New insights into the subterranean genus Speoplanes Müller, 1911 (Coleoptera, Leiodidae, Cholevinae) from the Dinarides.

In the present study morphological and molecular data from specimens of the troglobitic genus Speoplanes have been examined. The two subspecies Speoplanes giganteus giganteus Müller, 1911 and Speoplanes giganteus biocovensis Müller, 1934 are changed in rank from subspecies to species level and discrete identification characters for the separation of the two species are provided. Additionally, all known localities of the species (including new records) are listed and a distribution map of the genus is provided.

Phylogeny of the supertribe Nebriitae (Coleoptera, Carabidae) based on analyses of DNA sequence data.

The phylogeny of the carabid beetle supertribe Nebriitae is inferred from analyses of DNA sequence data from eight gene fragments including one nuclear ribosomal gene (28S), four nuclear-protein coding genes (CAD, topoisomerase 1, PEPCK, and wingless), and three mitochondrial gene fragments (16S + tRNA-Leu + ND1, COI ("barcode" region) and COI ("Pat/Jer" region)). Our taxon sample included 264 exemplars representing 241 species and subspecies (25% of the known nebriite fauna), 39 of 41 currently accepted genera and subgenera (all except Notiokasis and Archileistobrius), and eight outgroup taxa. Separate maximum likelihood (ML) analyses of individual genes, combined ML analyses of nuclear, nuclear protein-coding, and mitochondrial genes, and combined ML and Bayesian analyses of the eight-gene-fragment matrix resulted in a well-resolved phylogeny of the supertribe, with most nodes in the tree strongly supported. Within Nebriitae, 167 internal nodes of the tree (out of the maximum possible 255) are supported by maximum-likelihood bootstrap values of 90% or more. The tribes Notiophilini, Opisthiini, Pelophilini, and Nebriini are well supported as monophyletic but relationships among these are not well resolved. Nippononebria is a distinct genus more closely related to Leistus than Nebria. Archastes, Oreonebria, Spelaeonebria, and Eurynebria, previously treated as distinct genera by some authors, are all nested within a monophyletic genus Nebria. Within Nebria, four major clades are recognized: (1) the Oreonebria Series, including eight subgenera arrayed in two subgeneric complexes (the Eonebria and Oreonebria Complexes); (2) the Nebriola Series, including only subgenus Nebriola; (3) the Nebria Series, including ten subgenera arrayed in two subgeneric complexes, the Boreonebria and Nebria Complexes, with the latter further subdivided into three subgeneric subcomplexes (the Nebria, Epinebriola, and Eunebria Subcomplexes)); and (4) the Catonebria Series, including seven subgenera arrayed in two subgeneric complexes (the Reductonebria and Catonebria Complexes). A strong concordance of biogeography with the inferred phylogeny is noted and some evident vicariance patterns are highlighted. A revised classification, mainly within the Nebriini, is proposed to reflect the inferred phylogeny. Three genus-group taxa (Nippononebria, Vancouveria and Archastes) are given revised status and seven are recognized as new synonymies (Nebriorites Jeannel, 1941 and Marggia Huber, 2014 = Oreonebria Daniel, 1903; Pseudonebriola Ledoux & Roux, 1989 = Boreonebria Jeannel, 1937; Patrobonebria Bänninger, 1923, Paranebria Jeannel, 1937 and Barbonebriola Huber & Schmidt, 2017 = Epinebriola Daniel & Daniel, 1904; and Asionebria Shilenkov, 1982 = Psilonebria Andrewes, 1923). Six new subgenera are proposed and described for newly recognized clades: Parepinebriola Kavanaugh subgen. nov. (type species: Nebria delicata Huber & Schmidt, 2017), Insulanebria Kavanaugh subgen. nov. (type species: Nebria carbonaria Eschscholtz, 1829), Erwinebria Kavanaugh subgen. nov. (type species Nebria sahlbergii Fischer von Waldheim, 1828), Nivalonebria Kavanaugh subgen. nov. (type species: Nebria paradisi Darlington, 1931), Neaptenonebria Kavanaugh subgen. nov. (type species: Nebria ovipennis LeConte, 1878), and Palaptenonebria Kavanaugh subgen. nov. (type species: Nebria mellyi Gebler, 1847). Future efforts to better understand relationships within the supertribe should aim to expand the taxon sampling of DNA sequence data, particularly within subgenera Leistus and Evanoleistus of genus Leistus and the Nebria Complex of genus Nebria.

A new cavernicolous species of the genus Bergrothia Reitter, 1884 (Coleoptera, Staphylinidae, Pselaphinae) from Georgia.

During a biospeleological survey of caves of Imereti region (Georgia), a new species of the beetle genus Bergrothia Reitter, B. barbakadzei sp. nov. was discovered in three caves: Datvis (Bear), Prometheus and Melouri caves (Sataplia-Tskaltubo karst massif, Tskaltubo district). The new species is described, illustrated and differentiated from other morphologically similar species of the genus-B. mingrelica (Reitter). Bergrothia barbakadzei sp. nov. is the first cavernicolous species of the genus Bergrothia. A key to the species of this genus is provided.

A molecular phylogeny of Speonemadus Jeannel, 1922 with description of two new species from Morocco (Coleoptera: Leiodidae: Cholevinae: Anemadini).

The genus Speonemadus Jeannel, 1922 currently includes 12 species distributed in the Western Mediterranean (Iberian Peninsula, North Africa, Sicily and continental Italy). Two new species are described from Morocco, S. brusteli sp. n. from the Rif and S. comasi sp. n. from the High Atlas, the first in the S. vandalitiae-group of species and the second with an isolated position within the genus. Two species are also resurrected from synonymy, S. pulchellus (Reitter, 1885) stat. rest. (former synonym of S. orchesioides (Fairmaire, 1879)) and S. gracilis (Kraatz, 1870) stat. rest. (former synonym of S. vandalitiae (Heyden, 1870)), raising the total number of species of the genus to 16. Lectotypes are designated for Anemadus pulchellus Reitter, 1885, and Anemadus tenuipes Peyerimhoff, 1917. A molecular phylogeny of the genus Speonemadus is presented, based on a combination of mitochondrial and nuclear genes and including seven of the currently recognised species, one of the newly described (S. brusteli sp. n.) and the two reinstated taxa. The genus is estimated to have diversified from the late Miocene (Tortonian), with successive cladogenetic events related to the isolation of landmasses between Iberia and North Africa, including the opening of the Strait of Gibraltar at the end of the Messinian. The distribution of some of the species of Speonemadus remains still uncertain, due to their frequent misidentification and the problems with their taxonomic status.

Phylogeny of the beetle supertribe Trechitae (Coleoptera: Carabidae): Unexpected clades, isolated lineages, and morphological convergence.

Using data from two nuclear ribosomal genes and four nuclear protein-coding genes, we infer a well-resolved phylogeny of major lineages of the carabid beetle supertribe Trechitae, based upon a sampling of 259 species. Patrobini is the sister group of Trechitae, but the genus Lissopogonus appears to be outside of the Patrobini + Trechitae clade. We find that four enigmatic trechite genera from the Southern Hemisphere, Bembidarenas, Argentinatachoides, Andinodontis, and Tasmanitachoides, form a clade that is the sister group of Trechini; we describe this clade as a new tribe, Bembidarenini. Bembidarenini + Trechini form the sister group of remaining trechites. Within Trechini, subtribe Trechodina is not monophyletic, as three trechodine genera from Australia (Trechobembix, Paratrechodes, Cyphotrechodes) are the sister group of subtribe Trechina. Trechini appears to have originated in the continents of the Southern Hemisphere, with almost all Northern Hemisphere lineages representing a single radiation within the subtribe Trechina. We present moderate evidence that the geographically and phylogenetically isolated genera Sinozolus (six species in the mountains of China), Chaltenia (one species in Argentina and Chile), and Phrypeus (one species in western North America) also form a clade, the tribe Sinozolini. The traditionally recognized tribe Bembidiini sens. lat., diagnosed by the presence of a subulate terminal palpomere, is shown to be polyphyletic; subulate palpomeres have arisen five times within Trechitae. Anillini is monophyletic, and the sister group of Tachyini + Pogonini + Bembidiini + Zolini + Sinozolini; within anillines, we confirm earlier results indicating the eyed New Zealand genus Nesamblyops as the sister to the rest. Sampled New World Pogonini are monophyletic, rendering the genus Pogonus non-monophyletic. Tachyina and Xystosomina are sister groups. Within Xystosomina, the New World members are monophyletic, and are sister to an Australia-New Zealand clade. The latter consists of the genus Philipis as well as taxa not previously recognized as xystosomines: Kiwitachys, the "Tachys" ectromioides group, and "Tachys" mulwalensis. Within Tachyina, the subgenus Elaphropus is not closely related to other subgenera previously placed in the genus Elaphropus; we move the other subgenera into the genus Tachyura. Tachyina with a bifoveate mentum do not form a clade; in fact, a bifoveate mentum is found in Xystosomina, Sinozolini, Trechini, Trechitae and its sister group, Patrobini. Extensive homoplasy in the morphological characters previously used as key indicators of relationship is supported by our results: in addition to multiple origins of subulate palpomeres and bifoveate menta, a concave protibial notch has arisen independently in Anillina, Xystosomina, and Tachyina. Phylogenetically and geographically isolated, species-poor lineages in Trechini, Bembidarenini, and Sinozolini may be relicts of more widespread faunas; many of these are found today on gravel or sand shores of creeks and rivers, which may be an ancestral habitat for portions of Trechitae. In addition to the description of Bembidarenini, we present a diagnosis of the newly delimited Sinozolini, and keys to the tribes of Trechitae.

Contributions to the knowledge of the genus Tonkinaphaenops Deuve, 2013 from China and Vietnam (Coleoptera: Carabidae: Trechinae).

Cavernicolous trechine beetles of the genus Tonkinaphaenops Deuve, 2013, known so far only from Vietnam, are reported for the first time from southern China, and new data are also provided regarding this genus from northern Vietnam. Two new Chinese species, T. yinquanicus sp. nov. from cave Yinquan Dong and T. jingxicus sp. nov. from cave Nianluo Dong in Jingxi County, southwesternmost Guangxi Zhuang Autonomous Region, are described and illustrated, together with the descriptions of two new Vietnamese species: T. anthonyi sp. nov. from the cave Hang Dõi and T. impunctatus sp. nov. from the cave Hang Ran in Cao Bang province. A key to all known species of Tonkinaphaenops is also provided.

A remarkable new species of cavernicolous Collartidini from Madagascar (Hemiptera: Heteroptera: Reduviidae).

Mangabea troglodytes sp. nov. (Hemiptera: Heteroptera: Reduviidae: Emesinae) is described based on four specimens collected in a cave of the Namoroka Karstic System, Madagascar, and deposited in the Collection of the Muséum National d'Histoire Naturelle, Paris. The dorsal habitus as well as diagnostic characters of male and female genitalia are extensively illustrated and imaged. A key to species of the genus Mangabea Villiers, 1970 is provided and the degree of cave specialization of the new species is discussed.

On the genus Aphaobius Abeille de Perrin, 1878, with description of a new species from the mesovoid shallow substratum (MSS) of Austria (Coleoptera: Leiodidae: Cholevinae: Leptodirini).

Aphaobius haraldi sp. n. from the mesovoid shallow substratum (MSS) of the Austrian Alps is described, illustrated, and compared with the closest species of the genus. The new species belongs to the A. kraussi species group, formerly including five species. It can be readily separated from other species of the genus by the large parameres, with an enlarged apical part, a unique feature among species of Aphaobius. The phylogenetic position of the new species is clarified using mitochondrial and nuclear data of four related species of Aphaobius, plus some representatives of related genera from the same geographic area. The diversification of the group was estimated to be recent, dating from the Pleistocene. New records and molecular data are provided regarding the enigmatic monospecific genus of the Austrian Alps, Lotharia Mandl, 1944, which was found to be sister to the studied species of Aphaobius.

Thermal niche estimators and the capability of poor dispersal species to cope with climate change.

For management strategies in the context of global warming, accurate predictions of species response are mandatory. However, to date most predictions are based on niche (bioclimatic) models that usually overlook biotic interactions, behavioral adjustments or adaptive evolution, and assume that species can disperse freely without constraints. The deep subterranean environment minimises these uncertainties, as it is simple, homogeneous and with constant environmental conditions. It is thus an ideal model system to study the effect of global change in species with poor dispersal capabilities. We assess the potential fate of a lineage of troglobitic beetles under global change predictions using different approaches to estimate their thermal niche: bioclimatic models, rates of thermal niche change estimated from a molecular phylogeny, and data from physiological studies. Using bioclimatic models, at most 60% of the species were predicted to have suitable conditions in 2080. Considering the rates of thermal niche change did not improve this prediction. However, physiological data suggest that subterranean species have a broad thermal tolerance, allowing them to stand temperatures never experienced through their evolutionary history. These results stress the need of experimental approaches to assess the capability of poor dispersal species to cope with temperatures outside those they currently experience.

Gondwanian relicts and oceanic dispersal in a cosmopolitan radiation of euedaphic ground beetles.

Anillini are a tribe of minute, euedaphic ground beetles (Carabidae) characterized by the loss of eyes, loss of wings and high levels of local endemism. Despite their presumed low dispersal, they have a nearly cosmopolitan distribution, including isolated islands such as New Zealand and New Caledonia. We used a time calibrated molecular phylogeny to test, first, if the tribe as currently understood is monophyletic and, second, whether the time of divergence is compatible with an early vicariant diversification after the breakup of Gondwana. We sequenced portions of 6 mitochondrial and 3 nuclear genes for 66 specimens in 17 genera of Anillini plus 39 outgroups. The resulting phylogenetic tree was used to estimate the time of diversification using two independent calibration schemes, by applying molecular rates for the related genus Carabus or by dating the tree with fossil and geological information. Rates of molecular evolution and lineage ages were mostly concordant between both calibration schemes. The monophyly of Anillini was well-supported, and its age was consistent with a Gondwanian origin of the main lineages and an initial diversification at ca. 100Ma representing the split between the eyed Nesamblyops (New Zealand) and the remaining Anillini. The subsequent diversification, including the split of the Nearctic Anillinus and the subsequent splits of Palaearctic lineages, was dated to between 80 and 100Ma and thus was also compatible with a tectonic vicariant origin. On the contrary, the estimated age of the New Caledonian blind Orthotyphlus at ca. 30±20Ma was incompatible with a vicariant origin, suggesting the possibility of trans-oceanic dispersal in these endogean beetles.

One new species of Ptomaphaminus Perreau, 2000 (Coleoptera: Leiodidae: Cholevinae: Ptomaphagini) from a granitic subterranean environment in Vietnam.

Ptomaphaminus granophilus n. sp. from Vietnam is described and illustrated. The special subterranean biotope of this species, deep interstices in a granitic blockfield, is emphasized.

Confirmation of occurrence of Trechus Clairville, 1806 in the Baltic amber forests, with description of a flightless edaphic species, and remarks on Trechoides Motschulsky, 1856 (Coleoptera: Carabidae: Trechini).

The first reliably identified Trechini beetle species from Baltic amber, Trechus balticus sp. n., is described. This new fossil species is most similar to extant species of the Palaearctic Trechus sensu stricto, as well as to the Nearctic Microtrechus Jeannel, 1927. Since the fossil specimen is a female and some important character states cannot be investigated, its true systematic position within the genus Trechus sensu lato remains open. Trechoides fasciatus Motschulsky, 1856 is another fossil species that perhaps belongs to the Trechini clade. However, since the original description does not present any character of systematic value and, because the type is very probably lost, the true systematic position of this taxon remains unknown.

On the Way to Speciation: Shedding Light on the Karstic Phylogeography of the Microendemic Cave Beetle Aphaenops cerberus in the Pyrenees.

The highly modified morphology and ecological features of cave-dwelling organisms are a strong obstacle to dispersion. Hence, they represent ideal models for the study of historical biogeography at both large and fine timescales. Here, we study the phylogeography of Aphaenops cerberus, an endemic hypogean ground beetle with a fragmented distribution in the French Northern Pyrenees. We extracted 75 exemplars of 17 populations of A. cerberus and sequenced one mitochondrial and one nuclear marker to assess the geographic structuration as well as the recent biogeographic history of this species. We used Bayesian Inference and Maximum Likelihood to reconstruct the relationships among most of the extant populations of this species across its distributional range. We inferred divergence time estimates using carabid substitution rates and reconstructed haplotype networks to investigate the recent biogeographic history of this lineage. We recover a strong geographic structuration of the populations across the mountain range. The strong impact of geology on the structure of the populations is evidenced although geological continuity does not systematically lead to continual gene flow. The origin of the species is dated from the Early Pleistocene and the dispersal predates the main Last Glacial Maximum. Our results indicate broad similitudes between islands and karsts, which make cave organisms an excellent model for the study of evolution mechanisms.

Convergent Reduction of Ovariole Number Associated with Subterranean Life in Beetles.

BACKGROUND: Some species of obligate cavernicolous beetles are known to possess a unique feature-a contraction of the larval cycle. In contrast to many other subterranean beetles, life-cycle contraction in Trechini ground beetles (Carabidae) is correlated with a reduction in the number of eggs and a drastic reduction in the number of ovarioles. This remarkable peculiarity has only been reported for a small number of closely related species. RESULTS: We give a description of the female internal reproductive system for six species of Trechini, including five subterranean species, with a particular focus on the western Pyrenean radiation of Aphaenops, a group for which nothing is known regarding the early life stages. We redescribe the internal female genitalia of A. crypticola Linder. Study of the ovarioles allowed us to infer the postembryonic development of the larvae for each species examined. We then used a phylogenetic framework to recognize two independent reductions in the number of ovarioles in the Pyrenean lineage. We discuss the multiple convergent evolutions in ovariole number and the potential link between a reduction of ovariole number and troglobiomorphism in a phylogenetic context. CONCLUSIONS: There is an extreme reduction in ovariole number and size within the species studied; the eggs produced by small ovarioles have a remarkably large size. A reduction to one ovariole has occurred independently at least twice in this subterranean group. A reduction in the number of ovarioles in ground beetles is one of the striking consequences of subterranean specialization and it is correlated with another remarkable adaptation of subterranean beetles, a reduction in the number of larval instars.

A molecular phylogeny of Alpine subterranean Trechini (Coleoptera: Carabidae).

BACKGROUND: The Alpine region harbours one of the most diverse subterranean faunas in the world, with many species showing extreme morphological modifications. The ground beetles of tribe Trechini (Coleoptera, Carabidae) are among the best studied and widespread groups with abundance of troglobionts, but their origin and evolution is largely unknown. RESULTS: We sequenced 3.4 Kb of mitochondrial (cox1, rrnL, trnL, nad1) and nuclear (SSU, LSU) genes of 207 specimens of 173 mostly Alpine species, including examples of all subterranean genera but two plus a representation of epigean taxa. We applied Bayesian methods and maximum likelihood to reconstruct the topology and to estimate divergence times using a priori rates obtained for a related ground beetle genus. We found three main clades of late Eocene-early Oligocene origin: (1) the genus Doderotrechus and relatives; (2) the genus Trechus sensu lato, with most anisotopic subterranean genera, including the Pyrenean lineage and taxa from the Dinaric Alps; and (3) the genus Duvalius sensu lato, diversifying during the late Miocene and including all subterranean isotopic taxa. Most of the subterranean genera had an independent origin and were related to epigean taxa of the same geographical area, but there were three large monophyletic clades of exclusively subterranean species: the Pyrenean lineage, a lineage including subterranean taxa from the eastern Alps and the Dinarides, and the genus Anophthalmus from the northeastern Alps. Many lineages have developed similar phenotypes independently, showing extensive morphological convergence or parallelism. CONCLUSIONS: The Alpine Trechini do not form a homogeneous fauna, in contrast with the Pyrenees, and show a complex scenario of multiple colonisations of the subterranean environment at different geological periods and through different processes. Examples go from populations of an epigean widespread species going underground with little morphological modifications to ancient, geographically widespread lineages of exclusively subterranean species likely to have diversified once fully adapted to the subterranean environment.

Discovery of a new blind ground beetle in western French Pyrenees, and its relevance to the phylogeny of Pyrenean hypogean Trechini.

Geotrechus sarpedon sp. n., a new species of blind Trechini is described from Arbailles Massif, western Pyrenees, trance. The generic attribution of the new species was challenging as it shares morphological features of both genera Geotrechus Jeannel, 1919 and Aphaenops subgenus Hydraphaenops Jeannel, 1926, confirming their polyphyly, which has been highlighted by previous molecular studies on the group. Molecular data suggests unambiguously strong affinities between the new species and the clade Aphaenops vasconicus Jeannel, 1913-A. galani Espanol, 1968 occurring in the same geographic area, and not related with Aphaenops ehlersi (Abeille de Perrin, 1872), type species of the subgenus Hydraphaenops. The species is included in the paraphyletic genus Geotrechus until its affinities with the other species of Geotrechus and A. (Hydraphaenops) are clarified.

Molecular phylogeny of the Trechus brucki group, with description of two new species from the Pyreneo-Cantabrian area (France, Spain) (Coleoptera, Carabidae, Trechinae).

A molecular phylogeny of the species from the Trechus brucki clade (previously Trechus uhagoni group)based on fragments of four mitochondrial genes and one nuclear gene is given. We describe Trechus (Trechus) bouillonisp. n. from the western pre-Pyrenees: Sierras de Urbasa-Andía, Navarra, Spain. The species was collected in mesovoid shallow substratum (mss), a subterranean environment. Molecular as well as morphological evidences demonstrate that the new species belongs to the Trechus brucki clade. A narrow endemic species of high altitude in western French Pyrenees merged with Trechus brucki Fairmaire, 1862a, Trechus bruckoidessp. n., is described. A lectotype is designated for Trechus brucki and Trechus planiusculus Fairmaire, 1862b (junior synonym of Trechus brucki). The species group is redefined based on molecular and morphological characters, and renamed as the brucki group, as Trechus brucki was the first described species of the clade. A unique synapomorphy of the male genitalia, a characteristic secondary sclerotization of the sperm duct, which is shared by all the species of the brucki group sensu novo, is described and illustrated. The Trechus brucki group sensu novo is composed of Trechus beusti (Schaufuss, 1863), Trechus bouillonisp. n., Trechus brucki, Trechus bruckoidessp. n., Trechus grenieri Pandellé, 1867, T. uhagoni uhagoni Crotch, 1869, T. uhagoni ruteri Colas, 1935 and Trechus pieltaini Jeannel, 1920. We discuss the taxonomy of the group and provide illustrations of structures showing the differences between the species, along with distribution data and biogeographical comments.