Disentangling phylogenetic relations and biogeographic history within the Cucujus haematodes species group (Coleoptera: Cucujidae).

Recent progress in the taxonomy of flat bark beetles (Cucujidae), specifically, in the genus Cucujus, has revealed great diversity in subtropical Asia, but the seemingly well-known temperate and boreal taxa need further attention because of their conservation status. Here, we used an integrative approach using morphology, DNA, and species distribution modelling to disentangle phylogenetic relations, verify the number of species, and understand the historical biogeography of Palearctic and Nearctic Cucujus beetles, particularly the C. haematodes species group. Species distinctiveness was supported for C. cinnaberinus, but present-day C. haematodes turned out to be a species complex made up of separate lineages in the western, middle and eastern parts of its Palearctic range. Cucujus muelleri was a member of that complex, being sister to Asian C. haematodes. Moreover, C. haematodes caucasicus was found to be phylogenetically closely related to Italian C. tulliae, and both to be sister to European C. haematodes. North American C. clavipes clavipes and C. c. puniceus resulted to be enough divergent to be considered different species. Interestingly, western American C. puniceus turned out to be closely related to the C. haematodes complex, whereas eastern American C. clavipes constituted a separate lineage, being distantly related to both C. puniceus and C. cinnaberinus. These patterns suggest former trans-continental connections among the ancestors of extant flat bark beetle species. Moreover, a divergent lineage of C. cinnaberinus was found in Calabria, which should be regarded at the very least as a subspecies. The ancestor of C. hameatodes group originated in mid-Miocene, and next, ca. 6.2 Mya, a line leading to C. cinnaberinus had split. Speciation of the American lineages occurred during Pliocene (4.4 Mya for C. clavipes and 3.3 Mya for C. puniceus). Species classified as C. haematodes, C. tulliae and C. muelleri, as well as distinct lineages within C. cinnaberinus split during mid Pleistocene (ca. 1.5 Mya). A comparison of species climatic requirements and their present distribution allowed to identify glacial refugia in south-eastern areas of North America (C. clavipes), south-western areas of North America (C. puniceus), and the Mediterranean and Caspian Sea Basins (European Cucujus species), or south-eastern areas of Asia and the foothills of the central Asian mountains (eastern C. haematodes). Subsequent climatic changes in the Holocene forced these beetles to move their ranges northwards along the coasts of the Pacific (C. puniceus) or Atlantic (C. clavipes), north-eastwards to central, northern, and eastern Europe (C. cinnaberinus and European C. haematodes) or Siberia (Asian C. haematodes). The combined use of molecular, morphological and climatic data allows a comprehensive understanding of the phylogenetic relations and past distributions of Cucujus beetles, highlighting the complexity of C. haematodes species group evolution.

Reconstructed historical distribution and phylogeography unravels non-steppic origin of Caucasotachea vindobonensis (Gastropoda: Helicidae).

Existing data on the phylogeography of European taxa of steppic provenance suggests that species were widely distributed during glacial periods but underwent range contraction and fragmentation during interglacials into "warm-stage refugia." Among the steppe-related invertebrates that have been examined, the majority has been insects, but data on the phylogeography of snails is wholly missing. To begin to fill this gap, phylogeographic and niche modeling studies on the presumed steppic snail Caucasotachea vindobonensis were conducted. Surprisingly, reconstruction of ancestral areas suggests that extant C. vindobonensis probably originated in the Balkans and survived there during the Late Pleistocene glaciations, with a more recent colonization of the Carpatho-Pannonian and the Ponto-Caspian regions. In the Holocene, C. vindobonensis colonized between the Sudetes and the Carpathians to the north, where its recent and current distribution may have been facilitated by anthropogenic translocations. Together, these data suggest a possible non-steppic origin of C. vindobonensis. Further investigation may reveal the extent to which the steppic snail assemblages consist partly of Holocene newcomers.

Review of the Argentinean species of Pseudomicrocara Armstrong (Coleoptera: Scirtidae).

The Pseudomicrocara Armstrong from Argentina are reviewed. In total, seven species are present: Pseudomicrocara angusta sp. nov., P. antarctica (Fairmaire) comb. nov., P hieroglyphica sp. nov., P. inflexipenis sp. nov., P livida (Fabricius), P. obliquata (Solier) comb. nov., and P patagonica (Curtis) comb. nov. New provincial records are provided for several species. Pseudomicrocara obliquata, previously known only from Chile, is recorded from Argentina for the first time. Illustrations of habitus and genitalia as well as distributional data for all Argentinean species of Pseudomicrocara are provided.

A new marsh beetle from mid-Cretaceous amber of northern Myanmar (Coleoptera: Scirtidae).

As one of the earliest-diverging lineage of the megadiverse beetle suborder Polyphaga, marsh beetles (Scirtidae) are crucial for reconstructing the ancestor of all polyphagan beetles and the ecomorphological underpinnings of their remarkable evolutionary success. The phylogeny of marsh beetles has nonetheless remained challenging to infer, not least because of their fragmentary Mesozoic fossil record. Here we describe a new scirtid beetle genus and species, Varcalium lawrencei gen. et sp. nov., preserving internal tissue, from Albian-Cenomanian Kachin amber (ca 99 Ma), representing the second member of this family known from the deposit. Based on a formal morphological phylogenetic analysis, Varcalium is recovered within the crown-group of Scirtinae, forming a clade with other genera that possess subocular carinae. The finding suggests that the crown-group of Scirtinae has already diversified by the mid-Cretaceous.

The genus Scirtes Illiger of Fiji (Coleoptera: Scirtoidea: Scirtidae).

Scirtes Illiger is the only genus of Scirtidae known to inhabit Fiji. In the present revision, 10 species of Scirtes are recorded from Fiji, including seven newly described. Scirtes humeralis Pic, S. natovensis Champion, and S. veitchi Champion are redescribed and illustrated. Scirtes evansi Pic is a junior synonym of S. natovensis. Both males and females of Fijian Scirtes have foveae on abdominal ventrite 2, a character unknown in other members of the genus. Lectotypes of Scirtes natovensis Champion, S. evansi Pic, and S. humeralis Pic are designated.

Discovery of a complex sclerotized sperm pump in Monotomidae (Coleoptera): Functional morphology and phylogenetic value.

In insects, a long ductus ejaculatorius surrounded by a muscular sheath transfers seminal fluids and sperms from the testes into the male copulatory organ, the aedeagus. In several unrelated coleopteran families, a section of the ductus has evolved to form a specialized pumping organ. The most complex forms of this "sperm pump" was known in Lepiceridae and two subfamilies of evolutionary distant Staphylinidae; in these groups two sclerotized funnels situated on ends of the sperm pump provide attachment sites for a remarkably robust compressor muscle. A sperm pump with two funnels and a previously unknown spiral ridge, representing the most complex sperm pumping apparatus among Coleoptera, is for the first time reported to occur in the superfamily Cucujoidea, family Monotomidae. The compressor muscle is particularly large, and the pump shows features more advanced than those in any other beetle. Not the entire section between funnels is compressed, but only a pair of specialized thin-walled, unsclerotized and resilin-rich regions within the pump. The spiral ridge stores mechanical energy to decompress the pump when the muscle relaxes, and provides a sclerotized scaffold for the entire structure. The pumps do not occur in all Monotomidae, but seem to be restricted to the subfamily Monotominae. Two distinct forms were found, each in a different tribe (with one notable exception; Mimemodes having a sperm pump typical of Monotomini, not Europini). Moreover, the shapes of the pumps were found to be species-specific. The phylogenetic value of this structure is discussed, as sperm pumps may help defining monophyletic lineages within this poorly studied family. An evolutionary scenario for a development of sperm pumps in Coleoptera from an undifferentiated ductus ejaculatorius to the most complex "monotomid form" is proposed, assuming several distinct stages exemplified by structures found in other beetle families.

An overview of Scirtidae (Coleoptera) described by Antoine Joseph Jean Solier (1792-1851).

In 1849 a French entomologist, Antoine Solier, described 11 species of Scirtidae collected in Chile by Claude Gay. Based on examination of type specimens deposited in the Museum national d'Histoire naturelle (MNHN) in Paris, 9 species: Contacyphon luteolineatus, C. parvus, C. tristis, Pseudomicrocara lunata, P. obliquata, P. torquata, and Elodes rousseli are briefly described with emphasis on male or female genitalia. Morphological details are illustrated. Type specimens of two species: Contacyphon maculatus and Elodes velutina were not found in the collection of MNHN. Two new synonyms were established: Contacyphon maculicornis (Solier, 1849) is a junior synonym of Pseudomicrocara patagonica (Curtis, 1838), and Cyphon testaceus Solier, 1849 is a junior synonym of Pseudomicrocara livida (Fabricius, 1775). Lectotypes of most species described by Solier are designated.

A new species of Daploeuros Watts (Coleoptera: Scirtidae).

Daploeuros reichertae sp. nov. is described from New South Wales, Australia. The species is the smallest member of the genus, easy to distinguish on the basis of the morphology of the male copulatory organs. A short supplementary description of the female of Daploeuros tasmanicus Watts and a key to identify the species of Daploeuros are provided.

Calvariopsis gen. nov., a new genus of Neotropical Scirtidae (Coleoptera: Scirtoidea).

A new genus of Scirtidae, Calvariopsis gen. nov., is described to accommodate two species described by Maurice Pic (Prionocyphon brasiliensis Pic, 1916 and Cyphon sculptipenne Pic, 1931) and 14 newly described species: Calvariopsis bituberculata sp. nov. (French Guiana), C. borowieci sp. nov. (Ecuador), C. fourgassiense sp. nov. (French Guiana), C. guyanense sp. nov. (Guyana), C. kawense sp. nov. (French Guiana), C. nana sp. nov. (Brazil), C. panamense sp. nov. (Panama), C. peruviana sp. nov. (Peru), C. picta sp. nov. (Bolivia), C. pittieri sp. nov. (Venezuela), C. saopaulense sp. nov. (Brazil), C. venezuelense sp. nov. (Venezuela), C. wittmeri sp. nov. (Brazil), and C. yanayacuense sp. nov. (Ecuador) [type species of Calvariopsis]. The new genus can be distinguished from other Neotropical Scirtidae on the basis of the following characters: antennae filiform, subgenal ridge without buttonhole configuration, head with deep subantennal grooves, mesoventral process short, as long as wide or shorter, hind legs not saltatorial.

Meatopida gen. nov., a new genus to accommodate two species originally described in Atopida White, 1846 (Coleoptera: Scirtoidea: Scirtidae).

The New Zealand species of marsh beetles Atopida dorsale Broun, 1893 and Atopida testacea Broun, 1880 share several features that differentiate them from other members of Atopida White. Analysis of morphological characters and the structural anatomy of the genitalia justified the description of a new genus to accommodate these two species. Sensory structures of the apical maxillary palpomere are described for the first time in Scirtidae. The presence of lacinial hooks in Scirtidae is discussed. Morphology of female prehensors is described and observations of male and female genitalia during copulation are discussed. Lectotypes of both species are designated.

Description of females of Chilarboreus Ruta, 2011 (Coleoptera: Scirtidae: Scirtinae).

Chilarboreus is a small genus comprising three species occurring in Nothofagus forests of Chile. The female of a single species, Chilarboreus fossulatus, was described in the original description. In the present paper we provide descriptions of females of the remaining two species for the first time. Morphology of females supports the previous division of Chilarboreus into two subgenera. Pronotal excitators are described and illustrated for the first time in Scirtidae.

An illustrated catalogue of the New Zealand marsh beetles (Coleoptera: Scirtidae).

In this study we summarise the knowledge of the history and current state of research on the New Zealand Scirtidae to provide a base for further research on the family. Data on Tord Nyholm's research and collections are presented, based on a study of the archives and collection of Swedish Museum of Natural History. The main part of the paper is a catalogue of all described species of Scirtidae known to occur in New Zealand. A total of 11 genera and 126 species of Scirtidae is recorded for New Zealand, with 82% genera and 100% of species endemic to the New Zealand region. A reference to the original description, type locality, type depository and the known distribution within New Zealand is included for each species. Primary type specimens are illustrated for most species. Type species are designated in the present paper for Cyphanus Sharp, 1878 (type species: Cyphanus debilis Sharp, 1878), Mesocyphon Sharp, 1878 (type species: Mesocyphon marmoratus Sharp, 1878), and Veronatus Sharp, 1878 (type species: Anobium tricostellum White, 1846) as they were not fixed in the original descriptions or in subsequent works. Brounicyphon Pic, 1947 is considered a junior subjective synonym of Veronatus. Cyphon huttoni Sharp, 1878 is transferred to the genus Contacyphon Gozis, 1886.

Anticyphon gen. nov., a new genus of Scirtidae (Coleoptera: Scirtoidea) inhabiting high altitude Andean cloud forests and páramo formation.

A new genus of Scirtidae is described to accommodate six newly described species distributed in the northern part of the Andes. Three species are described from Ecuador (Anticyphon davidsoni sp. nov., A. ecuadorensis sp. nov., A. paramoensis sp. nov.), two from Peru (A. oyonensis sp. nov., A. peruvianus sp. nov.), and a single one from Colombia (A. santanderensis sp. nov.). Morphological characters are illustrated with focus on hind wings, mouthparts and male genitalia. An identification key to species is provided.

Redescriptions of Scirtidae (Coleoptera: Scirtoidea) described by Carl Henrik Boheman (1796-1868) with notes on Scirtes adustus iversenotatus Pic, 1930.

Boheman described four species of Scirtidae: one from Sweden (Cyphon pallidulus Boheman, 1850), two from South Africa (Elodes caffer Boheman, 1851 & Elodes marginipennis Boheman, 1851), and one from Argentina (Scirtes adustus Boheman, 1858). Cyphon pallidulus Boheman is a junior synonym of Contacyphon ochraceus (Stephens, 1830). Types of the remaining species were never re-examined, and none of them were redescribed. In the present paper, three species are redescribed on the basis of type specimens deposited in the Swedish Natural History Museum in Stockholm. Genitalia are illustrated for the first time and a new combination is proposed: Brachycyphon marginipennis (Boheman, 1851). The type specimen of a variety described by Pic, S. adustus var. diversenotatus, was examined and revealed to represent a distinct species, which is redescribed under the name of Scirtes diversenotatus Pic, 1930 stat. nov. in the present paper.

A supplement to the revision of the Scirtes flavoguttatus species-group (Coleoptera: Scirtidae: Scirtinae).

New species of the Scirtes flavoguttatus species-group are described from SE Asia. Altogether 34 species are newly described, including Scirtes beccus sp. nov. (Malaysia), S. bocakorum sp. nov. (Indonesia), S. crockerensis sp. nov. (Malaysia), S. decorus sp. nov. (Malaysia), S. dumogensis sp. nov. (Indonesia), S. gunongmulensis sp. nov. (Malaysia), S. ishikawai sp. nov. (Vietnam), S. kinabalensis sp. nov. (Malaysia), S. kundasangensis sp. nov. (Malaysia), S. lambriensis sp. nov. (Indonesia), S. leuserensis sp. nov. (Indonesia), S. luteus sp. nov. (Malaysia), S. malaisei sp. nov. (Myanmar), S. melinauensis sp. nov. (Malaysia), S. noonadan sp. nov. (Philippines), S. pallicolor sp. nov. (Malaysia), S. penampangensis sp. nov. (Malaysia), S. phoupanensis sp. nov. (Laos), S. prodigiosus sp. nov. (Malaysia), S. punctatus sp. nov. (Philippines), S. quasibalehensis sp. nov. (Malaysia), S. ranauensis sp. nov. (Malaysia), S. sarawakensis sp. nov. (Malaysia), S. seblatensis sp. nov. (Indonesia), S. sibayensis sp. nov. (Indonesia), S. sibolangitensis sp. nov. (Indonesia), S. sulawesicus sp. nov. (Indonesia), S. sulcigeroides sp. nov. (Malaysia), S. talinisensis sp. nov. (Philippines), S. ulukimanisensis sp. nov. (Malaysia), S. velutinus sp. nov. (Malaysia), S. vietnamicus sp. nov. (Vietnam), S. wallacei sp. nov. (Indonesia), S. yangsinensis sp. nov. (Vietnam). New localities of six species are provided. An updated identification key, checklist and a summary of distributional data are included. Probability of the occurrence of the Scirtes flavoguttatus species-group was evaluated with an analysis in MaxEnt software. It is highly plausible that members of the group occur in most mountainous rainforests of SE Asia. 

Erotylidae (Insecta, Coleoptera) of Poland - problematic taxa, updated keys and new records.

New data concerning the occurrence of pleasing fungus beetles (Coleoptera: Erotylidae) in Poland are given, with a focus on rare and difficult to identify Central European taxa. Cryptophilus cf. integer (Heer) (Cryptophilinae) is reported from the Polish territory for the first time based on adult and larval specimens collected in the Wielkopolska-Kujawy Lowland. Identification problems concerning species of Cryptophilus introduced to Europe are discussed. Triplax carpathica Reitter (Erotylinae) is recorded from the Bialowieza Primeval Forest, which is the first known non-Carpathian finding of this species, located in the close proximity of the Polish-Belarussian UNESCO World Heritage Site "Bialowieza Forest". Discussion of Triplax carpathica being conspecific with Siberian Triplax rufiventris Gebler is provided. New Polish localities of several other Erotylidae are reported, and an updated key to Central European species of Triplax is given. The Triplax key is supplemented with dorsal and ventral habitus images of all treated Triplax species. One of the rarest Central European erotyline species Combocerus glaber (Schaller) is recorded from xerothermic grasslands in North-West Poland.