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-View of the Tham Pha cave.

-View of the Tham Pha cave.

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The first stygobiontic diving beetle known from Laos, Laodytes lapiei n. gen., n. sp., is described from a cave located in the Vientiane province. Its morphological characters lead to its placement, among Hydroporinae, in Hydroporini. Inside these, the new species could not be assigned to an existing genus. As a result, a new genus has been defined...

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... a speleological expedition to Laos, which took place in March 2019, the third author (MF) collected three specimens of a small diving beetle in a cave of the Pha Lay mountain range, near Kasi. Since this was the dry season, she was able to progress in this cave along several fossil levels ( fig. 1) for more than 2 km under the range. At the intersection with a muddy system of galleries, she reached a more humid lower floor, climbing down a block chaos followed by a calcite flow; there, water from the last floods remained in residual pools and even flew along a small channel. The calcite flow was darkened by guano deposits: ...
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... flow was darkened by guano deposits: although no bats were seen during the visit, they should rest at times in the overhanging fault crack on in the ceiling. The beetles were found swimming in one of the pools ( fig. 2). More recently, during an expedition in March 2020 to the same cave, eight more specimens were found in the lower fossil level ( fig. 17). There also, a little water was flowing along calcite cascades, coming from the upper level where a lake ...
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... beetles, two males and one female, did not appear to fit any species mentioned in this book; he therefore asked the first author (PQ) to lend a helping hand. PQ succeeded in extracting the genitalia of one male. Given its tiny size (0.2 mm), we decided to mount it on a glass slide in DMHF resin; we thus could obtain a clear view of the organ ( fig. 14), at the cost of some flattening by the coverslip. Later, with eight more specimens available, JML mounted another male genitalia in a drop of mounting media DMHF without coverslip, to obtain more natural views of the organ ( fig. 15-16). These pictures were taken with an Olympus™ TG4 camera, adapted to an Olympus CX21 optical ...
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... (0.2 mm), we decided to mount it on a glass slide in DMHF resin; we thus could obtain a clear view of the organ ( fig. 14), at the cost of some flattening by the coverslip. Later, with eight more specimens available, JML mounted another male genitalia in a drop of mounting media DMHF without coverslip, to obtain more natural views of the organ ( fig. 15-16). These pictures were taken with an Olympus™ TG4 camera, adapted to an Olympus CX21 optical microscope through a LM-Scope™ adapter. High-resolution pictures of the external morphology ( fig. 4-12) have been kindly taken by Michel Perreau on a Keyence™ VHX5000 microscope equipped with a VH-Z250T camera lens. The lateral view ( fig. 13) ...
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... specimens available, JML mounted another male genitalia in a drop of mounting media DMHF without coverslip, to obtain more natural views of the organ ( fig. 15-16). These pictures were taken with an Olympus™ TG4 camera, adapted to an Olympus CX21 optical microscope through a LM-Scope™ adapter. High-resolution pictures of the external morphology ( fig. 4-12) have been kindly taken by Michel Perreau on a Keyence™ VHX5000 microscope equipped with a VH-Z250T camera lens. The lateral view ( fig. 13) was obtained with a Toupcam™ 14 Mp CMOS camera mounted on an Olympus™ SZX9 stereomicroscope, from several shots assembled with HeliconFocus™ v.7.6. ...
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... organ ( fig. 15-16). These pictures were taken with an Olympus™ TG4 camera, adapted to an Olympus CX21 optical microscope through a LM-Scope™ adapter. High-resolution pictures of the external morphology ( fig. 4-12) have been kindly taken by Michel Perreau on a Keyence™ VHX5000 microscope equipped with a VH-Z250T camera lens. The lateral view ( fig. 13) was obtained with a Toupcam™ 14 Mp CMOS camera mounted on an Olympus™ SZX9 stereomicroscope, from several shots assembled with HeliconFocus™ v.7.6. http://zoobank.org/374332F7-4F3C-444E-AEE4-B49AD388A795 Type species: Laodytes lapiei n. ...
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... with protruding front angles encasing the head to the basal third; front margin straight along the posterior part of head; base V-shaped in the middle, sinuated towards right posterior angles; sides with a rimmed gutter, the anterior two-thirds of which contain 6-7 setigerous punctures, provided with deciduous sensory bristles; in lateral view ( fig. 13), the side edge sinuated and not in continuity with the elytral edge. Surface strongly reticulated as on hind part of head, the meshes polygonal but a little wider, reticulation vanishing on sides; the posterior margin showing a short and faint notch near the external third, followed externally by Fig. 6-12. -Laodytes lapiei n. gen., n. ...
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... sensory bristles; in lateral view ( fig. 13), the side edge sinuated and not in continuity with the elytral edge. Surface strongly reticulated as on hind part of head, the meshes polygonal but a little wider, reticulation vanishing on sides; the posterior margin showing a short and faint notch near the external third, followed externally by Fig. 6-12. -Laodytes lapiei n. gen., n. sp., ♂. -6, Head and pronotum, dorsal view. -7, Head, ventral view. -8, Prosternal process. -9, Metaventrite and metacoxae. -10, Metacoxal process. -11, Protarsus. -12, ...
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... process (fig. 9); epipleuron getting regularly narrower to the metacoxae level, without oblique carina near shoulder; metepisternum in right triangle, about twice longer than wide; metaventral wings very narrow; metacoxal lines absent ( fig. 9): only a flattened raised plate visible on each side of the median groove; metacoxal process ( fig. 10) with median part extended backwards; hind margin obtusely emarginate medially, with each side obliquely cut and both margins converging forward; lateral lobes not rounded, deeply incised and flat; disk of metacoxae reticulated; metacoxal process with median part extended backwards slightly above the abdomen; the latter with five ...
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... Pro-and mesotarsi four-segmented, rather wide, pro-and mesotibiae short and dilated on the distal half ( fig. 5), pro-and mesotarsomeres 1-2 with adhesive setae but without distinct sucker cups ( fig. 11), hind legs thinner with oblong femora and large oval hind trochanters (fig. 10); mesotibiae densely adorned with spiny setae; metatrochanters reticulated with transversely elongated meshes; metatarsal claws of subequal length ( fig. ...
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... Pro-and mesotarsi four-segmented, rather wide, pro-and mesotibiae short and dilated on the distal half ( fig. 5), pro-and mesotarsomeres 1-2 with adhesive setae but without distinct sucker cups ( fig. 11), hind legs thinner with oblong femora and large oval hind trochanters (fig. 10); mesotibiae densely adorned with spiny setae; metatrochanters reticulated with transversely elongated meshes; metatarsal claws of subequal length ( fig. ...
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... on the distal half ( fig. 5), pro-and mesotarsomeres 1-2 with adhesive setae but without distinct sucker cups ( fig. 11), hind legs thinner with oblong femora and large oval hind trochanters (fig. 10); mesotibiae densely adorned with spiny setae; metatrochanters reticulated with transversely elongated meshes; metatarsal claws of subequal length ( fig. ...
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... (Figs. 14-16): as described in the genus diagnosis. Female. -External sexual dimorphism limited to the protarsi, thinner than those of the male and without visible adhesive ...

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Chapter
The phylogenetics and higher (family-group) classification of extant members of the beetle family Dytiscidae (Coleoptera), or predaceous diving beetles, is reviewed and reassessed. A phylogenetic analysis of the family is presented based on 168 species of diving beetles and 9 outgroup taxa from Gyrinidae, Noteridae, Amphizoidae, and Paelobiidae. All currently recognized dytiscid subfamilies and tribes are represented, most by multiple genera and species. Data include 104 morphological characters and approximately 6700 aligned bases from 9 DNA sequence fragments from cytochrome c oxidase I (COI) and II (COII), histone III (H3), 16S rRNA (16S), 12S rRNA (12S), arginine kinase (argkin), RNA polymerase II (RNA pol II), elongation factor 1 alpha (Ef1α), and wingless (wnt). Parsimony and Bayesian analyses were conducted. The topology of the parsimony tree (consensus of 13 equally-parsimonious solutions) exhibits numerous anomalies inconsistent with convincing morphological features and the Bayesian results and has, generally, relatively poor bootstrap support for major clades. The Bayesian topology is more consistent with major morphological features and has strong support for most clades, and conclusions are based primarily on this estimate. Major higher-level phylogenetic relationships with strong support include: (1) monophyly of Dytiscidae Leach, (2) Matinae Branden sister to the rest of Dytiscidae, (3) Agabinae Thomson + Colymbetinae Erichson, (4) Hydrodytinae Miller + Hydroporinae Aubé, (5) Dytiscinae Leach + Laccophilinae Gistel + Cybistrini Sharp + Copelatinae Branden, (6) monophyly of the subfamilies Matinae, Colymbetinae, Copelatinae, Coptotominae Branden, Lancetinae Branden, Laccophilinae (including Agabetes Crotch), Agabinae (support weaker than in other subfamilies) and Hydroporinae (monophyly of Hydrodytinae not tested), (7) paraphyly of Dytiscinae with Cybistrini sister to Laccophilinae (with strong support) and this clade sister to other Dytiscinae, and (8) monophyly of both Agabini (Agabus-group of genera) and Hydrotrupini Roughley (Hydrotrupes Sharp and the Platynectes-group of genera). Major conclusions regarding tribes within Hydroporinae include: (1) monophyly of the tribes Vatellini Sharp, Methlini Branden, Hydrovatini Sharp, Hygrotini Portevin, Hyphydrini Gistel (without Pachydrus Sharp) and Bidessini Sharp (including Peschetius Guignot, Hydrodessus J. Balfour-Browne and Amarodytes Régimbart) (monophyly of Laccornini Wolfe and Roughley and Pachydrini Biström, Nilsson and Wewalka not tested), (2) Pachydrini is a problematic, long-branched taxa resolved here as sister to Hydrovatini but with weak support, (3) Hydroporini monophyletic except for Laccornellus Roughley and Wolfe and Canthyporus Zimmermann, (4) Laccornellus and Canthyporus together monophyletic and sister to Hydroporinae except Laccornini. Four groups are resolved within Hydroporini exclusive of Laccornellus + Canthyporus corresponding to the Deronectes-, the Graptodytes-, the Necterosoma- and the Hydroporus-groups of genera. The classification of Dytiscidae is revised with the following taxonomic changes [2014]: (1) Hydrotrupini is recognized as a tribe of Agabinae including the genus Hydrotrupes and the Platynectes-group of genera, (2) the genus Rugosus García is moved from Colymbetinae to Copelatinae, (3) Cybistrini is elevated from tribe rank within Dytiscinae to subfamily of Dytiscidae, (4) Hyderodini Miller is placed as a junior synonym of Dytiscini, (5) Laccornellus and Canthyporus are removed from Hydroporini and placed in their own tribe, Laccornellini, (6) the following family-group names are resurrected from synonymy with Hydroporini and placed as subtribes within Hydroporini, Deronectina Galewski (for the Deronectes-group of genera), Siettitiina Smrž (for the Graptodytes-group of genera), Sternopriscina Branden (for the Necterosoma-group of genera), and Hydroporina (for the Hydroporus-group of genera), (7) Carabhydrini Watts is placed as a junior synonym of Sternopriscina, and (8) Hydrodessus, formerly incerta sedis with respect to tribe, is placed in Bidessini. Each subfamily, tribe and subtribe is diagnosed and its taxonomic history discussed.