The collembolan fauna of Maestrazgo caves (Teruel, Spain) with description of three new species

Three new species of cavernicolous Collembola belonging to genera Pygmarrhopalites and Oncopodura from five caves at El Maestrazgo (Teruel, Spain) are described: Pygmarrhopalites maestrazgoensis sp. nov., P. cantavetulae sp. nov. and Oncopodura fadriquei Jordana & Baquero sp. nov. In addition five other species have been found in the same caves: Heteromurus nitidus (Templeton, 1836), Pseudosinella encrusae Gisin & Gama, 1969 (second record of the species), Megalothorax minimus Willem, 1900, Protaphorura aconae Arbea & Jordana 1994 (second record of the species), and Schaefferia decemoculata (Stach, 1939) (sensu: Thibaud 1970). The explorations of cave fauna in these cavities has been carried out by the "Associació Catalana de Bioespeleologia” under the sponsorship of CEMAT (Centro de Estudios del Maestrazgo Turolense)

Flourishing in subterranean ecosystems: Euro-Mediterranean Plusiocampinae and tachycampoids (Diplura, Campodeidae)

Este artículo contiene 138 páginas, 213 figuras, 14 tablas.Diplura is a group of entognathous hexapods, often considered a sister group to insects. They play an important role in recycling organic matter in soil and subterranean terrestrial ecosystems. The Campodeidae is the most diverse family, divided into four subfamilies. The subfamily Plusiocampinae has a subterranean life-style with many species distributed in the Euro-Mediterranean area. The incertae sedis tachycampoids (“lignée Tachycampoïde”) is a group within the family Campodeidae that share with the Plusiocampinae a strong preference for subterranean habitats and several morphological characters, such as slender body shape, elongated appendages, considerable increment in the number of antennomeres and cercal articles, and complexity of sensorial structures. The present monograph provides a taxonomic revision of the subfamily Plusiocampinae and the genera belonging to the tachycampoid lineage from Europe and the Mediterranean region. It comprises detailed morphological descriptions and illustrations together with data on the habitats and distributions of 87 species, 10 subspecies and 11 affinis forms. Seven new species are described among those, namely: Plusiocampa (Plusiocampa) apollo Sendra, Giachino & Vailati sp. nov., P. (P.) chiosensis Sendra & Gasparo sp. nov., P. (P.) dublanskii Sendra & Turbanov sp. nov., P. (P.) hoffmanni Sendra & Paragamian sp. nov., P. (P.) rhea Sendra sp. nov., P. (P.) ternovensis Sendra & Borko sp. nov. and P. (Venetocampa) ferrani Sendra & Delić sp. nov.DS’s field trips were funded by the Serbian Ministry of Education, Science and Technology (Grant 173038); KP was partially funded by the HISR project “Conservation of the Cave Fauna of Greece” funded by the MAVA Foundation and WWF Greece; PMG and DV carried out sampling in Greece using permits from the Ministry of Environment no. 124085/1362/2015, 135366/373/2016 and 166238/248/2018; IT was funded within the framework of the state assignment of FASO Russia (themes no. АААА-А18-118012690106-7 and АААА-А18-118012690105), supported in part by RFBR (project no. 17-54-40017Абх_а); sampling efforts by AF and colleagues resulted from various projects on the evolution of cave fauna diversity, some of them funded by the German Research Foundation, DFG (DFG FA 1042/1-1 and DFG BA 2152/14-1); PMG and DV were partly supported by the program “Research Missions in the Mediterranean Basin” sponsored by the World Biodiversity Association onlus XLI contribution; ASPSR was supported by a research grant (15471) from Villum Fonden.Peer reviewe

Early Pliocene range expansion of a clade of subterranean Pyrenean beetles

Aim To investigate the possibility of range expansion and diversification within the subterranean environment in a genus of troglobiont beetles of the family Leiodidae (Troglocharinus), which have a disjunct distribution between the Pyrenees and the Catalonian coast. Location North-eastern Iberian Peninsula. Methods We sequenced 4 kb of five mitochondrial and two nuclear genes of 50 specimens of 12 of the 18 species of Troglocharinus, plus several outgroups. We reconstructed a phylogeny using Bayesian inference and maximum likelihood, estimated divergence times using Bayesian probabilities and an a priori evolutionary rate, compared the diversification of the main clades within the genus, and reconstructed their ancestral distribution using maximum likelihood. Results We found strong support for the monophyly of Troglocharinus and the clades in each of the geographical areas, which diverged in the early Pliocene. The coastal clade was further divided into geographically well-defined lineages, separated by Quaternary deposits. The origin of the coastal clade was a single colonization in the early Pliocene from the central Pyrenees. The diversification of the Pyrenean clade followed a constant rate, while the diversification rate of the coastal clade significantly decreased through the Plio-Pleistocene transition. Main conclusions Troglocharinus expanded its range from its ancestral area in the central Pyrenees to the coast of Catalonia and subsequently diversified, probably within the subterranean environment. Our favoured scenario is a stepping-stone migration, with possibly short-distance dispersals through the surface, along the eastern margin of the north-eastern Ebro basin. The range expansion took place in a narrow temporal window with favourable conditions between the early Pliocene and the onset of the Mediterranean climate by the mid-Pliocene. Surface dispersal was probably severely limited afterwards, as shown by the fragmentation of the coastal lineage

FIGURES 1–4 in The collembolan fauna of Maestrazgo caves (Teruel, Spain) with description of three new species

FIGURES 1–4. Pygmarrhopalites maestrazgoensis sp. nov.: 1, chaetatoxy of head; 2, labrum with prelabral setae (rows: pl, prelabral; p, posterior; m, medial; a, anterior); 3, labial palp (posterior papillae drawn separately); 4, maxillary outer lobe (mx.p., maxillary palp; b.s., basal seta; sl.p., sublobal plate; s.f., oral fold

Late Miocene origin of an Ibero-Maghrebian clade of ground beetles with multiple colonizations of the subterranean environment

[Aim] To test different biogeographical scenarios for the evolution of the ground beetles of the Trechus fulvus group, a lineage with many narrowly distributed flightless subterranean species, highly suitable for tracing their biogeographical history.[Location] The Western Palaearctic, focusing on the Betic-Rifean area between south-east Iberia and north Morocco.[Methods] We sequenced 3.3 kb of four mitochondrial and two nuclear genes of 30 individuals of 15 species of the T. fulvus group, plus 29 outgroups. We reconstructed their phylogeny and estimated divergence times using Bayesian probabilities and a priori evolutionary rates, and their ancestral distribution using maximum likelihood.[Results] The phylogenetic reconstruction uncovered multiple independent colonizations of the subterranean environment within the T. fulvus group, a scenario also supported by variation in troglomorphic characters. Most of the Moroccan and south-east Iberian species form a clade with strong geographical structure, including the former genus Antoinella. The biogeographical model best fitting the current distribution and phylogeny of the group was a late Miocene palaeogeographical scenario with isolated populations on the Betic and Rifean areas and a south-eastern Iberian origin of the north Moroccan species. The widespread T. fulvus was sister to a central Moroccan species, suggesting a second Iberian-North African vicariance event within the group. One of the species, T. lallemantii, expanded its range to the east (Algeria and Tunisia) and the north (extreme south of the Iberian Peninsula) during the Pleistocene.[Main conclusions] The T. fulvus group originated in the early Miocene, and the south-eastern Iberian lineage in the Tortonian. The lineage dispersed to Morocco during the Messinian, diversifying in rapid succession in the Atlas and the Rif and colonizing the subterranean environment multiple times. The geography of the Betic-Rifean region at the end of the Miocene can still be traced from the distribution of the extant species of the group.© 2014 John Wiley & Sons Ltd.A.F. was supported by a postdoctoral Research Fellowship from the Alexander von Humboldt Foundation in the ZSM. The German Research Foundation (project FA1042/1-1) and the Spanish Ministry of Science and Innovation (project CGL2010-15755) provided financial support.Peer Reviewe

Early Pliocene range expansion of a clade of subterranean Pyrenean beetles

Aim To investigate the possibility of range expansion and diversification within the subterranean environment in a genus of troglobiont beetles of the family Leiodidae (Troglocharinus), which have a disjunct distribution between the Pyrenees and the Catalonian coast. Location North-eastern Iberian Peninsula. Methods We sequenced 4 kb of five mitochondrial and two nuclear genes of 50 specimens of 12 of the 18 species of Troglocharinus, plus several outgroups. We reconstructed a phylogeny using Bayesian inference and maximum likelihood, estimated divergence times using Bayesian probabilities and an a priori evolutionary rate, compared the diversification of the main clades within the genus, and reconstructed their ancestral distribution using maximum likelihood. Results We found strong support for the monophyly of Troglocharinus and the clades in each of the geographical areas, which diverged in the early Pliocene. The coastal clade was further divided into geographically well-defined lineages, separated by Quaternary deposits. The origin of the coastal clade was a single colonization in the early Pliocene from the central Pyrenees. The diversification of the Pyrenean clade followed a constant rate, while the diversification rate of the coastal clade significantly decreased through the Plio-Pleistocene transition. Main conclusions Troglocharinus expanded its range from its ancestral area in the central Pyrenees to the coast of Catalonia and subsequently diversified, probably within the subterranean environment. Our favoured scenario is a stepping-stone migration, with possibly short-distance dispersals through the surface, along the eastern margin of the north-eastern Ebro basin. The range expansion took place in a narrow temporal window with favourable conditions between the early Pliocene and the onset of the Mediterranean climate by the mid-Pliocene. Surface dispersal was probably severely limited afterwards, as shown by the fragmentation of the coastal lineage

Early Pliocene range expansion of a clade of subterranean Pyrenean beetles

Aim: To investigate the possibility of range expansion and diversification within the subterranean environment in a genus of troglobiont beetles of the family Leiodidae (Troglocharinus), which have a disjunct distribution between the Pyrenees and the Catalonian coast. Location: North-eastern Iberian Peninsula. Methods: We sequenced 4 kb of five mitochondrial and two nuclear genes of 50 specimens of 12 of the 18 species of Troglocharinus, plus several outgroups. We reconstructed a phylogeny using Bayesian inference and maximum likelihood, estimated divergence times using Bayesian probabilities and an a priori evolutionary rate, compared the diversification of the main clades within the genus, and reconstructed their ancestral distribution using maximum likelihood. Results: We found strong support for the monophyly of Troglocharinus and the clades in each of the geographical areas, which diverged in the early Pliocene. The coastal clade was further divided into geographically well-defined lineages, separated by Quaternary deposits. The origin of the coastal clade was a single colonization in the early Pliocene from the central Pyrenees. The diversification of the Pyrenean clade followed a constant rate, while the diversification rate of the coastal clade significantly decreased through the Plio-Pleistocene transition. Main conclusions: Troglocharinus expanded its range from its ancestral area in the central Pyrenees to the coast of Catalonia and subsequently diversified, probably within the subterranean environment. Our favoured scenario is a stepping-stone migration, with possibly short-distance dispersals through the surface, along the eastern margin of the north-eastern Ebro basin. The range expansion took place in a narrow temporal window with favourable conditions between the early Pliocene and the onset of the Mediterranean climate by the mid-Pliocene. Surface dispersal was probably severely limited afterwards, as shown by the fragmentation of the coastal lineage. © 2013 John Wiley & Sons Ltd.This work was funded by a postgraduate grant to V.R. (Università La Sapienza, Roma) and projects CGL2007-61943 (to A. Cieslak) and CGL2010-15755 (to I.R.).Peer Reviewe

Range expansion throughout subterranean environment in the genus Troglocharinus (Coleoptera, Leiodidae, Cholevinae)

Trabajo presentado en el II Iberian Congress of Biological Systematics (CISA2013), celebrado en Barcelona del 25 al 27 de septiembre de 2013.The tribe Leptodirini (Coleoptera, Leiodidae, Cholevinae) includes some 240 genera and 1800 mostly subterranean species with morphological and physiological characters related to their habitat: blindness, depigmentation, modifications of the life cycle. This tribe is a monophyletic radiation, distributed in the north side of the Mediterranean area, from the Iberian peninsula to the Middle East. Despite continuous attention from entomologists for the last two centuries, their phylogenetic relationships and evolutionary origin remain controversial. Particularly puzzling are the cases in which apparently closely related subterranean taxa have disjunct distributions separated by areas not suitable for subterranean species, such as sedimentary soils older than their assumed separation. Prominent among these is the beetle genus Troglocharinus, part of the Pyrenean lineage of subterranean Leptodirini (the Speonomus series). This genus has a disjunt distribution, with twenty species distributed in the coastal ranges of Catalonia (Garraf, San Llorenç del Munt i Obac and Montserrat), and twelve in the pre-Pyrenees (Serra del Montsec de Rubies, Serra del Boumort, Alt Urgell and Serra de Lleràs), with a single isolated species in Alto Aragòn. We established a robust phylogeny to study the evolution of this subterranean species radiation, and to provide a temporal framework for the diversification of various lineages and the colonization of the geographical areas in which they occur. For that purpose we used molecular phylogenies of six mitochondrial (cox1, cob, rrnL, trnL and nad1) and two nuclear (SSU and LSU) genes, using eleven species and seven subspecies. We found the respective monophyly of the Pyrenean and the coastal clades, with a strong geographical structuring within each of them, suggesting that the ancestor of the extant coastal Troglocharinus expanded its range from the central Pyrenees to the coastal area of Catalonia in the Early Pliocene, where it subsequently diversified during the late Pliocene and the Pleistocene. Our results add to the growing evidence of the possibility of range expansions in lineages fully adapted to the subterranean environment, without the need to invoke extinct multiple surface ancestors to explain the extraordinary diversity and distribution of the subterranean fauna.Peer Reviewe