s PROC. BIOL. SOC. WASH. 106(2), 1993, pp. 332-338 THE IDENTITY OF TALITROIDES ALLUAUDI (CHEVREUX) (CRUSTACEA: AMPHIPODA: TALITRIDAE) WITH NOTES ON A NEW LOCALITY Hiroshi Morino and Reuven Ortal —A Abstract. lectotype of Talitroides alluaudi (Chevreux) is designated from the type-series and described to remove taxonomic confusion on the identity ofthis species. The discovery of this species in Israel is briefly discussed. Examination of terrestrial talitrids col- tail. The Paris material was described again & lected from southern parts ofIsrael revealed in 1925 (Chevreux Page 1925) with il- A two genera and three species, one ofwhich lustrations. study of these three descrip- proved to be Talitroides alluaudi. This spe- tions discloses the following apparent un- cies is one ofthe most widespread landhop- conformity in some characters: The Paris pers, known from the tropics to warm-tem- 1896 and Seychelles material have well-de- perate regions and hothouses in Europe and veloped brood plate on female gnathopod & NorthAmerica(Friend Richardson 1986), 2, though this is lacking in the Paris 1925 though so far not known from the Levant. material, and; the Seychelles material has Although some authors have remarked on 1-articulated and 3-articulated inner ramus various characters, especially concerning the on pleopods 1 and 2, respectively, though pleopods, ofthis species (Medcof1940, Pal- the Paris 1925 material has 4-articulated men 1949, Andersson 1962), no full de- and 1-articulated inner ramus, respectively scriptions or figures have been published (no description of pleopods for the Paris since the original description of Chevreux 1896 material). If these descriptions are (1896, 1901). Close examinationofmaterial taken authentically, we couldrecognizethree from Israel, with reference to Chevreux' species in the Chevreux's concept of T. al- descriptions, revealed a few minor but dis- luaudi: two from Paris and one from the tinct discrepancies between them as well as Seychelles. And if the Paris 1896 material between previous descriptions ofT. alluau- was from the Seychelles, as assumed by di. Shoemaker (1936), two species may be dis- Chevreux (1896) described this species cernible, from Paris and the Seychelles, re- three times from two localities. He gave a spectively. Medcof(1940) has also pointed brief original description ofthis species on out some confusion in the earlier descrip- the basis of material procured from hot- tions ofthis species, which he attributed to houses at the Paris Museum ("les serres du geographical variation ("based on exami- Museum de Paris"). In this paper he men- nations of specimens collected from areas tioned that the Paris material was identical sometimes widely separated"). Since Med- to that collected from the Seychelles. Al- cofs analysis of variation is limited to the though he did not specify the locality ofthe pleopod structure, he did not notice the type, the title ofhis paper suggests that Paris variation in the brood plate. In addition, is the probable type locality. Shoemaker material from Israel at hand displays a pe- (1936), however, assumed that the Sey- culiar feature ofthe pereopod dactyl, which chelles was the type locality. In 1901, Chev- was described in the Seychelles material dif- reux treated the Seychelles material in de- ferently, andwas not mentioned forthe Par- VOLUME 106, NUMBER 2 333 is material. Thus it is necessary to deter- awa, Molokai Is., Hawaii, 9 Mar 1968, Si- mine the original concept of T. alluaudi nonaga collector (Morino Cat. No. T.868); through direct reference to the type-series. 1 female (4.8 mm), from Kokee, Kauai Is., The type material ofTalitrus alluaudide- Hawaii, 11-12 Mar 1968, Sinonaga collec- posited in Museum National d'Histoire Na- tor (Morino Cat. No. T.869). turelle in Paris consists of preserved spec- Description of lectotype.—Eyt small in imens from four localities (MNHN Paris size, subround. Inferior antennal sinus me- Am 4500, 4501, 4502, 4503) and slides dium deep. Head longer than deep. (MNHN Paris Am 4504). The label on the Antenna 1 exceeding mid-point of pe- slides ("Serre du Museum") suggests that duncular article 5 ofantenna 2, peduncular they comprise the type specimen. Unfor- articles subequal in length; flagellum a little tunately the condition ofthe slides is so bad shorter than peduncle, 6-articulated. An- that it is not possible to discriminate the tenna 2: peduncle weakly spinose, article 5, pertinent characters, and all the preserved 1.79 times as long as article 4; flagellum specimens are from other than the Paris subequal to peduncle in length, 1 1-articu- Museum. Thus the lectotype is selected from lated. the localitynearesttothe Paris Museum and Upper lip: as wide as deep. Mandible: in- described to resolve the problem of T. al- cisor chitinized strongly, brown in color, luaudi. Material from Israel and Hawaii is 5-dentate; left lacinia chitinized as incisor, also examined. 4-dentate; right lacinia not strongly chitin- ized, with 3 cusps; spine rows of 2 broad plumosebristles. Lowerlip normal. Maxilla Systematic Account 1: innermost spine of outer plate leaning Talitroides alluaudi (Chevreux, 1896) medially, outermost 2 spines without den- Figs. 1, 2 ticles; palp small,jointofarticle 2 indistinct. Maxilla 2: inner plate distolaterally pro- Tal1i-t4r;us1A9l0l1u:a3u8d9i,Chfiegvs.re1u-x6,.-1C89h6e:v1r1e2,uxfig&s. dseutcaeedpilnutmoopsoei.ntMeadxitlilp,ipseedv:erianlnmeerdpiloa-tdeisatpail- Page, 1925:270, figs. 280-281. cally truncate, with 3 weak spine-teeth, in- Orchestia senni Menz—el, 191 1:438, figs. 4-9 creasing in size laterally, and with several Talitroidesalluaudi. Palmen, 1949:61, figs. plumose stiff" setae; outer plate weakly ar- 1-12.—Andersson, 1962:21 1, figs. 1-3.— cuate with rounded apex, weakly setose; palp Bousfield, 1984:210. articles 2 and 3 slender, with a few stiff"setae Material examined. —T^o females (5.5 distally, article 2 with remnant of medial mm— mm— lectotype, 5.0 paralectotype), lobe represented by elongate setae, article 4 from Serres de la Ville de Paris, Boulogne partially fused to article 3. sur Seine, France (MNHN Paris Am 4500); Gnathopod 1: coxa truncate distally, low- 1 female (5.0 mm), from lies Sechelles, er margin spinose, inner shelfweak, with a Mahe, Auct. det 1901 (MNHN Paris Am spine; basis broadened distally; carpus 1.50 mm 4502); 2 females (6 and 7 mm), from times as long as propod; propod simple, Ponta Delgada, lie S. Michel, Agores, Aug gradually narrowed to base of dactyl, pos- 1930, M. Mequignon collector, (MNHN terior margin with 3 stiff* long spines and Paris Am 4503); 5 females (up to 5.5 mm), several shorter submarginal spines on inner from Ben Gurion Univ. campus, Beersheba, and outer sides, anterior margin with 2 spine Israel, bamboo stands, 10 Dec 1987, Y. clusters; dactyl cuspate anteriorly, base with Margalit collector (Morino Cat. No. T.766), a stiff* seta posteriorly, nail as long as base. 4 Mar 1989, R. Ortal collector (Morino Cat. Gnathopod 2: coxa cuspate posteriorly, as No. T836); 1 female (4.5 mm), from Hal- deep as wide, lower margin rounded and 334 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON MNHN Am Fig. I. Talitroides alluaudi (Chevreux). Boulogne sur Seine, France. The lectotype, 4500-1 mm (female, 5.5 body length). A. lateral view; B, antenna 1; C, antenna 2; D, left mandible; E, right mandible; F. maxilla 2; G, maxilla 1: H. maxilliped; I. distal part ofright palp ofmaxilliped; J, upper lip; K, lower Up. weakly spinulose; basis slender, anterior a stiff seta posteroventrally, and a few spi- margin spinulose; merus and carpus with nules ventrally. Coxa 5 anterolobate, ante- tumescence posterioriy, carpus as long as rior lobe deeper than that of pereopod 4. propod; propod anterior margin weakly se- Coxa 6: posterior lobe truncate distally, an- tose, posterior tumescent lobe well devel- terior margin of the lobe vertical. Coxa 7 oped, exceeding dactyl by half of propod shallow. length. Pereopods 3-7 cuspidactylate, propod Coxae 3 and 4 shallower than wide, with lacking hinge spine, dactyl base with a stiff VOLUME 106, NUMBER 2 335 Fig. 2. Talitroides alluaudi (Chevreux). Boulogne sur Seine. France. A. gnathopod 1; B, dactyl ofgnathopod 1; C, gnathopod 2; D. coxal gill ofgnathopod 2; E. pereopod 3; F & G. dactyl ofpereopod 3; H, pereopod 4; I, dactyl of pereopod 4; J. pereopod 5: K. dactyl of pereopod 5; L, pereopod 6; M, coxal gill ofpereopod 6; N, dactyl ofpereopod 6; O, pereopod 7; P. dactyl of pereopod 7: Q. abdominal side plates 1-3; R, pleopod 1; S, pleopod 2; T, pleopod 3; U, uropod 3: V. uropod 2: W. uropod 1: X. telson. G = Azores, MNHN Am 4503-1 (female 6.0 mm); M = Boulogne sur Seine, France, paralectotype MNHN Am 4500-2 (female 5.0 mm); others = Boulogne sur Seine. France, lectotype MNHN AM 4500-1 (female 5.5 mm). 336 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON seta. Pereopod 3: dactyl base weakly ramus small, conical, partially fused to pe- pinched, nail with slight prominence at duncle, with a spinule at tip. Telson broad, middle on posterior margin. Pereopod 4 apically rounded with apical and 3-4 lateral shorter than pereopod 3, basis weakly ar- spines. cuate, dactyl base pinched, nail with sharp Male. —Not known. dentitionposteriorly. Pereopod 5 longerthan Variations. —Eye in the Israeli material is pereopod 4, basis narrow with a submar- somewhat larger than that ofthe lectotype. ginal spine at posterodistal comer, lacking Number of flagellar articles ranges from 4 posterodistal lobe. Pereopod 6 similar to to 6 on antenna 1 andfrom 8 to 9 onantenna but much shorter than pereopod 5, dactyl 2 in the material examined, tending to in- nailalmost straight. Pereopod 7 slightlylon- crease with body size. Marginal setae on ger than pereopod 6, basis expanded pos- propod anterior margin ofgnathopod 2 also teriorly, as deep as wide. show variation in number: Seychelles and Coxal gill ofgnathopod 2 as long as basis, Hawaiian Islands material lacks them, Is- distal margin with crenulations. Coxal gills raeli material with 1-2 setae, Azores ma- ofpereopods 3-5 small, constricted at mid- terialwith0-2. Swissmaterial (Menzel 1911, dle. Coxal gill of pereopod 6 longer than as Orchestia senni) also has one seta. Inner basis, reverse L-shaped. Brood plates con- rami of pleopods 1 and 2 fluctuate in the fined to pereopods 3-5, small and slender number of articles: from 1 to 4 (mostly 2) with 2-3 simple setae apically. articles in pleopodm1m; 1, exceptionally 3 (in Abdominal side plate 1 rounded postero- Azores material, 7 bodylength) articles ventrally. Plates 2 and 3 bluntly pointed in pleopod 2. Onlythis larger specimen from posteroventrally, posterior margins weakly the Azores has reduced ramus in pleopod spinulose. Pleopod 1: peduncle weakly ar- 3, otherwise lacking them. For each speci- cuate, with 3 plumose setae on outer mar- men examined, article number ofinner ra- gin, with 2 retinaculae; outer ramus mus ofpleopod 1 is higherthan, rarely same 7-articulated; inner ramus shorter than half as that ofpleopod 2. This range and pattern of outer ramus, 2-articulated. Pleopod 2 a of variation in pleopods is in accord with little shorterthan pleopod 1 peduncle outer those in the other geographical populations: , margin with a plumose seta, outer ramus Basel, Switzerland (Menzel 1911); Urbana, 6-articulated, inner ramus 1-articulatedwith Illinois, U.S.A. (Medcof 1940); Finland apical setae. Pleopod 3 small and slender, (Palmen 1949) and the Canary Islands (An- 0.4 times as long as peduncle ofpleopod 2, dersson 1962). Dactyl nail of pereopod 3 with 2 spinules subapically, lacking rami. has usually very weak prominence at the Uropod 1: peduncle with marginal spines posterior middle. The Azores material (6.0 on both edges, distolateral spine strong, mm)has, however, distinctprominence (Fig. closely set to distomarginal spine; rami sub- 2G) which is rather similar to the dentition equal in length and shorter than peduncle, ofpereopod 4. with elongate apical spines; outer ramus marginally bare; inner ramus with 3 mar- Discussion ginal spines. Uropod 2: peduncle with mar- ginal spines distally; rami subequal to each Most ofthe variations mentioned above other and to peduncle in length, with elon- occur within a given population rather than gate apical spines; outer ramus marginally between geographically separated popula- bare; inner ramus with spines proximally. tions. Thus it is not possible to recognize Uropod 3 very small, not reaching tip of more than one species amongstthe material telson, subtriangular in shape; peduncle examined. All the material lacks a brood broad at base with a strong spine ventrally; plate on female gnathopod 2, and has dac- VOLUME 106, NUMBER 2 337 tylar dentition on pereopod 4 (rarely also this country, most ofwhich were imported on pereopod 3). Chevreux (1901) men- from West Europe. Thus it is highly prob- tioned the dactylar dentition on pereopod able that the population of T. alluaudi in 5, not on pereopod 4, in his Seychelles ma- Beersheba was introduced accidentally from terial. At the same time, in our specimens, European countries with the bamboo and the inner ramus ofpleopod 1 has more ar- has survived on this small "island," where ticles than that ofpleopod 2, and never less, a humid microclimate may have prevailed. as by Chevreux (1901). Thus it is possible that Chevreux treated an exceptional spec- Acknowledgments imen or more probably some confusion oc- We are grateful to D. Defaye in Museum curred during the preparation of his mate- National d'Histoire Naturelle, Paris for the rial. loan of the type material treated here. The present speciesis most closely related Thanks to J. Aoki in Yokohama National to Talitroidestopitotum (see Bousfield 1984). University who provided us with compar- & Lam Friend (1985) listed several discrim- ative material. Thanks also to K. Iwatsuki inating characters between T. alluaudi and ofThe University ofTokyo who helped lo- T. topitotum. In addition, thefollowingones cate the type material. areto be mentioned: T. alluaudihas gnatho- pod 2 without brood plate, strongly chitin- ized mandible, maxilla 2 with distally Literature Cited pointed inner plate, and pereopod 4 with Andersson, A. 1962. On a collection ofAmphipoda dactylar dentition. These features are rather of the family Talitridae from the Canar>' Is- peculiar among landhoppers. Talitroides al- lands.-Arkiv for Zoologi 15(1 1):211-218. luaudialso resemblesArcitalitrussylvaticus, Bousfield, E. L. 1984. Recent advances in the sys- a non-cuspidactylate landhopper, in having tematicsandbiogeographyoflandhoppers(Am- phipoda: Talitridae) of the Indo-Pacific re- simple gnathopod weakly setose anterior 1, gion.-Pp. 171-210 in F. J. Radovsky, P. H. margin in gnathopod 2, reduced and plu- Raven & S. H. Sohmer. ed., Biogeography of mose-setose peduncle of pleopods, similar the tropical Pacific, Bishop Museum Special spination in uropods 1 and 2, etc. But the Publication 72. latter is distinguished from the former by Chevreux, E. 1896. Recherches zoologiques dans les serres du Museum de Paris IV.—Sur un Am- the possession of a strongly arcuate outer phipodeterrestreexotique. TalitrusAlluaudinow plate and less setose maxilliped. It is inter- sp., acclimate dans les serres du Jardin des esting that T. alluaudi has a weakly arcuate Plantes de Paris.—Feuille des Jeunes Natural- inner plate. istes 26:112-113. The distribution of T. alluaudi in conti- . 1901. Mission scientifique de M.Ch.Alluaud aux lies Sechelles. Crustaces Amphipodes.— nental areas is believed to be due to syn- anthropic dispersal (Friend & Lam 1985), M1e4:m3o8i8r-e4s38d.e la Societe Zoologique de France and the occurrence is thought to be limited . & L. Fage. 1925. Faune de France 9. Am- by the humidity (Andersson 1962). Thus it phipodes. Paul Lechevalier. Paris. 486 pp. may be astonishing that T. alluaudi has been Friend, J. A., & P. K. S. Lam. 1985. Occurrence of the terrestrial amphipod Talitroides topitotum discovered in an arid part ofIsrael with pre- mm (Burt) on Hong Kong Island.—Acta Zootaxon- cipitation as little as 200 annually. So omica Sinica 10(l):27-33. far in Israel the present species is known & A. M. M. Richardson. 1986. Biology of . solely from the bamboo stands growing on terrestrial amphipods.—Annual Review ofEn- tomology 31:25^8. a small "island" in the middle of a small pool in the University campus at Beersheba. Medcof. J. C. 1940. Variations in the pleopod struc- ture ofthe terrestrial amphipod Talitrusalluau- The bamboo, now commonly utilized for di Chevreux.-Lloydia 3(l):79-80. gardening in Israel, is an exotic plant for Menzel, R. 1911. Exotische Crustaceen im botan- 338 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ischenGartenzuBazel.—RevueSuissedeZool- (HM) Department of Biology, Ibaraki ogie 19:433-444. University, Mito 310, Japan; (RO) Depart- Palmen, E. 1949. Talitroides alluaudi (Chevreux) ment of Ecology, Systematics and Evolu- (Amphipoda, Talitridae) in Finnland gefun- den.—Archivum Societatis Zoologicae Botani- tion, The Hebrew University ofJerusalem, cae Fennicae 'Vanamo' 2:61-64. Givat Ram, Jerusalem 91904, Israel. Shoemaker, C. R. 1936. The occurrence ofthe ter- restrial amphipods, Talitrus alluaudi and Tali- trussylvaticus, inthe UnitedStates.—Journalof the WashingtonAcademy ofSciences 26(2):60- 64.