MIGUEL, AZORES, WITH DESCRIPTION OF TWO NEW...

THE TANAIDACEANS (ARTHROPODA: PERACARIDA: TANAIDACEA) OF SÃOMIGUEL, 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] CIBIO-Pólo Açores, Department of Biology, University of the Azores, 9501-801 Ponta Delgada, São Miguel,

Azores, Portugal

ABSTRACTDuring 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 arthropod fauna of this region, includingtanaidaceans. In the event, 338 tanaidacean specimens representing three species werecollected; 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 and1997 was analyzed, from which a further 272 specimens (three species, one not found in2006) were identified. No apseudomorph tanaidaceans were found. All of the material isdescribed, and an attempt is made to investigate the provenance of the Azoreantanaidacean fauna, although the general lack of data from the North-east Atlantic pre-cludes any reasonable interpretation other than some possible links with theMediterranean. However, recent material of Zeuxo exsargasso collected from Tenerife, inthe Canary Islands, does suggest Macaronesian links with the West Atlantic.

RESUMODurante 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 demodo a caracterizar a fauna de pequenos artrópodes marinhos dessa região, incluindo ostanaidáceos. Assim, recolheream-se 338 espécimens de tanaidáceos representando trêsespécies; duas dessas espécies, Leptochelia caldera e Paratanais martinsi, são novas para aciência. Para além disso, analisou-se material recolhido previamente à volta de São Miguelem 1996 e 1997, do qual foram identificados 272 especimens ( três espécies, uma das quaisnão encontrada em 2006). Não se encontraram tanaidáceos apseudomorfos. Todo omaterial foi descrito, e abordou-se a questão da proveniência da fauna tanaidáceaAçoreana, embora a ausência generalizada de dados do Nordeste Atlântico impeçaqualquer interpretação razoável para além de algumas possíveis ligações com oMediterrâneo. No entanto, material recente de Zeuxo exsargasso recolhido em Tenerife,Ilhas Canárias, sugere ligações Macaronésicas com o Atlântico Oeste.

INTRODUCTION

The Azores are a group of islandssomewhat isolated in the north-east

Atlantic, lying adjacent to the Mid-Atlantic Ridge some 1300 km west ofPortugal and 1730 km southeast ofNewfoundland. The main surface watercurrents reaching the archipelago bring

waters from two directions: the AzoresDrift, a diffuse southerly arm of the GulfStream breaking off from the NorthAtlantic Drift supplies water from theAmericas, while the somewhat less-sig-nificant western eddies of the CanaryCurrent bring waters from Spain andNorth Africa; below these, the midwatercurrent brings warm, hyperhaline water

AÇOREANA, Suplemento 6, Setembro 2009: 183-200

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from the Mediterranean outflow (Gofas,1990; Morton et al., 1998). This hydrogra-phy clearly has implications for the colo-nization of the islands by benthic marinespecies.

Tanaidaceans are a group of thearthropods with minimal dispersive abil-ity; the larvae are not planktonic, andthere are only limited examples of adultsswimming (Bamber, 1998). Some speciesare known to have spread in fouling com-munities on ship’s hulls (Bamber, 1977),and others are known to live in floatingalgae (Sieg, 1980) or ectoparasitically onturtle tests and on manatees (see Morales-Vela et al., 2008). It is therefore of someinterest to determine the suite of specieswhich has colonized the Azores archipel-ago, and their provenance.

Previous records of tanaidaceans fromthe Azores are very sparse, and largelyincidental, although Dollfus (1897)reported on the tanaidacean material col-lected around the Azores during thecruise of the Hirondelle in 1887 and 1888.All but two of the species recorded in litt.are from deep-water (>100 m; mostly>500 m). Dollfus (1897) reportedLeptochelia savignyi Krøyer, 1842 fromHorta; despite the controversy regardingearlier records of sibling species of thistaxon, Dollfus’ records appear to be sub-stantiated (see below); L. savignyi wasdescribed originally from Madeira, so itsoccurrence in the Azores is not surprising.Dollfus (1897) also recorded Tanaisdulongii (Audouin, 1826) (as Tanaiscavolinii Milne-Edwards, 1828) from Baïede Fayal, and described as newT. grimaldii from Horta, distinguishing thetwo on the shape of the cephalon (his newspecies having a shorter cephalon) andthe number of uropod articles; there isdoubt whether his T. dulongii specimenswere in fact of that species (see below);Morton et al. (1998) mention the occur-rence of Tanais in littoral algal mats.

Finally, the record of “Paratanais atlanti-cus” of Dollfus (1897) is incertae sedis, butnot attributable to Paratanais (see below).

The nine confirmed species recordedfrom around the Azores are:

Suborder ApseudomorphaLeviapseudes leptodactylus (Beddard,

1886), Azores 1830 m.

Suborder TanaidomorphaSuperfamily Tanaoidea

Tanais grimaldii Dollfus, 1897, Azores,littoral, 5-6 m.

Superfamily ParatanaoideaSiphonolabrum mirabile Lang, 1872,

Azores 3500-4165 m.Agathotanais hanseni Lang, 1971, E.

Pacific & Azores, 2861-4165 mLeptognathia abyssorum (Dollfus, 1897),

Azores, 1287 m.Paratyphlotanais richardi (Dollfus, 1897),

W Ireland, Azores, 699-1287 m.Typhlotanais spiniventris Dollfus, 1897,

Azores, 130-1287 m.Mesotanais dubius Dollfus, 1897,

Azores, 1287 m.Leptochelia savignyi Krøyer, 1842,

Azores, 5 to 6 m.

During the Third InternationalWorkshop of Malacology and MarineBiology at Vila Franca do Campo, SãoMiguel, in July 2006, sampling of the lit-toral and sublittoral benthos was under-taken in order to characterize the smallermarine arthropod fauna of this region,including tanaidaceans. In the event, 338specimens representing three specieswere collected; two of these species arenew to science. In addition, a previouscollection from around the island wasmade available, from which a further 272specimens (three species, one additionalto the above three) were identified. Noapseudomorph tanaidaceans were found.

BAMBER & COSTA: TANAIDACEANS FROM SÃO MIGUEL 185

In addition, recently collected materi-al from Tenerife, in the Canary Islands,kindly supplied to us by Brian Morton,revealed a new record of a tanaidacean,relevant to the origins of theMacaronesian fauna. All of the material isdescribed below.

MATERIAL AND METHODS

The present Azores material comesfrom two sources. During the Workshopat Vila Franca do Campo in July 2006, anumber of littoral and infralittoral habi-tats on the island of São Miguel were sam-pled for tanaidaceans, including crevicehabitats, macroalgae and soft sediments.The principal sampling areas were the lit-toral sediments, rocks and algae belowthe Clube Naval building (the old VilaFranca do Campo abattoir); the sedimentsand algae within the caldera of the Ilhéude Vila Franca; and the soft sediments offVila Franca do Campo (ca N37º 43’ W25º25’), from 12 to 200 m depth. Offshoresediments were sampled using a 0.025 m2

van Veen grab and various dredges. Allsamples were washed through a 0.5 mmmesh sieve, and specimens sorted alive.Some of these specimens were fixed inabsolute ethanol to allow DNA analysis.

Extensive material collected in 1996and 1997, from 11 (1996) and 20 (1997)infralittoral rocky-substratum sitesaround São Miguel, was also analysed indetail. Samples of algae (Stypocaulon sco-paria, Halopteris filicina and Zonaria tourne-fortii) were collected by SCUBA diving atdepths from between 5 and 16 m. Detailsof the sampling and protocols are givenby Costa & Ávila (2001). The samplingsites, anti-clockwise around the Islandfrom the north-west, were Mosteiros,Ponta da Ferraria, Santa Clara, Pesqueiro,Emissário, Sinaga, Atalhada, Ribeira daPraia, Caloura, Porto de Vila Franca doCampo, Ilhéu de Vila Franca, Ribeira

Quente, Faial da Terra, Nordeste, PortoFormoso, Ladeira de Velha, Ribeirinha,Lactoaçoreana, Cofaco and São Vicente.These sites variously represented exposedand sheltered shores, undisturbed, natu-rally disturbed (near shallow-water ventsites or stream mouths) and pollutedshores.

Finally, a collection of littoral algalturf was made from a rocky intertidalplatform at Playa de Fanabe, Costa Adeje,Tenerife, Canary Islands, in June 2007,from which a further species not found inthe São Miguel material was extracted.

Terminology used herein recognizesthe first pair of antennae as antennules,the second pair as the antennae. The firstmaxilla is termed the maxillule. The firstpair of pereopods (of six) is the pairimmediately posterior to the chelipeds.Serially repetitive body-parts, such as thepereonites and subdivisions of antennalflagella are segments, others (such as theparts of the limb) are articles. Total lengthis measured axially from the tip of therostrum to the posterior edge of the pleo-telson; measurements were made dorsal-ly on the body and antennules, and later-ally on the pereopods and antennae. Theterm ‘spines’ is used in the traditionalsense to distinguish between rigid ‘thorn-like’ structures and the more flexible‘hair-like’ setae (in keeping with their ety-mology and all historic literature); non-articulating spine-shaped extensions ofthe cuticle are considered to be apophy-ses; comb–rows of fine setules, occasion-ally present on maxillae and pereopodarticles, inter alia, are referred to asmicrotrichia.

Voucher and type-material has beenlodged in the collections of The NaturalHistory Museum, London (NHM). Thehigher taxonomy is based on Guţu & Sieg(1999).

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SYSTEMATICS

SÃO MIGUEL MATERIAL

Suborder TANAIDOMORPHA Sieg, 1980Superfamily Tanaoidea Dana, 1849Family Tanaidae Dana, 1849Subfamily Tanainae Dana, 1849Genus Tanais Latreille, 1831

Tanais grimaldii Dollfus, 1897Figure 1A, B

Sieg, 1980, pp. 84-91; Figure 31, 22.

Material: 1 female with oostegites, 1female with empty brood pouch, 2subadult females, 2 neuters, 1 manca(Registration Nos NHM.2007.764-771), Isl.24.2, drift algae within the flooded craterof the Ilhéu de Vila Franca, 24 July 2006,coll. A. Salvador, R. Robbins & R.B.; 12females with oostegites, 4 broodingfemales, 6 males, 148 neuters, 24 mancae(NHM.2007.772-781), Isl. 24.4, attachedlow-littoral algae, south wall of the flood-ed crater of the Ilhéu de Vila Franca, 24July 2006, coll. A. Salvador, R. Robbins &R.B.; 4 females with oostegites, 1 brood-ing female, 4 males, 31 subadult females,42 neuters, 28 mancae, Isl. 24.5, mid-lagoon algae, within the flooded crater ofthe Ilhéu de Vila Franca, 24 July 2006, coll.A. Salvador, R. Robbins & R.B.; 2 neuters,2 mancae (NHM.2007.782-785), WVF011,northeastern side of Ilhéu de Vila Francado Campo, 16 m, scuba dive collection byGonçalo Calado, José Pedro Borges, JoanaXavier, Paola Rachello, Patrícia Madeira,20 July 2006.

1 brooding female, 1 neuter, 1 mancaII, Mosteiros, 17 June 1997; 1 neuter, 1manca II, Ponta da Ferraria, 17 June 1997;3 females (1 brooding), 2 males, 1 manca,Emissário, 2 November 1996; 1 manca,Atalhada, 30 May 1997; 1 female, 4neuters, 15 mancae, Caloura, 26 June1997; 5 neuters, 1 manca, Ribeira Quente,

November 1996; 1 manca, Ribiera Quente,18 June 1997; 1 manca, Nordeste, 21November 1996; 1 male, 1 broodingfemale, 1 neuter, 2 mancae. Nordeste, 2May 1997; 1 female, Porto Formoso, 30October 1996; 4 males, 1 female with oost-egites, 3 neuters, 2 mancae, PortoFormoso, 19 June 1997; 2 males, 3 mancae,Ladeira da Velha, 19 June 1997; 4 females(1 brooding, 2 with oostegites), 1 male,4neuters, 1 manca, Ribeirinha, 8 October1996; 2 females with oostegites, 7 neuters,5 mancae, Ribeirinha, 25 June 1997; 4females (1 brooding), 8 neuters, 9 mancae,Lactoaçoreana, 25 June 1997; 1 male, 1female, Cofaco, 7 October 1996; 4 females(1brooding), 7 males, 6 neuters, 1 manca,São Vicente, 7 October 1996; 3 females, 3neuters, 6 mancae, São Vicente, 6 May1997. All coll. A.C.C. & João Brum.

Remarks: Tanais grimaldii is one of the onlytwo littoral tanaidacean species recordedpreviously from the Azores: the type-locality is Horta (Faial), at 5 to 6 m depth(Dollfus, 1897). T. grimaldii is distin-guished from its only northeast Atlanticcongener, T. dulongii (Audouin, 1826) bythe conformation of the laciniae mobili ofthe mandibles (see Sieg, 1980, Figure 31),but also adults of the present species haveone more article in the uropod (basis plusthree-segmented endopod: Figure 1B)than does T. dulongii (basis plus two-seg-mented endopod). Dollfus (1897) alsorecorded Tanais dulongii (Audouin, 1826)(as Tanais cavolinii Milne-Edwards, 1828)from Baie de Fayal, distinguishing it andT. grimaldii on the shape of the cephalon(his new species having a shortercephalon) and the number of uropod arti-cles; it is unclear whether his specimensof putative T. dulongii were fully mature,as only adult T. grimaldii have four seg-ments to the uropod endopod (T. dulongiican be distinguished from immatureT. grimaldii with three-segmented


endopods, as the penultimate segment istwice as long as the ultimate segment inthe former, only slightly longer in the lat-ter). Dollfus did not examine themandibular structure, and indeed wouldhave been unaware of its significance.

It is likely that all previous records ofTanais dulongii from the Azores in factrefer to T. grimaldii. The present species isalso recorded from the ItalianMediterranean (Gulf of Naples, Ischiaand Linosa); there are unconfirmed

records from the western Mediterraneanand Casablanca (Sieg, 1980).

All the material reported above wascollected from algae at less than 17 mdepth. Brooding females and mancaewere present in all the months sampled(May, June, July, October and November).

Superfamily Paratanaoidea Lang, 1949Family Anarthruridae Lang, 1971Subfamily Leptognathiinae Sieg, 1976Genus Leptognathia Sars, 1882

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).

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cf. Leptognathia breviremis (Lilljeborg,1864)

Figure 1C to E

Material: 1 headless specimen, Ponta daFerraria, 17 June 1997, coll. A.C.C. & JoãoBrum.

This damaged specimen is almostunidentifiable; the cheliped and uropods areappropriate to L. breviremis. This species isknown from the eastern North Atlantic, buthas been recorded (dubiously) from theNorth Pacific. A figure of L. breviremis (mod-ified from Sars, 1899) is given to aid possiblerecognition of this taxon in the Azores infuture (Figure 1, C to E).

Family Leptocheliidae Lang, 1973Genus Leptochelia Dana, 1849

Leptochelia caldera sp. nov. Figures 2, 3

Material: 1 female with oostegites, holo-type (NHM.2007.424), 1 male, allotype(NHM.2007.425), 13 females, paratypes(NHM.2007.426-435), Isl. 24.4, attachedlow-littoral algae, south wall of the flood-ed crater of the Ilhéu de Vila Franca, 24July 2006, coll. A. Salvador, R. Robbins &R.B.; 2 females, paratypes(NHM.2007.436-437), Isl. 24.5, mid-lagoon algae, within the flooded crater ofthe Ilhéu de Vila Franca, 24 July 2006, coll.A. Salvador, R. Robbins & R.B..

2 females, Pesqueiro, 7 November1997; 3 males, 32 females (3 brooding), 5neuters, Emissário, 2 November 1996; 3females, 1 male, Emissário, 12 June 1997; 1manca, Sinaga, 30 May 1997; 4 males, 40females (2 brooding), 9 neuters, 2 mancae,Atalhada, 25 October 1996; 1 male, 14females, 7 neuters, Atalhada, 30 May1997; 1 female, Caloura, 26 June 1997; 1neuter, Faial da Terra, 12 June 1997; 1female, 1 neuter, Nordeste, 12 May 1997; 1male, 1 neuter, Ribeirinha, 8 October 1996;all coll. A.C.C. & João Brum.

Description of female: body (Figure 2A)slender, holotype 3.6 mm long, 7.3 timesas long as wide. Cephalothorax subrec-tangular, 1.6 times as long as wide, aslong as pereonites 2 and 3 together, withslight rostrum, eyelobes rounded, eyespresent and black, single setae at posteri-or of eyelobes and posterolaterally. Sixfree pereonites; pereonite 1 shortest, pere-onites 2 and 3 subequal, 1.3 times as longas pereonite 1; pereonites 4 and 5 sube-qual (pereonite 4 longest) and 1.25 timesas long as pereonite 2; pereonite 6 justlonger than pereonite 1 (all pereonitesrespectively 1.9, 1.4, 1.3, 1.1, 1.0 and 1.4times as wide as long). Pleon of five freesubequal pleonites bearing pleopods;each pleonite about 3.5 times as wide aslong, with paired lateral setae. Pleotelson(Figure 2M) semicircular, 0.16 times aslong as pleon, 2.8 times as wide as long,with paired lateral setae, paired postero-lateral setae on each side and two distalsetae.

Antennule (Figure 2D) of four taper-ing articles, proximal article 3.6 times aslong as wide, 1.5 times as long as distalthree articles together, with two longouter and single short dorsal and innersetae; second article twice as long as wide,distal outer seta shorter than article; thirdarticle 1.3 times as long as second andwith one aesthetasc; fourth article minute,eccentric, with five distal setae.

Antenna (Figure 2G) of six articles,proximal article compact, naked; secondarticle as long as wide, with single ven-trodistal and dorsodistal slender spines;third article 1.3 times as long as wide,with dorsodistal slender spine; fourtharticle longest, three times as long aswide; fifth article half as long as fourth;sixth article minute.

Labrum (Figure 2H) rounded, setose,typical of genus. Left mandible (Figure2I) with crenulate lacinia mobilis widerthan pars incisiva, proximal crenulation


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, pleotelsonand 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.

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on pars incisiva, pars molaris with strongrugosity; right mandible similar but with-out lacinia mobilis. Labium typical ofgenus, distally finely setose, withoutpalp. Maxillule (Figure 2J) with sevenlong and two short distal spines andsetose margins; palp of two articles withtwo distal setae; maxilla simple, ovoid.Maxilliped (Figure 2K) palp first articlenaked, second article with one outer andthree inner setae, and distal seta exceed-ing distal margin of third palp article;third and fourth articles with filteringrows of six and seven setae respectively,third article with two further inner distalsetae, fourth article with submarginalouter seta; basis with three long setaeextending to third palp article; enditesdistally with single outer seta and oneinner rounded and two robust spatulatespines. Epignath (Figure 2L) elongate,arcuate, with setose margin distally andproximally.

Cheliped (Figure 3A) with rounded,compact basis 1.9 times as long as wide,with inner distal seta; merus subtriangu-lar with three ventral setae; carpus 1.9times as long as wide, with three midven-tral setae; propodus typical for the genus,fixed finger with three ventral and threeinner setae, cutting edge crenulate, setalrow at base of dactylus of three setae;dactylus with proximal seta.

Pereopod 1 (Figure 3C) longer thanother pereopods, coxa with seta androunded apophyses; basis slender, 4.1times as long as wide, with dorsoproxi-mal seta; ischium compact with one seta;merus just shorter than carpus; carpuswith three distal setae, longest of which is0.4 times length of propodus; propodusas long as carpus and merus together,with three dosrodistal setae on slightmound, one ventrodistal seta; dactylusslender, extending into shorter slenderunguis, the two together as long as propo-dus. Pereopod 2 (Figure 3D) more com-

pact than pereopod 1; basis 3 times aslong as wide; ischium with 2 setae, longerseta longer than ischium width; meruslonger than carpus, merus with strongventrodistal spine, carpus with shorterventrodistal spine; propodus with paireddorsodistal setae and ventrodistal spine;merus, carpus and propodus with ventralmicrotrichia; dactylus and short unguistogether 0.6 times as long as propodus;dactylus (Figure 3E) with collar of finesetules at half length and dorsodistalsetule. Pereopod 3 (Figure 3F) similar topereopod 2, including longer seta onischium and dactylus setulation, but car-pus longer than merus and with shortouter distal seta.

Pereopod 4 (Figure 3G) basis stout, 2.4times as long as wide; ischium with twoshort setae; merus shorter than carpus,merus with two short, ventrodistalspines, carpus with outer, ventral andinner curved distal spines; propoduslonger than carpus, with two ventrodistalshort spines, four dorsodistal setae, oneas long as dactylus; dactylus and unguispartially fused into a claw, curved.Pereopods 5 (Figure 3H) and 6 as pereo-pod 4, but with pereopod 5 propoduswith fewer distal setae.

Pleopods all alike, typical for thegenus, with single dorsal plumose seta onbasis.Uropod (Figure 2M) biramous, basisnaked; exopod of one segment, 0.7 timesas long as proximal endopod segment,outer distal seta longer than inner distalseta; endopod of six segments, distal seg-ments slender.

Description of male: typical primary male,half length of female (allotype length1.7 mm), body (Figure 2B, C) more com-pact, cephalon just longer than pereonites1 to 3 together, with large eyelobes bear-ing conspicuous black eyes; pereonite 1shortest, pereonites 2 to 6 progressively


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. Scaleline = 0.2 mm for A to D and F to H, 0.1 mm for E.

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longer, pereonite 4 1.7 times as long aspereonite 1. Five free pleonites, subequalin length, pleotelson just longer thanpleonite 5. Sexual dimorphism as fol-lows.

Antennule (Figure 2E) elongate, firstpeduncle article arcuate, 4 times as longas wide; second article 0.6 times as long asfirst with ventrodistal penicillate setaeand 2 midventral simple setae; third arti-cle 0.6 times as long as second, with dor-sodistal seta; flagellum of 7 segments,first with 4 proximal and 5 distal aes-thetascs, segments 2 to 5 with 4, 4, 3 and 3distal aesthetascs respectively; segments 6and 7 more slender than others. Antenna(Figure 2F) similar to that of female butmore compact. Mouthparts atrophied.

Cheliped (Figure 3B) larger than thatof female; carpus slender, 3.6 times aslong as wide, with ventrodistal invagina-tion to accommodate propodus on reflex-ion; propodus fixed finger shorter thanpalm, with two inner tooth-like apophy-ses on cutting edge; moveable finger withspinules along cutting edge.

Pleopods more setose than those offemale.

Etymology: caldera (from the Spanishcauldron) is a volcanic crater, the 2006type-material of this species having beencollected from the sea-water-floodedcrater of Ilhéu de Vila Franca.

Remarks: there are four recorded speciesof Leptochelia with only 3 maxillipedbasis setae, none of them occurring in theNorth Atlantic or Mediterranean, viz.L. itoi Ishimaru, 1982 (from Japan),L. lusei Bamber & Bird, 1997 (from HongKong), L. nobbi Bamber, 2005 (fromWestern Australia) and a species fromQueensland, Australia (Bamber, inpress). Of these four, only L. nobbi has aproximal antennule article more than 3times as long as wide, but that species

has a compact basis to pereopod 1 (onlythree times as long as wide) and a uro-pod exopod only half as long as theproximal endopod segment length.Despite its much more compact proxi-mal antennule peduncle article (2.5 timesas long as wide), L. itoi shows most sim-ilarity to L. caldera sp. nov., but the che-liped basis of the Japanese species ismore compact (1.5 times as long aswide), the distal seta of antennulepeduncle article 2 is as long as the article(shorter in L. caldera), and the dactylusplus claw of the first pereopod is muchlonger than the propodus (1.33 times aslong, compared with subequal in lengthin L. caldera).

The number of maxilliped basis setaein L. neapolitana Sars, 1882 is not known,but that species again differs fromL. caldera in that the proximal antennulepeduncle article is shorter (three times aslong as wide), the uropod exopod is lessthan half the length of the proximalendopod segment, the dactylus plusclaw of the first pereopod is much longerthan the propodus (1.36 times as long)and the cheliped basis more compact(1.35 times as long as wide).

The male of the present species isgenerally very similar to the male ofL. dubia sensu Sars, 1886 (non Krøyer,1842), but that species has longer anteri-or pereonites, and is without the degreeof ventrodistal invagination on the che-liped carpus shown by L. caldera (and thefemale of Sars’ species has five maxil-liped basis setae). The male of L. neapoli-tana has a similar cheliped to the presentspecies, but has fewer antennule flagel-lum segments and more attenuate pere-onites.

None of the species described previ-ously has the peculiar collar of setules athalf length on the dactylus of pereopodstwo and three shown by Leptocheliacaldera.


Dollfus (1897) recorded a male and afemale of Leptochelia savignyi Krøyer,1842 from Horta at 5 to 6 m depth. Thistaxon has been the subject of some con-fusion over the last 150 years (includingerroneous synonymy with L. dubiaKrøyer, 1842), and it is now apparentthat numerous species of Leptocheliaawait distinction based on morphologi-cal features of both genders which werenot examined in any detail beforeIshimaru (1985) (see Bamber, 2005 fordiscussion). From his subsequent reporton Mediterranean and Atlantic species(Dollfus, 1898), it is apparent thatDollfus distinguished L. savignyi correct-ly on the basis of four longer articles inthe antennule of the female (inter alia).There is debate whether the extra anten-nular article is an intermediate feature ofa female changing into a male in a genusknown to show progynous hermaphro-ditism (e.g. Smith, 1906); while this trendhas been observed in some taxa ofLeptochelia (R. Heard, pers. comm.), themale of L. savignyi sensu Sars, 1886 is aprimary male, while Larsen & Rayment(2002) found this antennular structure aconsistent feature of females of their newspecies L. elongata, including an oviger-ous paratype. Dollfus (1898) reported anumber of collections including femalesattributed to L. savignyi from theMediterranean and the eastern Atlantic,implying a frequency of this antennularmorphology unlikely to be shown onlyby transitional hermaphrodite speci-mens. His records from the Azores arethus accepted as valid, and L. savignyisensu Sars, 1886 (see Figure 6A, B) isaccepted as the same as L. savignyiKrøyer, 1842.

Krøyer (1842) named a third speciesof Leptochelia, “Tanais” edwardsii, alsofrom Madeira, but based only on themale. While it is possible that this taxonmay never be confirmed (it is assumed to

be the male of L. savignyi), from his fig-ure (Krøyer, 1842: pl.2: Figures 13-19) itdoes not have the same antennular orpleotelson proportions as L. caldera.

Family Paratanaidae Lang, 1949Sufamily Paratanaidinae Lang, 1949Genus Paratanais Dana, 1852

Paratanais martinsi sp. nov.Figures 4, 5

?Paratanais euelpis Monod, 1925, nonParatanais euelpis Barnard, 1920.Non- Paratanais atlanticus Dollfus,

1897 (incertae sedis)

Material: 1 female with brood pouch (intube), holotype (NHM.2007.438), 1 femaledissected, 2 females, 1 manca, 1 headlessfemale, paratypes,(NHM.2007.439-440),WVF040, off Amora, Ponta Garça, SãoMiguel, Azores, 37º42’720”N25º21’554”W, 37.8 m depth, small dredgesample, 26 July 2006, coll. António deFrias Martins & Jerry Harasewych.

Description of female: body (Figure 4A)elongate, slender, 4.2 mm long, eighttimes as long as wide, colour translucentwhite, eyes black. Cephalothorax subrec-tangular, 1.2 times as long as wide, withslight rounded rostrum, single mid-later-al setae; eyes present, pigmented. Six freecylindrical pereonites; pereonite 1 short-est, with single anterolateral setae; pere-onites 2 and 3 subequal, twice as long aspereonite 1, also with single anterolateralsetae; pereonites 4 and 5 subequal (pere-onite 4 longest), 1.2 times as long as pere-onite 2, pereonite 6 just shorter than pere-onite 2 (all pereonites respectively 2, 1.1,1.0, 0.9, 0.9 and 1.2 times as wide as long).Pleon of five free subequal pleonites bear-ing pleopods; pleonites 4.5 times as wideas long; pleonites 1 to 4 with oneplumose, articulating lateral seta on eachside, pleonite 5 with simple lateral seta.

194 2009, Sup. 6: 183-200A Ç O R E A N A

Pleotelson semicircular, short, 1.9 times aswide as long, distally with paired dorsaland paired terminal setae.

Antennule (Figure 4B) of four articles,proximal article 2,3 times as long as wide,second article 1.3 times as long as wide,about one-third length of first, with dor-sal seta longer than article length; thirdarticle nearly two-thirds length of second;distal article slender, longer than secondand third articles together, with five distalsetae and single aesthetasc.

Antenna (Figure 4C) of six articles,proximal article compact, naked; secondarticle with long ventrodistal and shortdorsodistal apophyses each bearing seta;third article as long as wide, naked, withdorsodistal spine; fourth article justlonger than second, with two distal sim-ple setae; fifth article half as long asfourth with two distal setae; sixth articleminute with five longer and one shorterdistal setae.

Labrum (Figure 4D) apically round-ed, setose. Left mandible (Figure 4E)with crenulate pars incisiva and wide,crenulate lacinia mobilis; pars molarisrobust with elaborate marginal “teeth”.Right mandible (Figure 4F) withoutlacinia mobilis, pars molaris less elabo-rate. Labium (Figure 4G) simple, finelysetose, with fine outer marginal spinule,without palp. Maxillule (Figure 4H)with seven longer and two shorter distalspines, palp slender with two long distalsetae; maxilla ovoid, naked. Maxilliped(Figure 4I) endites characteristic ofgenus, wide with denticulate outer mar-gin, two inner distal ovate tubercles andsingle inner seta; palp first article naked,second article inner margin with twosimple setae and shorter distal spine,outer margin with distal seta; third arti-cle with three inner bidenticulate spines,adjacent shorter simple spine; fourtharticle with four inner bidenticulatespines, single inner submarginal and

outer simple setae; single inner spine onbasis exceeding distal margin of endites.Epignath (Figure 4J) ribbon-like,glabrous but with two fine distal setae.

Cheliped (Figure 5A) compact, car-pus 1.2 times as long as wide; propoduswider than long, fixed finger short withlamellate apophyses on cutting edge, ter-minal spine indistinct; dactylus withdorsoproximal simple seta.

Pereopod 1 (figue 5B) longer thanothers, coxa simple with seta; basis slen-der, arcuate, five times as long as wide;ischium compact with single seta; merus1.5 times as long as carpus; propodus 1.2times as long as merus, with one ventraland three dorsal distal setae; dactyluswith distinct, slender claw, both togetheras long as propodus. Pereopod 2 (Figure5C) similar to pereopod 1, but more com-pact, basis 3.1 times as long as wide,ischium with tow setae, merus shorterthan carpus with ventral microtrichiaand ventrodistal spine, carpus with ven-tral microtrichia, two ventrodistal andone larger inner distal spines. Pereopod3 (Figure 5D) similar to pereopod 2.Pereopod 4 (Figure 5E) basis robust, 2.2times as long as wide; merus 0.8 times aslong as carpus, each with spination aspereopod 3; propodus longer than car-pus with mid-dorsal penicillate seta,dorsodistal slender spine and ventrodis-tal stout spine; dactylus and claw form-ing unguis, curved, two-thirds as long aspropodus. Pereopod 5 (Figure 5F) aspereopod 4. Pereopod 6 (Figure 5G) aspereopod 4, but propodus more compactwith two ventrodistal spines and threedorsodistal setae adjacent to slenderspine.

Pleopods (Figure 4K) all alike, withnaked basis, endopod with single innerplumose seta; exopod without setae oninner margin.

Uropod (Figure 5H) basis naked,endopod of two segments, exopod of one


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 2009, Sup. 6: 183-200A Ç O R E A N A

segment, shorter than proximal segmentof endopod.

Male unknown.

Etymology: named after António de FriasMartins, in gratitude for his assistance inattending the workshop, and exemplaryhospitality.

Remarks: the genus Paratanais has beencapably diagnosed by Lang (1973). Theonly species from the North Atlantic

attributed previously to Paratanais are allincertae sedis. Bate & Westwood (1868)described “Paratanais” rigidus from onespecimen collected from Laminaria hold-fasts off Glasgow, western Scotland: whiletheir four-articled antennule is appropri-ate, they carefully describe the uropodrami as both being of one segment, and anelongate, slender chela; their species isnot a member of the genus Paratanais, butwith the inadequate description and fig-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.


Harger, 1878 (see Harger, 1880, fordescription and figures) has a three-arti-cled antennule, and the uropod has twosegments in the exopod and five in theendopod: Harger himself (1880) movedthis species, apparently correctly, toLeptochelia. Dollfus (1897) described“Paratanais” atlanticus from 130 m depthoff the Azores, based on a male and twofemales; although the description issomewhat cursory, and the figures inade-quate, his species clearly had a three-arti-cled antennule in the female, and the uro-pod had a two-segmented exopod and athree-segmented endopod; Dollfus’species thus cannot be a member of thegenus Paratanais. Finally, Monod (1925)suspected a specimen from 110 m depthoff Morocco to be P. euelpis Barnard, 1920(a species adequately refigured by Lang,1973), but without full confidence (andwithout description or figure). It is possi-ble that his specimen, if indeed of thisgenus, was of the present species.

In having the setose apophyses on thesecond article of the antenna, more char-acteristic of species of Leptochelia,Paratanais martinsi sp. nov. is similar onlyto P. gaspodei Bamber, 2005, with which italso shares the elongate, slender uropodsegments, and the short chela. The pre-sent species differs from P. gaspodei in anumber of characters, including beinggenerally more slender (pleonites 3, 4 and5 as long as or longer than wide, all widerthan long in P. gaspodei), with more slen-der articles in the antennule and antenna(proximal peduncle article of antennuleless than twice as long as wide inP. gaspodei), in the inner spines on themaxilliped basis exceeding the distal mar-gin of the endites (not reaching the mar-gin in P. gaspodei), and with the merus ofpereopod 1 being 1.5 times as long as thecarpus (subequal in length in P. gaspodei).With regard to the possible Moroccanrecord of Monod (1925), P. euelpis also has

a shorter merus to pereopod 1, and iswithout the apophyses on the second arti-cle of the antenna, as well as differencesin mouthpart setation.

TENERIFE MATERIAL

Superfamily Tanaoidea Dana, 1849Family Tanaidae Dana, 1849Subfamily Pancolinae Sieg, 1980Genus Zeuxo Templeton, 1840

Zeuxo (Parazeuxo) exsargasso Sieg, 1980Figure 6 C, D

Zeuxo (Parazeuxo) exsargasso Sieg,1980, 217-221, figure 61.

Material: 3 males, 3 females withoostegites, 1 brooding female(NHM.2007.757-763), 1 female withoostegites (dissected), rocky intertidalplatform covered in algal turf, Playa deFanabe, Costa Adeje, Tenerife, 27 June2007. Coll. B Morton.

Remarks: Zeuxo exsargasso was onlyknown from the type collection fromfloating Sargassum natans, 20 miles south-east of Bermuda (Sieg, 1980). Its presencein the Canary Islands implies the possibil-ity of transport by drift from America viathe Gulf Stream and the Azores andCanary Currents. The only other record-ed species of Zeuxo in north-easternAtlantic waters is the only-distantly-relat-ed Zeuxo (Zeuxo) holdichi Bamber, 1990,known from the Atlantic shores of Franceand Portugal and the English Channel(Bamber, 1990; & unpubl. data). Zeuxospecies are distinguished from Tanaisspecies in their having five dorsally-demarcated pleonites (Tanais has onlyfour), and no dorsal rows of plumosesetae on the pleon (Tanais has conspi-cuous rows on pleonites 1 and 2); the uro-pod of Z. exsargasso has five segments(Figure 6D).

198 2009, Sup. 6: 183-200A Ç O R E A N A

DISCUSSION

There are now five species of shallow-water tanaidacean recorded from theAzores, Tanais grimaldii, Leptochelia savi-gnyi sensu stricto, Leptochelia caldera,Paratanais martinsi and the unconfirmedLeptognathia from Ponta da Ferraria.

The present data demonstrate thatAzorean tanaidacean fauna inhabits lit-toral to infralittoral algae, is generallysparse, but may show a relatively highdegree of endemism. For the only speciesoccurring in the 1996 and 1997 data in suf-ficient numbers for interpretation, Tanaisgrimaldii, no trends were detected in rela-tion to disturbance or exposure.

Unfortunately, these taxa give littleinformation on the origins of the Azoreanfauna. The Leptognathia specimen tellsnothing. The two species described as

new above are as yet unknown from else-where: Leptochelia is a worldwide genus,while Paratanais is predominantly south-ern hemisphere in distribution (Australia,Subantarctica, South Africa) but alsofound in the Indo-West Pacific, the KurileIslands and California.

T. grimaldii has been recorded fromthe Mediterranean (Adriatic) by Sars(1886, as T. cavolinii), his figure clearlyshowing the appropriate uropod struc-ture. Sieg (1980) attributes the record ofT. chevreuxi from the Moroccan coast byMonod (1925) to T. grimaldii, but regardsit as doubtful, as he did the record fromthe Bay of Naples by Smith (1906); neitherauthor gives a description or figure, andthe decision of Sieg (loc. cit.) is based ontheir attributing the authority for theirname to Sars, 1886. Both of these recordsare surely incertae sedis. There are no pub-

FIGURE 6. A and B, Leptochelia savignyi: A, dorsal; B, antennule (redrawn after Sars, 1886). C andD, Zeuxo (Parazeuxo) exsargasso (Tenerife specimen): A, dorsal; B, pleotelson and uropods, dorsal.


lished records of Tanais species from theCanary Islands or from Madeira.

While there has been confusion overthe years regarding Leptochelia savignyi(see above), the type locality is Madeira,and the only other valid records (otherthan those of Dollfus, 1897; 1898) arethose of Sars (1886) from theMediterranean, from the Ligurian Seaand off Sicily.

Thus there are a few indications thatthe Azorean tanaidacean fauna may havelinks with the Mediterranean, but notelsewhere. Morton & Britton (2000)found that most components of theAzorean marine fauna show affiliationwith the Mediterranean and southernEurope. However, the discovery of Zeuxoexsargasso in Tenerife strongly implies col-onization from the western Atlantic toMacaronesia via the Gulf Stream, theAzores Current and the Canary Current(see Timmermann, 1932, for a discussionon faunistic transport in Sargassum).

It is undoubtedly the case that thetanaidacean fauna of Macaronesia is veryunderstudied, and that of theMediterranean and Atlantic coasts ofEurope and North Africa is little betterknown, still relying heavily on 19th cen-tury information. At the same time thespeciation of such taxa as Leptocheliaaround the North-east Atlantic andMediterranean needs proper study.

ACKNOWLEDGEMENTS

We are indebted to Roni Robbins forassistance in the field collecting and sam-ple analysis in 2006, to Andreia Salvadorfor sampling assistance in Ilhéu de VilaFranca, to João Brum for assistance in thediver collections in 1996 and 1997, toBrian Morton for collecting the Tenerifematerial, to António de Frias Martins forthe organisation of, and inviting us to, the2006 Workshop, and to the other partici-

pants at the workshop for samples andfor entertainment.

LITERARURE CITED

BAMBER, R.N., 1977. On mobile littoralenvironments. Porcupine Newsletter,1(4): 62-63.

BAMBER, R.N., 1990. A new species ofZeuxo (Crustacea: Tanaidacea) fromthe French Atlantic coast. Journal ofNatural History, 24: 1587-1596.

BAMBER, R.N., 1998. Zoogeographictrends in some Hong Kong arthro-pods. In: MORTON, B. (ed.), TheMarine Biology of the South China Sea.Proceedings of the Third InternationalConference on the Marine Biology of theSouth China Sea. Hong Kong, 28 October- 1 November 1996. pp. 91-112. HongKong: Hong Kong University Press.

BAMBER, R.N., 2005. The Tanaidaceans(Arthropoda: Crustacea: Peracarida:Tanaidacea) of Esperance, WesternAustralia, Australia. In: WELLS, F.E.,D.J. WALKER & G.A. KENDRICK(eds), Proceedings of the Twelfth Inter-national Marine Biological Workshop:The Marine Flora and Fauna ofEsperance, Western Australia, pp. 613-728. Western Australia Museum,Perth.

BATE, C.S., & J.O. WESTWOOD, 1868. AHistory of the British Sessile-EyedCrustacea, 2: 117-154. John Van Voorst,London.

COSTA, A.C., & S.P. ÁVILA, 2001.Macrobenthic mollusc faunainhabiting Halopteris spp. subtidalfronds in São Miguel, Azores. ScientiaMarina, 63: 117-126.

DOLLFUS, A., 1897. Note préliminairesur les Tanaidæ recueillis aux Açorespendant les campagnes de l’Hirondelle(1887-1888). Bulletin de la SociétéZoologique, France, 21: 207-215.

DOLLFUS, A., 1898. Campagnes de la

200 2009, Sup. 6: 183-200A Ç O R E A N A

Melita. Tanaidae récoltes par M. Ed.Chevreux dans l’Atlantique et dans laMéditerranée. Memoires de la SociétéZoologique, France, 11: 33-47.

GOFAS, S., 1990. The littoral Rissoidaeand Anabathridae of São Miguel,Azores. In: MARTINS, A.M.F. (ed),The Marine Fauna and Flora of theAzores (Proceedings of the FirstInternational Workshop of Malacology,São Miguel, 1988). Açoreana, 1990Supplement: 97-134.

GUTU, M., & J. SIEG, 1999. OrdreTanaïdacés (Tanaidacea Hansen,1895). Mémoires de l’InstituteOcéanographique, Monaco, 19: 353-389.

HARGER, O., 1880. Appendix E. TheNatural History of Marine Animals,XIV. Report on the marine Isopoda ofNew England and adjacent waters.Report of the United States Commissionof Fish and Fisheries, 6: Report of theCommissioner for 1878, pp. 295-449,plates 1-13.

ISHIMARU, S-i., 1985. A new species ofLeptochelia (Crustacea, Tanaidacea)from Japan, with a redescription of L.savignyi (Kroyer, 1842). Publications ofthe Seto Marine Biological Laboratory,30(4/6): 241-267.

KRØYER, H., 1842. Nye Arter af SlægtenTanais. Naturhistorishe Tiddskrift, 4:167-188.

LANG, K., 1973. Taxonomische und phy-logenetische Untersuchungen überdie Tanaidaceen (Crustacea). 8. DieGattung Leptochelia Dana, ParatanaisDana, Heterotanais G.O. Sars undNototanais Richardson. Dazu einigeBemerkungen über die Mono-konophora und ein Nachtrag.Zoologica Scripta, 2: 197-229.

LARSEN, K., & H. RAYMENT, 2002. Newspecies of Leptochelia (Crustacea:Tanaidacea) from the Andaman Sea,north-eastern Indian Ocean. PhuketMarine Biological Center SpecialPublication, 32(1): 17-31.

MONOD, T., 1925. Tanaidacés et iso-

podes aquatiques de l’Afrique occi-dentale et septentrionale. 1er partie:Tanaidacea, Anthuridae, Valvifera.Bulletin de la Société des SciencesNaturelles du Maroc, 5(3): 61-85.

MORALES-VELA, B., E. SUAREZ-MORALES, J. PADILLA-SALDIVAR& R.W. HEARD, 2008. The tanaidHexapleomera robusta (Crustacea:Peracarida) from the Caribbean mana-tee, with comments on other crus-tacean epibionts. Journal of the MarineBiological Association of the UnitedKingdom, 88: 591-596.

MORTON, B., & J.C. BRITTON, 2000.The origins of the coastal and marineflora and fauna of the Azores.Oceanography and Marine Biology: anAnnual Review, 38: 13-84.

MORTON, B., J.C. BRITTON & A.M.F.MARTINS, 1998. Coastal Ecology of theAçores, 249 pp. Sociedade AfonsoChaves, Ponta Delgada.

SARS, G.O., 1886. Nye bidrag til kund-skaben om Middelhavets invertebrat-fauna. III. Middelhavets saxisopoder(Isopoda chelifera). Archiv forMathematik og Naturvidenskab, 11: 263-368.

SARS, G.O., 1899. An account of theCrustacea of Norway with short descrip-tions and figures of all the species. II.Isopoda, 265 pp. + pls. 1-18. BergenMuseum, Christiania.

SIEG, J., 1980. Taxonomische Mono-graphie der Tanaidae Dana 1849(Crustacea: Tanaidacea). Abhandlungender Senckenbergischen NaturforschendenGesellschaft, 537: 1-267.

SMITH, G., 1906. High and low dimor-phism, with an account of certainTanaidae of the Bay of Naples.Mitteilungen aus der ZoologischenStation zu Neapel, 17: 312-340.

TIMMERMANN, G., 1932. Biogeo-graphische Untersuchungen über dieLebensgemeinschaft des treibendenGolfkrautes. Zeitschrift für Morphologieund Oekologie des Tieres, 25: 288-355.

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