The genus Armadillo Latreille, 1802 (Oniscidea, Armadillidae) from Cyprus, with descriptions of two new species.

The present work aims to morphologically characterize the species of Armadillo from Cyprus. Moreover, two new species of the genus are described, A. konstantinoui sp. nov. from several places along the island, and A. karametae sp. nov. from Pafos region. The circum-Mediterranean species A. officinalis is also redescribed.

Intra-island patterns of cryptic diversity within an oceanic island: Insights from the differentiation of Schizidium Verhoeff, 1901 (Oniscidea, Armadillidiidae) within Cyprus, with descriptions of two new species.

Cyprus is a Mediterranean island of volcanic origin isolated for at least 5.3 Myr from surrounding continental areas. The present study focuses on the diversification of the isopod genus Schizidium within the island, including also specimens from surrounding continental areas. The genus Schizidium sensu lato is probably non monophyletic, comprising 26 species distributed from Greece to Iran. Up to date the only representative of the genus reported from Cyprus was Schizidium fissum. Aiming to investigate the patterns of genetic diversity within the focal island, to evaluate the morphology-based taxonomy of the species in the genus, and at the same time to explore phylogenetic relationships with mainland populations, we applied genome-wide ddRADseq as well as Sanger sequencing targeting three mitochondrial (16S, COI and 12S) and the nuclear NaK loci. Results of phylogenetic analyses support the existence of two distinct epigean Schizidium clades with well-defined geographic boundaries that conform to the known paleogeography of Cyprus, plus one endogean clade with restricted distribution within the island. Genetic data and morphology corroborate the assignment of this latter endogean clade to a new species, Schizidium myrrae n. sp. The two epigean clades are also considered as distinct species, one corresponding to the known S. fissum at the eastern part of the island (Pentadaktylos massif) and the other to the newly described S. christosi n. sp. distributed along the western part of the island (Troodos massif). Even though detailed examination of many specimens could not retrieve any morphological differences among representatives of these two clades, clado-chronological analysis indicates a long isolation between them, estimated at âˆ¼ 9 Mya, as well as the sharing of a common ancestor with S. tiberianum from Israel at âˆ¼ 15 Mya. Hence, we can consider these epigean Schizidium species as one more case of cryptic diversity on Cyprus, exhibiting similar patterns with the recently described case in the genus Armadillo.

Everything is similar, everything is different! Trichorhina (Oniscidea, Platyarthridae) from Brazilian caves, with descriptions of 11 new species / Tudo é parecido, tudo é diferente! Trichorhina (Oniscidea, Platyarthridae) de cavernas brasileiras, com descrições de 11 novas espécies

Abstract The present work describes 11 new species of the genus Trichorhina from Brazilian caves in the states of Bahia, Mato Grosso do Sul, Minas Gerais, Pará, and São Paulo. Moreover, the distribution of T. tomentosa has been expanded to include the states of Mato Grosso and Minas Gerais. Additionally, a distribution map and a key for all Brazilian species of Trichorhina, including epigean ones, are provided.

Resumo O presente trabalho descreve 11 novas espécies do gênero Trichorhina de cavernas brasileiras nos estados da Bahia, Mato Grosso do Sul, Minas Gerais, Pará e São Paulo. Além disso, a distribuição de T. tomentosa foi ampliada para incluir os estados de Mato Grosso e Minas Gerais. Adicionalmente, são fornecidos um mapa de distribuição e uma chave para todas as espécies brasileiras de Trichorhina, incluindo as epígeas.

Species conservation profiles of cave-adapted terrestrial isopods from Portugal.

Background: Terrestrial isopods (Oniscidea) are the most diverse group of troglobionts in caves of continental Portugal. They occur in all karst regions of Portugal, play a major role in decomposition of organic matter in caves and may act as umbrella species for the conservation of all other cave-adapted invertebrates. New information: We present the IUCN Red List profiles for the cave-adapted terrestrial isopods from continental Portugal, based on recent distribution data from caves.

Expanding the knowledge on the diversity of the cavernicolous Styloniscidae Vandel, 1952 (Oniscidea, Synocheta) from Brazil, with descriptions of two new species from the semiarid karst regions.

Two new species of Pectenoniscus from two caves in karst areas of the Brazilian semiarid region are described. Pectenoniscuspankaru Campos-Filho, Torres & Bichuette, sp. nov. from Gruna do Govi cave, Serra do Ramalho karst area, state of Bahia, and Pectenoniscusfervens Campos-Filho, Taiti & Bichuette, sp. nov. from Toca Coroa do Frade cave, Barra Bonita karst area, state of Piauí. In addition, specimens of Cylindroniscusflaviae from Gruta da Tapagem (= Caverna do Diabo), Açungui karst area were also recorded. An updated diagnosis of Pectenoniscus and a distribution map of the species examined herein are given.

Genetic evidence against monophyly of Oniscidea implies a need to revise scenarios for the origin of terrestrial isopods.

Among the few crustacean taxa that managed to inhabit terrestrial environments, Oniscidea includes the most successful colonizers in terms of species richness and abundance. However, neither morphological traits nor molecular markers have definitively resolved phylogenetic relationships among major Oniscidea clades or established the monophyly of the taxon. Herein, we employed the highly conserved, nuclear protein-coding genes Sodium-Potassium Pump (NAK) and Phosphoenolpyruvate Carboxykinase (PEPCK), along with the traditionally used 18 s and 28 s ribosomal RNA genes, in an attempt to clarify these questions. Our dataset included sequences representing all major Oniscidea clades and closely related aquatic taxa, as suggested by previous studies. We applied Bayesian Inference and Maximum Likelihood methods and produced a robust and fully resolved phylogenetic tree that offers strong evidence against the monophyly of Oniscidea. The amphibious genus Ligia appears to be more closely related to representatives of marine suborders, while the phylogenetic pattern of the remaining Oniscidea implies a complex history of the transition from the marine environment to land. With the exception of the basal clade, all other established major clades have been recovered as monophyletic, even though relationships within these clades call for a revised interpretation of morphological characters used in terrestrial isopod taxonomy.

Two new species and new records of terrestrial isopods (Crustacea, Isopoda, Oniscidea) from Brazilian caves.

To date, approximately 190 species of terrestrial isopods are known from Brazil and only 14 are considered troglobiotic. After the examination of a large collection from caves in Bambuí and Una geomorphological areas, along the states of Bahia, Minas Gerais and Goiás, two new troglobiotic species were recognized. Pectenoniscus liliae Campos-Filho, Bichuette Taiti sp. n. (Styloniscidae) is described from Serra do Ramalho karst area, and Benthana xiquinhoi Campos-Filho, Bichuette Taiti sp. n. (Philosciidae) from sandstone caves of Chapada Diamantina region. The latter constitutes the second troglomorphic species of the genus. Xangoniscus aganju (Styloniscidae) is also recorded from two caves in the Serra do Ramalho karst area. The systematic position of Iuiuniscus iuiuensis is briefly discussed and Venezillo congener (Armadillidae) is redescribed from São Domingos karst area.

Taxonomy of Alpioniscus (Illyrionethes): A. magnus and three new species from the Dinaric Karst (Isopoda: Oniscidea: Trichoniscidae).

Alpioniscus (Illyrionethes) is a widespread subgenus of Trichoniscidae in the Dinaric Karst with 15 nominal troglobiotic species. Here we redescribe Alpioniscus (Illyrionethes) magnus and describe three new species (A. lossinii n. sp., A. drazinai n. sp., and A. mandalinae n. sp.) restricted to coastal areas. As a result, 18 nominal Illyrionethes species are known from the Dinaric Karst.

A molecular phylogeny of Porcellionidae (Isopoda, Oniscidea) reveals inconsistencies with present taxonomy.

Porcellionidae is one of the richest families of Oniscidea globally distributed but we still lack a comprehensive and robust phylogeny of the taxa that are assigned to it. Employing five genetic markers (two mitochondrial and three nuclear) we inferred phylogenetic relationships among the majority of Porcellionidae genera. Phylogenetic analyses conducted via Maximum Likelihood and Bayesian Inference resulted in similar tree topologies. The mtDNA genes cytochrome oxidase I (COI) and 16s rRNA (16s) were used for clade dating using previously published mutation rates. Our results provide evidence against the monophyly of both Porcellionidae and the largest genus of the family Porcellio. These results are compared to previous published work based on morphological evidence. The genera Leptotrichus and Brevurus are not grouped with the rest of Porcellionidae whereas Agnaridae are grouped with part of Porcellionidae. Armadillidium and Schizidium (Armadillidiidae) occupy a basal position on the phylogenetic tree. Even though the African genera Tura and Uramba (distributed in East Africa) are grouped together there is no general geographical pattern in other sub-clades. Additional taxonomic issues that arise in this work such as the assignment of the recently described genus Levantoniscus, arealso discussed. The status of Porcellionidae should be further revised and morphological characters traditionally used in Oniscidea taxonomy should be reconsidered in view of molecular evidence. The origin of the monophyletic clade within Porcellionidae as indicated in the present work is dated back to the Oligocene (~32 mya).

The genus Alpioniscus Racovitza, 1908 in Sardinia: taxonomy and natural history (Isopoda, Oniscidea, Trichoniscidae).

The genus Alpioniscus Racovitza, 1908 (Trichoniscidae) from Sardinia is revised. Three new cave-dwelling species are described: A.onnisi Taiti & Argano, sp. n., A.stochi Taiti & Argano, sp. n., and A.sideralis Taiti & Argano, sp. n.. The genus Utopioniscus Schmalfuss, 2005 is considered to be a junior synonym of Alpioniscus, after morphological and molecular analyses. Alpioniscusfragilis (Budde-Lund, 1909) and A.kuehni from Grotta del Bue Marino are illustrated. With the new species, the genus Alpioniscus in Sardinia comprises six species: two troglobionts (A.fragilis and A.onnisi), one endogean and troglobiont (A.thanit Taiti & Argano, 2009), and three stygobionts (A.kuehni, A.stochi, and A.sideralis). All the species occur in karstic areas in the central-eastern and south-eastern part of the island. A key to all the Sardinian species of Alpioniscus is provided.

Tailored disorder in calcite organization in tergite cuticle of the supralittoral isopod Tylos europaeus Arcangeli, 1938.

The crustacean cuticle forms skeletal elements consisting of chitin-protein fibrils reinforced by amorphous and crystalline calcium carbonate and phosphate minerals. The edges of skeletal elements are of particular interest. They are subject to repeated strain and stress because they form transitions to the arthrodial membranes connecting them. These allow for relative movements of skeletal elements. In this study, we investigate structure, chemical composition, mineral organization and local mechanical properties of the anterior and posterior edges of the tergite cuticle in the conglobating beach isopod Tylos europaeus and compare these with the protective dorsal region of the tergites. The distribution of mineral phases at the edges resembles that of dorsal regions of the tergites. At the transition with the unmineralized arthrodial membrane the calcite containing distal exocuticle is replaced by epicuticular material and the subjacent cuticular layers containing amorphous calcium carbonate become enriched with amorphous calcium phosphate. At the edges, the local elastic modulus and hardness values are significantly lower compared to dorsal regions of the tergite cuticle, for both, the calcite and the amorphous mineral containing layers. The calcite within the tergite cuticle is assembled in different texture patterns: (i) almost random co-orientation, (ii) almost single crystalline calcite, and (iii) a graded organization. Calcite organization and co-orientation strength is highly variable, not only on very few tens of micrometres, but also between regions with different skeletal functionality. Our results show that besides structure and composition, patterns of calcite organization contribute to the hierarchical architecture and functionality of biological composites.

A new genus of Trachelipodidae Strouhal, 1953 (Crustacea, Isopoda, Oniscidea) from the eastern Mediterranean.

Levantoniscus n. gen. is erected for two new species from Cyprus: Levantoniscus bicostulatus n. sp. and Levantoniscus makrisi n. sp. Levantoniscus wahrmani (Strouhal, 1968) n. comb. from Israel and southern Turkey is transferred from the genus Bathytropa Budde-Lund, 1885 and family Bathytropidae. The new genus is included in the family Trachelipodidae and is characterized by distinct dorsal ornamentation, interlocking pleopods and uncovered pleopodal lungs which are located in invaginations on pleopod 3-5 exopodites.

Taxonomic revision of the genus Benthana Budde-Lund, 1908 (Isopoda: Oniscidea: Philosciidae).

The neotropical genus Benthana (Philosciidae) is reviewed. Fourteen species are redescribed: Benthana picta, B. olfersii, B. longicornis, B. schubarti, B. albomarginata, B. werneri, B. santosi, B. longipenis, B. bocainensis, B. moreirai, B. dimorpha, B. iporangensis, B. sulcata, and B. longicaudata. Additional characters were proposed for B. convexa, B. taeniata, B. serrana, B. trinodulata, B. araucariana, and B. cairensis. New records are provided for 13 species, and four new species are described: Benthana aimores sp. nov., Benthana goitacas sp. nov., Benthana tupinamba sp. nov. and Benthana canastraensis sp. nov.. Two Chilean species are considered incertae sedis. A pictorial key and distribution maps for the species are provided.

Patterns of taxonomic diversity among terrestrial isopods.

The publication of the world catalog of terrestrial isopods some ten years ago by Schmalfuss has facilitated research on isopod diversity patterns at a global scale. Furthermore, even though we still lack a comprehensive and robust phylogeny of Oniscidea, we do have some useful approaches to phylogenetic relationships among major clades which can offer additional insights into isopod evolutionary dynamics. Taxonomic diversity is one of many approaches to biodiversity and, despite its sensitiveness to biases in taxonomic practice, has proved useful in exploring diversification dynamics of various taxa. In the present work, we attempt an analysis of taxonomic diversity patterns among Oniscidea based on an updated world list of species containing 3,710 species belonging to 527 genera and 37 families (data till April 2014). The analysis explores species diversity at the genus and family level, as well as the relationships between species per genera, species per families, and genera per families. In addition, we consider the structure of isopod taxonomic system under the fractal perspective that has been proposed as a measure of a taxon's diversification. Finally, we check whether there is any phylogenetic signal behind taxonomic diversity patterns. The results can be useful in a more detailed elaboration of Oniscidea systematics.

The terrestrial Isopoda (Crustacea, Oniscidea) of Rapa Nui (Easter Island), with descriptions of two new species.

Nine species of terrestrial isopods are reported for the Polynesian island of Rapa Nui (Easter Island) based upon museum materials and recent collections from field sampling. Most of these animals are non-native species, but two are new to science: Styloniscusmanuvaka sp. n. and Hawaiiosciarapui sp. n. Of these, the former is believed to be a Polynesian endemic as it has been recorded from Rapa Iti, Austral Islands, while the latter is identified as a Rapa Nui island endemic. Both of these new species are considered 'disturbance relicts' and appear restricted to the cave environment on Rapa Nui. A short key to all the oniscidean species presently recorded from Rapa Nui is provided. We also offered conservation and management recommendations for the two new isopod species.

Global diversification at the harsh sea-land interface: mitochondrial phylogeny of the supralittoral isopod genus Tylos (Tylidae, Oniscidea).

The supralittoral environment, at the transition between sea and land, is characterized by harsh conditions for life. Nonetheless, evolution of terrestrial isopods (Oniscidea), the only group of Crustacea fully adapted to live on land, appears to have involved a transitional step within the supralittoral. The two most basal oniscidean lineages (Ligiidae and Tylidae) have representatives that successfully colonized the supralittoral. One of them is the genus Tylos, which is found exclusively in supralittoral sandy beaches from tropical and subtropical coasts around the world. Comprehensive phylogenetic hypotheses for this genus are lacking, which are necessary for understanding the evolution and biogeography of a lineage that successfully diversified in the harsh sea-land interface. Herein, we studied the phylogenetic relationships among 17 of the 21 currently recognized species of the genus Tylos, based on sequences from four mitochondrial genes (Cytochrome Oxidase I, Cytochrome b, 16S rDNA, and 12S rDNA). Maximum Likelihood and Bayesian phylogenetic analyses identified several lineages with deep divergences and discrete geographic distributions. Phylogenetic and distributional patterns of Tylos provide important clues on the biogeography and evolution of this group. Large divergences among the most basal clades are consistent with ancient splits. Due to the biological characteristics of Tylos, which likely prevent dispersal of these isopods across vast oceanic scales, we argue that tectonic events rather than trans-oceanic dispersal explain the distribution of Tylos in different continents. Overwater dispersal, however, likely enabled range expansions within some basins, and explains the colonization of volcanic oceanic islands. Present-day distributions were also likely influenced by sea level and climate changes. High levels of allopatric cryptic genetic differentiation are observed in different regions of the world, implying that the dispersal abilities of Tylos isopods are more limited than previously thought. Our results indicate that a taxonomic revision of this group is necessary.

A complex evolutionary history in a remote archipelago: phylogeography and morphometrics of the Hawaiian endemic Ligia isopods.

Compared to the striking diversification and levels of endemism observed in many terrestrial groups within the Hawaiian Archipelago, marine invertebrates exhibit remarkably lower rates of endemism and diversification. Supralittoral invertebrates restricted to specific coastal patchy habitats, however, have the potential for high levels of allopatric diversification. This is the case of Ligia isopods endemic to the Hawaiian Archipelago, which most likely arose from a rocky supralittoral ancestor that colonized the archipelago via rafting, and diversified into rocky supralittoral and inland lineages. A previous study on populations of this isopod from O'ahu and Kaua'i revealed high levels of allopatric differentiation, and suggested inter-island historical dispersal events have been rare. To gain a better understanding on the diversity and evolution of this group, we expanded prior phylogeographic work by incorporating populations from unsampled main Hawaiian Islands (Maui, Moloka'i, Lana'i, and Hawai'i), increasing the number of gene markers (four mitochondrial and two nuclear genes), and conducting Maximum likelihood and Bayesian phylogenetic analyses. Our study revealed new lineages and expanded the distribution range of several lineages. The phylogeographic patterns of Ligia in the study area are complex, with Hawai'i, O'ahu, and the Maui-Nui islands sharing major lineages, implying multiple inter-island historical dispersal events. In contrast, the oldest and most geographically distant of the major islands (Kaua'i) shares no lineages with the other islands. Our results did not support the monophyly of all the supralittoral lineages (currently grouped into L. hawaiensis), or the monophyly of the terrestrial lineages (currently grouped into L. perkinsi), implying more than one evolutionary transition between coastal and inland forms. Geometric-morphometric analyses of three supralittoral clades revealed significant body shape differences among them. A taxonomic revision of Hawaiian Ligia is warranted. Our results are relevant for the protection of biodiversity found in an environment subject to high pressure from disturbances.

Evolution in Hawaiian cave-adapted isopods (Oniscidea: Philosciidae): vicariant speciation or adaptive shifts?

We assessed evolutionary relationships among Hawaiian cave-adapted isopods using a maximum-likelihood criterion to analyze cytochrome oxidase I nucleotide sequences. Results support morphological data that two genera of philosciid isopods have invaded caves independently in the islands. In the genus Littorophiloscia, a sister relationship between a surface-dwelling species, L. hawaiiensis, and an undescribed cave species was corroborated. This evidence, along with the known parapatric distributions between species, supports a speciation event by an adaptive shift on the island of Hawaii from a marine littoral to a terrestrial subterranean habitat. The monophyletic genus Hawaiioscia contains four known obligate cave-dwelling species, each of which occurs on a separate island. However, despite present-day allopatric distributions between Hawaiioscia species, the geographic and phylogenetic patterns are not sufficient to support a vicariant mode of speciation. Instead, we believe that the known species of Hawaiioscia evolved from a widespread ancestral surface species or a group of closely related species through multiple, independent adaptive shifts on each of the islands of Kauai, Oahu, Molokai, and Maui. This is the first molecular investigation of evolutionary relationships between surface-dwelling and cavernicolous arthropods in Hawaii and it suggests that simple vicariance is insufficient to explain the evolution of troglobites in tropical zones.