New records and species of Halophilosciidae (Crustacea, Isopoda, Oniscidea) from the Canary Islands (Spain)

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1 New records and species of Halophilosciidae (Crustacea, Isopoda, Oniscidea) from the Canary Islands (Spain) Stefano Taiti 1, * Heriberto López 2 1 Istituto per lo Studio degli Ecosistemi, CNR, Via Madonna del Piano 10, Sesto Fiorentino (Florence), Italy 2 Departamento de Biología Animal, Universidad de La Laguna, La Laguna, Tenerife, Spain * stefano.taiti@ise.cnr.it Abstract To date the family Halophilosciidae from the Canary Islands included four species: Halophiloscia couchii from all the islands of the archipelago, H. canariensis from the banks of a subterranean lake on Lanzarote, Stenophiloscia glarearum from Lanzarote and La Gomera, and Littorophiloscia culebrae from Tenerife. We examined a large number of specimens of Halophilosciidae collected from the coasts of the Chinijo Archipelago, Lanzarote, Lobos, Fuerteventura, Gran Canaria, Tenerife, La Gomera, La Palma and El Hierro, and from some caves on La Palma. Six species of Halophilosciidae are now recorded from the Canary Islands, including two new species of Halophiloscia described herein (H. rodriguezi from the coasts of Tenerife, La Palma, La Gomera and El Hierro, and H. microphthalma from numerous caves on La Palma). The distribution of all the species in the archipelago is discussed. Halophiloscia canariensis appears to be widespread along the coasts of the eastern islands (Chinijo Archipelago, Lanzarote, Lobos, Fuerteventura and Gran Canaria), H. rodriguezi along the coasts of the western islands (Tenerife, La Palma, La Gomera and El Hierro), H. couchii from Tenerife and Gran Canaria, while H. microphthalma is endemic to caves on La Palma. Stenophiloscia glarearum and Littorophiloscia culebrae are newly recorded from Gran Canaria. Keywords Canary Islands, Crustacea, Halophilosciidae, new species, Oniscidea Introduction The family Halophilosciidae includes the genera Halophiloscia Verhoeff 1908, Stenophiloscia Verhoeff 1908, and, according to Leistikow (2001), also Littorophiloscia Hatch All species have a philosciid appearance and belong to the runner type (sensu Schmalfuss, 1984). They usually occur in marine littoral habitats, along rocky and sandy shores, lagoons, and on the banks of river mouths. Only Littorophiloscia alticola (Vandel 1977) occurs in montane forests of St. Helena Island (Vandel, 1977), while another species of Littorophiloscia from Hawaii Island, not yet described (Rivera et al., 2002), is a cavernicolous form closely related to the littoral L. hawaiiensis Taiti & Ferrara At present the genera Halophiloscia with six species and Stenophiloscia with three species occur along the Mediterranean and Atlantic coasts of Africa and Europe, while Littorophiloscia with 21 species has mainly a tropical distribution (Taiti & Ferrara, 1986). The family Halophilosciidae was apparently well known in the Canary Islands due to a revision by Rodríguez & Barrientos (1993). They recognized three genera and four species: Halophiloscia canariensis Dalens 1973, endemic to Lanzarote, H. couchii (Kinahan 1858), widespread throughout the coasts of the whole archipelago with two cave populations from La Palma, Stenophiloscia zosterae Verhoeff 1928 (junior synonym of S. glarearum Verhoeff 1908) from Lanzarote, and Littorophiloscia culebrae (Moore 1901) from Tenerife. During recent investigations carried out along the coasts of all the Canary Islands and in many lava tubes on La Palma, a large number of specimens of Halophilosciidae were collected. Their examination revealed a picture more complex than that proposed by Rodríguez & Barrientos (1993). Two new species of Halophiloscia could be recognised on the basis of morphological characters. Their validity is also confirmed by the results of a I. Biodiversity & Systematics: 37-52

2 44 Taiti & López molecular analysis which will be published separately. Some misidentifications by previous authors are discussed and many new records of Halophilosciidae species in the Canary Islands are presented herein. Abbreviations: GIET = Grupo de Investigaciones Espeleológicas de Tenerife DZUL = Departamento de Biología Animal, Universidad de La Laguna, La Laguna, Tenerife MCNT = Museo de Ciencias Naturales de Tenerife, Santa Cruz de Tenerife, Tenerife MZUF = Museo di Storia Naturale, Sezione di Zoologia La Specola, Florence Systematic Account Family Halophilosciidae Genus Halophiloscia Verhoeff 1908 Halophiloscia couchii (Kinahan 1858) Figs 1, 8A,B, 9 Philoscia couchi; Dollfus, 1893: 56 (? partim: nec Fuerteventura). Halophiloscia couchi; Vandel, 1954: 2, 57; Hoese, 1984a: 28, 29 (? partim: nec Lanzarote and Fuerteventura); Hoese, 1984b: 41, 43 (? partim: nec Lanzarote and Fuerteventura); Rodríguez & Barrientos, 1993: 185, figs 3-7 (? partim: nec Montaña Clara, La Graciosa, Lanzarote, Lobos, Fuerteventura, La Gomera, La Palma and El Hierro). Halophiloscia couchii couchii; Arcangeli, 1958: 57, 95 (? partim: nec Fuerteventura). Halophiloscia couchii; Rodríguez et al., 2004: 175 (? partim: nec Tenerife (San Roque), Fuerteventura, Lanzarote, El Hierro, La Palma and La Gomera ). Material examined Gran Canaria: 4, 10, mouth of Barranco de Guayadeque, N W, 7.IV.2004, leg. H. López (MZUF). Tenerife: 3, 8, Playa de los Tronches, Punta del Hidalgo, N W, 18.I.2005, leg. GIET (MZUF); 2, 2, same data (DZUL); 6, same locality, 28.X.2007, leg. H. López (MZUF). Previous records Gran Canaria: salt marshes of Juan Grande (Dollfus, 1893); Faro de Maspalomas (Vandel, 1954); Juan Grande; Playa de Ojos de Garza (Rodríguez & Barrientos, 1993). Tenerife: Playa del Médano (Rodríguez & Barrientos, 1993). Distribution Coasts of the Atlantic Ocean from Senegal to the British Isles, Azores, Madeira, Canary Islands, Cape Verde, Mediterranean and Black Sea. Introduced to North and South America, Hawaii and Australia. According to the original description by Radu (1985), H. pontica from the Rumanian coasts of the Black Sea is most probably a junior synonym of H. couchii (Schmalfuss, 2003). Remarks Halophiloscia couchii was fully illustrated by Schmidt (2003, figs 52-57) on the basis of specimens from Provence, France, and Cornwall, England. Despite its wide distribution, its morphology is very stable. We have examined several specimens from Portugal (Faro, II.2004, leg. N. Diaz), Morocco (Estuary of Oued Tahadart, N of Asilah, 2.X.2005, leg. S. Taiti & C. Rossano; Oued Laou, 1.V.2004, leg. S. Taiti & C. Rossano), Libya (Sabratah, 27.II.2005, leg. S. Taiti), Egypt (Abu Qir, 6.II.2003, leg. S. Taiti) and Italy (Tuscany, Maremma Regional Park, mouth of Ombrone River, 17.V.2003, leg. S. Taiti). All the populations examined showed identical morphological characters. According to Rodríguez & Barrientos (1993), H. couchii occurs along the coasts of all the islands of the Canary Archipelago. The same authors also identified some specimens from two caves on La Palma (Cueva de Tacande and Cueva de los Palmeros) as H. couchii, pointing out in the discussion that these specimens showed some distinct morphological characters from the epigean littoral populations due to their adaptation to cave life, such as loss of pigment and eyes, longer appendages, setose dorsal body surface and larger size. Since the male characters were identical in the epigean and cave populations, they concluded that the troglomorphic populations also belonged to H. couchii but

3 Halophilosciidae from the Canary Islands 45 Figure 1. Halophiloscia couchii from Gran Canaria, mouth of Barranco de Guayadeque, : A, pleonites 4, 5, telson and uropods; B, pleopod 1 and genital papilla; C, pleopod 2. were probably going through a speciation process due to their isolation. Our examination of a large number of specimens of Halophiloscia from all the Canary Islands showed that the typical morphology of H. couchii was present only in a few populations from Tenerife and Gran Canaria, while all the littoral populations from La Palma, La Gomera, El Hierro and some populations from Tenerife showed some constant differences in morphology from the typical H. couchii. Constant differences were also found in all

4 46 Taiti & López Figure 2. Halophiloscia canariensis from Lanzarote, Jameos del Agua, : A, pleonites 4, 5, telson and uropods; B, pleopod 1 and genital papilla; C, pleopod 2. cave populations from La Palma. Therefore, these two forms do not belong to H. couchii but to two distinct new species (see remarks under H. rodriguezi n. sp. and H. microphthalma n. sp.). Moreover, all the specimens examined from the Chinijo Archipelago (i.e. the islets north of Lanzarote: La Graciosa, Montaña Clara and Alegranza), Lanzarote, Lobos and Fuerteventura, were clearly referable to H. canariensis, while no H. couchii was present. Both species are certainly present in Gran Canaria. Halophiloscia canariensis Dalens 1973 Figs 2, 8C,D, 9 Halophiloscia canariensis Dalens, 1973: 248, figs 1, 2; Hoese, 1984a: 28, 30; Hoese, 1984b: 41, 43; Rodríguez & Barrientos, 1993: 185, figs 8-10; Schmalfuss, 2003: 116; Rodríguez et al., 2004: 175.

5 Halophilosciidae from the Canary Islands 47 Philoscia couchi; Dollfus, 1893: 56 (? partim: Fuerteventura) Halophiloscia couchi; Hoese, 1984a: 28, 29 (? partim: Lanzarote and Fuerteventura); Hoese, 1984b: 41, 43 (? partim: Lanzarote and Fuerteventura); Rodríguez & Barrientos, 1993: 185, figs 3-7 (? partim: Montaña Clara, La Graciosa, Lanzarote, Lobos and Fuerteventura). Halophiloscia couchii couchii; Arcangeli, 1958: 57, 95 (? partim: Fuerteventura). Halophiloscia couchii; Rodríguez et al., 2004: 175 (? partim: Fuerteventura and Lanzarote); Macías et al., 2004: 58. Material examined Chinijo Archipelago: 4, 3, Alegranza, Playa de El Trillo, N W, 8.XII.2004, leg. GIET (MZUF); 6, 2, 1 juv., Montaña Clara, El Veril, N W, 6.XII.2004, leg. GIET (MZUF); 2, 2, same data (DZUL); 2, 4, La Graciosa, Punta Corrales, N W, 3.XII.2004, leg. GIET (MZUF). Lanzarote: 4, Jameos del Agua, N W, 27.III.2004, leg. GIET (MZUF); 2, 1, Playa de la Madera, Timafaya, N W, 28.III.2004, leg. GIET (MZUF); 1, 2, Punta Pasitos, Mala, N W, 26.III.2004, leg. GIET (DZUL). Lobos: 4, 3, Las Salinas, N W, 30.III.2004, leg. GIET (MZUF); 2, El Muelle, N W, 1.IV.2004, GIET (DZUL). Fuerteventura: 4, 2, Playa de Esquinzo, La Oliva, N W, 3.IV.2004, leg. GIET (MZUF); 2, 2, same data (DZUL); 3, 3, mouth of Barranco de los Cuchillos, Matas Blancas, N W, 19.I.2007, leg. GIET (MZUF); 1, 2, same data (DZUL); 1, Punta del Morrete, Giniginámar, N W, 5.II.2005, leg. N. Macías (MZUF). Gran Canaria: 1, 9, Baja de los Matos, Pozo Izquierdo, N W, 2.XI.2007, leg. H. López & H. Morales (MZUF); 3, same data (DZUL). Previous records Lanzarote: Jameos del Agua (Dalens, 1973; Rodríguez & Barrientos, 1993). As H. couchii:? Montaña Clara: SW coast (Rodríguez & Barrientos, 1993); Veril (Macías et al., 2004).? La Graciosa: Playa de las Conchas (Rodríguez & Barrientos, 1993).? Lanzarote: (Hoese 1984a, 1984b); Playa Blanca; Los Lollos; Punta Pasito (Rodríguez & Barrientos, 1993).? Lobos: Saladar de Lobos (Rodríguez Barrientos, 1993).? Fuerteventura: beach North of Puerto Cabras [= Puerto del Rosario] (Dollfus, 1893); Punta del Tarajalito; Pozo Negro; Playa de Taca; Puerto Lajas (Rodríguez & Barrientos, 1993). Distribution This species seems to be endemic to the eastern Canary Islands: Chinijo Archipelago, Lanzarote, Lobos, Fuerteventura and Gran Canaria. Remarks This species was previously considered endemic to the littoral anchialine lava tube Jameos del Agua on Lanzarote, while the new records prove this species to be widespread along the coasts of all eastern Canary Islands. Since Rodríguez & Barrientos (1993) recorded only H. couchii from the coasts of these eastern islands, most probably their identification is incorrect and must refer to H. canariensis. Unfortunately the material studied by these authors could not be re-examined since it is no longer present in the collections of the Laboratorio de Zoología of the Universidad Autónoma de Barcelona where it was supposed to be deposited (T. Munilla, pers. com.). Also the record of Puerto Cabras, Fuerteventura by Dollfus (1893) most probably refers to H. canariensis. Halophiloscia rodriguezi n. sp. Figs 3-5, 8E,F, 9 Halophiloscia couchi; Rodríguez & Barrientos, 1993: 185 (? partim: Tenerife (San Roque), La Gomera, La Palma and El Hierro). Halophiloscia couchii; Rodríguez et al., 2004: 175 (? partim: El Hierro, La Palma and La Gomera).

6 48 Taiti & López Figure 3. Halophiloscia rodriguezi n. sp. from Tenerife, Playa de Lima, paratype : A, ovigerous specimen, dorsal. Paratype : B, dorsal scale-seta; C, cephalon, frontal; D, disposition of the noduli laterales on pereonites; E, pleonites 4, 5, telson and uropods; F, antennula; G, antenna.

7 Halophilosciidae from the Canary Islands 49 Material examined Holotype: Tenerife: 1, Playa de Lima, Güímar, N W, 17.XII.2004, leg. H. López & H. Morales (MZUF). Paratypes: Tenerife: 2, 2, Playa de Lima, Güímar, N W, 17.XII.2004, leg. H. López & H. Morales (MZUF); 4, 6, Playa de El Draguillo, Anaga, N W, 8.IV.2006, leg. H. López (MZUF); 2, 2, same data (DZUL); 1, Litoral del Malpaís de la Rasca, N W, 10.VII.2007, leg. A. J. Pérez, R. Castro & D. Hernández, (DZUL); 2, same locality, 12.X.2007, leg. D. Hernández, (MZUF); 5, 30, 8 juvs, Playa de los Tronches, Punta del Hidalgo, N W, 28.X.2007, leg. H. López (MZUF); 2, 4, same data (DZUL); 2, 4, same data (MCNT); 1, 4, 5 juvs, Playa de San Roque, Garachico, N W, 10.XI.2007, leg. H. López (MZUF). La Gomera: 1, 3, Playa de Hermigua, N W, 20.VI.2004, leg. H. López (MZUF). La Palma: 1, Playa de la Martina, N W, 4.IV.2004, leg. GIET. El Hierro: 1, 2, Playa de Tijereta, 27º46 39 N 17º54 26 W, 10.IV. 2004, leg. P. Oromí & H. López (MZUF). Additional material (used for molecular analysis): La Palma: 1 spec., Playa de la Fajanita, Don Pedro, 28º N 17º W, 1.IV.2004, leg. GIET; 2, Puerto Espíndola, N W, 12.VII.2004, leg. GIET. El Hierro: 1, Playa Timijiraque, N W, 10.VI.2004, leg P. Oromí & H. López; 1, Playa Arenas Blancas, N W, 13.V.2004, leg. GIET. Previous records As H. couchii:? Tenerife: Playa de San Roque (Rodríguez & Barrientos, 1993).? La Gomera: Punta de la Calera; Puntallana (Rodríguez & Barrientos, 1993).? La Palma: Playa de la Martina (Rodríguez & Barrientos, 1993).? El Hierro: Punta de Arenas Blancas (Rodríguez & Barrientos, 1993). Description Maximum length:, 4.5 mm;, 5.5 mm. Body outline as in Fig. 3A. Colour brown mottled with the usual pale muscle spots. Back smooth with numerous cordiform scale-setae (Fig. 3B); no visible gland pores along the lateral margin of pereonites; noduli laterales on the pereonites similar to those present in H. couchii, i.e. 3 or 4 lines per side (Fig. 3D). Cephalon with no frontal line and suprantennal line slightly bent downwards in the middle (Figs 3C, 8E,F); eyes with ommatidia. Pleon much narrower than pereon; pleonites with very small posterior points slightly visible in dorsal view (Fig. 3E). Telson with sinuous sides and broadly rounded apex (Fig. 3E). Antennula (Fig. 3F) with second article much longer than first and third; third article with two apical aesthetascs, one row of several aesthetascs in the middle and two aesthetascs in a more proximal position. Antenna (Fig. 3G) reaching the rear margin of pereonite 5; flagellum as long as fifth article of peduncle; first flagellar article slightly longer than second and third; three aesthetascs on second flagellar article and four on third. Mandible with molar penicil consisting of many setae and 2+1 free penicils in the left mandible (Fig. 4A) and 1+1 in the right mandible (Fig. 4B). Outer branch of maxillule with 6 (5 cleft)+5 teeth and a slender stalk among the outer group of teeth; inner branch with two long narrow penicils (Fig. 4C). Maxilla with bilobate apex, two setae between the lobes and setose distal part (Fig. 4D). Maxilliped endite setose with a large penicil near the inner corner; palp of three articles, first article with two strong setae (Fig. 4E). Pereopods with a very long ungual seta apically enlarged and a small dactylar seta (Fig. 4F). Uropod short, with exopod almost three times as long as endopod; protopod and exopod grooved on outer margin; insertion of endopod distinctly proximal to that of exopod (Fig. 3E). Male: Pereopod 1 (Fig. 4F) and, to a lesser extent, 2 with carpus and merus distinctly flattened, enlarged and covered with numerous short scales. Pereopod 7 (Fig. 5A) with no distinct sexual modifications, ischium with

8 Taiti & López 50 Figure 4. Halophiloscia rodriguezi n. sp. from Tenerife, Playa de Lima, paratype : A, left mandible; B, right mandible; C, maxillule; D, maxilla; E, maxilliped; F, pereopod 1. straight sternal margin. Genital papilla (Fig. 5B) with a quadrangular ventral shield apically rounded and genital orifices opening up at the end of two long triangular lobes. Pleopod 1 (Fig. 5B) exopod with a short triangular posterior point and deeply sinuous outer M. Zimmer, F. Charfi Cheikhrouha & S. Taiti (Eds)

9 Halophilosciidae from the Canary Islands 51 margin with one to six short setae; endopod with thickset distal part with almost parallel sides, a long pointed process at apex, some setae subapically and a large lobe on the inner margin of the distal part. Pleopod 2 (Fig. 5C) exopod with a long narrow distal part bent outwards; endopod distinctly longer than exopod with apical part narrow and without any lobes. Pleopod 3-5 exopods as in Fig. 5D- F. Etymology The new species is named after Dr. R. Rodríguez (Gran Canaria), who has greatly contributed to the knowledge of the Oniscidea from the Canary Islands. Figure 5. Halophiloscia rodriguezi n. sp. from Tenerife, Playa de Lima, paratype : A, pereopod 7; B, pleopod 1 and genital papilla; C, pleopod 2; D, pleopod 3 exopod; E, pleopod 4 exopod; F, pleopod 5 exopod.

10 52 Taiti & López Figure 6. Halophiloscia microphthalma n. sp. from La Palma, Cueva de la Fajanita, paratype : A, adult specimen, dorsal; B, dorsal scale-seta; C, cephalon, frontal; D, pleonites 4, 5, telson and uropods; E, antennula; F, antenna; G, pereopod 1.

11 Halophilosciidae from the Canary Islands 53 Remarks Halophiloscia rodriguezi is morphologically close to H. couchii and H. canariensis. It is distinguished from H. couchii by the smaller size (maximum length of female 5.5 mm vs mm), eye with a smaller number of ommatidia (13-15 vs , compare Fig. 8E,F and Fig. 8A,B), antennula with two rows of aesthetascs instead of three (compare Fig. 3F and Fig. 53A1 in Schmidt, 2003), relatively shorter uropodal exopods (compare Fig. 3E and Fig.1A), male pleopod 1 exopod with the triangular posterior point less developed and with a smaller number of setae on the outer margin, endopod with a more developed lobe on the outer margin of the distal part and lacking the apical quadrangular lobe (compare Fig. 5B and Fig. 1B), and pleopod 2 endopod narrow and lacking the subapical rounded lobe typical of H. couchii (compare Fig. 5C and Fig. 1C). It differs from H. canariensis in the relatively longer uropodal exopods (compare Fig. 3E and Fig. 2A), the male pleopod 1 endopod with distal part straight instead of bent outwards and the modifications of its apex (compare Fig. 5B and Fig.2B), and the male pleopod 2 exopod with distal point narrower and less bent outwards (compare Fig. 5C and Fig. 2C). The validity of the new species is also confirmed by the high level of nucleotide sequence divergence for COI from both H. couchii and H. canariensis (M. Arnedo, pers. comm.). Halophiloscia rodriguezi seems to be endemic to the western Canary Islands from Tenerife to El Hierro. According to Rodríguez & Barrientos (1993), only H. couchii was present along the coasts of the western islands. Most probably their records from the littoral sites of La Palma, La Gomera and El Hierro refer to H. rodriguezi. Halophiloscia microphthalma n. sp. Figs 6, 7, 8G,H, 9 Halophiloscia couchi; Rodríguez & Barrientos, 1993: 185 (partim: cave populations from La Palma), figs 1, 2; García et al., 1995: 674, 675; García & Govantes, 1996: 134; González & Govantes, 1996: 138. Material examined Holoptype: La Palma: 1, Cueva de la Fajanita, N W, 12.VII.2004, leg. GIET (MZUF). Paratypes: La Palma: 4, 6, Cueva de la Fajanita, N W, 12.VII.2004, leg. GIET (MZUF); 2, same locality, 29.III.2000, leg. GIET (DZUL); 1, 2, same locality, 23.XI.2000, leg. GIET (MZUF); 1, same locality, 30.XI.2000, leg. GIET (MZUF); 5, 10, 1 juv., same locality, pitfall trap, 22.II.2005, leg. P. Oromí (DZUL); 2, 4, same data (MCNT); 2, 1, same locality, 12.VII.2004, leg. GIET (MZUF); 3, 5, Cueva de los Palmeros, 28º30 27 N 17º51 34 W, 14.XI.2005, pitfall trap, leg. P. Oromí (MZUF); 1, Cueva del Ratón, 28º27 46 N 17º50 47 W, 7.V.2000, leg. GIET (MZUF); 2, 2, same locality, 7.XII.2000, leg. GIET (DZUL); 1, same locality, 21.II.2005, leg. C. Villacorta (MZUF); 1, Cueva Machacadora, 28º30 30 N 17º49 42 W, 24.II.2005, leg. C. Villacorta (MZUF). Additional material: La Palma: 8, 2, Cueva Virgen de Fátima, N W, 17.X.1995, leg. R. García Becerra; 5, 3, Cueva los Arreboles, Fuencaliente, 5.XI.1995, leg. R. García Becerra; 3, 5, Cueva de Gallegos, Barlovento, N W, 21.XII.1990, leg. R. García Becerra; 3, Cueva de Salvatierra, Garafía, 28. IX.1995, leg. R. García Becerra; 1, Cueva de Nagy, Breña Baja, N W, 1.II.1997, leg. R. García Becerra; 1, Cueva de los Palmeros, 28º30 27 N 17º51 34 W, 5.X.1996, leg. R. García Becerra. Previous records As H. couchii: La Palma: Cueva Tacande; Cueva de Los Palmeros (Rodríguez & Barrientos, 1993); Cueva A del Salto de Tigalate (García et al. 1995); Cueva Virgen de Fátima (García & Govantes, 1996); Cueva de la Mamona (González & Govantes, 1996). Description Maximum length:, 7.0 mm;, 7.5 mm. Body outline as in Fig. 6A. Colourless body. Back smooth with numerous long triangular scale-setae (Fig. 6B); no visible gland pores

12 54 Taiti & López Figure 7. Halophiloscia microphthalma n. sp. from La Palma, Cueva de la Fajanita, paratype : A, pereopod 7; B, pleopod 1 and genital papilla; C, pleopod 2; D, pleopod 3 exopod; E, pleopod 4 exopod; F, pleopod 5 exopod. along the lateral margin of pereonites; noduli laterales as in H. rodriguezi. Cephalon with no frontal line and suprantennal line slightly bent downwards in the middle (Figs 6C, 8G); eyes

13 Halophilosciidae from the Canary Islands 55 reduced with 4-5 ommatidia (Fig. 8H) visible as small spots of dark pigment. Pleon much narrower than pereon; pleonites with small posterior points visible in dorsal view (Fig. 6D). Telson with straight sides and broadly rounded apex (Fig. 6D). Antennula (Fig. 6E) with second article slightly longer than first and distinctly longer than third; third article with two long slender apical aesthetascs, one row of several aesthetascs in the middle and one row close to the proximal margin. Antenna (Fig. 6F) very long and slender, reaching the rear margin of pereonite 7; flagellum slightly longer than fifth article of peduncle; first flagellar article more than twice as long as second, second and third article subequal in length; a row of five aesthetascs on second flagellar article and one of four aesthetascs on third. Buccal pieces as in H. rodriguezi. Pereopods very elongated, with an ungual seta not apically enlarged and a small dactylar seta (Fig. 6G). Uropod very long, with exopod almost four times as long as endopod; protopod and exopod grooved on outer margin; insertion of endopod distinctly proximal to that of exopod. Male: Pereopod 1 (Fig. 6 G) and, to a lesser extent, 2 with carpus and merus distinctly flattened, enlarged and covered with numerous short scales. Pereopod 7 (Fig. 7A) with no distinct sexual modifications, ischium with straight sternal margin. Genital papilla (Fig. 7B) with a short ovoidal ventral shield and genital orifices opening up at the end of two long triangular lobes. Pleopod 1 (Fig. 7B) exopod with a short triangular posterior point and deeply sinuous outer margin with some short setae; endopod with distal part narrow, with almost parallel sides, a long pointed process at apex, some setae subapically and a small lobe on the inner margin of the distal part. Pleopod 2 (Fig. 7C) exopod with a long narrow distal part bent outwards; endopod distinctly longer than exopod with apical part narrow and without any lobe. Pleopod 3-5 exopods as in Fig. 7D-F. Etymology From the Greek mikrós = small + ophthalmós = eye. The name refers to the reduced eye with only 4-5 ommatidia. Figure 8. Halophiloscia couchii from Gran Canaria, mouth of Barranco Guayadeque, : A, cephalon, lateral; B, eye. Halophiloscia canariensis from Lanzarote, Jameos del Agua, : C, cephalon, lateral; D, eye. Halophiloscia rodriguezi n. sp. from Tenerife, Playa de Lima, paratype : E, cephalon, lateral; F, eye. Halophiloscia microphthalma n. sp. from La Palma, Cueva de la Fajanita, paratype : G, cephalon, lateral; H, eye. Remarks Halophiloscia microphthalma is readily distinguishable from all the other species of Halophiloscia by its troglomorphic characters such as colourless body, very reduced eyes and elongated antennae, pereopods and uropods, and by the longer and narrower dorsal scalesetae. It also differs from H. couchii by an antennula with two rows of aesthetascs instead of three (compare Fig. 6E and Fig. 53A1 in

14 56 Taiti & López CANARY ISLANDS CHINIJO ARCHIPELAGO LA PALMA LANZAROTE LOBOS TENERIFE FUERTEVENTURA LA GOMERA GRAN CANARIA EL HIERRO 100 Km Halophiloscia couchii Halophiloscia canariensis Halophiloscia rodriguezi Halophiloscia microphthalma Stenophiloscia glarearum Littorophiloscia culebrae Figure 9. Recorded distribution of Halophilosciidae species on the Canary Islands (doubtful records not included). Schmidt, 2003), male pleopod 1 exopod with the triangular posterior point less developed and endopod with narrower distal part, longer pointed apical process and lacking the apical quadrangular lobe (compare Fig. 7B and Fig. 1B), and male pleopod 2 endopod narrow and lacking the subapical rounded lobe. It also differs from H. canariensis in the larger size (maximum length 7.5 mm vs. 5.5 mm), the male pleopod 1 endopod with distal part straight instead of bent outwards and the modifications of its apex (compare Fig. 7B and Fig. 2B), and the male pleopod 2 exopod with distal point narrower and less bent outwards (compare Fig. 7C and Fig. 2C). In the structure of the male pleopod 2, Halophiloscia microphthalma is morphologically close to H. rodriguezi from which it differs in the above mentioned troglomorphic characters and also in the narrower distal part of the male pleopod 1 endopod with a longer pointed apical process. The high level of nucleotide sequence divergence for COI from H. couchii, H. canariensis and H. rodriguezi (M. Arnedo, pers. comm.) confirms that the cave populations from La Palma belong to a distinct species and not to H. couchii as supposed by Rodríguez & Barrientos (1993). Genus Stenophiloscia Verhoeff, 1908 Stenophiloscia glarearum Verhoeff, 1908 Fig. 9 Stenophiloscia zosterae; Rodríguez & Barrientos, 1993: 185, figs Stenophiloscia glarearum; Schmalfuss, 2003: 283; Rodríguez et al., 2004: 175. Material examined Lanzarote: 2, 2, Playa de la Madera, Timanfaya, N W, 28.III.2004, leg. GIET (MZUF). Gran Canaria: 2, Baja de los Matos, Pozo Izquierdo, N W, 2.XI.2007, leg. H. López & H. Morales (DZUL). Previous records Lanzarote: Punta del Pasito; Playa de las Salinas (Rodríguez & Barrientos, 1993). La Gomera: Puntallana (Rodríguez & Barrientos, 1993).

15 Halophilosciidae from the Canary Islands 57 Distribution Southern England, Canary Islands, eastern Spain, Balearic Islands, south-eastern France, Italy, Malta, Croatia, and Ionian coasts of Greece (Schmalfuss, 2003). Remarks Stenophiloscia zosterae Verhoeff, 1928 was recognised to be a junior synonym of S. glarearum by Schmalfuss (2003). The specimens from Gran Canaria represent a new island record of this species for the Canary Islands. Genus Littorophiloscia Hatch, 1947 Littorophiloscia culebrae (Moore, 1901) Fig. 9 Littorophiloscia culebrae; Rodríguez & Barrientos, 1993: 190, figs 23-26; Schmalfuss, 2003: 151; Rodríguez et al., 2004: 175. Material examined Gran Canaria: many and, Faro de Maspalomas, ,50 N ,50 W, 2.I.1993, leg. S. Taiti & C. Manicastri (MZUF); many and, mouth of Barranco Draguillo, Ojos de Garza, N W, 2.XI.2007, leg. H. López & H. Morales (MZUF); 2, 4, same data (DZUL); many and, mouth of Barranco Las Palmas, Juan Grande, N W and N W, 29.XII.2007, leg. H. López (MZUF). Tenerife: 2, 4, Playa de Samarines, Candelaria, N W, 11.III.2004, leg. GIET (MZUF); 5, 8, Playa de Lima, Güímar, N W, 17.XII.2004, leg. H. López & H. Morales (MZUF); many and, Playa del Médano, N W, 11.XI.2007, leg. H. López & S. de la Cruz (MZUF); 2, 4, same data (DZUL). Previous records Tenerife: Playa del Médano (Rodríguez & Barrientos, 1993). Distribution Littorophiloscia culebrae has a wide distribution along the tropical coasts. It has been recorded from the Hawaiian Islands, Socotra Archipelago, Madagascar, Angola, Canary Islands, Florida, Cuba, Puerto Rico and the Virgin Islands. Discussion Six species in three genera of Halophilosciidae are now recorded from the Canary Islands: Halophiloscia couchii from Tenerife and Gran Canaria, H. canariensis from the eastern islands (Chinijo Archipelago, Lanzarote, Lobos, Fuerteventura and Gran Canaria), H. rodriguezi n. sp. from the western islands (Tenerife, La Palma, La Gomera and El Hierro), H. microphthalma n. sp. from some caves on La Palma, Stenophiloscia glarearum from Lanzarote, Gran Canaria and La Gomera, and Littorophiloscia culebrae from Gran Canaria and Tenerife. With the exception of Halophiloscia couchii, which has a wide distribution along the Mediterranean and Atlantic coasts of Europe and northern Africa, the other three species of Halophiloscia are endemic to the Canary Archipelago. Our results provide strong evidence for the colonization of the Canary Islands by independent lineages of Halophiloscia. One lineage corresponds to the littoral H. rodriguezi, occurring on the western islands which are the youngest of the archipelago, from 1.12 Myr of El Hierro to 11.6 Myr of Tenerife (Carracedo et al., 2002). The cave-dwelling troglomorphic H. microphthalma from La Palma appears morphologically close and related to H. rodriguezi. This is the second documented case of a cavernicolous species related to a littoral species in the family Halophilosciidae: the first case involved two species of Littorophiloscia (L. hawaiiensis Taiti & Ferrara, 1986, littoral, and Littorophiloscia sp., cavernicolous) from Hawaii Island (Rivera et al., 2002). Another lineage gave rise to H. canariensis in the eastern islands which are the oldest of the archipelago, from 14.5 Myr of Gran Canaria to 20.6 Myr of Fuerteventura (Carracedo et al., 2002). Finally, H. couchii from Gran Canaria and Tenerife is most likely a recent introduction to the archipelago from Atlantic populations of this species. On both Gran Canaria and Tenerife two littoral species of Halophiloscia are present: H. couchii and H. canariensis on the former, and H. couchii

16 58 Taiti & López and H. rodriguezi on the latter. A more detailed discussion on the phylogeny, phylogeography and colonization of Halophiloscia species in the Canary Islands will be presented in a separate paper on the basis of molecular data. Acknowledgments We appreciate the assistance of many people in obtaining material, especially: Pedro Oromí, Salvador de la Cruz, Elena Morales, Nuria Macías, Antonio Pérez (all from La Laguna, Tenerife), R. García (La Palma), and Natália Dias (Faro, Portugal). Dr. C. Giordano (CeME-CNR, Florence) and Dr. G. A. Gruber (ISE- CNR, Florence) are particularly acknowledged for helping us with SEM and drawings, respectively. Research funded by ISE-CNR, Florence, MEDCORE Project contract UE-INCOMED no. CA3 CT , Proyecto Atlantico INTERREG-IIIB and Project 2005SGR00045 from the Catalan Autonomous Government. References Arcangeli A La fauna isopodologica terrestre degli arcipelaghi di Madera e delle Canarie: la sua importanza per la sistematica e la biogeografia. Mem Estud Mus Zool Univ Coimbra 255: Carracedo JC, Pérez FJ, Ancochea E, Meco J, Hernán F, Cubas CR, Casillas R, Rodriguez E, Ahijando A Cenozoic volcanism II: The Canary Islands. In: Gibbons W, Moreno T (eds). The Geology of Spain The Geological Society of London. Dalens H Sur une Halophiloscia nouvelle des Iles Canaries: (H. Halophiloscia) canariensis n. sp. (Isopoda, Oniscoïdea). Bull Soc Hist Nat Toulouse 109: Dollfus A Voyage de M. Ch. Alluaud aux Îles Canaries (Novembre Juin 1890). Isopodes terrestres. Mém Soc Zool France 6: García R, González AJ, Govantes F Distribución de artrópodos en las cavidades A y B del Sistema de tubos lávicos del Salto de Tigalate en la isla de La Palma (Islas Canarias). Vieraea 24: García R, Govantes F La Cueva Virgen de Fátima, un tubo volcánico en la colada histórica del Volcán de San Juan (La Palma). In: Oromí P (ed.). 7th International Symposium on Vulcanospeleology Los Libros de la Frontera: St. Climent de Llobregat. González AJ, Govantes F La Cueva de la Mamona, primera descripción de un tubo volcánico del nordeste de La Palma (Islas Canarias). In: Oromí P (ed.). 7th International Symposium on Vulcanospeleology Los Libros de la Frontera: St. Climent de Llobregat. Hoese B. 1984a. Checkliste der terrestrischen Isopoden der Kanarischen Inseln (Crustacea: Isopoda: Oniscoidea). Cour Forsch-Inst Senckenberg 71: Hoese B. 1984b. Ein Beitrag zur Tiergeographie der terrestrischen Isopoden der Kanarischen Inseln (Crustacea: Isopoda: Oniscoidea). Cour Forsch-Inst Senckenberg 71: Leistikow A Phylogeny and biogeography of South American Crinocheta, traditionally placed in the family Philosciidae (Crustacea: Isopoda: Oniscidea). Org Div Evol 1 (E-Suppl 4): Macías N, Pérez AJ, López H, Oromí P Fauna de artrópodos de Montaña Clara (Islas Canarias) III: arácnidos, miriápodos y crustáceos terrestres. Rev Acad Canar Cienc 15: Radu VG Crustacea, Ordinul Isopoda, Subordinul Oniscoidea, Crinochaeta. Fauna Rep Soc România 4: Rivera MAJ, Howarth FG, Taiti S, Roderick GK Evolution in Hawaiian cave-adapted isopods (Oniscidea: Philosciidae): vicariant speciation or adaptive shifts? Mol Phylogenet Evol 25: 1-9. Rodríguez R, Barrientos J Las familias Halophilosciidae y Philosciidae en el Archipiélago Canario (Crustacea: Isopoda: Oniscidea). Bol Asoc Esp Ent 17: Rodríguez R, Oromí P, Zurita N Orden Isopoda. In: Izquierdo Zamora I, Martín Esquivel JL, Zurita Pérez N, Arechavaleta Hernández M (eds). Lista de especies silvestres de Canarias (hongos, plantas y animales terrestres) Litografia A. Romero SL: Tenerife. Schmalfuss H Eco-morphological strategies in terrestrial isopods. Symp Zool Soc Lond 53: Schmalfuss H World catalog of terrestrial isopods (Isopoda: Oniscidea). Stuttg Beitr Natkde (A) 654: Schmidt C Contribution to the phylogenetic system of the Crinocheta (Crustacea, Isopoda). Part 2 (Oniscoidea to Armadillidiidae). Mitt Mus Natkd Berl, Zool Reihe 79: Taiti S, Ferrara F Taxonomic revision of the genus Littorophiloscia Hatch, 1947 (Crustacea, Isopoda, Oniscidea) with description of six new species. J Nat Hist 20: Vandel A Étude des isopodes terrestres recueillis aux Îles Canaries par J. Mateu en Mars-Avril Mém Mus Natn Hist Nat Paris (A) 8: Vandel A La faune terrestre de l île de Sainte- Hélène. Quatrième Partie. 1. Isopodes terrestres. Ann Mus Afr Cent, Sér 8vo (Sci Zool) 220: , pl. II.