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Nouvelle classification des Bathyscnnae (Coléoptères, Catopidae), tableau des sous tribus, groupes de genres et genres
... In , the group was revised as a whole (under the name 'Bathysciinae Horn, 1880') for the first time, and, in Jeannel (1955), what is still considered by some authors to be the main division within the lineage was established, based on the structure of the basal region of the internal sac of the aedeagus: the 'supraflagellates' (with the internal sac of the aedeagus with a basal, dorsal flagellum) and the 'infraflagellates' (with the internal sac of the aedeagus with a basal, Y-shaped ventral piece, the 'Y-piece') (see also Giachino et al., 1998). Laneyrie (1967Laneyrie ( , 1969Laneyrie ( , 1978 proposed an alternative ordination in four main lineages, also based on the structures of the basal region of the internal sac of the aedeagus. Gue´orguiev (1974aGue´orguiev ( , b, 1976) formalized Jeannel's system in seven subtribes, followed by Newton (1998) (although with the recognition of their non-monophyly), and partly by Perreau (2004), defined on external morphology (insertion of antennae, tarsal formula, number, type and arrangement of the tibial spines). ...
... Species with mesocoxal cavities fused (character 8 ¼ 0) are coded as missing data. This character was mentioned by Laneyrie (1967) for some eastern Mediterranean species of Leptodirini. One change. ...
Abstract The tribe Leptodirini (Leiodidae: Cholevinae) is one of the largest radiations of Coleoptera in the subterranean environment. Although subjected to systematic and evolutionary studies, the phylogeny remains poorly understood. We assessed the phylogeny of the western Mediterranean lineages (Iberian Peninsula, Pyrenees and Sardinia) based on a cladistic analysis of fourteen characters of external morphology and twenty characters of the male and female genitalia, studied in 182 species belonging to thirty-nine genera. We tested the monophyly of the traditional two main divisions of the group (infraflagellates and supraflagellates), as well as that of some ‘phyletic series’. The final matrix contained fifty-eight terminal taxa, twenty-four of which had different character state combinations. The strict consensus of the sixty most parsimonious trees recovered a monophyletic Leptodirini, but not their separation into infraflagellates and supraflagellates. The supraflagellates formed a paraphyletic group with respect to the infraflagellates (corresponding to our sampled ‘Speonomus’ series), with Notidocharis sister to all other included Leptodirini, and Speonomidius sister to Leptodirini excluding Notidocharis. The series ‘Spelaeochlamys’, including the Sardinian genera but excluding Pseudochlamys, was recovered as monophyletic with weak support. The ‘Quaestus’ series formed a polytomy with Pseudochlamys plus the ‘Speonomus’ series (including Bathysciola), which was recovered as monophyletic with strong support. Speonomus, Bathysciola, Quaestus and Troglophyes were para- or polyphyletic. Our results suggested the respective monophyletic origin of the Leptodirini from the Pyrenees (Pseudochlamys plus the ‘Speonomus’ series) and the Mediterranean coast plus Sardinia (series ‘Spelaeochlamys’). On the contrary, the Leptodirini of the Atlantic north coast of the Iberian Peninsula (series ‘Quaestus’ and ‘Speonomidius’) were not monophyletic.
... Finally, Jeannel (1955) established what is considered to be the most primitive division of the Bathysciinae: "supraflagellates" and "infraflagellates". A decade later Laneyrie (1967Laneyrie ( , 1969Laneyrie ( and 1978 proposed a new ordenation based on the copulatory shield of the internal sac, basically the structures in the basal region. Guéorguiev (1974aGuéorguiev ( , 1974bGuéorguiev ( and 1976 proposed a new ordenation using the external morphology: the series of combs and spurs on the tibiae. ...
... All the insects in this study should be included in the Leptodirinae infraflagellates. Contributions by Jeannel (1924aJeannel ( , 1955, Laneyrie (1967), Guéorguiev (1974a, b and1976), Casale et al. (1991) and Giachino et al. (1998) are considered, but the meaning given to the term infraflagellates is that contributed by Dupré (1992) who redefines this group based on the study of histological sections of the internal sac of the aedeagus: models of internal sac with a Y-shaped piece in the basal region, and dorsal phanerae in the median region. The spermathecal complex in all these genera correspond to "type 1" of the spermatheca defined by Perreau (1989). ...
Se realiza una revisión taxonómica de las secciones IV, VI y VII (Jeannel, 1924) con las especies Ibéricas y de los Pirineos del género Bathysciola. Esta propuesta de ordenación tiene como base el examen de las estructuras genitales de los dos sexos, y de un modo especial el estudio de la armadura esclerotizada del saco interno del edeago. Se han diferenciado los siguientes grupos de especies en el género Bathysciola Jeannel, seccion IV: grupo aubei de Peyerimhoff (1 taxon): B. aubei (Kiesenwetter); madoni nuevo grupo (2 táxones): B. madoni Jeannel y B. penicillata Jeannel; zariquieyi nuevo grupo (2 táxones): B. zariquieyi zariquieyi Bolívar y B. zariquieyi serratensis Coiffait; sección VII o grupo ovata de Perreau (8 táxones): B. asperula asperula (Fairmaire), B. asperula subasperata (Saulcy) (= intermedia Jeannel), B. ovata ovata (Kiesenwetter), B. ovata aragonica Coiffait, B. ovata catalana Coiffait, B. ovata gabasensis Hustache, B. simonis (Abeille de Perrin) y B. talpa (Normand); sección VI: lapidicola nuevo grupo (4 táxones): B. arcuatipes Jeannel, B. lapidicola lapidicola (Saulcy), B. lapidicola rectipes Coiffait y B. lapidicola simplex Coiffait; meridionalis nuevo grupo (3 táxones): B. finismillennii sp. n., B. grenieri (Saulcy) y B. meridionalis (J. Du Val) (= nitidula Normand); larcennei nuevo grupo (2 táxones): B. convena Jeannel, nueva posición en el grupo, y B. larcennei (Abeille de Perrin); grupo schiodtei de Perreau (9 táxones): B. bigerrica Jeannel, nueva posición en el grupo (= convexa Coiffait n. syn.), B. breuili Bolívar (= azuai Bolívar n. syn.), B. diegoi Salgado & Fresneda, B. fauveli Jeannel, B. grandis (Fairmaire), B. obermaieri Bolívar, B. parallela (Jeannel), B. rugosa (Sharp) y B. schiodtei (Kiesenwetter) (= navarica Coiffait). Incertae sedis: Bathysciola aranensis Coiffait y Bathysciola minuscula (Abeille de Perrin).
... (Jeannel, 1914); Cavernicole, Herzégovine: grottes du Vysočica planina (C. Setnik) (Jeannel, 1924); Yougoslavie (Herzégowine) (Laneyrie, 1967); BH (H: Visočica planina), endem. (Pretner, 1968); Bosnia Herzégovine (Guéorguiev, 1976); v jamach na Visočici v Hercegovini (Pretner, 1970); Bosnia, Herzegovina (Newton, 1998); Bosnie-Herzégovine: cavités du Visočica planina (Perreau, 2000); BH (Visočica planina) (Perreau, 2015); BiH (Visočica planina) (Hlaváč, Perreau & Čeplík, 2017); Buca del Viglie, Luka, Zavodnica, Visočica pl. ...
A redescription of the genus Antrosedes Reitter, 1912 is given. An identification key for both species of the genus, as well as data on their distribution is provided.
... The classification of Cholevinae, and in particular of the Leptodirini tribes has been the subject of much controversy (Giachino et al. 1998;Newton 1998;Perreau 2015;Fresneda et al. 2007). Jeannel (1910aJeannel ( , b, 1911Jeannel ( , 1924Jeannel ( , 1955) established a system of informal groups ('phyletic series') based on external morphological characters and male genitalia, later on, other authors proposed alternative ordinations (Laneyrie 1967;Guéorguiev 1976;Giachino et al. 1998). The phyletic series "Bathysciola" (sensu Zoia & Rampini 1994) of the Leptodirini tribe includes four genera: Bathysciola Jeannel, 1910, Besuchetiola Rampini andZoia, 1991 (one species), Parabathyscia Jeannel, 1908 (forty-one species) and Sengletiola Zoia & Rampini, 1994 (one species) (Perreau 2000). ...
The genus Bathysciola is widely distributed in the northern Mediterranean region, although its range extends east to the Caucasus and Iran. More than 130 species belonging to this genus are actually known in the whole geographic distribution area and 45 species are distributed in continental and insular Italy. The species belonging to the Bathysciola sisernica Cerruti and Patrizi, 1952 species group occur in the Central-Southern Italian Apennines and Pre-Apennines. This group consists of seven species, four of which (B. sisernica, B. delayi Latella and Rampini, 1994, B. rampinii Latella, 2002, B. sbordoni Rampini and Latella, 1993) were already known to science and three are described herein, Bathysciola fabiolae sp. nov., Bathysciola octaviani sp. nov., and Bathysciola valeriae sp. nov., markedly increasing the knowledge on the distribution of this genus in Central Italy. A morphological analysis was carried out based on diagnostic characters usually used to distinguish different taxa, and including both genitalia and external traits. Based on morphological characters, we reconstructed the phylogeny of this group of species, comparing them with the species belonging to other phyletic lineages, such as B. derosasi Jeannel, 1914, B. georgi Cerruti, Patrizi, 1952, B. vignai Sbordoni and Rampini, 1978, and B. sarteanensis sarteanensis (Bargagli, 1870). Results suggested that morphological traits show a clear taxonomic signal but a poor phylogenetic signal. To better understand the relationships within this group of species, we performed a molecular analysis by sequencing three mitochondrial genes, 12S rRNA, 16S rRNA, partially sequenced and the entire gene of COI. Molecular markers were used to infer phylogenetic relationships among the Bathysciola sisernica species group and to reconstruct the historical processes that shaped their current geographic distribution. Results showed that these species became isolated in very ancient times, showing very high genetic differentiation.
... Three species and one subspecies of the genus Aphaobiella Pretner, 1949, all from Slovenia, are presently known: A. budnarlipoglavseki budnarlipoglavseki Pretner, 1949, A. budnarlipoglavseki mozirjensis Pretner, 1949, A. tisnicensis Pretner, 1949and A. mlejneki J. Moravec, 1996(Pretner 1949Laneyrie 1967;Guéorguiev 1976;Moravec,1996;Perreau 2000Perreau , 2004. ...
Aphaobiella kofleri sp. n. from Grintavec, Zgornje, Ravni (Slovenia) is described and illustrated. The description of this new species, closely related to A. budnarlipoglavseki budnarlipoglavseki Pretner, 1949, increases the zoogeographical knowledge of this genus endemic to Slovenia.
... Nuoro prov.: Ulassai, Su Marmuri, Grotta di Su Marmori or Grutta de su Marmuri (55 Sa/ NU) (loc. typ.) (Fairmaire 1872, as Adelops Gestroi n. sp.;Gestro 1904;Jeannel 1924;Müller 1930;Barajon 1955;Laneyrie 1967;Altara 1968;Cerruti 1968;Cassola 1982;Sbordoni et al. 1982;Puddu 1983;De Waele 1996;Grafitti 1999b;De Waele & Grafitti 2000;Grafitti 2001a;De Waele & Grafitti 2004;Grafitti , 2009); Ulassai, Grotta sa Foxi 'e s'Abba (728 Sa/NU) (Gestro 1904;Grafitti 1999bGrafitti , 2009); Ulassai, Grutta de su Segretariu or "Reale" or "de is Quaddus" (661 Sa/ NU) (Grafitti 1999b(Grafitti , 2009); Lecorci, Grotta di Lecorci (660 Sa/NU) (Grafitti 1999b(Grafitti , 2009); Taccu de Ulassai, Truculu, Grutta de Is Lianas or Grotta Truculu (193 Sa/NU) (Grafitti 1999b(Grafitti , 2009); Ulassai, Monte Tisiddu, Grotta del Porcellino (692 Sa/NU) (Grafitti 1999b(Grafitti , 2009, Monte Tisiddu, Grutta de is Janas (715 Sa/NU) (Grafitti 1999b(Grafitti , 2009, Monte Tisiddu, Grutta de is Chillottis (727 Sa/ NU) (Grafitti 1999b(Grafitti , 2009). Gairo, Genna 'e Ua, Grotta di Genna 'e Ua (43 Sa/NU) (Gestro 1904;Jeannel 1911Jeannel , 1924, as "Grotte près de la gare de Gairo";Müller 1930;Cerruti 1968;Puddu 1970;); Gairo, Taccu Arba, Taquisara, Grotta di Taquisara o del Marmo (86 Sa/NU) (Puddu 1970;Grafitti 2004;); Gairo, Taquisara ); Gairo, Taccu de Ulassai, Serbissi, Grotta di Serbissi (669 Sa/NU, on the border with the municipality of Osini) (Grafitti 1999b(Grafitti , 2009); Serbissi, Grutta de su Coloru (670 Sa/NU) (Grafitti 1999b(Grafitti , 2009); Serbissi, Grotta del Macigno or "degli Pseudoscorpioni" (2350 Sa/NU) (Grafitti 1999b(Grafitti , 2009); Serbissi, Grotta delle Vaschette (2352 Sa/NU) (Grafitti 1999b(Grafitti , 2009. ...
We present a synthesis of the knowledge on the cholevid fauna of Sardinia, with an annotated checklist and an updated list of both published and unpublished collecting localities. The Cholevidae (Coleoptera, Staphylinoidea) are a group of small to medium-sized beetles, most of which are saprophagous; a significant percentage of Cholevidae are cave-dwellers, and some are highly specialized troglobitic elements. In Sardinia, twenty-five species have been recorded so far. More than half of them are endemic to the island, while some are sub-endemic (i.e. present also in Corsica or in Sicily and/or the Balearic Islands); nine species, belonging to the genera Ovobathysciola and Patriziella, are specialized hypogean elements. Four more species, belonging to the genus Bathysciola, are soil-inhabiting or cave-dwellers. The authors stress the high biogeographic and biospeleological interest of Sardinian cholevids. In particular, the colonization processes appear to be in good agreement with the current hypotheses about the origin and composition of the existing fauna of Sardinia. The hypogean species of the genera Ovobathysciola, Patriziella and species of Speonomus of the subgenus Batinoscelis are currently accepted as being related to the Baetic and Pyrenean lineages of Anillochlamys and Speonomus, respectively; therefore, they should belong to the most ancient stock of elements, originated during the Miocene drift of the Corso-Sardinian massif in the western Mediterranean. Some other endemic or sub-endemic taxa, with W-Mediterranean affinities, are probably derived from more recent colonizers as a consequence of the several connection/isolation phases which existed between Sardinia, Corsica, and other Mediterranean areas during the late Miocene (Messinian) salinity crisis, and during the Plio-Pleistocene ice-phase-related strong variations in the sea level. Finally, some widespread taxa, all epigean and winged, reached the island in more recent times, through active and passive dispersal.
... Horn (1880) was evidently the first to use the name Bathysciae, based on Bathyscia Schiodte, for this group; he gave no reason for not using a name based on Leptoderus or Leptodirus, which he included in the group. Most subsequent authors, including virtually all systematists and ecologists working on this taxon, have followed Horn, using a name based on Bathyscia as the valid name for the group (e.g., Jeannel, 1910Jeannel, , 1911Jeannel, , 1914Jeannel, , 1936Laneyrie, 1967Laneyrie, , 1978Gueorguiev, 1974Gueorguiev, , 1976Peck, 1973Peck, , 1990. Reitter (1884Reitter ( , 1886, using group names based on both Bathyscia and Leptoderus, adopted one based on Bathyscia for the larger group including both, but later (Reitter, 1891, 1 906) chose Leptoderini for the larger group, and still later ( 1 909) returned to Bathysciini. ...
The current classification of the beetle series Staphyliniformia is discussed, and a classification is presented that includes 455 taxa above the genus level (ranging in rank from subtribe through superfamily). This classification is used as the framework for a catalog of earliest use and synonyms of the valid names of these taxa, with explanatory discussions as needed. The main provisions of the International Code of Zoological Nomenclature relating to family-group names are reviewed. Application of the Code to such names in Staphyliniformia requires changes (summarized in a table) in about 60 commonly used names. A third of these needed changes have already been pointed out and implemented by others, but 39 are first noted here, required by priority (20), homonymy (2), homonymy of type genus (6), or incorrect stem formation (11, including unjustified emendations).
The most significant of the changes adopted here, grouped by family, are Hydrophilidae: Acidocerina (for Helocharina); Leiodidae: Leptodirini (for Bathysciini), Platypsyllinae (for Leptininae), Sogdini (for Hydnobiini); Pselaphidae: Iniocyphini (for Tanypleurini), Natypleurina, nom. nov. (for Tanypleurina); Ptiliidae: Cephaloplectinae (for Limulodinae); Scydmaenidae: Cyrtoscydmini (for Euconnini, Neuraphini, or Stenichnini), Mastiginae (for Clidicinae); Staphylinidae: Athetini (for Callicerini), Clavilispinina, nom. nov. (for Paralispinina), Dorylophilini (for Deremini), Geostibina (for Callicerina), Glyptomina, nom. nov. (for Calocerina), Homalotini (for Bolitocharini or Gyrophaenini), Hypocyphtini (for Oligotini), Hyptiomina (for Holisina), Leptanillophilini (for Mimecitonini), Lomechusini (for Myrmedoniini or Zyrasini), Mimanommatini (for Dorylomimini), Mycetoporini (for Bolitobiini), Paglini, nom. nov. (for Pachyglossini), Solieriinae, nom. nov. (for Physognathinae), Strigotina (for Acrotonina), and Thoracophorini (for Lispinini). A few required changes are not implemented for reasons discussed in the text, which include pending applications to the ICZN. Ten cases apparently involving homonymy of family-group names (but not their type genera) were discovered and are discussed. Those in which the Staphyliniformia name is junior are Leiodidae: Triarthrini Jeannel, 1962 (not Ulrich, 1930, Trilobita); Pselaphidae: Metopiini Raffray, 1904 (not Foerster, 1868, Hymenoptera; senior to Townsend, 1908, Diptera); Staphylinidae: Callicerini Jacobson, 1908 (not Rondani, 1856, Diptera), Cyphini Lohse, 1974 [nom. nud.?] (not Leng, 1920, Coleoptera: Curculionidae), and Toxoderina Bernhauer & Schubert, 1911 (not Saussure, 1869, Mantodea). Those in which the Staphyliniformia name is senior are Hydrophilidae: Helocharina Orchymont, 1919 (predates Metcalf, 1 965, Homoptera [nom. nud.?]), Hydrobiini Mulsant, 1844 (predates Troschel, 1857, Mollusca); Staphylinidae: Cryptobiina Casey, 1905 (predates Hollande, 1952, Protozoa [nom. nud.?]), Steninae MacLeay, 1825 (predates Fraser & Purves, 1 960, Mammalia [nom. nud.?]), and Tachininae Fleming, 1 82 1 (predates Robineau-Desvoidy, 1830, Diptera).
The name Empelinae, subfam. nov. (type genus, Empelus LeConte; Staphylinidae), is made available for the first time; diagnoses are provided for this and for Solieriinae. In Pselaphidae, the replacement name Natypleurus, nom. nov., is proposed for the genus Tanypleurus Raffray (not Steenstrup & Luetken).
Many common family-group names in Coleoptera have been attributed to "Leach, 1817"; evidence is presented that they should actually be treated as "Fleming, 1821." Relative priority of four works published by Erichson and by Heer in 1839 is discussed.
... The Bathysciinae beetles contain c. 600 species in c. 135 genera, most of which are cave-dwelling and endemic and occur in Europe (Vandel 1965, Laneyrie 1967). Among the many species of troglobitic beetles in the Pyrenees, the large genus Speonomus contains many cave-dwelling species from the more primitive to the more specialized (Jeannel 1924). ...
The population ecology of the beetle Speonomus hydrophilus, occurring both in caves (reduced fluctuations in many abiotic parameters) and under the deepest layer of soil in mountains (MSS, more exposed to climatic variations), was studied in four habitats in the French central Pyrenees We have assessed some of the characteristics of the environment where these populations occur c g physical data (altitude and exposure), geologic data (nature of the parent-rock) and abiotic parameters (temperature with its seasonal fluctuations) and we investigated the relative importance of environmental structure and ecological characteristics on the temporal organization of S hydrophilus and the troglobitic fauna which cohabits The climatic study shows the existence of an annual thermal cycle which is regular and well marked for the MSS habitats but slightly out of phase with the surface cycle These periodic variations however slight may be stressful for troglobitic species In the MSS populations, the phenology of the entire community is reflected in the pattern seen in Speonomus The analysis of faunal profiles shows that samples follow the same seasonal succession during the annual cycle A potential seasonal rhythm of emergence may reflect a seasonal rhythm of vitellogenesis which produces a rhythm of egg-laying
... With further evolution and In the Bathysciina (Bathysciini of hiding. Laneyrie, 1967) the humicoles Adelopsella Appendages. Figure 8 is a series of an-( which has the largest eyes of the subtribe tennae demonstrating the length-to-width as noted above) and Sciaphyes have a low proportions of species of different species carina. ...
Submitted to: Dept. of Biology. Thesis (Ph. D.)--Harvard University, 1971.
Oryotus tragoniae Müller, 1934 è un coleottero leptodirino descritto di una piccola cavità nei pressi del Monte Bivera (Alpi Carniche, Forni di Sopra, UD) e noto su pochi esemplari. Viene descritta una seconda località di raccol-ta che amplia verso Est l'areale distributivo della specie. Viene inoltre proposta una ridescrizione della specie basata su un esemplare topotipico.
A leptodirine beetle Oryotus tragoniae Müller, 1934 was described for a small cavity near Bivera Mt. (Carnic Alps, Forni di Sopra, UD) and presently known for a few specimens. A second sampling site is described, which expands towards East the distribution area of the species. A redescription of the species based on a topotypical specimen is also proposed.
The first description of a larva and pupa of Platycholeus Horn, 1880; the only American representative of the otherwise Palaearctic tribe Leptodirini, is provided, illustrated and discussed in a systematic context. A combination of larval characters is consistent with the conclusion of recent molecular phylogenetic studies that have placed Platycholeus as the sister group of the rest of Leptodirini. Platycholeus represents most of the characters of the least specialised, classic type of Leptodirini larvae and pupae, according to the classification of Deleurance-Glaçon (1963).
A checklist of the troglobitic coleoptera species of the genus Cytodromus Abeille, 1876 is presented with an iconography. Distribution maps, descriptions and a key to the species are provided along with detailed pictures of males’ genitalia and some morphological characters.
The monospecific genus Speophyes Jeannel, 1910 (troglobitic coleoptera) is presented with a detailed description of both adult and larval stages, a distribution map and pictures of males' genitalia and morphological characters.
Investigations of preimaginal development in Tiphodytes gerriphagus Marchal reveal two larval instars, based on counting exuviae and comparing mandible length during development within Limnoporus dissortis Drake & Harris eggs. Tiphodytes gerriphagus eggs are stalked, as is typical of scelionids, and are 282.6 ± 3.48 μm (mean ± SE) long. The chorion ruptures at 8–9 h postoviposition and releases a nonfeeding embryo into the host. Feeding begins at 18–20 h postoviposition, after the embryonic cuticle is shed and a fully differentiated and active larva is released. The first larval stage is teleaform and lasted up to 5 days postoviposition, and its total length increased from 183.6 ± 3.35 to 517.0 ± 14.67 μm. The second larval stage is hymenopteriform and lasted from 5 to 13 days postoviposition, and grew from 920.2 ± 24.65 to 1352.4 ± 11.89 μm total length before pupating. The pupal period lasted about 11 days, with male pupae being shorter and thinner than female pupae. These findings differ from previous descriptions of T. gerriphagus, and it is suspected that the first instar was mistakenly divided into two stadia. The sex ratio under laboratory conditions was female biased (22% males), and males were smaller but did not emerge significantly earlier than females.
We present a synthesis of the knowledge on the cholevid fauna of Sardinia, with an annotated checklist and an updated list of both published and unpublished collecting localities. The Cholevidae (Coleoptera, Staphylinoidea) are a group of small to medium-sized beetles, most of which are saprophagous; a significant percentage of Cholevidae are cave-dwellers, and some are highly specialized troglobitic elements. In Sardinia, twenty-five species have been recorded so far. More than half of them are endemic to the island, while some are sub-endemic (i.e. present also in Corsica or in Sicily and/or the Balearic Islands); nine species, belonging to the genera Ovobathysciola and Patriziella, are specialized hypogean elements. Four more species, belonging to the genus Bathysciola, are soil-inhabiting or cave-dwellers. The authors stress the high biogeographic and biospeleological interest of Sardinian cholevids. In particular, the colonization processes appear to be in good agreement with the current hypotheses about the origin and composition of the existing fauna of Sardinia. The hypogean species of the genera Ovobathysciola, Patriziella and species of Speonomus of the subgenus Batinoscelis are currently accepted as being related to the Baetic and Pyrenean lineages of Anillochlamys and Speonomus, respectively; therefore, they should belong to the most ancient stock of elements, originated during the Miocene drift of the Corso-Sardinian massif in the western Mediterranean. Some other endemic or sub-endemic taxa, with W-Mediterranean affinities, are probably derived from more recent colonizers as a consequence of the several connection/isolation phases which existed between Sardinia, Corsica, and other Mediterranean areas during the late Miocene (Messinian) salinity crisis, and during the Plio-Pleistocene ice-phase-related strong variations in the sea level. Finally, some widespread taxa, all epigean and winged, reached the island in more recent times, through active and passive dispersal.
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