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Terrestrial Isopoda (Crustacea, Oniscidea) from the coasts of Costa Rica, with descriptions of three new species/Isopoda terrestre (Crustacea, Oniscidea) de las costas de Costa Rica, con descripciones de tres nuevas especies.

Up to date, the diversity of Costa Rican terrestrial isopods is poorly known. Only 25 species in 16 genera and 10 families are presently recorded, numbers that are certainly very small for a tropical country like Costa Rica. Many records are very old, mainly dated in the first half of 1900 (Richardson, 1910, 1913; Arcangeli, 1927, 1930, 1957; Van Name, 1936), while only the families Philosciidae and Scleropactidae have been recently revised by Leistikow (1997a, 1997b, 1998, 2000a, b, 2001) and Schmidt (2007). However, all the records come from sporadic collections. Only few records are known for most of the forested areas and only two species (Ligia baudiniana Milne Edwards, 1840 and Tylos wegeneri Vandel, 1952) were reported from the coasts of the country.

This paper deals with recent collections of Oniscidea from both the Pacific and Caribbean coastal areas of Costa Rica, and includes the descriptions of three new species and two new records for the country.

MATERIALS AND METHODS

The specimens included in this paper have been collected in November 2015 on both sandy and rocky coasts of the Pacific side of Costa Rica. Also included are littoral terrestrial isopods deposited in the collection of the Museo de Zoologia, Universidad de Costa Rica (MZUCR), San Jose. Specimens were collected by hand and stored in 75 % ethanol. The geographic co-ordinates of the locations were taken using WGS84 datum. Identifications are based on morphological characters. For each new species the material examined, description, etymology and remarks are given. For each species already recorded from Costa Rica the bibliographic references, material examined, distribution and remarks (when necessary) are included. Some poorly known species have been illustrated to facilitate future recognition. The taxa are illustrated with figures prepared with the aid of a camera lucida mounted on Wild M5 and M20 microscopes and digitally drawn using the method by Montesanto (2015, 2016). For some species pictures were taken with a Scanning Electron Microscope Hitachi S-3700N.

Abbreviations: BM = The Natural History Museum, London, MZUCR = Museo de Zoologia, Escuela de Biologia, Universidad de Costa Rica, San Jose, Costa Rica, MZUF = Museo di Storia Naturale dell'Universita, Sezione di Zoologia "La Specola", Florence, Italy.

SYSTEMATIC ACCOUNT

Family Ligiidae Leach, 1814

Genus Ligia Fabricius, 1798

Ligia baudiniana Milne Edwards, 1840

Ligia (Hirtiligia) baudiniana; Schultz, 1974: 167, figs 122-3.

Ligia baudiniana; Leistikow, 1997a: 1417, figs. 2-7; Leistikow & Wagele, 1999: 2; Schmalfuss, 2003: 141.

Material examined: 1 [male], 2 [female][female] (MZUF 9684), Playa Pita, S of Tarcoles, Puntarenas, 9[degrees]44'32.9" N and 84[degrees]37'53.0" W, under logs on a sandy beach, 27.XI.2015, leg. S. Taiti, R. Vargas & J.A. Vargas; 1 [male] (MZUCR 2580-02), Isla San Jose, Islas Murcielago, Guanacaste, 21.IV.2003; 13 [male][male], 18 [female][female] (MZUCR 2865-01), Reserva de Cabo Blanco, sector San Miguel, 10.VII. 1994, leg. L.A. Mena; 1 [female] (MZUCR 3154-03), Playa Blanca, Punta Morales, sandy beach, 29.XI. 2011, leg. J.A.Vargas & J. Sibaja, det. R. Vargas; 1 [male] (MZUCR 1870), Isla Cabuya, Cabo Blanco, 16.XII.1992, leg. L. Mena; 6 [female][female] (MZUCR 2069-01), Manuel Antonio, Quepos, Aguirre, Puntarenas, 3.IX.1984, leg. B. Morera, det. A. Leistikow; 1 [male], 1[female] (MZUCR 2070-01), same locality, 2.IX. 1984, leg. B. Morera, det. A. Leistikow; 3 [male][male] (MZUCR 2076-01), Samara, Guanacaste, date?, leg. O. Breedy.

Costa Rican records: Puntarenas (Schultz, 1974). Puntarenas Prov., Corcovado, Punta Salsipuedes; Limon Prov., Punta Cahuita; Isla San Lucas (Leistikow, 1997a).

Distribution: Atlantic and Pacific shores of the Americas from Florida to Brazil and from California to Ecuador, including Galapagos Islands (Schmalfuss, 2003).

Remarks: At present, this is the only species of Ligia recorded from both coasts of Costa Rica. The species has been fully redescribed and illustrated by Leistikow (1997a).

Family Tylidae Milne Edwards, 1840

Genus Tylos Audouin, 1826

Tylos wegeneri Vandel, 1952

Tylos wegeneri; Schultz, 1983: 682; Leistikow & Wagele, 1999: 4; Schmalfuss & Vergara, 2000: 10; Schmalfuss, 2003: 324.

Costa Rican records: Puntarenas (Schultz, 1983).

Distribution: USA: Florida; Costa Rica: Pacific Coast; Lesser Antilles: Saint Martin; Trinidad; Tobago; Venezuela (Schmalfuss, 2003).

Remarks: Tylos wegeneri was recorded from Puntarenas by Schultz (1983). This is the only record of this species for the Pacific coast, while the species seems to be widespread in the Caribbean Sea. However, no records from the Caribbean coast of Costa Rica are known. For a description and figures of this species see Vandel (1952) and Schultz (1970).

Tylos niveus Budde-Lund, 1885 Figs. 1, 2

Material examined: 3 [male][male] (MZUCR 2867-02), Boca Coronado, manglar TerrabaSierpe, 3.IX.2008, leg. L. Rolier Lara.

Distribution: Florida; Bahamas; Bermudas; Cuba; Tobago; Bonaire; Dominica; Curazao; Virgin Islands; Puerto Rico; Mexico; Belize; Venezuela; Colombia; Brazil (Schmalfuss, 2003; Carpio-Diaz, Lopez-Orozco, Herrera-Medina, Navas-S., & Bermudez, 2016). New record for Costa Rica.

Remarks: The main characters of this species are illustrated in Figs. 1 and 2 to confirm its identification and facilitate future recognition. For a complete list of synonyms of this species see Schmalfuss & Vergara (2000). This is the first record for the Pacific coast.

Incertae sedis Genus Buchnerillo Verhoeff, 1942 Buchnerillo neotropicalis n. sp. Figs. 3-6

Holotype: [male] (MZUF 9685), Playa Pita, [male] of Tarcoles, Puntarenas, 9[degrees]44'32.9" N and 84[degrees]37'53.0" W, sandy beach, under logs, 27.XI.2015, leg. S. Taiti, J.A.Vargas & R. Vargas.

Paratypes: 6 [female][female] (MZUF 9685), 4 [female][female] (MZUCR-3529-04), same data as holotype.

Additional material: 2 [female][female] used for scanning microscope analysis, same data as holotype.

Description: Maximum size: [male] 15 x 0.8 mm; [female] 1.9 x 1 mm. Colour pale. Animals able to roll up into a perfect ball (Fig. 3A, B). Dorsal body surface covered with tubercles disposed as follows: five rows on cephalon; three rows on pereonite 1; two rows on pereonites 2-7; two rows on pleonites 3 and 4; one row on pleonite 5; four tubercles on telson; each dorsal tubercle apically with one scaleseta (Fig. 3C, D). Cephalon (Fig. 4A, B) with frontal shield grooved, upper margin regularly convex, lower margin sinuous on both sides; eye consisting of 4 ommatidia. Pereonite 1 (Fig. 4B) with semicircular hollow on anterior margin in which ocular lobe fits; postero-lateral corner with schisma, inner lobe rounded and distinctly protruting backwards; posterior margin straight. Pereonites 2-4 with triangular epimera. Pereonites 5-7 with quadrangular epimera. Pleonites 1 and 2 not visible in dorsal view, pleonite 3 with no visible epimera, pleonite 4 twice as long as pleonite 5 (Fig. 4C). Telson (Fig. 4D) semicircular. Antennula (Fig. 5A) of two articles; first article about twice as long as second; second article with two large petaliform aesthetascs at apex. Antenna (Fig. 5B) short and thickset; flagellum of 3 articles, second article bearing 2 aesthetascs. Mandibles (Fig. 5C, D) with molar penicil semidichotomized, left mandible with 2 + 1 free penicils; right mandible with 1 + 1 free penicils. Maxillula (Fig. 5E) inner branch with a few setae at apex but no penicils; outer branch with 4 + 4 teeth, all apically entire, and stalk among outer group of teeth. Maxilla (Fig. 5F) with triangular distal part bearing thick setae. Maxilliped (Fig. 5G) endite triangular with two subapical stout setae and apical penicil; palp with two setae on first article. Pereopods (Fig. 6A) with one large trifid seta on carpus, long setose dactylar seta and flagelliform ungual seta; pereopod 7 with water conducting system on basis. Uropod (Fig. 5H) protopod quadrangular; endopod much longer than exopod, both with very long apical setae.

Male: Pereopod 7 (Fig. 6B) with no distinct modifications; ischium sternal margin straight. Pleopod 1 (Fig. 6C) exopod small, ovoidal; endopod very elongated with distal part triangular, straight. Pleopod 2 (Fig. 6D) exopod triangular with one distal hairy seta and some short thin setae on inner and outer margin; endopod of two articles, distal article flagelliform, about 5 times longer than first. Ppleopod 3-5 exopods as in Fig. 6E-G.

Etymology: The name of the species refers to the localities of collection of the species, the Neotropical Region.

Remarks: The new species is included in Buchnerillo since it shows all the characters of the genus: small size; animal able to toll up into a perfect ball; endoantennal conglobation; dorsal surface tuberculated; cephalon with a wide frontal shield; pleonite 3 with epimera reduced; telson semicircular covering the uropods in dorsal view; antenna short and stout with a flagellum of three articles; male pleopod 2 with distal article flagelliform. Up to date, only two species of Buchnerillo are known: B. litoralis Verhoeff, 1942 and B. oceanicus Ferrara, 1974. The former is known from the shores of the Mediterranean Sea and Madeira (Schmalfuss, 2003); a record of a female specimen from Florida Keys (Paoletti & Stinner, 1989) is very doubtful and the identification needs confirmation. The latter is presently known from Somalia (Ferrara, 1974) and the Maldives (Taiti, 2014). Buchnerillo neotropicalis n. sp. differs from both species in the presence of a schisma on the pereonite 1 with inner lobe distinctly protruding backwards, whereas in B. litoralis and B. oceanicus a small rounded ventral lobe is present at the postero-lateral corners, not protruding backwards. It also differs from B. litoralis (see redescription in Vandel, 1960) in having the cephalon with frontal shield grooved, larger eyes (4 ommatidia instead of 1-2), dorsal tubercles more prominent and male pleopod 1 endopod with distal part thicker and straight; from B. oceanicus in the frontal shield with lower margin sinuous on both sides instead of regularly curved.

The systematic position of genus Buchnerillo is still uncertain. It was included by Vandel (1960) in the section Synocheta and family Buddelundiellidae (now a subfamily of Trichoniscidae). Tabacaru (1993) recognized that the genus could not belong to the Synocheta and Schmalfuss (2003) included it into the higher Oniscidea (the Crinocheta) and hypothesised that the genus might belong to the family Detonidae, close to the genus Armadilloniscus. According to the maxillular endite bearing only some apical setae without penicils, it might also be related to the family Olibrinidae. However, since no safe conclusion can be reached with morphological characters, we still maintain the genus as incertae sedis as proposed by Taiti & Ferrara (1991). A molecular analysis might be useful to clarify the family placement of Buchnerillo.

Family Detonidae Budde-Lund, 1904

Genus Armadilloniscus Uljanin, 1875

Armadilloniscus cf. caraibicus Paoletti & Stinner, 1989 Fig. 7

Material examined: 1 [female] (MZUF 9687), 1 [female] used for scanning microscope analysis, Playa Pita, S of Tarcoles, Puntarenas, 9[degrees]44'32.9" N and 84[degrees]37'53.0" W, beach under logs, 27.XI.2015, leg. S. Taiti, J.A. Vargas & R. Vargas.

Distribution: Venezuela (Paoletti & Stinner (1989). New record for Costa Rica.

Remarks: The two female specimens here examined are morphologically very similar to Armadilloniscus caraibicus described by Paoletti & Stinner (1989) for the Caribbean coast of Venezuela. The main characters of the Costa Rican specimens are shown in Fig. 7. They show the same disposition of dorsal tubercles as those from Venezuela (see Figs. 10 and 11 in the original description by Paoletti & Stinner, 1989 and Figs. 32 and 33 in Schmidt, 2002) but they are less developed. Since we have examined only 2 females, we only tentatively identify them as A. cf. caraibicus.

Family Philosciidae Kinahan, 1957

Genus Hawaiioscia Schultz, 1973

Hawaiioscia nicoyaensis n. sp. Figs. 8-11

Holotype: [male] (MZUCR-3529-05), Playa Pita, south of Tarcoles, Puntarenas, 9[degrees]44'32.9" N and 84[degrees]37'53.0" W, beach under logs, 27.XI.2015, leg. S. Taiti, J.A. Vargas & R. Vargas.

Paratypes: 5 [male][male], 8 [female][female] (MZUCR-3529-06), 5 [male][male], 15 [female][female], 12 juvs (MZUF 9688), same data as holotype.

Additional material examined: 1 [male], 1 [female], used for scanning microscope analysis, same data as holotype.

Description: Maximum size: [male], 5.8 x 2.5 mm; [female], 6 x 3 mm. Dorsum dark brown with the usual muscle spots; a large pale spot at base of pereon epimera. Body flat, ovoidal, with pleon narrower than pereon (Fig. 8A, B). Dorsal body surface finely granulated with small triangular scale-setae (Fig. 9A). Pereonites with no sulcus marginalis, gland pores absent. Noduli laterales small (Fig. 9B) but clearly visible, inserted on small tubercles and disposed as follows: two on cephalic vertex, one per side on pereonites 1-6 with that on fourth pereonite much more distant from lateral margin of segment, and two per side on pereonite 7 (Fig. 9C-E). Cephalon (Figs. 8C, 9E) with short triangular lateral lobes not protruding frontwards compared with obtuse middle lobe; frontal and supra-antennal lines absent; eyes consisting of 19-20 ommatidia. Pleon epimera reduced but with distinct posterior points (Fig. 8A, D). Telson (Fig. 8D) triangular, about twice as wide as long, with slightly concave sides and broadly rounded apex. Antennula (Fig. 9F) of 3 articles, second article slightly shorter than first and third; third article bearing two rows of 3 and 5 aesthetascs each, and 2 apical aesthetascs. Antenna (Fig. 9G) long and thin, reaching back rear margin of pereonite 3; flagellum slightly longer than fifth peduncular article, first flagellar article as long as third, second article shortest; one row of 1-2 and 3 aesthetascs on each second and third article. Mandibles (Fig. 10A, B) with molar penicil semidichotomized, i.e. consisting of 4 setae on a common stem; left mandible with 2+1 and right mandible with 1+1 free penicils. Maxillula (Fig. 10C) outer branch with 4+6 teeth, all simple and stalk among the outer teeth; inner branch with two stout subequal penicils. Maxilla (Fig. 10D) apically setose and bilobate. Maxilliped (Fig. 10E) endite apically setose and bearing large penicil at medial corner, proximal article of palp bearing 2 strong setae. Pereopods with flagelliform dactylar seta apically slightly enlarged; ungual seta flagelliform almost reaching tip of outer claw (Fig. 10F). Pleopodal exopods with no trace of respiratory structures. Uropodal protopod grooved on outer margin; insertion of endopod slightly proximal to that of exopod.

Male: Pereopods 1-4 (Figs. 8B, 10F) merus and carpus with a brush of trifid spines on sternal margin. Pereopod 7 (Figs. 8E, 11A) ischium with sternal margin slightly convex in middle. Genital papilla as in Fig. 11B. Pleopod 1 (Fig. 11C) exopod cordiform, with broadly rounded apex; endopod with thickset distal part, with bilobed apex. Pleopod 2 (Fig. 11D) exopod triangular, with straight outer margin and very convex medial margin, and bearing 4 setae on outer margin near apex. Pleopods 3-5 exopod as in Fig. 11E-G.

Etymology: The name of the species refers to the Gulf of Nicoya, where Playa Pita is located.

Remarks: The new species is included in the genus Hawaiioscia since all the most important characters (number and position of noduli laterales, maxillular teeth, penicil on maxillipedal endite, uropod and shape of male pleopod) correspond to the definition of that genus (see diagnosis in Taiti & Howarth 1997). The genus Hawaiioscia comprises five species from the Hawaiian Islands and Easter Island (Schultz 1973; Taiti & Howarth 1997; Taiti & Wynne, 2015): H. parvituberculata Schultz, 1973 from Maui, H. microphthalma Taiti & Howarth, 1997 from Oahu, H. paeninsulae Taiti & Howarth, 1997 from Molokai, H. rotundata Taiti & Howarth, 1997 from Kauai, and H. rapui Taiti & Wynne, 2015 from Easter Island. The Hawaiian species are all cave-dwelling and show troglomorphic traits, such as body depigmentation and eye reduction, while H. rapui inhabits cave entrances and does not show distict troglomorphic traits. Hawaiioscia nicoyaensis is the only littoral halophilic species known so far. The new species is readily distinguishable from the Hawaiian species by the pigmented body, the eye well developed and in having the molar penicil of the mandible semidichotomized, instead of simple. For this last character, the new species shows closest affinities with H. rapui from which it mainly differs in having larger eyes (19-20 instead of 8 ommatidia) and in the shape of the male pleopods 1 and 2.

Family Platyarthridae Verhoeff, 1949

Genus Trichorhina Budde-Lund, 1908 Trichorhina biocellata n. sp. Figs. 12-16

Holotype: [male] (MZUCR-3529-01), Playa Pita, S of Tarcoles, Puntarenas, 9[degrees]44'32.9" N and 84[degrees]37'53.0" W, beach under logs, 27.XI.2015, leg. S. Taiti, R. Vargas & J.A. Vargas.

Paratypes: 15 [male][male], 25 [female][female], 8 juvs (MZUF 9693), same data as holotype; 14 [male][male], 17 [female][female] (MZUCR-3529-02), same locality and date, leg. R. Vargas; 7 [male][male], 10 [female][female] (MZUCR-3529-03), 8 [male][male], 13 [female][female] (MZUF 9694), Bahia de Caldera, Puntarenas, 9[degrees]55'26.7" N and 84[degrees]42'53.7" W, beach under logs, 27.XI.2015, leg. S. Taiti, R. Vargas & J.A. Vargas; 2 [male][male] (MZUF 9695), Manuel Antonio, Playa Espadilla, Puntarenas, 9[degrees]23'19" N and 84[degrees]08'48" W, beach margin, under dead wood, 18.VII.2013, leg. S. Taiti.

Additional material examined: 1 [male], 1 [female], used for scanning microscope analysis, same data as holotype.

Description: Maximum size: S, 3.5 x 1.5 mm; [female], 3.2 x 1.5 mm. Dorsum pale brown. Body flat, ovoidal, without interruption between pereon and pleon (Fig. 12A, B). Dorsal body surface covered with spatuliform scale-setae (Fig. 12C). Pereonites with no sulcus marginalis, gland pores absent. Noduli laterales small (Fig. 14A, C), inserted on posterior margin of pereonites, one per side on pereonites 1-6 and two per side on pereonite 7. Cephalon (Figs. 12D, 14B) with short rounded lateral lobes not protruding frontwards compared with obtuse middle lobe; frontal and supra-antennal lines absent; eyes consisting of two ommatidia of same size (Fig. 13 A). Pleon epimera well developed, falciform, directed backwards (Fig. 13B). Telson (Fig.13B) triangular, about twice as wide as long, with slightly concave sides and narrowly rounded apex. Antennula (Fig. 14D) of 3 articles, second article much shorter than first and third; third article bearing tuft of about 8 aesthetascs at apex. Antenna (Figs. 13C, 14E) short, reaching back rear margin of pereonite 2; flagellum as long as fifth peduncular article, second flagellar article almost 3 times longer than first, bearing 2 aesthetascs. Mandibles (Fig. 14F, G) with molar penicil semidichotomized, i.e. consisting of 3-4 setae on a common stem; left mandible with 2+1 and right mandible with 1+1 free penicils. Maxillula (Fig. 14H) outer branch with 4+5 (4 slightly cleft) teeth; inner branch with two thin subequal penicils. Maxilla (Fig. 14I) distinctly bilobed, outer lobe slightly smaller than inner lobe, bearing a row of sensilla. Maxilliped (Fig. 15A) endite apically with 2 triangular setae, proximal article of palp bearing 2 strong setae. Pereopods with flagelliform dactylar seta apically slightly enlarged; ungual seta flagelliform reaching tip of outer claw (Fig. 15B). Pleopodal exopods with no trace of respiratory structures. Uropod: insertion of endopod slightly proximal to that of exopod.

Male: Pereopods 1-4 (Figs. 12B, 15B) merus and carpus with a brush of trifid spines on sternal margin. Pereopod 7 (Fig. 15C) ischium with sternal margin slightly sinuous. Genital papilla as in Fig. 16A. Pleopod 1 (Fig. 16B) exopod rounded, about twice as wide as long; endopod with distal part slightly bent medially, with triangular apex. Pleopod 2 (Fig. 16C) exopod triangular, with straight outer margin bearing 3 setae; endopod distinctly longer than exopod. Pleopods 3-5 exopod as in Fig. 16D-F.

Etymology: Latin: bi = double + ocellatus = having eyes. The name refers to the eye consisting of two ocelli of the same size.

Remarks: The new species belongs to the tomentosa-group of Trichorhina characterized by the presence of two noduli laterales per side on the pereonite 7. This group includes with certainty T. tomentosa (Budde-Lund, 1893), T. heterophthalma Lemos de Castro, 1964, both widespread in the tropics, and T. guanophila Souza-Kury, 1993 from Brazil. The new species is readily distinguished from all these species by the eye consisting of two ommatidia of equal size (one in T. tomentosa, two unequal ommatidia in T. heterophthalma, and five in T. guanophila); from T. guanophila also in the male pleopod 1 exopod wider than long.

Two more species of Trichorhina are recorded from Costa Rica by Arcangeli (1930): T. giannellii Arcangeli, 1929, known also from Cuba, and T. marianii Arcangeli, 1930. From their original descriptions no information is present on the number and position of the noduli laterales, so we do not know if they belong to the tomentosa-group. Trichorhina biocellata n. sp. differs from these two species in the eye with only two ommatidia (four or five in T. giannellii and 10 in T. marianii).

DISCUSSION

In the present study seven species of terrestrial isopods are recorded from sandy and rocky shores of both coasts of Costa Rica. Three species (Buchnerillo neotropicalis, Hawaiioscia nicoyaensis and Trichorhina biocellata) are described as new and two species (Tylos niveus and Armadilloniscus cf. caraibicus) represent new records for Costa Rica. The total number of Oniscidean species presently known from Costa Rica increases from 25 to 30 (Table 1).

Six species are strictly littoral, halophilic: L. baudiniana, T. wegeneri, T. niveus, B. neotropicalis n. sp., A. cf. caraibicus, and H. nicoyaensis n. sp. All these species, with the exception of H. nicoyaensis and B. neotropicalis, occur on both the Pacific and Caribbean coasts of Costa Rica or in other countries along the Atlantic coast of the Americas. No morphological differences could be detected from the Pacific and Caribbean populations of these species; only A. cf. caraibicus from Playa Pita on the Pacific coast showed small differences in the less developed dorsal ornamentation, even if of the same type, from the original specimens described from the Caribbean coast of Venezuela. It will be quite interesting to check both the Pacific and Caribbean populations of these five species with molecular markers to see if there is a criptic diversity between them, as revealed in other littoral isopods, e.g. in Excirolana braziliensis Richardson, 1912, Cirolanidae (Hurtado et al., 2016), considering that the isthmus of Panama was definitely closed 2.8 Ma (O'Dea et al., 2016).

An interesting discovery was the new species of Hawaiioscia inhabiting the sandy shore of the Pacific side. The genus was previously known only from troglomorphic cave-dwelling species in the Hawaiian Islands and a troglophilic species from Easter Island. The number of species of Oniscidea from littoral areas of Costa Rica will certainly increase as soon as all the coasts on both sides of the country will be properly investigated.

ACKNOWLEDGEMENTS

We express our sincere thanks to Rita Vargas, Curator of Crustaceans (MZUCR), and to Jeffrey Sibaja for their invaluable help in collecting part of the material here treated. We also thank Rafael Loaiciga for technical assistance with the Scanning Electron Microscope housed at CIEMIC, UCR. S.T. wishes to thank the Centro de Investigacion en Ciencias del Mar y Limnlogia (CIMAR) and the University of Costa Rica for their invitation to give a talk and do research in Costa Rica at the end of 2015. Field trips and SEM access were facilitated by projects UCR-VI-808-B3-113 The benthos of Punta Morales, and UCR-VI-808-B4-117 Ecology of beaches and rocky shores of Costa Rica, both with J. Sibaja as Principal Investigator.

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Richardson, H. (1910). Terrestrial isopods collected in Costa Rica by J. F. Tristan, with descriptions of a new genus and species. Proceedings of the United States National Museum, 39, 93-95.

Richardson, H. (1913). Terrestrial isopods collected in Costa Rica by Mr. Picado, with the description of a new genus and species. Proceedings of the United States National Museum, 44, 337-340.

Schmalfuss, H. (2003). World catalog of terrestrial isopods (Isopoda: Oniscidea). Stuttgarter Beitrage zur Naturkunde, Serie A, 654, 1-341.

Schmalfuss, H., & Vergara, K. (2000). The isopod genus Tylos (Oniscidea: Tylidae) in Chile, with bibliographies of all described species of the genus. Stuttgarter Beitrage zur Naturkunde, Serie A, 612, 1-42.

Schmidt, C. (2002). Contribution to the phylogenetic system of the Crinocheta (Crustacea, Isopoda). Part 1. (Olibrinidae to Scyphacidae s. str.). Mitteilungen aus dem Museum fur Naturkunde in Berlin (Zoologische Reihe), 78, 275-352.

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Schultz, G. A. (1970). A review of the spieces of the genus Tylos Latreille from the New World (Isopoda, Oniscoidea). Crustaceana, 19, 297-305.

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Schultz, G. A. (1983). Two species of Tylos from Chile, with notes on species of Tylos with three flagellar articles (Isopoda, Oniscoidea: Tylidae). Proceedings of the biological Society of Washington, 96, 675-683.

Tabacaru, I. (1993). Sur la classification des Trichoniscidae et la position systematique de Thaumatoniscellus orghidani Tabacaru, 1973 (Crustacea, Isopoda, Oniscidea). Travaux de l'Institut de Speologie <<Emile Racovitza>>, 32, 43-85.

Taiti, S. (2014). The terrestrial Isopoda (Crustacea, Oniscidea) of the Maldives. Tropical Zoology, 27, 9-33.

Taiti, S., & Ferrara, F. (1991). Terrestrial Isopods (Crustacea) from the Hawaiian Islands. Occasional Papers of the Bishop Museum, 31, 202-227.

Taiti, S., & Howarth, F. (1997). Terrestrial isopods (Crustacea, Oniscidea) from Hawaiian caves. Memoires de Biospeologie, 24, 97-118.

Taiti, S., & Wynne, J. J. (2015). The terrestrial Isopoda (Crustacea, Oniscidea) of Rapa Nui (Easter Island), with descriptions of two new species. In S. Taiti, E. Hornung, J. Strus & D. Bouchon (Eds.). Trends in Terrestrial Isopod Biology. ZooKeys, 515, 27-49.

Vandel, A. (1952). Etude des isopodes terrestres recoltes au Venezuela par le Dr. G. Marcuzzi suivie de considerations sur le peuplement du Continent de Gondwana. Memorie del Museo civico di Storia naturale di Verona, 3, 59-203.

Vandel, A. (1960). 64. Isopodes terrestres (premiere partie). Faune de France, 64, 1-416.

Van Name, W. G. (1936). The American land and freshwater isopod Crustacea. Bulletin of the American Museum of Natural History, 71, 1-535.

Stefano Taiti (1,2), Giuseppe Montesanto (3) & Jose A. Vargas (4,5)

(1.) Istituto per lo Studio degli Ecosistemi, Consiglio Nazionale delle Ricerche, Via Madonna del Piano 10, 50019 Sesto Fiorentino (Florence), Italy; stefano.taiti@ise.cnr.it

(2.) Museo di Storia Naturale dell'Universita di Firenze, Sezione di Zoologia "La Specola", Via Romana 17, 50125 Florence, Italy

(3.) Dipartimento di Biologia, Universita degli Studi di Pisa, via A. Volta 4bis, 56126 Pisa, Italy; giuseppe.montesanto@unipi.it

(4.) Centro de Investigacion en Ciencias del Mar y Limnologia (CIMAR), Universidad de Costa Rica, 11501-2060, San Jose, Costa Rica; jose.vargas@ucr.ac.cr

(5.) Escuela de Biologia, Universidad de Costa Rica, 11501-2060, San Jose, Costa Rica.

Received 06-II-2017. Corrected 02-VIII-2017. Accepted 03-I-2018.

Caption: Fig. 1. Tylos niveus Budde-Lund, 1885, [male] from Boca Coronado. A. Adult specimen in lateral view; B. Cephalon, frontal; C. Cephalon, dorsal; D. Pleonites 4, 5 and telson, dorsal; E. Pleon and uropods, ventral;. F. Antennula; G. Antenna.

Caption: Fig. 2. Tylos niveus Budde-Lund, 1885, [male] from Boca Coronado. A. Pereopod 1; B. Pereopod 7; C. Pleopod 2.

Caption: Fig. 3. Buchnerillo neotropicalis n. sp. [female] from Playa Pita. A. Adult specimen in lateral view; B. Adult specimen in ventral view; C. Dorsal tubercles with scale-setae; D. Dorsal scale-seta.

Caption: Fig. 4. Buchnerillo neotropicalis n. sp. [female] from Playa Pita. A. Cephalon, dorsal; B. Cephalon, lateral; C. Pereonite 7, pleon and telson, lateral; D. Pleon and telson, dorsal.

Caption: Fig. 5. Buchnerillo neotropicalis n. sp. [female] from Playa Pita. A. Antennula; B. Antenna; C. Left mandible; D. Right mandible; E. Maxillula; F. Maxilla; G. Maxilliped; H. Uropod.

Caption: Fig. 6. Buchnerillo neotropicalis n. sp. [male] from Playa Pita. A. Pereopod 1; B. Pereopod 7; C. Pleopod 1; D. Pleopod 2; E. Pleopod 3 exopod; F. Pleopod 4 exopod; G. Pleopod 5 exopod.

Caption: Fig. 7. Armadilloniscus cf. caraibicus Paoletti & Stinner, 1989. [female] from Playa Pita. A. Adult specimen in dorsal view; B. Cephalon and pereonites 1-3, dorsal; C. Pleonites 3-5, telson and uropods, dorsal; D. Antennal flagellum.

Caption: Fig. 8. Hawaiioscia nicoyaensis n. sp. specimens from Playa Pita. A. Adult female in dorsal view; B. Adult male in ventral view; C. Cephalon and pereonite 1, dorsal ([female]); D. Pleonite 5, telson and uropods, dorsal ([female]); E. Pleon, ventral ([male]).

Caption: Fig. 9. Hawaiioscia nicoyaensis n. sp. [female] from Playa Pita. A. Dorsal scale-seta; B. Nodulus lateralis; C. B/c and d/c coordinates of noduli laterales; D. Pereonites showing position of noduli laterales; E. Cephalon, frontal; F. Antennula; G. Antenna.

Caption: Fig. 10. Hawaiioscia nicoyaensis n. sp. [male] from Playa Pita. A. Left mandible; B. Right mandible; C. Maxillula; D. Maxilla; E. Maxilliped; F. Pereopod 1.

Caption: Fig. 11. Hawaiioscia nicoyaensis n. sp. [male] from Playa Pita. A. Pereopod 7; B. Genital papilla; C. Pleopod 1; D. Pleopod 2; E. Pleopod 3 exopod; F. Pleopod 4 exopod; G. Pleopod 5 exopod.

Caption: Fig. 12. Trichorhina biocellata n. sp. specimens from Playa Pita. A. Adult female in dorsal view; B. Adult male in ventral view; C. Dorsal scale-setae ([female]); D. Cephalon, dorsal ([female]).

Caption: Fig. 13. Trichorhina biocellata n. sp. [female] from Playa Pita. A. Eye; B. Pleon, telson and uropods, dorsal; C. Antennal flagellum.

Caption: Fig. 14. Trichorhina biocellata n. sp. [male] from Playa Pita. A. B/c and d/c coordinates of noduli laterales; B. Cephalon, frontal; C. Epimeron of pereonite 7 showing noduli laterales (n.l.); D. Antennula; E. Antenna; F. Left mandible; G. Right mandible; H. Maxillula; I. Maxilla.

Caption: Fig. 15. Trichorhina biocellata n. sp. [male] from Playa Pita. A. Maxilliped; B. Pereopod 1; C. Pereopod 7.

Caption: Fig. 16. Trichorhina biocellata n. sp. [male] from Playa Pita. A. Genital papilla; B. Pleopod 1; C. Pleopod 2; D. Pleopod 3 exopod; E. Pleopod 4 exopod; F. Pleopod 5 exopod.
TABLE 1
Species of terrestrial isopods recorded from Costa Rica

           Species              Family           Costa Rican records

1    Ligia baudiniana          Ligiidae       Puntarenas (Schultz,
     Milne-Edwards, 1840                      1974). Corcovado, Punta
                                              Salsipuedes; Punta
                                              Cahuita; Isla San Lucas
                                              (Leistikow, 1997a).
                                              Manuel Antonio, Playa
                                              Espadilla, Puntarenas;
                                              Manuel Antonio, Quepos,
                                              Aguirre, Puntarenas;
                                              Playa Pita, S of
                                              Tarcoles, Puntarenas;
                                              Isla San Jose, Islas
                                              Murcielago, Guanacaste;
                                              Samara, Guanacaste;
                                              Reserva de Cabo Blanco,
                                              sector San Mguel; Playa
                                              Blanca, Punta Morales,
                                              playa Arenosa; Isla
                                              Cabuya, Cabo Blanco (new
                                              records)

2    Tylos wegeneri             Tylidae       Puntarenas (Schultz,
     Vandel, 1952                             1983)

3    Tylos niveus Budde-        Tylidae       Boca Coronado, manglar
     Lund, 1885                               Terraba-Sierpe (new
                                              record)

4    Clavigeroniscus         Styloniscidae    Puente de Las Mulas;
     riquieri (Arcangeli,                     Orijuaco; San Jose
     1930)                                    (Arcangeli, 1930)

5    Buchnerillo            Incertae sedis    Playa Pita, S of
     neotropicalis n. sp.                     Tarcoles, Puntarenas;
                                              Tortuguero, Limon (new
                                              records)

6    Armadilloniscus cf.       Detonidae      Playa Pita, S of
     caraibicus Paoletti                      Tarcoles, Puntarenas (new
     & Stinner, 1989                          record)

7    "Chaetophiloscia"       Philosciidae     Apaican (Arcangeli, 1930)
     gatunensis (Van
     Name, 1926)

8    Hawaiioscia             Philosciidae     Playa Pita, S of
     nicoyaensis n. sp.                       Tarcoles, Puntarenas (new
                                              record)

9    Ischioscia              Philosciidae     Heredia, Caida el Angel,
     cadoangelis                              Vara Blanca; Cartago,
     Leistikow, 2000                          Tapanti (Leistikow,
                                              2000a)

10   Ischioscia elongata     Philosciidae     Planton (Richardson 1913;
     Leistikow, 1997                          Leistikow, 1997a).
                                              Alajuela, Volcan Poas
                                              (Leistikow, 2000a)

11   Ischioscia marmorata    Philosciidae     Guanacaste, Volcan
     Leistikow, 2000                          Tenorio (Leistikow,
                                              2000a)

12   Ischioscia martinae     Philosciidae     Cordillera de Talamanca,
     Leistikow, 1997                          Valle de Coto Bruz,
                                              Altamira; Cordillera de
                                              Tilaran, Monteverde Cloud
                                              Forest Reserve; San
                                              Isidro de El General,
                                              Coronado (Leistikow,
                                              1997a). Monteverde
                                              (Leistikow, 2000a)

13   Ischioscia muelleri     Philosciidae     Cervantes (Leistikow,
     Leistikow, 1997                          1997a). San Isidro,
                                              Coronado; Heredia, Ceno
                                              Zurqui; Alajuela, Volcan
                                              Poas (Leistikow, 2000a)

14   Ischioscia              Philosciidae     Guanacaste, Santa Rosa,
     plurimaculata                            Braulio Carrillo;
     Leistikow, 2000                          Cartago, Orosi, El
                                              Cedral; Cartago,
                                              Turrialba (Leistikow,
                                              2000a)

15   Ischioscia              Philosciidae     Heredia, Cerro Zurqui
     quadrispinis                             (Leistikow, 1997b)
     Leistikow, 2000

16   Mirtana                 Philosciidae     Heredia, Cerro Zurqui
     costaricensis                            (Leistikow, 1997b)
     Leistikow, 1997

17   Pentoniscus             Philosciidae     La Estrella; La Mica,
     pruinosus                                south of Orosi; Pitahaya,
     Richardson, 1913                         south of Cartago
                                              (Richardson, 1913). San
                                              Jose; San Juan; Apaican;
                                              Vulcan Irazu; Puente de
                                              las Mulas; Tuis
                                              (Arcangeli, 1930)

18   Pentoniscus vargasae    Philosciidae     San Jose, Parque Nacional
     Leistikow, 1998                          Braulio Carillo
                                              (Leistikow, 1998)

19   Colomboniscus          Scleropactidae    Puente de Las Mulas
     tristani (Arcangeli,                     (Arcangeli, 1930; Van
     1930)                                    Name, 1936). San Jose;
                                              San Juan; La Palma;
                                              Orijuaco; Apaican, Volcan
                                              Irazu; faldas Volcan
                                              Irazu (Arcangeli, 1930).
                                              San Jose; Originaco (=
                                              Orijuaco); "Fed. Hort.
                                              Board" (Schmidt, 2007)

20   Globopactes            Scleropactidae    Cordillera de Talamanca,
     talamancensis                            Valle de Coto Bruz,
     (Leistikow,                              Altamira (Leistikow,
                                              1997a; Schmidt, 2007).
                                              Corte Punta Burica;
                                              Corcovado, Sirena
                                              (Leistikow, 1997a)

21   "Scleropactes"         Scleropactidae    La Palma (Arcangeli,
     estherae Arcangeli,                      1930)
     1930

22   Navamundoniscus         Dubioniscidae    San Jose (Arcangeli,
     baldonii (Arcangeli,                     1930)

23   Trichorhina            Platyarthridae    Puente de Las Mulas; San
     giannellii                               Jose; Orijuaco; Apaican;
     Arcangeli, 1929                          faldas Volcan Irazu; San
                                              Juan (Arcangeli, 1930)

24   Trichorhina marianii   Platyarthridae    Puente de Las Mulas; San
     Arcangeli, 1930                          Jose; faldas Volcan
                                              Irazu; San Juan
                                              (Arcangeli, 1930)

25   Trichorhina            Platyarthridae    Playa Pita, S of
     biocellata n. sp.                        Tarcoles, Puntarenas (new
                                              record)

26   Nagurus cristatus      Trachelipodidae   Puerto Limon (Budde-
     (Dollfus, 1889)                          Lund, 1908; Arcangeli,
                                              1930). San Jose; Puente
                                              de Las Mulas (Arcangeli,
                                              1930)

27   Porcellionides         Porcellionidae    Turrubales (Richardson,
     pruinosus (Brandt,                       1910). San Jose; Apaican
     1833)                                    (Arcangeli 1930)

28   Synarmadillo            Armadillidae     Road between Juan Vinas
     tristani                                 and Reventazon; Turrialba
     (Richardson, 1910)                       (Richardson, 1910). San
                                              Jose (Arcangeli, 1930)

29   Venezillo               Armadillidae     San Jose (Arcangeli,
     grenadensis (Budde-                      1930)
     Lund, 1893)

30   Venezillo gigas         Armadillidae     San Jose (Leistikow,
     (Miers, 1877)                            1997a)

           Species                   Distribution

1    Ligia baudiniana       Atlantic and Pacific shores
     Milne-Edwards, 1840    of America from Florida to
                            Brazil and from California to
                            Ecuador, including Galapagos
                            Islands

2    Tylos wegeneri         Florida; Costa Rica; Lesser
     Vandel, 1952           Antilles; Trinidad; Tobago;
                            Venezuela

3    Tylos niveus Budde-    Florida; Bahamas; Bermudas;
     Lund, 1885             Cuba and other Caribbean
                            islands; Mexico; Belize;
                            Venezuela; Puerto Rico;
                            Colombia; Brazil

4    Clavigeroniscus        Pantropical
     riquieri (Arcangeli,
     1930)

5    Buchnerillo            Costa Rica
     neotropicalis n. sp.

6    Armadilloniscus cf.    Venezuela; Costa Rica
     caraibicus Paoletti
     & Stinner, 1989

7    "Chaetophiloscia"      Costa Rica; Panama;? Brazil
     gatunensis (Van
     Name, 1926)

8    Hawaiioscia            Costa Rica
     nicoyaensis n. sp.

9    Ischioscia             Costa Rica
     cadoangelis
     Leistikow, 2000

10   Ischioscia elongata    Costa Rica
     Leistikow, 1997

11   Ischioscia marmorata   Costa Rica
     Leistikow, 2000

12   Ischioscia martinae    Costa Rica
     Leistikow, 1997

13   Ischioscia muelleri    Costa Rica
     Leistikow, 1997

14   Ischioscia             Costa Rica
     plurimaculata
     Leistikow, 2000

15   Ischioscia             Costa Rica
     quadrispinis
     Leistikow, 2000

16   Mirtana                Costa Rica
     costaricensis
     Leistikow, 1997

17   Pentoniscus            Costa Rica
     pruinosus
     Richardson, 1913

18   Pentoniscus vargasae   Costa Rica
     Leistikow, 1998

19   Colomboniscus          Costa Rica; Venezuela
     tristani (Arcangeli,
     1930)

20   Globopactes            Costa Rica
     talamancensis
     (Leistikow,

21   "Scleropactes"         Costa Rica
     estherae Arcangeli,
     1930

22   Navamundoniscus        Costa Rica
     baldonii (Arcangeli,

23   Trichorhina            Cuba; Costa Rica
     giannellii
     Arcangeli, 1929

24   Trichorhina marianii   Costa Rica
     Arcangeli, 1930

25   Trichorhina            Costa Rica
     biocellata n. sp.

26   Nagurus cristatus      Pantropical; greenhouses in
     (Dollfus, 1889)        temperate climates

27   Porcellionides         Cosmopolitan species of
     pruinosus (Brandt,     Mediterranean origin
     1833)

28   Synarmadillo           Costa Rica
     tristani
     (Richardson, 1910)

29   Venezillo              Grenada; Costa Rica; Cuba;
     grenadensis (Budde-    Colombia; Venezuela
     Lund, 1893)

30   Venezillo gigas        Nicaragua; Costa Rica;?
     (Miers, 1877)          Colombia
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Author:Taiti, Stefano; Montesanto, Giuseppe; Vargas, Jose A.
Publication:Revista de Biologia Tropical
Date:Apr 1, 2018
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