Army ant middens - Home and nursery of a diverse beetle fauna.

Army ants provide nourishment to a large variety of animals. This includes birds that feed on animals flushed out by army ant raids, symbiotic arthropods that consume the ants' prey or their brood, and other arthropods that scavenge on army ant refuse deposits. The latter have not received much attention, and the few published studies lack detailed species identifications. Here we provide a first systematic inventory of the beetle fauna associated with refuse deposits of Eciton army ants, with a focus on Eciton burchellii. We collected 8364 adult beetles, 511 larvae, and 24 eggs from 34 deposits at La Selva Biological Station, Costa Rica. We used a combination of DNA barcoding and morphology to identify a subset of 436 specimens to species level. The samples included several new species, and we here formally describe two water scavenger beetles (Hydrophilidae). Refuse deposits harbored a diverse beetle fauna. The identified subset consisted of 91 beetle species from 12 families, with rove beetles being the most abundant and diverse visitors. Of the 85 species found with E. burchellii, 50 species were collected from only one or two refuse deposits. Conversely, seven species were found in 10 or more refuse deposits, indicating a certain level of habitat specialization. We matched adults and immatures for 22 beetle species via DNA barcodes, demonstrating that army ant middens also serve as a beetle nursery. The present survey highlights the significant ecological function of army ants as promoters of biodiversity and their status as keystone species in tropical rainforests.

The Dung Beetle Oxyomus of Taiwan (Coleoptera: Scarabaeidae): Review of the Fauna, a New Species and its Larva Associated by DNA Barcoding.

The Taiwanese fauna of the dung beetle genus Oxyomus Dejean, 1833 (Coleoptera: Scarabaeidae: Aphodiinae) is reviewed based on museum specimens and newly collected material. Four species, all endemic to Taiwan, are recognized, one of which is newly described here: O. alligator sp. nov. Remaining species are diagnosed, compared with similar relatives from outside of Taiwan, and their distribution is mapped. We show that Taiwanese Oxyomus species form three distinct morphological groups, similar to species from Japan, SE Asia and Malay Archipelago, respectively, indicating a possible composite origin of Taiwanese fauna. The species occur in submontane and montane forests at altitudes of 700-2550 m including the secondary Cryptomeria ones. Available data confirm their association with dung of various forest mammals (monkeys, muntjacs and serows), although the discovery of larvae in sifted forest leaf litter may indicate they can also develop in nutrient-rich substrate around the dung. The larva of O. alligator sp. nov. is described in detail, based on the larval specimens associated with adults by DNA barcodes. Larvae of Oxyomus alligator sp. nov. are similar to those of the European O. sylvestris (Scopoli, 1763), with important differences only found on maxilla and abdominal apex.

The genus Cangshanaltica (Coleoptera: Chrysomelidae: Alticinae): overview, new species, and notes on species complexes.

Moss-inhabiting flea beetles form a diversified ecological group of beetles with a number of recently described new species. Here, we present an overview of known species of the moss-inhabiting flea beetle genus Cangshanaltica Konstantinov, Chamorro, Prathapan, Ge & Yang, 2013 which is distributed in Eastern and South-Eastern Asia. We describe three new species: Cangshanaltica guanxiensis sp. nov. from Guanxi, China, C. marginata sp. nov. from Yunnan, China, and C. javana sp. nov., from Java, Indonesia. Additionally, we redescribe Cangshanaltica castanea (Gruev, 1985) comb. nov., a species formerly placed in Ivalia. Based on our studies of the morphological characters, we assume that some of the currently recognized species may in fact represent species complexes. Our findings also extend the known distribution of Cangshanaltica to Indonesia. A key of all described Cangshanaltica species, and an annotated checklist are provided.

Ancient relicts or recent immigrants? Different dating strategies alter diversification scenarios of New Zealand aquatic beetles (Coleoptera: Hydrophilidae: Berosus).

Dated species-level phylogenies are crucial for understanding the origin and evolutionary history of modern faunas, yet difficult to obtain due to the frequent absence of suitable age calibrations at species level. Substitution rates of related or more inclusive clades are often used to overcome this limitation but the accuracy of this approach remains untested. We compared tree dating based on substitution rates with analyses implementing fossil data by direct node-dating and indirect root-age constraints for the New Zealand endemic Berosus water beetles (Coleoptera: Hydrophilidae). The analysis based solely on substitution rates indicated a Miocene colonization of New Zealand and Pleistocene origin of species. By contrast, all analyses that implemented fossil data resulted in significantly older age estimates, indicating an ancient early Cenozoic origin of the New Zealand clade, diversification of species during or after the Oligocene transgression and Miocene-Pliocene origin of within-species population structure. Rate-calibrated time trees were incongruent with recently published Coleoptera time trees, the fossil record of Berosus and the distribution of outgroup species. Strong variation of substitution rates among Coleoptera lineages, as well as among lineages within the family Hydrophilidae, was identified as the principal reason for low accuracy of rate-calibrated analyses, resulting in underestimated node ages in Berosus. We provide evidence that Oligocene to Pliocene events, rather than the Pleistocene Glacial cycles, played an essential role in the formation of the modern New Zealand insect fauna.

Exceptionally preserved beetles in a Triassic coprolite of putative dinosauriform origin.

The Triassic was a crucial period for the early evolution and diversification of insects, including Coleoptera1-3-the most diverse order of organisms on Earth. The study of Triassic beetles, however, relies almost exclusively on flattened fossils with limited character preservation. Using synchrotron microtomography, we investigated a fragmentary Upper Triassic coprolite, which contains a rich record of 3D-preserved minute beetle remains of Triamyxa coprolithica gen. et sp. nov. Some specimens are nearly complete, preserving delicate structures of the legs and antennae. Most of them are congruent morphologically, implying that they are conspecific. Phylogenetic analyses suggest that T. coprolithica is a member of Myxophaga, a small suborder of beetles with a sparse fossil record, and that it represents the only member of the extinct family Triamyxidae fam. nov. Our findings highlight that coprolites can contain insect remains, which are almost as well preserved as in amber. They are thus an important source of information for exploring insect evolution before the Cretaceous-Neogene "amber time window." Treated as food residues, insect remains preserved in coprolites also have important implications for the paleoecology of insectivores, in this case, likely the dinosauriform Silesaurus opolensis.

Review of the Taiwanese Laccobius/ Erichson, 1837 (Coleoptera: Hydrophilidae), with description of a new species.

Taiwanese species of the genus Laccobius Erichson, 1837 (Coleoptera: Hydrophilidae) are reviewed based on newly collected material and published records. Five species classified in two subgenera are recorded for the region, including L. (Glyptolaccobius) politus Gentili, 1979, L. (Microlaccobius) formosus Gentili, 1979, L. (M.) hammondi Gentili, 1984, L. (M.) roseiceps Régimbart, 1903 and a new species from central and southern Taiwan: L. (M.) hoi. sp.nov. Laccobius philipinus Gentili, 2005 is excluded from Taiwanese fauna. We failed to re-collect Laccobius fragilis Nakane, 1966 despite sampling the area from which it was recorded, and its occurrence in Taiwan hence needs re-confirmation. Distribution data of all Taiwanese species is summarized, and a key of Taiwanese Laccobius and photographs of all species are provided.

Moss-inhabiting flea beetles in the Philippines (Coleoptera, Chrysomelidae, Alticinae).

The Philippine islands are one of the key biodiversity hotspots in the Indo-Pacific area. Knowledge of moss-inhabiting flea beetles (Coleoptera: Chrysomelidae: Alticinae), a diverse and ecologically and morphologically enigmatic group in the Philippines is described. Six species from the Philippines are recorded, belonging to three genera: Benedictus luzonicus Sprecher-Uebersax et al., 2009 (recorded from the Philippines previously), Ivalia antennata sp. nov., I. caligulata sp. nov. and I. postfasciata (Chen, 1934), comb. nov. (transferred from Chabria Jacoby, 1887), Cangshanaltica mindanaoensis sp. nov., and C. luzonica sp. nov. Cox1 barcode sequences of Ivalia antennata and Cangshanaltica mindanaoensis are presented. Biogeography and diversity of moss-inhabiting flea beetles in the Philippines are discussed.

Two new species and additional faunistic records of Cercyon Leach, 1817 from China (Coleoptera: Hydrophilidae).

Two new species of Cercyon Leach, 1817 are described from China: Cercyon (s.str.) bellus sp. nov. from Hebei, Hubei, and Jiangxi Provinces, and Cercyon (s.str.) biltoni sp. nov. from Sichuan. Five species are recorded from China for the first time: Cercyon (Acycreon) punctiger Knisch, 1921, Cercyon (Paracercyon) analis Seidlitz, 1888, Cercyon (s.str.) algarum Sharp, 1973, Cercyon (s.str.) borealis Baranowski, 1985, Cercyon (s.str.) tristis (Illiger, 1801). Additional faunistic data from China are provided for Cercyon (s.str.) alinae Ryndevich, 2004, Cercyon (s.str.) inquinatus Wollaston, 1854, Cercyon (s.str.) kabaki Ryndevich, 2004, Cercyon (s.str.) olibrus Sharp, 1874, Cercyon (s.str.) ovillus Motschulsky, 1860, Cercyon (Clinocercyon) incretus d'Orchymont, 1941, Cercyon (Clinocercyon) hanseni Jia, Fikácek Ryndevich, 2011 and Cercyon (Paracercyon) honorabilis Shatrovskiy, 1999. A checklist to all species of the genus known from China is provided.

A new species of Laccobius Erichson, 1837 (Hydrophilidae, Coleoptera) from the Chinese Himalaya, with comments on taxonomic status of subgenera Glyptolaccobius Gentili, 1989 and Cyclolaccobius Gentili, 1991 and additional faunistic records from China.

A new species of the water scavenger beetle, Laccobius (Glyptolaccobius) motuoensissp. nov., is described from Motuo County, Xizang, China and its diagnostic characters are illustrated. Examination of this new species and re-examination of previously described species revealed that the separation of the subgenus Glyptolaccobius Gentili, 1989 and Cyclolaccobius Gentili, 1991 is artificial: both subgenera are hence combined here. Cyclolaccobius syn. nov. is synonymized with Glyptolaccobius, and the latter is shown to be diagnosed by 7-segmented antennae as a unique synapomorphy. All species treated until now under Cyclolaccobius are here transferred to Glyptolaccobius, with the only exception of L. hingstoni Orchymont, 1926, L. jumlanus Gentili, 2015 and L. zugmayeri Knisch, 1910 which are tentatively transferred to the subgenus Hydroxenus Wollaston, 1867, as their antennae bear eight antennomeres. Three species are recorded for the first time from China: L. (Microlaccobius) orientalis Knisch, 1924 from Xizang, Laccobius (M.) exilis Gentili,1974 from Xinjiang, and Laccobius (M.) sublaevis J. Sahlberg, 1900 from Xinjiang. Additional faunistic data from China are provided for the following species: L. (Microlaccobius) hammondi Gentili, 1984, Laccobius (M.) formosus Gentili, 1979, Laccobius (Hydroxenus) hingstoni d'Orchymont, 1926, Laccobius (Glyptolaccobius) yunnanensis Gentili, 2003, Laccobius (Compsolaccobius) decorus (Gyllenhal, 1827), Laccobius (Dimorpholaccobius) bipunctatus (Fabricius, 1775), Laccobius (D.) striatulus (Fabricius, 1775), Laccobius (s. str.) bedeli Sharp, 1884, L. (s. str.) binotatus d'Orchymont, 1934, Laccobius (s. str.) cinereus Motschulsky, 1860, and Laccobius (s. str.) minutus (Linnaeus, 1758).

A review of the Cercyon Leach (Coleoptera, Hydrophilidae, Sphaeridiinae) of the Greater Antilles.

The representatives of the genus Cercyon Leach occurring in the Greater Antilles are reviewed. Ten species are recorded, of which five are described here as new: C. gimmelisp. n. (Dominican Republic), C. armatipenissp. n. (Dominican Republic), C. tainosp. n. (Dominican Republic), C. sklodowskaesp. n. (Jamaica) and C. spiniventrissp. n. (Dominican Republic). Diagnoses and detailed distributional data are also provided for C. floridanus Horn, 1890 (distributed in southeastern United States of America and Cayman Islands), C. insularis Chevrolat, 1863 (endemic to the Antilles) C. praetextatus (Say, 1825) (widely distributed in the New World incl. Greater Antilles), C. quisquilius (Linnaeus, 1761) (an adventive species of Paleartic origin) and C. nigriceps (Marshall, 1802) (an adventive species probably of Oriental origin). Cercyon armatipenis, C. gimmeli, C. taino form a group of closely related species only distinguishable by male genitalia and DNA sequences. A key to the Great Antillean Cercyon is provided and important diagnostic characters are illustrated. The larvae of C. insularis and C. taino were associated with adults using COI barcode sequences, illustrated and diagnosed. Full occurrence data, additional images and COI barcode sequences were submitted to open access on-line depositories in an effort to provide access to complete data.

The Hydraenidae of Cuba (Insecta: Coleoptera) III. Description of two new hygropetric species of Hydraena Kugelann.

Two new species of the genus Hydraena Kugelann, 1794 collected from hygropetric habitats in eastern Cuba are described: Hydraena (Hydraenopsis) blancae sp. nov. from the Nipe-Sagua-Baracoa mountain range, and Hydraena (Hydraenopsis) matthiasi sp. nov. from the Sierra Maestra mountain range. Both species, especially the latter, are closely related to Hydraena (Hydraenopsis) franklyni Deler-Hernández & Delgado, 2012. Diagnostic characters for both new species are provided and illustrated; habitat information and distributional data are also included. An updated key to Cuban species of Hydraena is provided. With this study, the number of species of Hydraena known from Cuba raises to six.

On the identity of Sternosternus grouvellei Guillebeau, 1894 (Coleoptera: Hydrophilidae, Phalacridae).

The genus Sternosternus Guillebeau, 1894 was described as an aberrant taxon of the family Phalacridae from Sumatra, based on the unique morphology of its meso- and metaventrite resembling those of Cetonia aurata Linnaeus, 1758 (Guillebeau 1894). It contained the single species, S. grouvellei Guillebeau, 1894, known from a single specimen. The identity of Sternosternus was long unclear, and nothing was published on the genus until Gimmel (2013) studied the type specimen and recognized it actually belonged to the family Hydrophilidae, likely being a member of the genus Dactylosternum Wollaston, 1854.

Two new species of Coelostoma Brullé, 1835 from China and additional faunistic records of the genus from the Oriental Region (Coleoptera: Hydrophilidae: Sphaeridiinae: Coelostomatini).

Two new species of Coelostoma Brullé, 1835 are described from China: C. (Lachnocoelostoma) jaechi sp. nov. (Hong Kong) and C. (Lachnocoelostoma) tangliangi sp. nov. (Hainan). Coelostoma (s. str.) subditum Orchymont, 1936 is reported for the first time from continental Asia (China: Hong Kong, Yunnan). The first country or province records are provided for an additional 10 species: C. (s. str.) vividum Orchymont, 1936 from Cambodia and Pakistan, C. (s. str.) vitalisi Orchymont, 1923 from Cambodia and Thailand; C. (Holocoelostoma) stultum (Walker, 1858) from Myanmar and China: Hainan, Chongqing; C. (Lachnocoelostoma) coomani Orchymont, 1932 from China: Guizhou, Sichuan; C. (L.) hongkongense Jia, Aston & Fikácek, 2014 from Thailand and China: Yunnan; C. (L.) horni (Régimbart, 1902) from Thailand; C. (L.) huangi Jia, Aston & Fikácek, 2014 from Thailand; C. (L.) phallicum Orchymont, 1940 from China: Yunnan; C. (L.) vagum Orchymont, 1940 from China: Fujian; and C. (L.) wui Orchymont, 1940 from China: Fujian, Henan. Coelostoma coomani diversum Orchymont, 1932 is elevated to species rank (as C. diversum stat. nov.) based on the study of its type specimens and comparison with species treated here. An updated key to the species of Chinese Coelostoma is provided.

A review of Elocomosta Hansen with a description of a new species with reduced eyes from China (Coleoptera, Hydrophilidae, Sphaeridiinae).

A new species of the genus Elocomosta Hansen, 1989 (Coleoptera: Hydrophilidae: Sphaeridiinae: Coelostomatini), Elocomosta lilizheni sp. n., is described from Guangxi Province, China. It is compared in detail with the only other known species of the genus, Elocomosta nigra Hansen, 1989 from Borneo, and the genus is diagnosed from the remaining coelostomatine genera. The new species is unusual among Hydrophilidae by having extremely reduced eyes.

Redescription and lectotype designation of Chasmogenus abnormalis (Sharp), with notes on its distribution.

Chasmogenus Sharp, 1882 is a relatively small genus of 43 described species of aquatic beetles assigned to the subfamily Acidocerinae of Hydrophilidae (Hansen 1999; Short & Hebauer 2006; Short & Fikácek 2011, 2013; Clarkson & Ferreira-Jr. 2014). It is distributed in all zoogeographic regions except the Nearctic, with the highest species richness known from the Afrotropical (18 spp.) and Neotropical (15 spp.) Regions. Only four species have been described from the Oriental Region: C. abnormalis (Sharp, 1890), C. rubricollis (Régimbart, 1903), C. larsi Hebauer, 1995, and C. cattienus Hebauer, 2002. Of these, only C. abnormalis was recorded multiple times after its description.

Redescriptions and lectotype designations of Central American species of Phaenonotum Sharp (Coleoptera, Hydrophilidae) based on the type material from the David Sharp collection.

In order to understand the identity of the Central American species of the genus Phaenonotum Sharp, 1882, the type specimens of the species described by Sharp (1882) deposited in the David Sharp collection in the Natural History Museum in London have been re-examined. The following species are redescribed: Phaenonotum apicale Sharp, 1882, Phaenonotum collare Sharp, 1882, Phaenonotum dubium Sharp, 1882 (confirmed as junior synonym of Phaenonotum exstriatum (Say, 1835)), Phaenonotum laevicolle Sharp, 1882, Phaenonotum rotundulum Sharp, 1882 and Phaenonotum tarsale Sharp, 1882. Lectotypes are designated for Phaenonotum apicale, Phaenonotum collare, Phaenonotum rotundulum and Phaenonotum tarsale. External diagnostic characters and morphology of male genitalia are illustrated. A table summarizing diagnostic characters allowing the identification of the species is provided.

ResumenPara entender la identidad de las especies del género Phaenonotum Sharp, 1882 de América Central, se han reexaminado los especímenes tipo de las especies descritas por Sharp (1882) depositadas en la colección de David Sharp, del Museo de Historia Natural en Londres. Las especies redescritas son: Phaenonotum apicale Sharp, 1882, Phaenonotum collare Sharp, 1882, Phaenonotum dubium Sharp, 1882 (confirmado como sinónimo más reciente de Phaenonotum exstriatum (Say, 1835)), Phaenonotum laevicolle Sharp, 1882, Phaenonotum rotundulum Sharp, 1882 y Phaenonotum tarsale Sharp, 1882. Se designan lectotipos para las especies Phaenonotum apicale, Phaenonotum collare, Phaenonotum rotundulum y Phaenonotum tarsale. Se ilustran los caracteres diagnósticos y la morfología de los genitales. Una tabla resumen con los caracteres diagnósticos para facilitar la identificación de las especies es ofrecida.

Notes on the Australian Anacaenini (Coleoptera: Hydrophilidae): description of male of Phelea breviceps Hansen and unravelling the identity of Crenitis neogallica Gentili.

The holotype and additional specimens of the Tasmanian endemic Phelea breviceps Hansen, 1999, and the holotype of Crenitis neogallica Gentili, 1996 were examined in order to explore their generic and tribal placement. The morphology of Phelea is illustrated in detail, its male genitalia are described for the first time, and it is confirmed as a member of Anacaenini, sharing numerous characters with Crenitis Bedel, 1881 and the New Zealand endemic Horelophus walkeri Orchymont, 1913. Crenitis neogallica is removed from Anacaenini and transferred to the rygmodine genus Pseudohydrobius Blackburn, 1898 (as Pseudohydrobius neogallicus (Gentili, 1996) comb. nov.) and its holotype is illustrated. The genus Crenitis is hence removed from the Australian fauna.

A review of the omicrine genera Omicrogiton, Mircogioton and Peratogonus of China (Coleoptera, Hydrophilidae, Sphaeridiinae).

The Chinese species of the genera Omicrogiton Orchymont, 1919, Peratogonus Sharp, 1884 and Mircogioton Orchymont, 1937 are reviewed, diagnosed and keyed. Mircogioton and Omicrogiton are reported for the first time from China, Peratogonus for the first time for mainland China. Five species are recognized: Omicrogitoncoomani Balfour-Browne, 1939 (Guangdong, Hongkong), Omicrogitonhainanensis sp. n. (Hainan), Omicrogitonroberti sp. n. (Hainan), Mircogiotoncoomani Orchymont, 1937 (Yunnan), and Peratogonusreversus Sharp, 1884 (Guangdong, Jiangxi, Taiwan). Lectotype of Omicrogitoncoomani is designated. Mircogiotoncognitus (Malcolm, 1981), syn. n. is considered a junior subjective synonym of Mircogiotoncoomani Orchymont, 1939. Species of Mircogioton and Omicrogiton inhabit decaying banana trunks, whereas Peratogonusreversus was always collected from moist forest leaf litter.

A new aberrant species of Nipponocercyon from the mountains of southeastern China (Coleoptera: Hydrophilidae: Sphaeridiinae).

Nipponocercyon satoi sp. nov. is described from the mountains of Zhenjiang and Jiangxi Provinces in Southwest China. Although the species bears all synapomorphies of Nipponocercyon Satô, 1963, it differs substantially from the other two species of the genus by its Pacrillum-like habitus (i.e. small, compact and strongly convex body with reduced surface sculpturing). The new species is compared in detail with the other two Nipponocercyon species as well as with Pacrillum Orchymont, 1941 and Megasternum Mulsant, 1844. The generic diagnosis of Nipponocercyon is adapted, and reasons for assigning the new species to Nipponocercyon are discussed.

Review of the Chinese species of the genus Coelostoma Brullé, 1835 (Coleoptera: Hydrophilidae: Sphaeridiinae).

The Chinese species of the genus Coelostoma Brullé, 1835 are revised and diagnosed. Five new species are described: Coelostoma bifidum sp. nov. (Jiangxi, Guangdong, Hong Kong), C. hajeki sp. nov. (Guangdong, Guangxi), C. hongkongense sp. nov. (Hong Kong), C. huangi sp. nov. (Guangxi, Jiangxi), and C. gentilii sp. nov. (Xizang). Coelostoma coomani Orchymont, 1932, C. orbiculare (Fabricius, 1775) and C. vividum Orchymont, 1936 are reported for the first time from China. Coelostoma sulcatum Pu, 1963, syn. nov. is synonymized with C. stultum (Walker, 1858). Additional faunistic data from China are provided for C. phallicum Orchymont, 1940, C. vagum Orchymont, 1940, C. turnai Hebauer, 2006, C. wui Orchymont, 1940, C. stultum (Walker, 1858), C. vitalisi Orchymont, 1923, C. fallaciosum Orchymont, 1936 and C. horni (Régimbart, 1920). Coelostoma fabricii (Montrouzier, 1860) is not considered as member of Chinese fauna as all previous records are dubious and no new material was found during our study. The distribution of C. horni in China is discussed in detail, with records from Hong Kong here considered dubious. An updated identification key to the Chinese species of Coelostoma is provided.