Description of Gnathophylleptum Tellei Gen. Nov., Sp. Nov., a Remarkable New Gnathophyllid Shrimp from Canary Islands (Crustacea

'' BULLETIN DE L'INSTITUT ROYAL DES SCIENCES NATURELLES DE BELGIQUE, BIOLOGIE, 71: 113-125, 2001 BULLETIN VAN HET KONI NKLIJK BELGISCH INSTITUUT VOOR NATUURWETENS CH APPE N, BIOLOGIE, 71: 113-1 25,200 1

Description of Gnathophylleptum tellei gen. nov., sp. nov., a remarkable new gnathophyllid shrimp from Canary Islands (Crustacea, Decapoda, Ca~idea) by Cedric d'UDEKEM d' ACOZ

Abstract is here erected for this remarkable new species, which has been found by Mr. Arthur TELLE (Gran Canaria). It may be A new genus Cnathophy/leptum gen. nov. is erected for a hi ghly di stinguished by'many characters, the two most conspicuous characteristic new species of gnathophyllid shrimp from the Canary being the considerable length and slenderness of the second Islands : Cnathophy/leptum tellei gen. nov., sp. no v. This species , pereiopod, and the complex tridimensional structure of the which has been found in the coastal waters of Gran Can aria, is unu third maxilliped. sually slender for a member of the Gn athophyllidae and possesses quite di stincti ve third maxillipeds. The relationship between Cnathophylleptum and related genera is discu ssed in detail. An identification key to all gnathophyllid genera is given. Systematics Key-words: Cnathophylleptwn, Gn athophyllidae, Palaemonoide a, Caridea, Decapoda, shrimp , taxonomy, phylogeny, corneal papilla, Gnathophylleptum gen. nov. Canary Island s, Easte rn Atlantic TYPE SPECIES Gnathophylleptum tellei gen. nov. , sp. nov. Resume ETYMOLOGY Un nou veau genre Cnathophy/leptum gen. no v. est propose pour The name derives from Gnathophyllum, the type genus of the une crevette Gn athophyllidae hautement caracteri stique des lies family Gn athophyllidae, and from A£1!1:0£, slender. The Canaries : Cllathophy/leptum tellei gen. nov., sp. no v. Cette es pece genus is neutral. qui a ete reco ltee dans les eaux c6tieres de Gran Canaria prese nte une silhouette singuli erement grac il e pour une Gnathophyllidae, et DESCRIPTION Ia morphologie de ses maxillipedes de Ia troisieme paire est tout a fait parti culiere. Les relations entre C!lathophylleptwn gen. nov. et Rostrum moderately long, armed both dorsally and ventrally. les genres ap parentes so nt discutees d'une mani ere ap profondie. Antenna! spine present and well developed. Cornea with well Une cle d'identi fica ti on est donnee pQur tous les ge nres de developed distal papilla, withou.t ocellum. First segment of Gnathophyllidae. antennular segment long and narrow, with short styliform Mots-des: Cnathophy/leptullz , Gnath op hyllidae, Palae monoidea, stylocerite. Basicerite with well developed lateral tooth. Caridea, Decapoda, crevette, taxo nomie, phylogenie, papille cor Scaphocerite narrow. Mandible very small , devoid of palp neenn e, lies Canaries, Atlantique oriental and incisor process. Upper lacinia of maxillula much larger than palp and lower lacinia. Maxi lla fairly small, with well developed palp , with basal endite entire and reduced to a Introduction short blunt lobe. First maxilliped very large, with well devel oped epipod and exopod, well developed palp; basal endite Our current knowledge of the decapod fauna of Europe and tri angul ar, narrowin g gradu all y towards its tip which is angu neighbouring areas is usuall y considered fairly good but is by lar; coxal endite protruding and completely fused to basal no mean complete. Each year, one or two new species are endite. Second max illiped unmodified, with well developed described from the area (D' UDEKEM D' Acoz, 1999), but in epipod. Third max illiped with well developed exopod; ulti mos t cases, these new species are closely related to already mate and penultimate segment somewhat fl attened; described taxa. The di scovery of new genu s remains excep ischiomerus strongly flattened, with outer face concave; ti onal , especiall y in the coastal waters. So, it was a real sur max imal width of ischiomerus similar to that of basis, these pri se to discover a new shall ow-water Canari an two segments being well separated. Carpus of first pereiopod gnathophyllid shrimp with a spectac ular colour pattern and very long and much longer than propodus; dactylus much not closely related to any member of thi s famil y. A new ge nu s shorter than palm. Second pereiopod slender and extremely ''

114 CEDRIC D' UDEKEM D' ACOZ

long; carpus longer th an merus and about as long as with fu sed rami consisting of 6 broad segments; long ramus propodus; dactylus considerably shorter than palm; chela with 6 narrow segments; short ramus with I segment. toothless. Dactylus of third, fo urth and fifth pereiopod robust Scaphocerite narrow, 3.5 times as long as wide, with lateral and bifid. margin sli ghtly concave; lateral tooth nearl y reaching tip of bl ade; tooth and bl ade separated by wide space; tip of blade regul arl y rounded, and barely oven eaching tip of antennular Gnathophylleptum tellei gen. nov. , sp. nov. peduncle. Antenn a! peduncle short and moderately slender, (Figs. 1-7) reaching 0.27 of scaphocerite. Basicerite with well devel MATERIAL oped sharp tooth. Canary Islands, Gran Canari a, Sardina del Norte, Caulerpa M andible very small , devoid of palp and incisor process; racenzosa (FORSSKA L) J. AG ARD H meadow, under a hand molar process armed with long sharp teeth. sized stone, 15m depth, SCUBA diving, Arthur TELLE coli. , Upper lacinia of maxillula much larger than palp and lower 18/I/200 1: I female holotype (collection Institut royal des lacinia, with double row of stout spines; palp and lower Sciences Naturelles de Belgique, Brussels, N° I.G. 29411 ). lacinia normall y developed; lower angle of palp with a ETYMOLOGY straight spine pointing laterall y. It is a pleasure to dedicate the species to Mr. Arthur TELLE M axill a: fairl y small , with well developed palp, with basal who discovered the species. The name is a genitive. endi te entire and reduced to a short blunt lobe without setae.

DESCRIPTION First maxilliped very large, with large epipod consisting of two well separated rounded lobes (upper one largest), long Rostrum straight, slender, moderately long, overreaching exopod, well developed palp nearl y as long as basal endite mesial margin of first segment of antennular peduncle but and longer th an caridean lobe; basal endite large, triangul ar, not reaching tip of distolateral tooth , 0.8 times as long as narrowing graduall y towards its tip whi ch is angular, with carapace; dorsal margin gently sloping downwards, with 7 very long slender and non-setulose marginal setae; coxal dorsal teeth spread evenl y over its length, posterionnost situ endite well developed, protruding, bluntly subquadrate and ated on carapace behind level of orbital margin ; ventral mar completely fused to basal endite, with only one seta in upper gin hori zontal, with 2 ventral teeth on distal 0.3; tip entire position; caridean lobe with tip broadly rounded, not protrud and styli fo rm. Posterior 0.9 of carapace unarmed, fa intly ing, much shorter than basal endite. convex and nearl y hori zontal. Antenna! spine situated above inferi or orbi tal angle, sharp and fa irl y large but not reaching Second maxilliped unmodified, with fairl y small unilobed tip of inferior orbital lobe. Pterygostomian area bluntly trian epipod and with very long exopod. gular, overreaching tip of antenna! spine. Third pleonite moderately convex in lateral view. Pleura of first three Third maxill iped without arthrobranch, without epipod, with pl eonites rounded ; pl eura of fo urth and fifth pleonites with long exopod overreaching ischiomerus, with well developed eral plate; ultimate and penultimate segment subequal, postero-ventral tooth , that of fifth pleonite bein g quite long. lat Posterolateral angle of sixth pl eonite with a sharp tooth. Dor much shorter than ischiomerus, somewhat fl attened but sal length of sixth pleonite 1.6 x as long as fifth. Ratio dorsal much longer than broad; penultimate segment with an outer length I height of sixth pleonite = 1. 6. Telson robust, armed subquadrate distal projecti on; outer angle of this proj ection with 2 pai rs of strong dorsolateral spines at 0.4 and 0.7 of its with a tin y but sharp tooth ; inner angle with a large blunt length. Tip of tel son triangul ar with 2 pairs of long spines, the tooth . Ischi omeru s fl attened and curved both longitudinall y inner spines bein g twice as long as outer spines; with a pair of (upwards) and laterall y, outer face transversall y concave and setulose, robust short setae between the inner teeth; with 2 inner face transversally convex, well developed distal outer pairs of short thin non- setulose setae, both situated between tooth, low tri angul ar median projecti on on tip of inner face; the inner spines and the setulose setae. outer border broadened (rounded in cross secti on); mesial border thin with a row of spines. Ischiomeru s and basis quite Eye large, oveiTeaching styloceri te; cornea considerably distinct (not fused), with m aximal width similar, but proxi shorter th an stalk, with dorsal surface sli ghtly convex, with mal part of ischiomerus narrower than tip of basis. ventral surface strong ly convex, and bearing a well devel oped distal papilla at the confluence of the dorsal and ventral First pereiopod with sparse setae; cutting edges of chela surfaces; no ocellum. toothless; carpus 1. 2 x as long as merus, 1.7 x as long as propodus; propodus 1.6 x as long as ischium; dactylus 0.33 x First segment of antennular peduncle very narrow, 3 times as as long as propodus; carpus 6.2 x as long as wide. long as wide, with a long and sharp distolateral tooth sli ghtl y overreaching second segment, with well developed Second left pereiopod extremely long and slender, with ventromesial tooth reaching tip of styloceri te, with short but sparse setae; cutting edges of chela toothless; merus over sharp stylocerite reaching 0.4 of fi rst segment (distal tooth reaching short ramus of upper antennular fl agellum; merus included). Second and third segments of antennular peduncle 1.1 x as long as ischium; carpus 1.3 x as long as merus, 0.9 x subequal, sli ghtly longer than broad, 0.2 x as long as first as long as propodus ; dactylu s 0.27 x as long as propodus; segment (d istal tooth included). Upper ante nnular fl agellum carpus 7.0 x as long as wide. Second ri ght pereiopod lac king. I I

Description of Gnathophylleptum tellei gen. nov., sp. nov. 115

Fig. I. Gnmhophyl/eplum Lellei gen. nov., sp. no v. Shrimp in lateral view. Scale bar 3.0 mm. I I

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Fig. 2. Gnathophy/leplum lellei gen. nov., sp. nov. Anterior part of shrimp in dorsal view. Scale bar 3.0 mm

Last three pereiopods morphologically similar, although P3 with an anterior V-shaped marking pointing posteriorly; each propodus < P4 propodus < P5 propodus, long, fairly robust branch of the "V" made by a row of white dots on a blood red (P3 merus = 5.5 x as long as wide), with sparse setae; background ; branches of "V" arising from anterior lower propodus > merus > ischium >carpus > dactylus; propodus part of carapace and dorsally converging at anterior 0.25 of with 4 widely spaced small spines on distal 0.6 of fl exor bor carapace midline; a few white dots within the space com der; dactylus short and robust, with terminal unguis and pri sed between the "V" and the anterior part of the carapace. sharp triangular accessory tooth on fl exor border; anterior Posterior 0.2 of carapace with a dorsal transverse stripe made border of accessory tooth perpendicular to fl exor border of up of white dots on a whitish pink background. Third tem1inal unguis. pleonite with a large dorsal semi-circular spot, anteri orly rounded and posteriorly straight, made up of white dots on a whitish pink background. Space between the transverse COLOUR PAITERN (ON THE BASIS OF COLOUR SLIDES) stripe and the semi-circular spot, as well as dorsal part of Most parts of body blood red with fai ntly contrasted dots of a fourth and fifth pleoni tes made of a paler red than the rest of darker red. Rostrum transparent, tinged with white. Carapace the body. Sixth pleonite and tail fa n transparent with a few I I

Description of Gnathophylleptwn tellei gen. nov. , sp. no v. 117

Fig. 3. Gnathophylleptum re /lei gen. nov., sp. nov. A, left eyestalk in late ral view ; B, first left antenna; C, second left antenna ; D, telson. Scale bar: A, B, C 1.0 mm ; D 0.68 mm .

discrete milky-white dots. Eyestalks transparent with white parent with a few white dots; transition between the red and dots and a few faint red marks; co rn ea grey. Antennulae and the transparent part ab rupt. Pleopods transparent, with a few antenn ae transparent with a few white dots. Proximal 0.4 of milky-whi te dots. third maxilliped red ; distal 0.6 transparent with white dots. Proximal part of pereiopods up to tip of ischium red; distal Size.- Carapace length= 2.8 mm; total length= 12 mm. part of pereiopods from basal part of merus onw ards trans- I I

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Ecology palp and . the lower lacinia, but these size differences are much more pronounced in Gnathophyllwn. In The unique specimen of Gnathophylleptum tellei gen. nov., Gnathophylleptwn gen. nov., the lower distal spine of the sp. nov. studied has been found in a Caule1pa racemosa palp is directed outwards, while in Gnathophylhun it is (FORSSKAL) J. AGARDH meadow, under a stone, at 15 m curved inwards. depth. According to Mr. Arthur TELLE who collected it dur The basal endite of the maxilla of Gnathophylleptum. gen. ing SCUBA diving, there was no evidence of any association nov. forms a reduced but distinct blunt protrusion, while in between the shrimp and any other organism. However, its Gnathophyllum, it is absent. strongly modified mouthparts suggest a specialized feeding habit. In Gnathophylleptum gen. nov., the exopod and the endopod of the first maxilliped are close to each other, while they are separated by a large space in Gnathophyllum. In Morphological affinities of Gnathophylleptum gen. nov. Gnathophylleptum gen. nov., the tip of the caridean lobe is not protruding ; in Gnathophyllum it is always long. In The Gnathophyllidae are a small family of palaemonoid Gnathophylleptum gen. nov., the palp considerably over shrimps, which have a highly speciali zed morphology, espe reaches the caridean lobe and nearly reaches the tip of the cially their mouthparts, and which are usuall y bri ghtl y col basal endite, while in Gnathophyllum it is shorter than the oured. Gnathophylleptum gen. nov. shows some affinities, caridean lobe· and is much shorter than the basal endite. In especially regarding the morphology of the second Gnathophylleptum gen. nov., the basal endite is triangular, maxilliped, with one member of the very heterogenous genus gradually narrowing towards tip which is angular ; in Gnathophylloides SCHMITT, 1933: G. mineri SCHMITT, 1933. Gnathophyllwn, it is elliptic with a rounded tip. In However, the latter species can be distinguished at first Gnathophylleptwn gen. nov., the coxal endite is large, while glance from Gnathophylleptum gen. nov. by its very short it is quite small in Gnathophyllum. and robust walking legs, whose dactylus are devoid of acces sory tooth. The general appearance of Gnathophylleptum In Gnathophylleptwn gen. nov., the second maxilliped shows gen. nov. is similar to that of the very homogenous genus little specialization in contrast with Gnathophyllum. In Gnathophyllum LATREILLE, 1819, but a closer examination Gnathophylleptum gen. nov., half of the exopod overreaches reveals profound differences, and both genera are obviously the endopod (tenninal setae not considered), wh ile in not closely related. A detailed comparison between Gnathophyllum, the exopod and the endopod are subequal ; Gnathophylleptum gen. nov. and Gnathophyllum is given the proximal part of the endopod is considerably broadened here below, as well as a key to all gnathophyllid genera. The in Gnathophyllwn, but not so in Gnathophylleptwn gen. nov. data on the morphology of Gnathophyllum are based on a ; in G11.athophylleptum gen. nov., the ultimate segment is Sicilian Gnathophyllum elegans (RISSO, 1816) which has smaller than the antepenul timate and both segments have a been dissected and on the detai led illustrated accounts of normal morphology ; in Gnathophyllum, the ultimate seg HOLTHUIS (1949), MANN ING (1963), CHACE & FULLER ment is very large and very elongate in a downwards direc ( 197 1), TITGEN ( 1989) and MANNING & CHACE (1990). tion, and is much larger than the antepenultimate segment which is reduced. Specificall y, the fo llowing differences are apparent between Gnathophylleptum gen. nov. and Gn.athophyllum. In Gnathophylleptum gen. nov., the third maxilliped has 2 distal teeth on the outer border of its penultimate segment; In Gnathophylleptwn gen. nov., the rostrum is slender, espe these teeth are lacking in Gnathophyllum. In ciall y in its distal part, while in Gnathophyllwn it is fai rly Gnathophylleptwn gen. no v., the ischiomerus is quite distinct robust. from the basis, while these segments are fused and scarcely In Gnathophylleptum gen. nov., the pleura of the fourth and distinct in Gnathophyllum. In Gnathophylleptum gen. nov., the fifth pleonites have a posterior tooth, that of the fifth be the outer face of the ischiomerus is strongly concave in the ing very sharp. In Gnathophyllum, the fourth pleonite is transverse plane, while it is slightly convex in regularly rounded and the fifth is rounded or bluntly angular. Gnathophyllum. In Gnathophylleptwn gen. nov., the outer border of the ischiomerus is thickened, whilst it is thin in In Gnathophylleptum gen. nov., the eyestalks are more slen Gnathophyllum. In Gnathophylleptum gen. nov., the outer der, with a more conical cornea than in Gnathophyllwn. In border has a strong distal tooth which is absent in Gnathophylleptum gen. nov., the first segment of the GJrathophyllunr . antennular peduncle is very slender with a short stylocerite, wh ile it is very broad with a very long styloceri te in The second left pereiopod in Gnathophylleptwn gen. nov. is Gnathophyllum. The scaphocerite in Gnathophylleptum gen. extremely long and slender, with a carpus nearly as long as nov. is slender, while it is very broad in Gnathophyllum. the propodus, and without teeth on the cutting edges of its chela (the second ri ght pereiopod is Jacking in the unique No incisor process is present on the mandible of specimen examined); in Gnathophyllum, the second left Gnathophylleptum gen. nov., whilst in Gnathophyllum, it is pereiopod is much shorter, not especiall y slender, has its car either absent or vesti gial, depending on the species. pus much shorter than propodus, and the cutting edges of its Both Gnathophylleptum gen. nov. and Gnathophyllum. have chelae are toothed. the upper lacinia of the maxillula considerably larger than the II

Description of Gnathophylleptum tellei gen. nov., sp. nov. 119

Fig. 4. Gnathophy/!eptum tel lei ge n. nov. , sp. nov. A, ri ght mandible; tip of ri ght mandible; C, first right maxilla; 0 , palp of first ri ght maxilla; E, second ri ght maxilla; F, first left max illiped ; G, second left max illi ped. Scale bar: A, C, E, F, G 0.50 mm ; B 0.083 m111 ; 0 0.17111111. II

120 CEDRIC D' UDEKEM D' ACOZ

Fig. 5. Gnathophylleptum tell ei gen. nov., sp. nov., third left maxilliped. A, lateral outer view; B, lateral inner view; C, facial inner view. Scale bar 1.0 mm. II

Description of Gnathophylleptum tellei gen. nov., sp. nov. 121

Fig. 6. Gnathophyl/eptwll rellei gen. nov. , sp. nov., A, first left pereiopod; B, second left pereiopod; C, dactylus of third left pereiopod. Scale bar: A, B 1.0 mm; C 0.17 mm. II

122 CEDRIC D' UDEKEM D' ACOZ

Fi g. 7. Gnathophy/leptumtellei gen. no v., sp. nov., A, third left pereiopod; 8 , fourth left pereiopod; C, fifth le ft pereiopod. Scale bar 1.0 mm . '' Description of Cnathophylleptwn tellei gen. nov. , sp. nov. 123

Key to the genera of Gnathophyllidae

I. • Dactylus of the last three pairs of pereiopods with a terminal unguis and a well developed accessory tooth on the distal part of the flexor border ...... 2 • Dactylus of the last three pairs of pereiopods with a terminal unguis and a blunt granulated lobe instead of an accessory tooth on the fl exor border ...... 4 2. • Rostrum with dorsal teeth. Body not depressed. Cornea broader than unpigmented part of eyestalk, with papilla (some times weakly developed). Last three periopods of n01mal robustness or slender; accessory tooth of their dactylus shorter than terminal unguis ...... 3 • Rostrum without dorsal teeth. Body depressed. Cornea less broad than unpigmented part of eyestalk, without papilla. Last three pereiopods very short and robust; accessory tooth of their dactylus longer than terminal unguis ... genus Pycnocaris BRUCE, 1972 3. • Basal antennular segment and scaphocerite very broad. Basal endi te of first maxilliped elliptic. Second maxilliped with basal part very broadened and with distal segment strongly elongated downwards. Ischiomerus of third maxilliped with outer face slightly convex. Second pereiopod neither especiall y long nor slender, with carpus much shorter than propodus, with fingers toothed ...... genus Cnathophyllum LATREI LLE, 1819 • Basal antennular segment and scaphocerite slender. Basal endite of fi rst maxilliped triangular. Second maxilliped with basal part not especially broadened and with distal segment not especiall y elongated. Ischiomerus of third maxilliped broadened wi th outer face concave. Second pereiopod very long and slender, with carpus nearly as long as propodus, with fingers toothless ...... : ...... genus Cnathophylleptum gen. nov. 4. • Second maxi lliped extremely long and slender, with very elongate cylindrical merus and carpus, exceeding third maxiliped with more than the last 3 segments ...... Levicaris BRUCE, 1973 • Second maxilliped short, with very robust merus and carpus, not reaching beyond third maxilliped .... Cnathophylloides SCHMITT, 1933

Evolutionary considerations on Gnathophylleptum gen. segment of the third maxilliped. The Hymenoceridae have nov. and other palaemonoid shrimps recently been removed from the Gnathophyllidae (BRUCE, 1986; CHACE, 1992; HOLTH UIS, 1993; CHACE & BRUCE, 1993), on the basis of the unique morphology of their third With the exception of the second pereiopod and the maxilliped. Indeed, whilst in other palaemonoid shrimps the ischiomerus of its third maxilliped, whi ch both exhibit ischium and the merus of the third maxilliped are completely apomorphic characters, Cnathophylleptwn gen. nov. seems fus ed, in the Hymenoceridae these two segments are quite to be a primitive Gnathophyllidae, with few morphological distinct and freely articulated with each other (BARNARD, specializations. The upper laci nia of the maxillula is only 1950; FUJINO, 1973; BRUCE, 1986, 1988b), and therefore it is slightly enlarged as in Pycnocaris, whilst in the three other a priori difficult to derive the hymenocerid disposition fro m genera : Cnathophylloides, Cnathophyllum and Levicaris, it the normal palaemonoid condition. is considerably enl arged. CnathoP,hylleptum gen. nov. is the only genus in wh ich the maxilla has a distinct basal endite. However, both families have- similar larvae, which are also The second max illiped of Cnathophylleptum gen. nov. is closely related to those of the Pontoniinae (Bruce, 1986), and unspecialized, as in Cnathophylloides mineri (SCHMITT, the Gnathophyllidae and the Hymenoceridae have some 1933) (see SCHM ITT, 1933), whilst it is moderately modified other features in common. In all gnathophyllid and in Cnathophylloides robustus BRUCE, 1973 (see BRUCE, hymenocerid genera, the body is fairly robust, the mandible 1973) and Pycnocaris chagoae BRUCE, 1972 (see BR UCE, is very small , and is devoid of an incisor process or has only 1972), and deeply modified in Cnathophyllum spp. (see a vestigial one; the maxilla has a basal endite entire, very HOLTHUIS, 1947; CHACE & FULLER, 197 1; TITGEN, 1989; reduced to absent; and the third maxilliped is broadened and MANN ING & CHACE, 1990) and Levicaris mammillata flattened. Furthermore, the gnathophyllid genera (EDMONDSON, 193 l)(see EDMONDSON, 193 1; BRUCE, 1973; Cnathophylleptum gen. nov. and Cnathophyllwn (material FUJI NO & TAKEDA , 1977). Finall y, in Cnathophylleptum, the examined), as well as the hymenocerid genera Hym.enocera separation between the ischiomerus and the basis is more (see BARNARD, 1950) and Phyllognathia (see BALSS , 1950; evident than in any other gnathophyllid genus. FUJINO, 1973; MINEMIZU, 2000) have a more or less devel oped corneal papilla, a rare character amongst palaemonoid Interestingly, both the gnathophyllid Cnathophylleptum shrimps. tellei gen. nov. , sp. nov. and the hymenocerid Phyllognathia ceratophthalma (BALSS , 19 13) (see BALSS, 1913) and Interestingly, the few other palaemonoid shrimps wh ich ex Phyllognathia simplex FUJINO, 1973 (see Bruce, 1988b) hibit a corneal papill a : Parapontonia nudirostris BRUCE, have two distal teeth on the outer border of the penultimate 1968 (BRUCE, 1968, 1994) and species of the Periclimenes I I

124 CEDRIC D' UDEKEM D' ACOZ

ceratophthalmus group (BRUCE, 1986) are echinoderm asso- BRUCE, A.J. , 1988b. Further records of the Hymenocerine shrimp ciates, just like several gnathophyllid species Phyllognathia simplex FUJINO (Crustacea: Decapoda). The Beagle, Gnathophyllum america11wn GUERIN-MENEVILLE, 1855 Records of the Northern TerritO!)' Museum ofArls and Sciences, 5 (MANN ING , 1963; TITGEN, 1989; PEREZ SANCHEZ & (1): 101-104. MORENO BATET, 1991; GONZALEZ PEREZ, 1995), BRUCE, A.J., 1994. A synopsis of the Indo-Pacific genera of the Gnathophyllwn elegans (RISSO, 18 16) (GONZALEZ PEREZ, Pontoniinae (Crustacea: Decapoda: Palaernonidae). Theses 1995), Gnathophylloides mineri SCHMITT, 1933 (LEWIS, Zoologicae Vol. 25, Konigstein, Koeltz Scienti fic Books, 172 pp. 1956; CHACE, 1972; BRUCE, 1974, 1988a; DEBELIUS, 1999; CHACE, F.A., Jr., 1972. The Shrimps of the Smithsonian-Bredin HICKM AN & ZIMMERMAN, 2000; DE GRAVE, 200 I; KATO & Caribbean Expeditions with a summary of the West Indian shallow OKUNO, 200 I), Gnathophylloides robust us (BRUCE, 1973) water species (Crustacea: Decapoda: Natantia). Smiihsonian Con (BR UCE, 1973), Levicaris mammillata (EDMONDSON, 1931) Iribwion Io Zoology, 98: i-x + 1-179. (EDMONDSON, 1931 ; BRUCE, 1973; FUJI NO & TAKEDA, CHACE, F.A. , Jr. , 1992. On the classification of the Caridea 1977; TITGEN, 1989) and the hymenocerid Hymenocera (Decapoda). Crustaceana, 63 ( I ): 70-80. picta DANA , 1852 (WICKLER, 1973). Although no symbiotic CHACE, F. A., Jr. & BRUCE, A.J ., 1993. The Caridean Shrimps (Crus relationship was recorded for Gnathophylleptum tellei gen. tacea: Decapoda) of the Albatross Philippine Expedition 1907- nov., sp. nov., the presence of a corneal papilla may indicate a 1910, Part 6: Superfamily Palaemonoidea. Smithsonian Coniribu similar echinoderm association. tion to Zoology, 543: i-vii + 1-152. CHACE, F. A., Jr. & FULLER, S.L.H., 197 1. A new shrimp of the ge nus Gnathophyllum (Decapoda, Caridea) from Puerto Rico. Pro Acknowledgements ceedings of the Biological Society of Washing/on, 83 (44): 493-504. DEBEUUS, H., 1999. Crustacea Guide Of The World. IKAN - I am deeply indebted to Mr. Arthur TELLE for giving me the Unterwasserarchiv, Frankfurt, 321 pp. opportunity to describe this handsome and remarkable new shrimp. I also would like to thank Dr. Sammy DE GRAVE for DE GRAVE, S., 200 I . Caridean shrim ps (Crustacea, Decapod a) from critical reading of the manuscript and for his assistance in Hansa Bay, Papua New Guinea: Palaemonidae and Gnathophyllidae. Bulletin de l'fnstitut Royal des Sciences tracing some relevant papers. Nawrelles de Belgique I Bulletin van het Koninklijk Belgisch lnsliluul voor Natuunve/enschappen, Biologie, 70 [year 2000]: 11 9- 148. 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