antenna; H, labrum, lateral; I, left mandible; J, maxillule and maxilla; K, maxilliped; L, epignath; M, pleotelson and left uropod. Scale line = 1 mm for A, B and C, 0.2 mm for D to G and M, 0.1 mm for H to L. 190 AÇOREANA 2009, Sup. 6: 183-200 on pars incisiva, pars molaris with strong pact than pereopod 1; basis 3 times as rugosity; right mandible similar but with- long as wide; ischium with 2 setae, longer out lacinia mobilis. Labium typical of seta longer than ischium width; merus genus, distally finely setose, without longer than carpus, merus with strong palp. Maxillule (Figure 2J) with seven ventrodistal spine, carpus with shorter long and two short distal spines and ventrodistal spine; propodus with paired setose margins; palp of two articles with dorsodistal setae and ventrodistal spine; two distal setae; maxilla simple, ovoid. merus, carpus and propodus with ventral Maxilliped (Figure 2K) palp first article microtrichia; dactylus and short unguis naked, second article with one outer and together 0.6 times as long as propodus; three inner setae, and distal seta exceed- dactylus (Figure 3E) with collar of fine ing distal margin of third palp article; setules at half length and dorsodistal third and fourth articles with filtering setule. Pereopod 3 (Figure 3F) similar to rows of six and seven setae respectively, pereopod 2, including longer seta on third article with two further inner distal ischium and dactylus setulation, but car- setae, fourth article with submarginal pus longer than merus and with short outer seta; basis with three long setae outer distal seta. extending to third palp article; endites Pereopod 4 (Figure 3G) basis stout, 2.4 distally with single outer seta and one times as long as wide; ischium with two inner rounded and two robust spatulate short setae; merus shorter than carpus, spines. Epignath (Figure 2L) elongate, merus with two short, ventrodistal arcuate, with setose margin distally and spines, carpus with outer, ventral and proximally. inner curved distal spines; propodus Cheliped (Figure 3A) with rounded, longer than carpus, with two ventrodistal compact basis 1.9 times as long as wide, short spines, four dorsodistal setae, one with inner distal seta; merus subtriangu- as long as dactylus; dactylus and unguis lar with three ventral setae; carpus 1.9 partially fused into a claw, curved. times as long as wide, with three midven- Pereopods 5 (Figure 3H) and 6 as pereo- tral setae; propodus typical for the genus, pod 4, but with pereopod 5 propodus fixed finger with three ventral and three with fewer distal setae. inner setae, cutting edge crenulate, setal Pleopods all alike, typical for the row at base of dactylus of three setae; genus, with single dorsal plumose seta on dactylus with proximal seta. basis. Pereopod 1 (Figure 3C) longer than Uropod (Figure 2M) biramous, basis other pereopods, coxa with seta and naked; exopod of one segment, 0.7 times rounded apophyses; basis slender, 4.1 as long as proximal endopod segment, times as long as wide, with dorsoproxi- outer distal seta longer than inner distal mal seta; ischium compact with one seta; seta; endopod of six segments, distal seg- merus just shorter than carpus; carpus ments slender. with three distal setae, longest of which is 0.4 times length of propodus; propodus Description of male: typical primary male, as long as carpus and merus together, half length of female (allotype length with three dosrodistal setae on slight 1.7 mm), body (Figure 2B, C) more com- mound, one ventrodistal seta; dactylus pact, cephalon just longer than pereonites slender, extending into shorter slender 1 to 3 together, with large eyelobes bear- unguis, the two together as long as propo- ing conspicuous black eyes; pereonite 1 dus. Pereopod 2 (Figure 3D) more com- shortest, pereonites 2 to 6 progressively BAMBER & COSTA: TANAIDACEANS FROM SÃO MIGUEL 191

FIGURE 3. Leptochelia caldera sp. nov., A, female cheliped; B, male cheliped; C, pereopod 1; D, pere- opod 2; E, detail of distal articles of pereopod 2; F to H, pereopods 3, 4 and 5 respectively. Scale line = 0.2 mm for A to D and F to H, 0.1 mm for E. 192 AÇOREANA 2009, Sup. 6: 183-200 longer, pereonite 4 1.7 times as long as has a compact basis to pereopod 1 (only pereonite 1. Five free pleonites, subequal three times as long as wide) and a uro- in length, pleotelson just longer than pod exopod only half as long as the pleonite 5. Sexual dimorphism as fol- proximal endopod segment length. lows. Despite its much more compact proxi- Antennule (Figure 2E) elongate, first mal antennule peduncle article (2.5 times peduncle article arcuate, 4 times as long as long as wide), L. itoi shows most sim- as wide; second article 0.6 times as long as ilarity to L. caldera sp. nov., but the che- first with ventrodistal penicillate setae liped basis of the Japanese species is and 2 midventral simple setae; third arti- more compact (1.5 times as long as cle 0.6 times as long as second, with dor- wide), the distal seta of antennule sodistal seta; flagellum of 7 segments, peduncle article 2 is as long as the article first with 4 proximal and 5 distal aes- (shorter in L. caldera), and the dactylus thetascs, segments 2 to 5 with 4, 4, 3 and 3 plus claw of the first pereopod is much distal aesthetascs respectively; segments 6 longer than the propodus (1.33 times as and 7 more slender than others. Antenna long, compared with subequal in length (Figure 2F) similar to that of female but in L. caldera). more compact. Mouthparts atrophied. The number of maxilliped basis setae Cheliped (Figure 3B) larger than that in L. neapolitana Sars, 1882 is not known, of female; carpus slender, 3.6 times as but that species again differs from long as wide, with ventrodistal invagina- L. caldera in that the proximal antennule tion to accommodate propodus on reflex- peduncle article is shorter (three times as ion; propodus fixed finger shorter than long as wide), the uropod exopod is less palm, with two inner tooth-like apophy- than half the length of the proximal ses on cutting edge; moveable finger with endopod segment, the dactylus plus spinules along cutting edge. claw of the first pereopod is much longer Pleopods more setose than those of than the propodus (1.36 times as long) female. and the cheliped basis more compact (1.35 times as long as wide). Etymology: caldera (from the Spanish The male of the present species is cauldron) is a volcanic crater, the 2006 generally very similar to the male of type-material of this species having been L. dubia sensu Sars, 1886 (non Krøyer, collected from the sea-water-flooded 1842), but that species has longer anteri- crater of Ilhéu de Vila Franca. or pereonites, and is without the degree of ventrodistal invagination on the che- Remarks: there are four recorded species liped carpus shown by L. caldera (and the of Leptochelia with only 3 maxilliped female of Sars’ species has five maxil- basis setae, none of them occurring in the liped basis setae). The male of L. neapoli- North Atlantic or Mediterranean, viz. tana has a similar cheliped to the present L. itoi Ishimaru, 1982 (from Japan), species, but has fewer antennule flagel- L. lusei Bamber & Bird, 1997 (from Hong lum segments and more attenuate pere- Kong), L. nobbi Bamber, 2005 (from onites. Western Australia) and a species from None of the species described previ- Queensland, Australia (Bamber, in ously has the peculiar collar of setules at press). Of these four, only L. nobbi has a half length on the dactylus of pereopods proximal antennule article more than 3 two and three shown by Leptochelia times as long as wide, but that species caldera. BAMBER & COSTA: TANAIDACEANS FROM SÃO MIGUEL 193

Dollfus (1897) recorded a male and a be the male of L. savignyi), from his fig- female of Leptochelia savignyi Krøyer, ure (Krøyer, 1842: pl.2: Figures 13-19) it 1842 from Horta at 5 to 6 m depth. This does not have the same antennular or taxon has been the subject of some con- pleotelson proportions as L. caldera. fusion over the last 150 years (including erroneous synonymy with L. dubia Family Paratanaidae Lang, 1949 Krøyer, 1842), and it is now apparent Sufamily Paratanaidinae Lang, 1949 that numerous species of Leptochelia Genus Paratanais Dana, 1852 await distinction based on morphologi- Paratanais martinsi sp. nov. cal features of both genders which were Figures 4, 5 not examined in any detail before Ishimaru (1985) (see Bamber, 2005 for ?Paratanais euelpis Monod, 1925, non discussion). From his subsequent report Paratanais euelpis Barnard, 1920. on Mediterranean and Atlantic species Non- Paratanais atlanticus Dollfus, (Dollfus, 1898), it is apparent that 1897 (incertae sedis) Dollfus distinguished L. savignyi correct- ly on the basis of four longer articles in Material: 1 female with brood pouch (in the antennule of the female (inter alia). tube), holotype (NHM.2007.438), 1 female There is debate whether the extra anten- dissected, 2 females, 1 manca, 1 headless nular article is an intermediate feature of female, paratypes,(NHM.2007.439-440), a female changing into a male in a genus WVF040, off Amora, Ponta Garça, São known to show progynous hermaphro- Miguel, Azores, 37º42’720”N ditism (e.g. Smith, 1906); while this trend 25º21’554”W, 37.8 m depth, small dredge has been observed in some taxa of sample, 26 July 2006, coll. António de Leptochelia (R. Heard, pers. comm.), the Frias Martins & Jerry Harasewych. male of L. savignyi sensu Sars, 1886 is a primary male, while Larsen & Rayment Description of female: body (Figure 4A) (2002) found this antennular structure a elongate, slender, 4.2 mm long, eight consistent feature of females of their new times as long as wide, colour translucent species L. elongata, including an oviger- white, eyes black. Cephalothorax subrec- ous paratype. Dollfus (1898) reported a tangular, 1.2 times as long as wide, with number of collections including females slight rounded rostrum, single mid-later- attributed to L. savignyi from the al setae; eyes present, pigmented. Six free Mediterranean and the eastern Atlantic, cylindrical pereonites; pereonite 1 short- implying a frequency of this antennular est, with single anterolateral setae; pere- morphology unlikely to be shown only onites 2 and 3 subequal, twice as long as by transitional hermaphrodite speci- pereonite 1, also with single anterolateral mens. His records from the Azores are setae; pereonites 4 and 5 subequal (pere- thus accepted as valid, and L. savignyi onite 4 longest), 1.2 times as long as pere- sensu Sars, 1886 (see Figure 6A, B) is onite 2, pereonite 6 just shorter than pere- accepted as the same as L. savignyi onite 2 (all pereonites respectively 2, 1.1, Krøyer, 1842. 1.0, 0.9, 0.9 and 1.2 times as wide as long). Krøyer (1842) named a third species Pleon of five free subequal pleonites bear- of Leptochelia, “Tanais” edwardsii, also ing pleopods; pleonites 4.5 times as wide from Madeira, but based only on the as long; pleonites 1 to 4 with one male. While it is possible that this taxon plumose, articulating lateral seta on each may never be confirmed (it is assumed to side, pleonite 5 with simple lateral seta. 194 AÇOREANA 2009, Sup. 6: 183-200

Pleotelson semicircular, short, 1.9 times as outer simple setae; single inner spine on wide as long, distally with paired dorsal basis exceeding distal margin of endites. and paired terminal setae. Epignath (Figure 4J) ribbon-like, Antennule (Figure 4B) of four articles, glabrous but with two fine distal setae. proximal article 2,3 times as long as wide, Cheliped (Figure 5A) compact, car- second article 1.3 times as long as wide, pus 1.2 times as long as wide; propodus about one-third length of first, with dor- wider than long, fixed finger short with sal seta longer than article length; third lamellate apophyses on cutting edge, ter- article nearly two-thirds length of second; minal spine indistinct; dactylus with distal article slender, longer than second dorsoproximal simple seta. and third articles together, with five distal Pereopod 1 (figue 5B) longer than setae and single aesthetasc. others, coxa simple with seta; basis slen- Antenna (Figure 4C) of six articles, der, arcuate, five times as long as wide; proximal article compact, naked; second ischium compact with single seta; merus article with long ventrodistal and short 1.5 times as long as carpus; propodus 1.2 dorsodistal apophyses each bearing seta; times as long as merus, with one ventral third article as long as wide, naked, with and three dorsal distal setae; dactylus dorsodistal spine; fourth article just with distinct, slender claw, both together longer than second, with two distal sim- as long as propodus. Pereopod 2 (Figure ple setae; fifth article half as long as 5C) similar to pereopod 1, but more com- fourth with two distal setae; sixth article pact, basis 3.1 times as long as wide, minute with five longer and one shorter ischium with tow setae, merus shorter distal setae. than carpus with ventral microtrichia Labrum (Figure 4D) apically round- and ventrodistal spine, carpus with ven- ed, setose. Left mandible (Figure 4E) tral microtrichia, two ventrodistal and with crenulate pars incisiva and wide, one larger inner distal spines. Pereopod crenulate lacinia mobilis; pars molaris 3 (Figure 5D) similar to pereopod 2. robust with elaborate marginal “teeth”. Pereopod 4 (Figure 5E) basis robust, 2.2 Right mandible (Figure 4F) without times as long as wide; merus 0.8 times as lacinia mobilis, pars molaris less elabo- long as carpus, each with spination as rate. Labium (Figure 4G) simple, finely pereopod 3; propodus longer than car- setose, with fine outer marginal spinule, pus with mid-dorsal penicillate seta, without palp. Maxillule (Figure 4H) dorsodistal slender spine and ventrodis- with seven longer and two shorter distal tal stout spine; dactylus and claw form- spines, palp slender with two long distal ing unguis, curved, two-thirds as long as setae; maxilla ovoid, naked. Maxilliped propodus. Pereopod 5 (Figure 5F) as (Figure 4I) endites characteristic of pereopod 4. Pereopod 6 (Figure 5G) as genus, wide with denticulate outer mar- pereopod 4, but propodus more compact gin, two inner distal ovate tubercles and with two ventrodistal spines and three single inner seta; palp first article naked, dorsodistal setae adjacent to slender second article inner margin with two spine. simple setae and shorter distal spine, Pleopods (Figure 4K) all alike, with outer margin with distal seta; third arti- naked basis, endopod with single inner cle with three inner bidenticulate spines, plumose seta; exopod without setae on adjacent shorter simple spine; fourth inner margin. article with four inner bidenticulate Uropod (Figure 5H) basis naked, spines, single inner submarginal and endopod of two segments, exopod of one BAMBER & COSTA: TANAIDACEANS FROM SÃO MIGUEL 195

FIGURE 4. Paratanais martinsi sp. nov., A, holotype, dorsal; B, antennule; C, antenna; D, labrum; E, left mandible; F, right mandible; G, labium; H, maxillule and maxilla; I, maxilliped; J, epignath: K, pleopod (plumose nature of all setae not shown). Scale line = 1 mm for A, 0.2 mm for B and C, 0.1 for D to J. 196 AÇOREANA 2009, Sup. 6: 183-200 segment, shorter than proximal segment attributed previously to Paratanais are all of endopod. incertae sedis. Bate & Westwood (1868) Male unknown. described “Paratanais” rigidus from one specimen collected from Laminaria hold- Etymology: named after António de Frias fasts off Glasgow, western Scotland: while Martins, in gratitude for his assistance in their four-articled antennule is appropri- attending the workshop, and exemplary ate, they carefully describe the uropod hospitality. rami as both being of one segment, and an elongate, slender chela; their species is Remarks: the genus Paratanais has been not a member of the genus Paratanais, but capably diagnosed by Lang (1973). The with the inadequate description and fig- only species from the North Atlantic ures must remain incertae sedis. P. limicola

FIGURE 5. Paratanais martinsi sp. nov., A, cheliped; B to G, pereopods 1 to 6 respectively; H, uro- pod. Scale line = 0.2 mm for A to G, 0.15 mm for H. BAMBER & COSTA: TANAIDACEANS FROM SÃO MIGUEL 197

Harger, 1878 (see Harger, 1880, for a shorter merus to pereopod 1, and is description and figures) has a three-arti- without the apophyses on the second arti- cled antennule, and the uropod has two cle of the antenna, as well as differences segments in the exopod and five in the in mouthpart setation. endopod: Harger himself (1880) moved this species, apparently correctly, to TENERIFE MATERIAL Leptochelia. Dollfus (1897) described “Paratanais” atlanticus from 130 m depth Superfamily Tanaoidea Dana, 1849 off the Azores, based on a male and two Family Tanaidae Dana, 1849 females; although the description is Subfamily Pancolinae Sieg, 1980 somewhat cursory, and the figures inade- Genus Zeuxo Templeton, 1840 quate, his species clearly had a three-arti- Zeuxo (Parazeuxo) exsargasso Sieg, 1980 cled antennule in the female, and the uro- Figure 6 C, D pod had a two-segmented exopod and a three-segmented endopod; Dollfus’ Zeuxo (Parazeuxo) exsargasso Sieg, species thus cannot be a member of the 1980, 217-221, figure 61. genus Paratanais. Finally, Monod (1925) suspected a specimen from 110 m depth Material: 3 males, 3 females with off Morocco to be P. euelpis Barnard, 1920 oostegites, 1 brooding female (a species adequately refigured by Lang, (NHM.2007.757-763), 1 female with 1973), but without full confidence (and oostegites (dissected), rocky intertidal without description or figure). It is possi- platform covered in algal turf, Playa de ble that his specimen, if indeed of this Fanabe, Costa Adeje, Tenerife, 27 June genus, was of the present species. 2007. Coll. B Morton. In having the setose apophyses on the second article of the antenna, more char- Remarks: Zeuxo exsargasso was only acteristic of species of Leptochelia, known from the type collection from Paratanais martinsi sp. nov. is similar only floating Sargassum natans, 20 miles south- to P. gaspodei Bamber, 2005, with which it east of Bermuda (Sieg, 1980). Its presence also shares the elongate, slender uropod in the Canary Islands implies the possibil- segments, and the short chela. The pre- ity of transport by drift from America via sent species differs from P. gaspodei in a the Gulf Stream and the Azores and number of characters, including being Canary Currents. The only other record- generally more slender (pleonites 3, 4 and ed species of Zeuxo in north-eastern 5 as long as or longer than wide, all wider Atlantic waters is the only-distantly-relat- than long in P. gaspodei), with more slen- ed Zeuxo (Zeuxo) holdichi Bamber, 1990, der articles in the antennule and antenna known from the Atlantic shores of France (proximal peduncle article of antennule and Portugal and the English Channel less than twice as long as wide in (Bamber, 1990; & unpubl. data). Zeuxo P. gaspodei), in the inner spines on the species are distinguished from Tanais maxilliped basis exceeding the distal mar- species in their having five dorsally- gin of the endites (not reaching the mar- demarcated pleonites (Tanais has only gin in P. gaspodei), and with the merus of four), and no dorsal rows of plumose pereopod 1 being 1.5 times as long as the setae on the pleon (Tanais has conspi- carpus (subequal in length in P. gaspodei). cuous rows on pleonites 1 and 2); the uro- With regard to the possible Moroccan pod of Z. exsargasso has five segments record of Monod (1925), P. euelpis also has (Figure 6D). 198 AÇOREANA 2009, Sup. 6: 183-200

DISCUSSION new above are as yet unknown from else- where: Leptochelia is a worldwide genus, There are now five species of shallow- while Paratanais is predominantly south- water tanaidacean recorded from the ern hemisphere in distribution (Australia, Azores, Tanais grimaldii, Leptochelia savi- Subantarctica, South Africa) but also gnyi sensu stricto, Leptochelia caldera, found in the Indo-West Pacific, the Kurile Paratanais martinsi and the unconfirmed Islands and California. Leptognathia from Ponta da Ferraria. T. grimaldii has been recorded from The present data demonstrate that the Mediterranean (Adriatic) by Sars Azorean tanaidacean fauna inhabits lit- (1886, as T. cavolinii), his figure clearly toral to infralittoral algae, is generally showing the appropriate uropod struc- sparse, but may show a relatively high ture. Sieg (1980) attributes the record of degree of endemism. For the only species T. chevreuxi from the Moroccan coast by occurring in the 1996 and 1997 data in suf- Monod (1925) to T. grimaldii, but regards ficient numbers for interpretation, Tanais it as doubtful, as he did the record from grimaldii, no trends were detected in rela- the Bay of Naples by Smith (1906); neither tion to disturbance or exposure. author gives a description or figure, and Unfortunately, these taxa give little the decision of Sieg (loc. cit.) is based on information on the origins of the Azorean their attributing the authority for their fauna. The Leptognathia specimen tells name to Sars, 1886. Both of these records nothing. The two species described as are surely incertae sedis. There are no pub-

FIGURE 6. A and B, Leptochelia savignyi: A, dorsal; B, antennule (redrawn after Sars, 1886). C and D, Zeuxo (Parazeuxo) exsargasso (Tenerife specimen): A, dorsal; B, pleotelson and uropods, dorsal. BAMBER & COSTA: TANAIDACEANS FROM SÃO MIGUEL 199 lished records of Tanais species from the pants at the workshop for samples and Canary Islands or from Madeira. for entertainment. While there has been confusion over the years regarding Leptochelia savignyi LITERARURE CITED (see above), the type locality is Madeira, and the only other valid records (other BAMBER, R.N., 1977. On mobile littoral than those of Dollfus, 1897; 1898) are environments. Porcupine Newsletter, those of Sars (1886) from the 1(4): 62-63. Mediterranean, from the Ligurian Sea BAMBER, R.N., 1990. A new species of and off Sicily. Zeuxo (Crustacea: Tanaidacea) from Thus there are a few indications that the French Atlantic coast. Journal of the Azorean tanaidacean fauna may have Natural History, 24: 1587-1596. links with the Mediterranean, but not BAMBER, R.N., 1998. Zoogeographic elsewhere. Morton & Britton (2000) trends in some Hong Kong arthro- found that most components of the pods. In: MORTON, B. (ed.), The Azorean marine fauna show affiliation Marine Biology of the South China Sea. with the Mediterranean and southern Proceedings of the Third International Europe. However, the discovery of Zeuxo Conference on the Marine Biology of the exsargasso in Tenerife strongly implies col- South China Sea. Hong Kong, 28 October onization from the western Atlantic to - 1 November 1996. pp. 91-112. Hong Macaronesia via the Gulf Stream, the Kong: Hong Kong University Press. Azores Current and the Canary Current BAMBER, R.N., 2005. The Tanaidaceans (see Timmermann, 1932, for a discussion (Arthropoda: Crustacea: Peracarida: on faunistic transport in Sargassum). Tanaidacea) of Esperance, Western It is undoubtedly the case that the Australia, Australia. In: WELLS, F.E., tanaidacean fauna of Macaronesia is very D.J. WALKER & G.A. KENDRICK understudied, and that of the (eds), Proceedings of the Twelfth Inter- Mediterranean and Atlantic coasts of national Marine Biological Workshop: Europe and North Africa is little better The Marine Flora and Fauna of known, still relying heavily on 19th cen- Esperance, Western Australia, pp. 613- tury information. At the same time the 728. Western Australia Museum, speciation of such taxa as Leptochelia Perth. around the North-east Atlantic and BATE, C.S., & J.O. WESTWOOD, 1868. A Mediterranean needs proper study. History of the British Sessile-Eyed Crustacea, 2: 117-154. John Van Voorst, ACKNOWLEDGEMENTS London. COSTA, A.C., & S.P. ÁVILA, 2001. We are indebted to Roni Robbins for Macrobenthic mollusc fauna assistance in the field collecting and sam- inhabiting Halopteris spp. subtidal ple analysis in 2006, to Andreia Salvador fronds in São Miguel, Azores. Scientia for sampling assistance in Ilhéu de Vila Marina, 63: 117-126. Franca, to João Brum for assistance in the DOLLFUS, A., 1897. Note préliminaire diver collections in 1996 and 1997, to sur les Tanaidæ recueillis aux Açores Brian Morton for collecting the Tenerife pendant les campagnes de l’Hirondelle material, to António de Frias Martins for (1887-1888). Bulletin de la Société the organisation of, and inviting us to, the Zoologique, France, 21: 207-215. 2006 Workshop, and to the other partici- DOLLFUS, A., 1898. Campagnes de la 200 AÇOREANA 2009, Sup. 6: 183-200

Melita. Tanaidae récoltes par M. Ed. podes aquatiques de l’Afrique occi- Chevreux dans l’Atlantique et dans la dentale et septentrionale. 1er partie: Méditerranée. Memoires de la Société Tanaidacea, Anthuridae, Valvifera. Zoologique, France, 11: 33-47. Bulletin de la Société des Sciences GOFAS, S., 1990. The littoral Rissoidae Naturelles du Maroc, 5(3): 61-85. and Anabathridae of São Miguel, MORALES-VELA, B., E. SUAREZ- Azores. In: MARTINS, A.M.F. (ed), MORALES, J. PADILLA-SALDIVAR The Marine Fauna and Flora of the & R.W. HEARD, 2008. The tanaid Azores (Proceedings of the First Hexapleomera robusta (Crustacea: International Workshop of Malacology, Peracarida) from the Caribbean mana- São Miguel, 1988). Açoreana, 1990 tee, with comments on other crus- Supplement: 97-134. tacean epibionts. Journal of the Marine GUTU, M., & J. SIEG, 1999. Ordre Biological Association of the United Tanaïdacés (Tanaidacea Hansen, Kingdom, 88: 591-596. 1895). Mémoires de l’Institute MORTON, B., & J.C. BRITTON, 2000. Océanographique, Monaco, 19: 353-389. The origins of the coastal and marine HARGER, O., 1880. Appendix E. The flora and fauna of the Azores. Natural History of Marine , Oceanography and Marine Biology: an XIV. Report on the marine Isopoda of Annual Review, 38: 13-84. New England and adjacent waters. MORTON, B., J.C. BRITTON & A.M.F. Report of the United States Commission MARTINS, 1998. Coastal Ecology of the of Fish and Fisheries, 6: Report of the Açores, 249 pp. Sociedade Afonso Commissioner for 1878, pp. 295-449, Chaves, Ponta Delgada. plates 1-13. SARS, G.O., 1886. Nye bidrag til kund- ISHIMARU, S-i., 1985. A new species of skaben om Middelhavets invertebrat- Leptochelia (Crustacea, Tanaidacea) fauna. III. Middelhavets saxisopoder from Japan, with a redescription of L. (Isopoda chelifera). Archiv for savignyi (Kroyer, 1842). Publications of Mathematik og Naturvidenskab, 11: 263- the Seto Marine Biological Laboratory, 368. 30(4/6): 241-267. SARS, G.O., 1899. An account of the KRØYER, H., 1842. Nye Arter af Slægten Crustacea of Norway with short descrip- Tanais. Naturhistorishe Tiddskrift, 4: tions and figures of all the species. II. 167-188. Isopoda, 265 pp. + pls. 1-18. Bergen LANG, K., 1973. Taxonomische und phy- Museum, Christiania. logenetische Untersuchungen über SIEG, J., 1980. Taxonomische Mono- die Tanaidaceen (Crustacea). 8. Die graphie der Tanaidae Dana 1849 Gattung Leptochelia Dana, Paratanais (Crustacea: Tanaidacea). Abhandlungen Dana, Heterotanais G.O. Sars und der Senckenbergischen Naturforschenden Nototanais Richardson. Dazu einige Gesellschaft, 537: 1-267. Bemerkungen über die Mono- SMITH, G., 1906. High and low dimor- konophora und ein Nachtrag. phism, with an account of certain Zoologica Scripta, 2: 197-229. Tanaidae of the Bay of Naples. LARSEN, K., & H. RAYMENT, 2002. New Mitteilungen aus der Zoologischen species of Leptochelia (Crustacea: Station zu Neapel, 17: 312-340. Tanaidacea) from the Andaman Sea, TIMMERMANN, G., 1932. Biogeo- north-eastern Indian Ocean. Phuket graphische Untersuchungen über die Marine Biological Center Special Lebensgemeinschaft des treibenden Publication, 32(1): 17-31. Golfkrautes. Zeitschrift für Morphologie MONOD, T., 1925. Tanaidacés et iso- und Oekologie des Tieres, 25: 288-355.