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Bijdragen Tot De Dierkunde Contributions to Zoology, 65 (2) 79-99 (1995) SPB Academic Publishing bv, Amsterdam New perspectives on the evolution of the genus Typhlatya (Crustacea, Decapoda): first record of a cavernicolous atyid in the Iberian Peninsula, Typhlatya miravetensis n. sp. Sebastián Sanz & Dirk Platvoet 1 Unitat d'Ecologia, Facultat de Ciències Biologiques, Universitat de Valencia, E-46100 Burjassot, 2 Valencia, Spain; Institutefor Systematics and Population Biology (Zoological Museum, Amsterdam), University of Amsterdam, P.O. Box 94766, 1090 GT Amsterdam, The Netherlands Keywords: Typhlatya, Decapoda, Spain, subterranean waters, systematics, zoogeography, vicariance, evolution, key to genus Abstract historia geológica de la zona y la distribución mundial del género, del grupo de géneros, y la familia. On several occasions, shrimps belonging to a new species ofthe genus Typhlatya were collected in a cave in the province of Castellón, Spain. This is the first record of the in the genus Introduction Iberian Peninsula. The species is described and the validity, dis- tribution, and zoogeography of the genus, as well as the status In 1993 and were on several of the discussed. 1994, shrimps caught genus Spelaeocaris, are Former models for the occasions in in the evolution of the genus Typhlatya and its genus group are re- a cave near Cabanes, province viewed, as well asthe system ofinner classification of the Atyidae of Castellón, eastern Spain. The specimens belong and its For the and evolution of biogeographical meaning. age the to genus Typhlatya Creaser, 1936, a genus the genus we developed a new model based on vicariance prin- with members known from the Galápagos Islands, ciples that involves further evolution of each species after the Ascension and the Caribbean of the ancestral This allows estimations Island, Bermuda, disruption range. new for the of area Dominican Mona age the genus. Accordingly, we suppose that other (Mexico, Cuba, Republic, proposals, such as recent dispersal through the sea, should be Island, Barbuda, Puerto Rico, Caicos Islands, disregardedfor this genus. Theevolutionary developmentofthis Bonaire, and Curaçao) (cf. Holthuis, 1986; species is discussed in the context of the geologicalhistory ofthe Banarescu, 1990; Stock, 1993). Up to the present, and the world distribution ofthe the area genus, genus group, members of the have not been recorded in and the family. genus Eurasia. Several works have dealt with the origin and bio- Resumen geography of the genus group, the genus, or some of its species (Monod & Cals, 1970; Croizat et al., En diversas ocasiones se capturaron camarones pertenecientes a 1974; Monod, 1975; Rosen, 1976; Iliffe et al., 1983; una nueva especie del género Typhlatya en una cueva de la Banarescu, 1990; Stock, provincia de Castellón (España). Esta constituye la primera cita 1993). del la Península Ibérica. In of the coherent géneroen La especie es descrita y se dis- spite vicariance models sug- cute sobre la validez, la distribución la del y zoogeografía gested by several authors cited above, a general género, así como sobre el estatus del género Spelaeocaris. Se agreement on the evolutionary history of the genus revisan anteriores modelos sobre la evolución del género y la is far from being reached. In other works, a recent serie, así como el sistema de clasificación interna de los atiídos the sea has been considered su la edad evolución dispersal through y significado biogeográfico. Respecto a y & Hart del género, desarrollamos un nuevo modelo basado en princi- (Chace Hobbs, 1969; Peck, 1974; et al., pios de vicariancia considera la evolución de cada que posterior 1985; Stock, 1986). The occurrence of a member of especie después de la del ancestral. Esto disrupción rango per- the genus in the Iberian Peninsula made us recon- mite aportar nuevas estimaciones respecto a la edad del género. sider the differenttheories and formulateour ideas De acuerdo con este modelo, otras propuestas, como dispersión about the reciente evolution of both this new species and the por el mar, deberían desestimarse para este género. Se comenta whole el desarrollo evolutivo de esta especie en base a la genus. Downloaded from Brill.com10/01/2021 11:12:48AM via free access & D. Platvoet - Evolution 80 S. Sanz of Typhlatya ischium shorter than merus. Exopodite of Systematic part always fifth pereiopod reaching beyond ischium (Fig. 4a). 4. Epipodites present on pereiopods 1 to Dactylus Typhlatya miravetensis n. sp. of fifth pereiopod distinctly longer than that of (Figs. 1-4) pereiopods 3 and 4. Pereiopod 5 longest. First pleopod (Fig. 4b) with short inner ramus; Material examined. - Male holotype, 20.7 mm, female allotype, no interna Second pleopod (Fig. 18.5mm,and two female paratypes,11.9 and 13.3 mm (coll. no. appendix present. Rambla de and short ZMA De. 201475, a, b, c). Cave "Ullal de la 4e) with appendix interna appendix mas- Miravet" between the towns of Cabanes and Orpesa, province culina with nine setules implanted near apex. Pleo- ofCastellón, eastern Spain (UTM coordinates: 30T YK 504447), pods 3 (Fig. 4c) to 5 with subequal rami and appen- chloride 25 30 June 1993. Conductivity 615.2 nmhos; mg/1, pH dix interna. fauna: 7.01; water temp. 24°C. Accompanying Typhlocirolana Uropods (Fig. 4d) long and slender. Diaeresis sp. (Isopoda: Flabellifera). March 1994. Other material: same locality, 10 specimens, with short spine on posterodistal corner and three setules. Telson (Fig. 2d) with one pair of lateral inter- Description of holotype. - Habitus illustrated(Fig. spines and 13 spines on posterior margin, mixed with small Two la). Length 20.7 mm. Rostrum short, not reaching two setae. longitudinal rows beyond eyestalks, almost absent. Eyes without pig- of setules on dorsal surface. ment. First peduncular segment of antennule as and third sub- Female: for the second where the long as second combined, flagella Except pleopod, of females lack the sexual equal in length, betweenhalfand two-thirds body appendix masculina, no length (Figs, la, b), outer flagellum with proximal dimorphism was found. fourteen segments swollen. Stylocerite slender, not Remarks. - This fits well within the defini- reaching beyond first peduncular segment. Anten- species nal scaphocerite (Fig. le) reaching well beyond tionof the genus(see Creaser 1936; Monod & Cals, Hobbs absence of on peduncle; distal tooth on straight outer margin at 1970; et al., 1977): spines antennular 80% of total length. Inner flagellum 35% longer carapace, rostrum never overreaching first fifth than body. Outer flagellum one-segmented and peduncle and lacking spines, through with first to fourth pereio- extremely reduced. pereiopods exopodites, Mandibles with row of hairs between molar and pods with epipodites. incisor (Figs, lc, d). Palp absent. Small projection The species diagnosis is based on a combination of first maxilla of characters that makes it differentfrom all other present at base of upper lobe (Fig. of setules base of of the short If), inner margin with row near species genus(see below): very rostrum, of teeth. Inner of lower lobe with absence of ischium and merus strong edge long eye pigment, of fifth pereiopod setae of varying length. Palp one-segmented with pereiopods not fused, exopodite well A to the of three small setae near apex. Membranous structure developed. key species Typhlatya second maxilla is in the at mid-length on palp of (Fig. 2b), given Appendix. near connection between scaphognathite and palp. First maxilliped (Fig. 2a) with long flagellar lobe, Derivatio nominis. - The specific name, mirave- de distal lobe of exopodite reduced. Epipodite small tensis, refers to the mediaevalname of "Tinença located. but distinct, thumb-shaped. Second maxilliped as Miravet", where the cave is illustrated (Fig. 2c). Flagellum of exopodite of third maxilliped (Fig. 3a) with podobranch and small Ischium-merus fusion. - In some species of the epipodite. Segmentation between basis and ischium Typhlatya, fusion of the ischium and merus in such indistinct. pereiopods occurs (Table I). In many groups, like this is Exopodites of pereiopods decreasing in relative as amphipods, a character considered to length from anterior to posterior (Figs. 3b-e, 4a). be of generic value. This could imply that the genus be dividedinto six sub- Ischium and merus separate in all pereiopods, Typhlatya might as many as Downloaded from Brill.com10/01/2021 11:12:48AM via free access Contributions to Zoology, 65 (2) - 1995 81 Fig. 1. Typhlatya miravetensis n. sp.: a, habitus; b, first antenna with enlarged seta from segment 1 (scales III and V); e, left mandible (I); d, right mandible (I); e, second antenna with enlarged seta from antennal scale and detail of distal part of peduncle (III); f, first maxilla (IV). Downloaded from Brill.com10/01/2021 11:12:48AM via free access 82 S. Sanz & D. Platvoet - Evolution of Typhlatya Fig. 2. Typhlatya miravetensis n. sp.: a, first maxilliped (scale IV); b, second maxilla (IV); c, second maxilliped (IV); d, telson (II). Downloaded from Brill.com10/01/2021 11:12:48AM via free access Contributions to Zoology, 65 (2) - 1995 83 Fig. 3. Typhlatya miravetensis n. sp.: a, third maxilliped (scale II); b, first pereiopod(II); c, second pereiopod (II); d, third pereiopod (II); e, fourth pereiopod (II). since characters rank groups. However, other (rostrum, Typhlatya pretneri (Matjasic, 1956) new eye pigmentation, exopodite P5 reduction) do not support such a division we consider the genus Typh- The genus Spelaeocaris (S. pretneri Matjasic, for workable which is found in latya, the time being, as a taxon. 1956), Hercegovina (former Downloaded from Brill.com10/01/2021 11:12:48AM via free access 84 S. Sanz & D. Platvoet - Evolution of Typhlatya second Fig. 4. Typhlatya miravetensis n. sp.: a, fifth pereiopod(scale II); b, first pleopod (II); c, third pleopod (II); d, uropod (II); e, pleopod with details (II, IV, V). Yugoslavia), is distinguished from Typhlatya by the T. pearsei Creaser, 1936, the exopodites of the pe- of the from P5.
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