The cavernicolous Oniscidea (Crustacea: Isopoda) of Portugal. European Journal of Taxonomy


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The cavernicolous Oniscidea (Crustacea: Isopoda) of Portugal. European Journal of Taxonomy

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Title:
The cavernicolous Oniscidea (Crustacea: Isopoda) of Portugal. European Journal of Taxonomy
Series Title:
European Journal of Taxonomy
Creator:
Reboleira, Ana Sofia P.S.
Fernando Gonçalves
Oromí, Pedro
Taiti, Stefano
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EJT Consortium
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English

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Subjects / Keywords:
Isopoda ( local )
New Species ( local )
Troglobiont ( local )
Subterranean Ecosystems ( local )
Iberian Peninsula ( local )
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serial ( sobekcm )

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Abstract:
The study of subterranean Oniscidea in Portugal has been neglected for nearly 70 years, but recent investigations have revealed high diversity. All the terrestrial isopods from caves of mainland Portugal, including data from the literature and new material, are listed, revealing new biodiversity patterns. Twenty-seven species, belonging to 15 genera and six families, are known, of which 16 species are considered to be exclusively from subterranean ecosystems, i.e., troglobionts. Trichoniscidae is the most diverse family. Seven species in the family Trichoniscidae and one species in the family Styloniscidae are described as new (all with Reboleira & Taiti as authors): Trichoniscoides bellesi sp. nov. from the Montejunto Massif; T. sicoensis sp. nov. from the Sicó Massif; Metatrichoniscoides salirensis sp. nov. from the typhonic valley of Caldas da Rainha; Troglonethes olissipoensis sp. nov. from the Lisbon Peninsula; T. arrabidaensis sp. nov. from the Arrábida Massif; Miktoniscus longispina sp. nov. from the Sicó Massif and Cesaredas Plateau; Moserius inexpectatus sp. nov. from the Estremenho Massif; and Cordioniscus lusitanicus sp. nov. from Alentejo and Algarve, the southernmost provinces of Portugal. The subgenus Trogleluma Vandel, 1946 (Armadillidiidae) is raised to genus level. In this geographic region terrestrial isopods are the richest group of cave-adapted animals.
Original Version:
European Journal of Taxonomy, Vol. 161 (2015).

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University of South Florida Library
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K26-05227 ( USFLDC: LOCAL DOI )
k26.5227 ( USFLDC: LOCAL Handle )

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European Journal of Taxonomy 161: 1 ISSN 2118-9773 http://dx.doi.org/10.5852/ejt.2015.161 www.europeanjournaloftaxonomy.eu 2015 Reboleira A.S.P.S. et al. This work is licensed under a Creative Commons Attribution 3.0 License. Monograph urn:lsid:zoobank.org:pub:2297E4A3-D279-4D0A-923C-D5E0D5DCB3C0 1 The cavernicolous Oniscidea (Crustacea: Isopoda) of Portugal 1,* 2 , 3 & Stefano TAITI 4 1,2 1,3 38206 La Laguna, Tenerife, Islas Canarias, Spain. 1 4 Istituto per lo Studio degli Ecosistemi, Consiglio Nazionale delle Ricerche, * Corresponding author: sreboleira@ua.pt 2 Email: fjmg@ua.pt 3 Email: poromi@ull.es 4 Email: stefano.taiti@ise.cnr.it 1 urn:lsid:zoobank.org:author:338DE845-4839-4EF5-B684-587C021F076C 2 urn:lsid:zoobank.org:author:4B8A2E04-3543-465F-9949-B7E1745DA931 3 urn:lsid:zoobank.org:author:00F2ACC4-308B-4BEE-BBA4-DE6AC8905449 4 urn:lsid:zoobank.org:author:62E97059-6AE5-4984-9ABB-7FB6F7358BD6 Abstract. but recent investigations have revealed high diversity. All the terrestrial isopods known from caves of mainland Portugal, including data from the literature and new material, are listed, revealing new biodiversity patterns. Twenty-seven species, belonging to 15 genera and six families, are known, of which 16 species are considered to be exclusively from subterranean ecosystems, i.e., troglobionts. Trichoniscidae is the most diverse family. Seven species in the family Trichoniscidae and one species in the Trichoniscoides bellesi T. sicoensis Metatrichoniscoides salirensis Troglonethes olissipoensis sp. nov. from T. arrabidaensis Miktoniscus longispina sp. Moserius inexpectatus sp. nov. from the Estremenho Cordioniscus lusitanicus sp. nov. from Alentejo and Algarve, the southernmost provinces of Portugal. The subgenus Trogleluma geographic region terrestrial isopods are the richest group of cave-adapted animals. Key words. Isopoda, new species, troglobiont, subterranean ecosystems, Iberian Peninsula. Portugal. European Journal of Taxonomy 161: 1. http://dx.doi.org/10.58552/ejt. 2015.161

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2 Table of contents Introduction .............................................................................................................................................3 Results .....................................................................................................................................................6 Family Trichoniscidae Sars, 1899 .........................................................................................................6 Genus Trichoniscoides Sars, 1899 ................................................................................................... ...6 T. broteroi ............................................................................................................ ...6 T. machadoi .......................................................................................................... ...6 T. ouremensis ........................................................................................................ ...7 T. serrai Cruz, 1993 .................................................................................................................... ...7 T. subterraneus ..................................................................................................... ...8 T. meridionalis ...................................................................................................... ...8 T. bellesi Reboleira & Taiti sp. nov. ............................................................................................ ...9 T. sicoensis Reboleira & Taiti sp. nov. ......................................................................................... ...13 Genus Metatrichoniscoides .................................................................................... ...17 M. salirensis Reboleira & Taiti sp. nov. .................................................................................... ...17 Genus Troglonethes Cruz, 1989 ................................................................................................... ...21 T. olissipoensis Reboleira & Taiti sp. nov. ............................................................................. ...21 T. arrabidensis Reboleira & Taiti sp. nov. ........................................................................... ...25 Genus Miktoniscus ........................................................................................... ...28 M. longispina Reboleira & Taiti sp. nov. ................................................................................. ...28 Genus Haplophthalmus Schbl, 1860 ......................................................................................... ...32 H. siculus ............................................................................................................ ...32 Genus Moserius Strouhal, 1940 ................................................................................................... ...32 M. inexpectatus Reboleira & Taiti sp. nov. ............................................................................... ...32 ................................................................................................... ...35 Genus Cordioniscus Grve, 1914 ................................................................................................. ...35 C. lusitanicus Reboleira & Taiti sp. nov. .................................................................................. ...35 Family Philosciidae Kinahan, 1947 .................................................................................................. ...39 Genus Anaphiloscia Racovitza, 1907 ......................................................................................... ...39 A. sicula Arcangeli, 1934 ......................................................................................................... ...39 Genus Ctenoscia .................................................................................................. ...40 C. minima ....................................................................................................... ...40 ............................................................................................... ...40 Genus Trichorhina Budde-Lund, 1908 ........................................................................................ ...40 T. anophthalma Arcangeli, 1935 ............................................................................................. ...40 Family Porcellionidae Brandt, 1831 ................................................................................................. ...42 Genus Porcellionides ............................................................................................... ...42 P. pruinosus .................................................................................................... ...42 P. sexfasciatus ........................................................................................ ...43 P. cingendus ................................................................................................. ...43 Genus Porcellio Latreille, 1804 ................................................................................................... ...43 P. cavernicolus ................................................................................................... ...43 P. dilatatus dilatatus Brandt, 1831 .......................................................................................... ...46 P. dispar P. echinatus Lucas, 1849 ............................................................................................................47 Family Armadillidiidae Brandt, 1833 .............................................................................................. ...47 Genus Eluma Budde-Lund, 1885 ................................................................................................. ...47 E. caelata .......................................................................................................... ...47 E. tuberculata Cruz, 1991 ....................................................................................................... ...49

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3 Genus Paraschidium ........................................................................................... ...50 Paraschizidium sp. .................................................................................................................. ...50 Genus Trogleluma .................................................................................................. ...50 T. machadoi .................................................................................................... ...51 Discussion ......................................................................................................................................... ...54 Acknowledgements ........................................................................................................................... ...57 References ......................................................................................................................................... ...57 Introduction et al. Terrestrial isopods are successful land colonizers, but a great majority of species have a high dependence Several species of oniscideans have adapted to the aphotic environment, where they can be found in the et al. et al. et al. et al. Trichoniscoides serrai Portugal, i.e., Trichoniscoides broteroi T. subterraneus T. meridionalis T. ouremensis massif. The family Porcellionidae was represented by the species Porcellio cavernicolus known from Sic, and the family Armadillidiidae by Troglarmadillium Trogleluma machadoi Recent investigations in caves of karst areas from Portugal revealed the presence of several new species of terrestrial isopods. The oniscidean fauna from the subterranean environment of mainland Portugal is here discussed based on literature data and the new material examined. Material and methods Field work was performed between 2007 and 2014 in many karst caves from mainland Portugal and names for caves are maintained throughout the text, i.e., “Gruta” for a horizontal cave, “Algar” for a vertical cave and “Lapa” for a small cave. Each cave was monitored in two different zones: just beyond the twilight zone and in its deeper parts. Specimens were collected using baited pitfall traps and active search in each cave, and most were preserved in 70% ethanol, while some specimens were preserved in 100% ethanol for future molecular

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4 subterranean environment using the Schiner-Racovitza system and following the same criteria as in Reboleira et al. species that exhibit a combination of depigmentation and lack or reduction of eyes, and are found exclusively in subterranean ecosystems. Table 1. Studied caves in Portugal. temperature at soil level in C. Cave Karst area Coordinates E T Gruta de Santo Adrio 580 12.0 Cantanhede 70 13.2 Gruta da Arrifana Sic 300 16.7 Gruta do Algarinho Sic 208 14.9 Gruta do Soprador do Carvalho Sic 20 15.1 Gruta da Senhora da Estrela Sic 380 13.8 Gruta da Cermica Sic 355 15.0 Algar da Ervilha Sic 335 15.1 Gruta de So Simo Sic 335 14.6 Gruta Abrigo Tomar I Estremenho 92 14.6 Gruta do Papagaio Estremenho 419 15.6 Gruta do Burro Estremenho 390 15.4 Algar do Ladoeiro Estremenho 485 15.5 Estremenho 307 18.0 Gruta do Almonda Estremenho 95 18.0 Algar do Z de Braga Estremenho 255 17.0 Estremenho 294 14.3 Gruta de Salir Caldas da Rainha 60 17.3 Gruta dos Bolhos Cesaredas 145 16.3 Algar do Javali 380 14.4 Gruta da Assafora Lisbon Peninsula 89 16.7 Gruta da Alvide Lisbon Peninsula 42 18.3 Gruta do Frade 0 21.5 Lapa da Furada 159 18.1 Algar de Santo Antnio Alandroal 370 17.6 Gruta de Ibne Ammar Algarve 10 18.9 Algaro do Remexido Algarve 131 18.1 Algarve 239 17.0 Gruta da Senhora Algarve 85 17.8

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5 All material was collected by A.S.P.S. Reboleira and has been deposited in the following collections: Fig. 1. Location of the main karst areas of Portugal. 10. Algarve.

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6 Results Family Trichoniscidae Sars, 1899 Genus Trichoniscoides Sars, 1899 Trichoniscoides broteroi Trichoniscoides broteroi Trichoniscoides broteroi – et al. 2011a: 7, table 1. Records Remarks No specimens of this species were collected during our research. The type locality is now closed by the municipality and the whole surrounding area is urbanized. Distribution Presently known only from the type locality. Trichoniscoides machadoi Fig. 2A–C Trichoniscoides machadoi Trichoniscoides machadoi – Schmlzer Schmalfuss 2003: 303. Material examined Previous records Remarks The diagnostic features of this species, i.e., the male pereopod 7 and male pleopods 1 and 2, are illustrated Distribution Species endemic to Portugal where it is widely distributed, from Serra do Gers in the north to the Algarve in the south.

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7 Ecological notes This species is both cavernicolous and endogean. Trichoniscoides ouremensis Trichoniscoides pseudomixtus ouremensis Trichoniscoides ouremensis – 76, 139, map 4. Schmalfuss 2003: 304. Reboleira et al. 2011a: 7, table 1. Records Distribution Species endemic to Portugal, presently only known from the type locality. Trichoniscoides serrai Cruz, 1993 Trichoniscoides serrai Trichoniscoides serrai – Schmalfuss 2003: 304. Fig. 2. Trichoniscoides machadoi A . Pereopod 7. B . Pleopod 1. C . Pleopod 2.

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8 Records Distribution Species endemic to northern Portugal, known only from the type locality. Trichoniscoides subterraneus Trichoniscoides machadoi subterraneus Trichoniscoides subterraneus – 1971: 5, 77, 140, map 4. Schmalfuss 2003: 304. Reboleira et al. 2011a: 7, table 1. Records Distribution Species endemic to central Portugal, known only from the type locality. Trichoniscoides meridionalis Fig. 3A–C Trichoniscoides pseudomixtus meridionalis Fig. 3. Trichoniscoides meridionalis A . Pereopod 7. B . Pleopod 1. C . Pleopod 2.

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9 Trichoniscoides meridionalis – 75, 139, map 4. Reboleira et al. 2011a: 7, table 1. Material examined Previous records Distribution This species is restricted to the Estremenho karst massif, in central-western Portugal. Remarks Trichoniscoides pseudomixtus . species level. The male pereopod 7 and male pleopods 1 and 2 are illustrated in Fig. 3 from specimens Ecological notes This species can be found from the entrances to the deepest parts of the caves, normally associated with the only troglobiotic oniscidean so far known from this large karst area. Contrary to other troglobiotic arthropods from the same massif, it does not present closely related species along the three subunits et al. i.e., the spider Nesticus lusitanicus Fage, 1931, the campodeid Podocampa cf. fragiloides Silvestri, 1932, and the ground beetles Trechus gamae Reboleira & Serrano, 2009, T. lunai Reboleira & Serrano, 2009 and T. machadoi et al. Trichoniscoides bellesi Reboleira & Taiti sp. nov. urn:lsid:zoobank.org:act:FE37F766-5E77-430F-87C2-C5CB28A78D5D Figs 4A –G, 5A–E, 6A –D Diagnosis A blind and colourless Trichoniscoides characterised by the male pleopod 1 exopod having a broadly rounded outer margin and two equal distal lobes, and the male pleopod 2 endopod having a distal article Etymology knowledge on the subterranean fauna from the Iberian Peninsula.

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10 Fig. 4. Trichoniscoides bellesi A . Adult specimen in dorsal view. B . Dorsal scale-seta. C . Cephalon in frontal view. D . Cephalon in E . Pleonites 4, 5, telson and right uropod. F . Antennula. G . Antenna.

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11 Material examined Holotype Paratypes Description Fig. 5. Trichoniscoides bellesi A . Right mandible. B . Left mandible. C D E

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12 Fig. 6. Trichoniscoides bellesi A . Pereopod 1. B . Pereopod 7. C . Pleopod 1. D . Pleopod 2 endopod.

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13 and endopod with a long and a short seta at apex. MALE. article. Pleopod 2 Remarks In the shape of the male pleopod 2 exopod, with a thickset distal article ending in a narrow point, the new species is similar to Trichoniscoides lusitanus T. broteroi and T. heroldi 1952. It differs from T. lusitanus , an epigean species from north-western Spain and northern Portugal, in lacking eyes and pigment, in the male pleopod 1 endopod having a triangular basal article, and in T. broteroi in lacking the hook on the male pereopod 7 merus, and in the triangular shape of the basal article of the T. heroldi , an epigean species from eastern France, in the lack of eyes and the shape of the male pleopod 1. Ecological notes Trichoniscoides bellesi sp. nov. is typically a troglobiotic species, lacking eyes and body pigment. It was discovered in the deepest and most thermally insulated parts of Algar do Javali, around 10 m deep. Javali without recovering any specimens of T. bellesi . The subterranean community of Algar do Javali includes other troglobiotic species, such as the campodeid Podocampa cf. fragiloides , a species of the oniscidean genus Paraschizidium Trechus tatai pselaphid beetle, and the pseudoscorpions Chthonius cardosoi Zaragoza, 2012 and Roncocreagris occidentalis et al. Trichoniscoides sicoensis Reboleira & Taiti sp. nov. urn:lsid:zoobank.org:act:FCBE373B-42CE-4305-A932-28D88355FFB8 Figs 7A–G, 8A–E, 9A–E Diagnosis A blind and colourless Trichoniscoides characterised by the male pereopod 7 merus having a lobe on the mid-sternal margin, the male pleopod 1 exopod having a broadly rounded outer margin and two unequal setae, the endopod having a fusiform distal article with a distinct circular suture in the middle, and the male pleopod 2 endopod having thickset distal article bearing two short triangular lobes and two setae at the apex. Etymology The new species is named after the Sic karst area, where the species occurs. Material examined Holotype Paratypes

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14 Fig. 7. Trichoniscoides sicoensis Reboleira & Taiti sp. nov. from Gruta da Cermica, Sic. Paratype, A . Specimen in dorsal view. B . Dorsal scale-seta. C . Cephalon in frontal view. D . Cephalon in dorsal view. E . Pleonites 4, 5, telson and right uropod. F . Antennula. G . Antenna.

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15 Description third article with distinct spine and three long aesthetascs at apex. Antenna Fig. 8. Trichoniscoides sicoensis A . Right mandible. B . Left mandible. C D E

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16 Fig. 9. Trichoniscoides sicoensis A . Pereopod 1. B . Pereopod 7. C . Genital papilla. D . Pleopod 1. E . Pleopod 2.

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17 seta at apex. MALE. with an area of short quadrangular scales on sternal margin of carpus and with convex inner margin and concave outer margin, distal article fusiform with distinct suture in middle and setose apex. Pleopod 2 distal article about 1.5 times as long as basal one, thickset, with two short triangular lobes and setae at apex. Remarks This new species is tentatively included in the genus Trichoniscoides since it has all the characters of the genus, except for the distal article of the male pleopod 1 endopod, which shows a distinct circular suture in the middle, and the thickset distal part of the male pleopod 2 endopod. These characters are present also in an epigean species described by Gregory et al. Trichoniscoides species C” northern Portugal. The new species differs from the one recorded by Gregory et al. of the basal article of the male pleopod 1 endopod and of the distal part of the male pleopod 2 endopod. Trichoniscoides sicoensis Reboleira & Taiti sp. nov., together with the species recorded by Gregory et al. characters are found. For the presence of a lobe on the mid-sternal margin of the male pereopod 7 merus, the new species also resembles T. broteroi , from which it is distinguishable by the shape of the male pleopods 1 and 2. A hook-like lobe on the sternal margin of the male pereopod 7 merus is present also in sensu lobe is near the base instead of being on the mid-part of the sternal margin. Ecological notes This species seems to be endemic to caves of the Sic karst area. In Gruta do Soprador do Carvalho one specimen was collected under a stone completely submerged in the cave stream. Despite the amphibian where roots are abundant. Several troglobiotic species are known to share this habitat with T. sicoensis Reboleira &Taiti sp. nov.: the pseudoscorpions Roncocreagris blothroides , R. borgesi Zaragoza & Reboleira, 2013 and R. cavernicola Scutogona minor Enghoff T. broteroi, Miktoniscus longispina Reboleira & Taiti sp. nov. and Porcellio cavernicolus Domene lusitanica et al. Genus Metatrichoniscoides Metatrichoniscoides salirensis Reboleira & Taiti sp. nov. urn:lsid:zoobank.org:act:7E09D1C5-19F8-44DF-A1AE-B2D741C411AE Figs 10A–G, 11A–E, 12A–E Diagnosis A blind and colourless Metatrichoniscoides characterised by the male pleopod 1 exopod having two long distal setae subequal in length, and the male pleopod 2 endopod having a thickset distal article, ending in a thinner sinuous part with a beak-like small lobe medially directed.

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18 Etymology The new species is named after Gruta de Salir, the cave where the specimens were collected, in the municipality of Caldas da Rainha. Fig. 10. Metatrichoniscoides salirensis Reboleira & Taiti sp. nov. from Gruta de Salir, Caldas da Rainha typhonic valley. A . Adult specimen in dorsal view. B . Dorsal scale-seta. C . Cephalon in frontal view. D . Cephalon in dorsal view. E . Pleonites 4, 5, telson and right uropod. F . Antennula. G . Antenna.

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19 Material examined Holotype Paratypes Fig. 11. Metatrichoniscoides salirensis Reboleira & Taiti sp. nov. from Gruta de Salir, Caldas da Rainha, A . Right mandible. B . Left mandible. C D E

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20 Description with epimera directed backwards. Pleon epimera and short posterior points. Telson Fig. 12. Metatrichoniscoides salirensis Reboleira & Taiti sp. nov. from Gruta de Salir, Caldas da Rainha, A . Pereopod 1. B . Pereopod 7. C . Genital papilla. D . Pleopod 1. E . Pleopod 2.

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21 and second and bearing four long aesthetascs at apex. Antenna exopod with several long, pointed setae and endopod with long seta at apex. MALE. like distal article inserted on medial margin of basal article. Pleopod 2 beak-like small lobe medially directed. Remarks Metatrichoniscoides M. leydigii M. nemausiensis M. fouresi M. celticus Metatrichoniscoides salirensis Reboleira & Taiti sp. nov. is readily distinguished from all the other species in the genus by the male pleopod 1 exopod bearing two long setae and the peculiar shape of Peninsula. Ecological notes Gruta de Salir is a peculiar, beautiful cave located near the sea, with a genesis related to previous sea known from this cave and this karst area. Genus Troglonethes Cruz, 1989 Troglonethes olissipoensis Reboleira & Taiti sp. nov. urn:lsid:zoobank.org:act:7E09D1C5-19F8-44DF-A1AE-B2D741C411AE Figs 13A–G, 14A–E, 15A–D Diagnosis A species of Troglonethes exopod triangular, as wide as long, and the male pleopod 2 endopod with the distal article bearing a basal and a distal hook-like process. Etymology de Alvide is located. Material examined Holotype

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22 Paratypes Fig. 13. Troglonethes olissipoensis Reboleira & Taiti sp. nov. from Gruta de Alvide, Lisbon Peninsula. A . Adult specimen in dorsal view. B . Dorsal scale-seta. C . Cephalon in frontal view. D . Cephalon in dorsal view. E . Pleonites 4, 5, telson and left uropod. F . Antennula. G . Antenna.

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23 Description three long aesthetascs at apex. Antenna Fig. 14. Troglonethes olissipoensis Reboleira & Taiti sp. nov. from Gruta de Alvide, Lisbon Peninsula, A . Right mandible. B . Left mandible. C D E

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24 several long pointed setae and endopod with a long and a short seta at apex. Fig. 15. Troglonethes olissipoensis Reboleira & Taiti sp. nov. from Gruta de Alvide, Lisbon Peninsula, A . Pereopod 1. B . Pereopod 7. C . Genital papilla and pleopod 1. D . Pleopod 2.

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25 MALE. straight sternal margin, carpus with row of scales on distal margin, and propodus with long, thin setae with setose apex. Pleopod 2 distal article with outer margin bearing hook-like process at base and one at apex. Remarks The genus Troglonethes T. aurouxi from a cave north of Portugal is included in the genus since it has all the characters listed in the diagnosis. Troglonethes olissipoensis Reboleira & Taiti sp. nov. differs from T. aurouxi and the male pleopod 2 endopod with a basal hook-like process on the outer margin of the distal article. Ecological notes This species was only found in Gruta de Alvide, located in an overurbanized area, with part of the cave ceiling used as the base of a residential building. The specimens were collected in the deepest parts of the cave, the so-called third level. It is the second troglobiotic species from karst caves in the Lisbon Peninsula, after the record of the bristletail Coletinia et al. Troglonethes arrabidaensis Reboleira & Taiti sp. nov. urn:lsid:zoobank.org:act:78A12550-F7CA-4892-9E90-7F85446423D3 Figs 16A–G, 17A–D Diagnosis A species of Troglonethes pod 7 carpus enlarged in the basal part, the pleopod 1 exopod triangular, as wide as long, and the male pleopod 2 endopod with the distal article bearing an apical hook-like process. Etymology Material examined Holotype Paratypes Description .

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26 Fig. 16. Troglonethes arrabidaensis A . Adult specimen in dorsal view. B . Dorsal scale-seta. C . Cephalon in frontal view. D . Cephalon in dorsal view. E . Pleonites 4, 5, telson and left uropod. F . Antennula. G . Antenna.

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27 MALE. distal article with hook-like process at apex. Fig. 17. Troglonethes arrabidaensis A . Pereopod 1. B . Pereopod 7. C . Genital papilla and pleopod 1. D . Pleopod 2.

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28 Remarks The new species is very similar to Troglonethes aurouxi in the dorsal granulation and shape of the male male pleopod 1 exopod. It is readily distinguishable from T. olissipoensis Reboleira & Taiti sp. nov. in the larger and less numerous dorsal granules, the enlarged basal part of the male pereopod 7 carpus, the male pleopod 1 endopod with the basal article shorter than the exopod, and the distal article of the male pleopod 2 endopod lacking the hook-like process at the base. Ecological notes Genus Miktoniscus Miktoniscus longispina Reboleira & Taiti sp. nov. urn:lsid:zoobank.org:act:6B9A5E8B-53F8-408D-B3BA-8E10BCE20717 Figs 18A–F, 19A–E, 20A–D Diagnosis A species of Miktoniscus characterised by a depigmented body, the male pereopod 7 having a long and stout seta on the distal corner of the ischium, and a triangular male pleopod 1 exopod. Etymology From Latin longus spina the distal part of the male pereopod 7. Material examined Holotype Paratype Additional material Description 3 with reduced epimera and very short posterior points. Telson

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29 Fig. 18. Miktoniscus longispina Reboleira & Taiti sp. nov. from Gruta dos Bolhos, Cesaredas Plateau. A B . Dorsal scale-seta. C . Cephalon in dorsal view. D . Pleonite 5, telson and left uropod. E . Antennula. F . Antenna.

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30 Fig. 19. Miktoniscus longispina Reboleira & Taiti sp. nov. from Gruta dos Bolhos, Cesaredas Plateau, A . Right mandible. B . Left mandible. C D E

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31 MALE. Fig. 20. Miktoniscus longispina Reboleira & Taiti sp. nov. from Gruta dos Bolhos, Cesaredas Plateau, A . Pereopod 1. B . Pereopod 7. C . Pleopod 1. D . Pleopod 2.

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32 enlarged subapicaly, with triangular and crenulated apical part. Pleopod 2 Remarks At present, the genus Miktoniscus species in the genus by the presence of a strong and long seta at the distal corner of the male pereopod 7 ischium. In continental Portugal and northwestern Spain only one species was previously known, Miktoniscis bisetosus M. longispina Reboleira & Taiti sp. nov. also differs in lacking a recurved seta on the male pereopod 7 carpus and in having the male pleopod 1 exopod with an acute, rather than broadly rounded, apex. The new species occurs in two distinct massifs of central Portugal, on the west border of Cesaredas Plateau near the Atlantic coast, and in Gruta da Cermica Sic compared with the type specimens from Cesaredas, i.e., the lack of visible eyes and the different populations, we consider them as belonging to the same species. Genus Haplophthalmus Schbl, 1860 Haplophthalmus siculus Dollfus, 1896 Haplophthalmus siculus Haplophthalmus siculus Records Distribution Portugal, Spain, southern Italy and Sicily. Genus Moserius Strouhal, 1940 Moserius inexpectatus Reboleira & Taiti sp. nov. urn:lsid:zoobank.org:act:18B5B016-F7A6-4047-A2CB-28B4394797CF Figs 21A–E, 22A–C Diagnosis A species of Moserius characterised by the male pereopod 7 carpus having a distal lobe on the sternal margin, and the pleopod 1 exopod having a truncate and sinuous distal point. Etymology From Latin inexpectatus known distribution range of the genus. Material examined Holotype

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33 Description with short, quadrangular frontal lateral lobes, directed outwards and not protruding compared with median lobe. Eyes absent. Pereonites with rounded Fig. 21. Moserius inexpectatus Reboleira & Taiti sp. nov. from Gruta do Almonda, Estremenho, A . Specimen in dorsal view. B . Cephalon in dorsal view. C . Pleonite 5, telson and uropod. D . Antenna. E . Pereopod 1.

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34 about twice as wide as long, with distal part trapezoidal, with concave sides and truncate apex. Antenna distal point, sinuous outer margin and straight medial margin. Fig. 22. Moserius inexpectatus Reboleira & Taiti sp. nov. from Gruta do Almonda, Estremenho, EstremozA . Pereopod 7, caudal view. B . Pereopod 7, rostral view. C . Pleopod 1.

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35 Remarks Moserius since tubercles on pleonite 3. The genus Moserius was previously known only for two species: M. percoi Strouhal, 1940, originally described from the cave Belinca Jama, Slovenia, and later recorded from M. elbanus Taiti & Ferrara, 1995, from a small cave on Elba Island, Tuscany, Italy. The new species is readily distinguished from the other two members of the genus by the peculiar shape of the male pleopod 1 exopod, with a truncate and sinuous, rather than triangular, distal point. The location of M. inexpectatus Ecological notes This species occurs in the Gruta do Almonda, the largest Portuguese cave, with at least 10 km of mapped subterranean galleries. It shares habitat with other troglobiotic species, namely the spider Nesticus lusitanicus Fage, 1931, the woodlouse Trichoniscoides meridionalis , and the beetle Trechus lunai Reboleira & Serrano, et al. Genus Cordioniscus Grve, 1914 Cordioniscus lusitanicus Reboleira & Taiti sp. nov. urn:lsid:zoobank.org:act:B1F2D756-D90F-43BF-9F42-4F8EAE6C79F0 Figs 23A–G, 24A–E, 25A–E Diagnosis A blind species of Cordioniscus characterised by a colourless body, the male pereopod 7 ischium having a rounded hyaline basal lobe, the triangular male pleopod 1 exopod, as long as the endopod, and the complex apical part of the male pleopod 2 endopod. Etymology From Latin lusitanicus Material examined Holotype Paratypes Description

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36 Pleon Fig. 23. Cordioniscus lusitanicus Reboleira & Taiti sp. nov. from Algar de Santo Antnio, EstremozA B . Dorsal scale-seta. C . Cephalon in frontal view. D . Cephalon in dorsal view. E . Pleonites 4, 5, telson and uropods. F . Antennula. G . Antenna.

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37 posterior point, and with one subapical and four apical, long aesthetascs. Antenna Fig. 24. Cordioniscus lusitanicus Reboleira & Taiti sp. nov. from Algar de Santo Antnio, EstremozA . Right mandible. B . Left mandible. C D E

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38 MALE. Fig. 25. Cordioniscus lusitanicus Reboleira & Taiti sp. nov. from Algar de Santo Antnio, EstremozA . Pereopod 1. B . Pereopod 7. C . Genital papilla. D . Pleopod 1. E . Pleopod 2.

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39 article, with parallel sides and complex apical part. Remarks At present the genus Cordioniscus Cordioniscus are mainly distinguishable by the shape of the male pleopod 2 endopod. In the Iberian Peninsula, only Cordioniscus stebbingi from C. stebbingi lobe on the male pereopod 7 ischium, and the thickset and complex, rather than tapering, distal part of the male pleopod 2 endopod. In the presence of a lobe at the base of the male pereopod 7 ischium, C. lusitanus Reboleira & Taiti sp. nov. is similar to C. bulgaricus Andreev, 1986 from Bulgaria and C. andreevi Schmalfuss & Erhard, 1998, C. beroni C. graecus C. kithnosi Ecological notes This species was collected in two isolated karst areas, Alentejo and Algarve. These two areas are more with the troglobiotic millipede Sireuma nobile Family Philosciidae Kinahan, 1857 Genus Anaphiloscia Racovitza, 1907 Anaphiloscia sicula Arcangeli, 1934 Anaphiloscia sicula – 19. Anaphiloscia sicula – – 59. Records Distribution Remarks Anaphiloscia simoni Racovitza, 1907, described on the basis of a female specimen from a cave on characters of A. simoni exist, it is impossible to clarify this taxonomic problem without a re-examination of the type material. No specimens of this genus have been collected during our investigations. Ecological notes Epigean species.

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40 Genus Ctenoscia Ctenoscia minima Philoscia minima Ctenoscia minima Material examined Previous records Distribution Remarks considered it to be a senior synonym of Ctenoscia dorsalis Ecological notes Epigean species. Genus Trichorhina Budde-Lund, 1908 Trichorhina anophthalma Arcangeli, 1935 Figs 26A–G, 27A–C Trichorhina anophthalma Trichorhina anophthalma – 89, 146. Schmalfuss 2003: 315. Material examined Previous records Distribution

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41 Remarks Trichorhina anophthalma T. hoestlandti of Trichorhina hoestlandti species. The specimens of Trichorhina Ecological notes This species is probably endogean and troglophile rather than a true troglobiont. Fig. 26. Trichorhina anophthalma A . Adult specimen in dorsal view. B . Scale-seta. C . Cephalon in frontal view. D . Pleonite 5, telson and right uropod. E . Antennula. F . Antenna. G . Pereopod 1.

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42 Family Porcellionidae Brandt, 1831 Genus Porcellionides Porcellionides pruinosus Porcellio pruinosus Brandt, 1833: 481. Metoponorthus Metoponorthus pruinosus – Fig. 27. Trichorhina anophthalma A . Pereopod 7. B . Pleopod 1. C . Pleopod 2.

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43 Material examined Previous records Distribution Ecological notes Epigean species, occasionally found at cave entrances. Porcellionides sexfasciatus Metoponorthus sexfasciatus Budde-Lund, 1885: 167. Metoponorthus Polytretus sexfasciatus lusitanus – Records Distribution It has also been introduced to many other parts of the world. Ecological notes Epigean species, trogloxene. Porcellionides cingendus Porcellio cingendus Metoponorthus Lusitanoniscus cingendus – Records Distribution Southern British Isles, Atlantic coasts of France, Portugal and Spain. Ecological notes Epigean species, trogloxene. Genus Porcellio Latreille, 1804 Porcellio cavernicolus Figs 28A–G, 29A–F Porcellio cavernicolus Porcellio cavernicolus – Schmalfuss 2003: 220. Reboleira et al

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44 Material examined Fig. 28. Porcellio cavernicolus A . Adult specimen in dorsal view. B . Cephalon in dorsal view. C . Epimeron of pereonite 1. D . Pleonites 3, telson and uropods. E . Antennula. F G . Pleonite 4, 5, telson and uropods.

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45 Fig. 29. Porcellio cavernicolus A . Pereopod 1. B . Pereopod 7. C . Pleopod 1. D . Pleopod 2. E . Pereopod 1. F . Pereopod 7.

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46 Previous records Distribution Remarks localities from where it was described, considerably enlarges the distribution of this species. Populations from the two distinct karst areas show some small differences which, however, do not seem to justify Ecological notes from the ceiling, from which specimens can easily be distinguished by their whitish coloration. Porcellio dilatatus dilatatus Brandt, 1831 Porcellio dilatatus – d. Porcellio Porcellio dilatatus – Arcangeli 1935b: 12. Porcellio dilatatus – Material examined Previous records a

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47 Distribution Species widely distributed in Europe and introduced to many other parts of the world. Ecological notes This species is a common inhabitant of caves and other subterranean habitats. It has no morphological adaptations to cave life and can be considered as a troglophile. Porcellio dispar Porcellio dispar Porcellio dispar – Records Distribution Portugal. Ecological notes Epigean species, trogloxene. Porcellio echinatus Lucas, 1849 Porcellio echinatus Porcellio echinatus Records Distribution Ecological notes Epigean species, trogloxene. Family Armadillidiidae Brandt, 1833 Genus Eluma Budde-Lund, 1885 Eluma caelata Armadillidium caelatum Eluma purpurascens – Records

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48 Distribution north-western Africa. It has been introduced to Tasmania and French Guyana. Fig. 30. Eluma tuberculata Cruz, 1991 from Gruta do Soprador do Carvalho, Sic. A . Specimen in lateral view. B . Cephalon in frontal view. C . Pleonites 4, 5, telson and uropods. D . Antenna. E . Pereopod 1. F . Pereopod 7.

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49 Remarks The genus Eluma includes three epigean species: E. caelata , E. tuberculata Cruz, 1991 from Portugal E. praticola Eluma caelata has Ecological notes Epigean species, trogloxene. Eluma tuberculata Cruz, 1991 Figs 30A–F, 31A –B Eluma tuberculata Eluma tuberculata – Schmalfuss 2003: 107. Material examined Fig. 31. Eluma tuberculata A . Pleopod 1. B . Pleopod 2.

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50 Previous records Distribution This species seems to be endemic to central Portugal. Remarks Eluma tuberculata The numerous specimens collected during our research allow us to illustrate the species with its most morphology concerning the dorsal tubercles, which may be more or less developed. The presence of several adult males permits the description of the male characters which were previously unknown: half having parallel sides and small recurved and pointed apical part not protruding outwards. Pleopod 2 Ecological notes sensu Juberthie et al. Genus Paraschizidium Paraschizidium sp. Fig. 32A –E Material examined Remarks At present, the genus Paraschizidium includes with certainty only three species: P. coeculum P. hispanum Arcangeli, 1935 from southern Spain, and P. roubali a junior synonym of P. coeculum included in the genus Paraschizidium transferred to the genus Schizidium belong to the genus Paraschizidium : they are depigmented and blind, have no schisma at the posteroGenus Trogleluma Remarks Trogleluma Troglarmadillidium include the new species Troglarmadillidium Trogleluma machadoi from two caves in South Portugal Troglarmadillidium

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51 Troglarmadillidium and Trogleluma , with the species T. Troglarmadillidium stygium T. Trogleluma machadoi. Both these species have a depigmented body, are blind, have no schisma at the postero-lateral corner of pereonite 1, a triangular telson, and uropods subgenera show quite distinct morphological traits in the cephalic structure: in Troglarmadillidium the triangular frontal scutellum of the cephalon, frontal line and antennal lobes are missing, while in Trogleluma these structures are present. These T. Troglarmadillidium stygium Trogleluma as a distinct genus from Troglarmadillidium . Trogleluma includes the type species, Trogleluma machadoi , and a few undescribed Trogleluma machadoi Figs 33A–J, 34A –E Troglarmadillidium Trogleluma machadoi Troglarmadillidium Trogleluma machadoi – Troglarmadillidium machadoi – Schmlzer 1971: 106, 155. Schmalfuss 2003: 319. Reboleira et al. 2011a: 7, table 1. Fig. 32. Paraschizidium A . Specimen in lateral view. B . Cephalon in frontal view. C . Cephalon in dorsal view. D . Pleonite 5, telson and uropods. E . Antennula.

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52 Fig. 33. Trogleluma machadoi A . Specimen in lateral view. B C . Cephalon in frontal view. D . Cephalon in dorsal view. E . Cephalon in lateral view. F . Epimeron of pereonite 1 in dorsal view. G . Epimeron of pereonite 2 in ventral view. H . Pleonite 5, telson and right uropod. I . Antennula. J . Antenna.

PAGE 53

53 Fig. 34. Trogleluma machadoi A B . Pereopod 1. C . Pereopod 7. D . Pleopod 1. E . Pleopod 2.

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54 Material examined Previous records Distribution Endemic to caves of the Algarve, the southernmost province of Portugal. Remarks Ecological notes Troglobiotic species. The specimens were mostly found inside the clay, which often covers their walking on cave walls and completely clean of clay. In the caves of the Algarve, this species occurs together with other remarkable troglobiotic species: the pseudoscorpions Titanobochica magna Zaragoza & Reboleira, 2010 and Lusoblothrus aenigmaticus Harpactea stalitoides Ribera, 1993 and Teloleptoneta synthetica Boreviulisoma barrocalense Reboleira & Enghoff, 2013, Acipes machadoi Enghoff & Reboleira, 2013, and Litocampa mendesi thysanuran Squamatinia algharbica et al. Discussion et al caves of Portugal, with the number of known species doubled with the present work. The Portuguese oniscidean fauna is now composed of 16 troglobiotic species in mainland Portugal, 5 troglophiles and Atlantic islands: Trichoniscus bassoti et al. et al. Trichoniscidae is also the richest family in Portugal, with 14 species frequently found in caves of mainland Portugal, 12 of which are considered to be true troglobionts. Trichoniscoides is the most diverse genus. Trichoniscoides machadoi is an endogean and cavernicolous species occurring in different regions of Portugal, while the remaining species are restricted to single karst areas: T. serrai to the palaeokarst of T. broteroi T. sicoensis to the T. meridionalis T. subterraneus in the Aljubarrota Plateau at

PAGE 55

55 the western part of the massif, and T. ouremensis Despite the sampling effort at their type localities, the troglobiotic species Trichoniscoides broteroi , T. serrai , T. ouremensis and T. subterraneus were not found during our research. The Iberian genus Troglonethes now also includes two species from central Portugal: T. olissipoensis Reboleira & Taiti sp. nov. from a cave of the Lisbon Peninsula, and T. arrabidaensis Reboleira & Taiti sp. nov. from a Miktoniscus longispina Reboleira & Taiti sp. nov. was collected in two different karst areas: Sic and the Cesaredas Plateau, the specimens from these localities presenting slight morphological differences from each other. The styloniscid Cordioniscus lusitanicus Reboleira & Taiti sp. nov. is described from caves of the Alentejo and Algarve karst areas, more than 200 km apart in the south of Portugal. Table 2. Terrestrial isopod species recorded in subterranean habitats of karst areas from Portugal, with Family Species ssssss Ecology Karst area Trichoniscidae Trichoniscoides broteroi Tb Sic Trichoniscoides machadoi Tph widespread Trichoniscoides ouremensis Tb Estremenho Trichoniscoides serrai Tb Trichoniscoides subterraneus Tb Estremenho Trichoniscoides meridionalis Tb Estremenho Trichoniscoides bellesi Tb Sic Trichoniscoides sicoensis Tb Sic Metatrichoniscoides salirensis Tb Caldas da Rainha Troglonethes olissipoensis Tb Lisbon Peninsula Troglonethes arrabidaensis Tb Miktoniscus longispina Tb Cesaredas and Sic Haplophthalmus siculus Tx Lisbon Peninsula Moserius inexpectatus Tb Estremenho Styloniscidae Cordioniscus lusitanicus Tb Alentejo Philosciidae Anaphiloscia sicula Tph Algarve Ctenoscia minima Tx Algarve and Lisbon Peninsula Platyarthridae Trichorhina anophthalma Tph Alentejo Porcellionidae Porcellionides pruinosus Tx widespread Porcellionides sexfasciatus Tx Algarve Porcellionides cingendus Tx Estremenho Porcellio cavernicolus Tb Sic and Catanhede Porcellio dilatatus Tph widespread Armadillidiidae Eluma caelata Tx Eluma tuberculata Tph Sic and Estremenho Paraschizidium sp. Tb Trogleluma machadoi Tb Algarve

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56 The family Platyarthridae is represented by the species Trichorhina anophthalma , described from found in karst caves of Alandroal in central Alentejo and in the Algarve. The family Porcellionidae has one troglobiotic species, Porcellio cavernicolus , described from the distribution. Slight morphological differences can easily be observed between the extremes of its area of distribution, and molecular studies may clarify the taxonomy of these isolated populations. This species is particularly abundant in shallow caves with roots pending from the ceiling. The troglophilic Porcellio dilatatus dilatatus The family Armadillidiidae is represented by four species belonging to the genera Eluma , Paraschizidium and Trogleluma . The genus Eluma includes the epigean E. caelata and E. tuberculata, originally Paraschizidium Trogleluma machadoi is widely distributed in caves along the Barrocal area in the Algarve karst massif. From a biospeleological point of view, mainland Portugal is traditionally divided into two main areas, the Lusitanian district in central Portugal, and the Baetic district in the south, including the Algarve and et al. for terrestrial isopods reinforces this separation, with the Lusitanic district clearly rich in troglobiotic species of the genus Trichoniscoides and the Baetic district characterised by the presence of the genera Trogleluma and Cordioniscus . The recent rise in the number of known subterranean species is a result of the extensive use of a standard sampling methodology in a large number of karst caves, which has also provided new taxa of other caveet al. 2009, 2010a, 2010b, 2010c, 2011b, 2012a, ecosystems of Portugal, and by extension of the Iberian Peninsula, requires a consistent sampling effort to reach a complete level of evaluation. Terrestrial isopods have been used as bioindicators in land ecosystems, since they are extremely et al. in support of this is their basal trophic position in cave communities. Decomposing detritus and their strong dependence on water makes them more vulnerable to contamination. The presence of troglobiotic oniscidean species is generally related to deep, stable and undisturbed parts of caves, while in the most disturbed areas only trogloxenes or, at most, troglophile isopods are present, suggesting that they can be used as indicators of the quality of the subterranean ecosystems that they occupy. An example of this troglophile Porcellio dilatatus dilatatus is abundant, while the troglobiont Trichoniscoides meridionalis et al. considered for sustainable management of karst areas in such an important world biodiversity hotspot.

PAGE 57

57 Acknowledgements CEEAA, NEL, AES, ECLER, NALGA, Desnvel, Alto relevo, DIR-SPE, CIES, SAGA and NEC for Trogloarmadillidium stygium one month at the Istituto per lo Studio degli Ecosistemi, CNR, Florence. All specimens were collected under permits of the Instituto de Conservao da Natureza e das Florestas. References Bryologist 57: 242. http://dx.doi.org/10.2307/3240091 Andreev S. 1986a. Cordioniscus bulgaricus la famille Styloniscidae en Bulgarie. Acta zoologica bulgarica 31: 67. Cordioniscus , Alpioniscus et Trichoniscus et nouvelles donnes sur les isopodes terr. Biologia gallo-hellenica 11: 153. Cordioniscus et Trichoniscus et nouvelles donnes sur les isopodes terrestres de la Bulgarie. Historia naturalis bulgarica 15: 55. Arcangeli A. 1934. Anaphiloscia sicula , nuova specie di isopodo umicolo della Sicilia. Bollettino del Laboratorio di Zoologia generale e agraria della Reale Scuola superiore d’Agricoltura in Portici 28: 152. Arcangeli A. 1935a. Gli isopodi terrestri del Portogallo. Bollettino del Laboratorio di Zoologia generale e agraria della Reale Scuola superiore d’Agricoltura in Portici 29: 1. Eos 10: 171. Fauna Caverncola i Intersticial de la Pennsula Ibrica i les Illes Balears . Editorial troglobiont species. International Journal of Speleology http://dx.doi.org/10.5038/1827Bulletin de la Socit impriale des Naturalistes de Moscou 6: 171 – 193. Brandt J.F. & Ratzeburg J.T.C. 1831. Isopoda. In Medizinische Zoologie oder getreue Darstellung und Beschreibung der Thiere, die in der Arzneimittellehre in Betracht kommen, in systematischer Folge herausgegeben Brian A. 1963. grotte della Liguria. Doriana 3: 11. Evolutionary Ecology http://dx.doi.org/10.1007/s10682-012-9625-8

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58 Budde-Lund G. 1885. Crustacea Isopoda Terrestria per Familias et Genera et Species Descripta. Nielsen & Lydiche, Copenhagen. . Zoological Journal of the Linnean Society 172: 360. http://dx.doi.org/10.1111/zoj.12172 Cruz A. 1989. Troglonethes aurouxi Miscelnea Zoolgica 13: 51. Cruz A. 1990. Contribucin al Conocimiento de los Ispodos Terrestres (Oniscidea) de la Pennsula Ibrica y Baleares. PhD Thesis , Barcelona University, Spain. Cruz A. 1991. Especies nuevas o poco conocidas de ispodos terrestres de la Peninsula Ibrica. II. Bulletin de la Socit d’Histoire naturelle de Toulouse 127: 71. Cruz A. 1993. Especies nuevas o poco conocidas de isopodos terrestres de la Pennsula Ibrica. III. Trichoniscoides pitarquensis sp. n. y T. serrai Bulletin de la Socit d’Histoire naturelle de Toulouse 127: 15. Anales de la Sociedad espaola de Historia natural 21: 161. Dollfus A. 1896. Crustacs isopodes de la Sicile. Notes faunistiques Scutogona from Zootaxa http:// dx.doi.org/10.11646/zootaxa.3736.2.5 Subterranean species of Acipes Zootaxa http://dx.doi.org/10.11646/zootaxa.3652.4.6 Frankenberger Z. 1940. Paraschizidium roubali 8: 71. Bulletin of the British Myriapod and Isopod Group 26: 6. Courier Forschungsinstitut Senckenberg 71: 27. and behavior. Terrestrial Arthropod Reviews non calcaire. Comptes Rendus de l’Acadmie des Sciences 290: 49. genus, and a detailed list of the British species of Ligia , Philougria , Philoscia , Porcellio , Oniscus and Armadillium [sic]. Natural History Review 4: 258. de l’Algerie, pendant les annes 1840, 1841, 1842. Sciences physiques, Zoologie 1: 67. Available from http://biodiversitylibrary.org/page/46116745 [accessed 1 Dec. 2015] Biogeographia 17: 125.

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59 with descriptions of new genera and species. Proceedings of the Zoological Society of London 1877: 653. Avalialbe from http://biodiversitylibrary.org/page/28518607 [accessed 1 Dec. 2015] Metatrichoniscoides Journal of Natural History 15: 525. http://dx.doi. org/10.1080/00222938100770371 sustainability and use as bioindicators. Agriculture, Ecosystems & Environment http:// Animal Production Science Archives de Zoologie exprimentale et gnrale , 4 e Srie 7: 145. Reboleira A.S.P.S. 2007. Os Colepteros (Insecta, Coleoptera) Caverncolas do Macio Calcrio Estremenho: uma Aproximao sua Biodiversidade University of Aveiro, Portugal . Reboleira A.S.P.S. 2012. Biodiversity and Conservation of Subterranean Fauna of Portuguese Karst . PhD Thesis, University of Aveiro, Portugal . The genus Boreviulisoma Zootaxa http://dx.doi.org/10.11646/zootaxa.3646.5.2 Journal of Cave and Karst Studies http://dx.doi.org/10.4311/2013LSC0113 Sireuma , a new genus of subterranean millipedes from the Zootaxa http:// dx.doi.org/10.11646/zootaxa.3785.1.6 Description of the larva and female genitalia of Trechus gamae with data on its ecology. Bulletin of Insectology Reboleira A.S.P.S., Gonalves F. & Serrano A. 2009. Two new species of cave dwelling Trechus Clairville, 1806 of the fulvus Deutsche Entomologische Zeitschrift http://dx.doi.org/10.1002/mmnd.200900009 Trechus T. fulvus species group. Zootaxa 2689: 15. Portugal: description of a new species of the genus Litocampa Zootaxa 2728: 50. Titanobochica , surprising discovery Zootaxa 2681: 1. of a biodiversity hotspot region – Portugal: an overview and its conservation. International Journal of Speleology

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60 Lathromene Domene Fauvel, 1872 from Portugal. Zootaxa 2780: 48. Squamatinia algharbica gen. n. sp. Zootaxa 3260: 33. Lusoblothrus , a new syarinid fauna. Zootaxa 3544: 52. Acute toxicity of copper sulfate and potassium dichromate on stygobiont Proasellus : general aspects of groundwater ecotoxicology and future perspectives. Water, Air & Soil Pollution http://dx.doi.org/10.1007/s11270013-1550-0 Literature survey, bibliographic analysis and a taxonomic catalogue of subterranean fauna from Portugal. Subterranean Biology 10: 51. http:// dx.doi.org/10.3897/subtbiol.10.4025 Roncocreagris Zootaxa Journal of Zoology http://dx.doi.org/10.1111/j.1469-7998.2007.00408.x Stuttgarter Beitrge zur Naturkunde , Serie A 343: 1. Stuttgarter Beitrge zur Naturkunde , Serie A 654: 1. Schizidium distribution, morphology. Stuttgarter Beitrge zur Naturkunde A (New Series) 1: 143. Cordioniscus Stuttgarter Beitrge zur Naturkunde , Serie A 582: 1. Schmlzer K. 1965. Ordnung Isopoda Landasseln Monografas de Ciencia moderna 80: 1. Crustaceana 63: 199. Stuttgarter Beitrge zur Naturkunde , Serie A 519: 1. Moserius percoi Zoologischer Anzeiger 129: 13. Thaumatoniscellus orghidani Travaux de l’Institut de Spologie “mile Racovitza” 32: 43. Taiti S. 2007. 10. Gli Isopodi terrestri. In Il Parco Regionale della Maremma e il suo Territorio : 91. Pacini Editore, Pisa.

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61 Mmoires de Biospologie 22: 169. Taiti S. & Rossano C. 2015. Journal of Natural History http://dx.doi.org/10.1080/00222933.2015.1009512 Trogloniscus nomen novum, with descriptions of Tropical Zoology 183. http://dx.doi.org/10.1080/03946975.2012.751240 Bulletin de la Socit zoologique de France 67: 133. Anais da Faculdade de Cincias do Porto 30: 135. isopodes terrestres. Bulletin de la Socit d’Histoire naturelle de Toulouse 82: 263. origines et son histoire. Revue franaise d’Entomologie 15: 101. Archives de Zoologie exprimentale et gnrale 88: 231. Mmoires du Musum national d’Histoire naturelle, Srie A 6: 116. Notes biospologiques 13: 131. Faune de France 64: 1. Mmoires du Musum national d’Histoire naturelle , Srie A 22: 1. Faune de France 66: 417. Cordioniscus Annales de Splologie 23: 621. Miscelnea Zoolgica 3: 1. Zoologischer Anzeiger 24: 403, 417. Available from http://biodiversitylibrary.org/page/9745364 [accessed 1 Dec. 2015] Manuscript received: 26 September 2015 Manuscript accepted: 3 November 2015 Published on: 11 December 2015 Topic editor: Rudy Jocqu Desk editor: Danny Eibye-Jacobsen Printed versions of all papers are also deposited in the libraries of the institutes that are members of the EJT Denmark, Copenhagen, Denmark.


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