Ground-beetles of the subgenus Cryptophonus Brandm. et Z. Brandm., genus Harpalus Latr. (Coleoptera, Carabidae)

A taxonomic review of ten species of the subgenus Cryptophonus Brandmayr et Zetto Brandmayr, 1982, the genus Harpalus Latreille, 1802 is given. In addition to nine Palaearctic species, the Ethiopian H. agnatus Reiche, 1849 is also included in Cryptophonus. Harpalus cyrenaicus Koch, 1939, stat. n. from Libya, which was originally described as a subspecies of H. litigiosus Dejean, 1829, is treated as a distinct species. Harpalus tenebrosus Dejean, 1829 is treated as a polytypical species with two subspecies: H. t. tenebrosus (West Palaearctic) and H. t. paivanus Wollaston, 1867, stat. n. (Cabo Verde). A new substitute name H. melancholicus reicheianus nom. n. is proposed to replace the objective homonyms H. reichei Jacobson, 1907 (non Desbrochers des Loges, 1867) and H. ovalis Reiche, 1861 (non Motschulsky, 1844). Lectotypes are designated for H. tenebrosus Dejean, 1829, H. paivanus Wollaston, 1867, H. litigiosus Dejean, 1829, H. agnatus Reiche, 1849, H. fulvus Dejean, 1829, H. melancholicus Dejean, 1829, and H. ineditus Dejean, 1829. The status and diagnosis of Cryptophonus are discussed and a key to all the species of this subgenus is provided. Data on distribution of each species are provided. The following species are recorded for the first time: H. tenebrosus from Afghanistan, H. grilli Kataev, 2002 from India (Uttarhand), H. agnatus Reiche, 1849 from Tanzania, and H. fulvus Dejean, 1829 from Portugal.     

Antennal sensilla of the ground beetle Bembidion lampros Hbst (Coleoptera, Carabidae)

Antennal sensilla typology, number and distribution pattern were studied in the ground beetle Bembidion lampros Hbst (Coleoptera, Carabidae) using scanning electron microscopy. The 1.6–1.8 mm long filiform antennae of both sexes consist of the scape, pedicel and of the flagellum composed of nine flagellomeres. In both sexes, three types of sensilla chaetica, two types of sensilla trichodea, five types of sensilla basiconica, one type of sensilla coeloconica, one type of sensilla campaniformia and Böhm sensilla were distinguished. The possible function of the sensilla was discussed and three types of sensilla were considered as olfactory, sensilla trichodea type 2 and sensilla basiconica types 1 and 2. Olfactory sensilla occupy dorsal and/or ventral areas of the flagellomeres and occur sparsely (sensilla basiconica type 1) or not at all (sensilla basiconica type 2 and sensilla trichodea type 2) outside these areas. No remarkable sexual differences in the types, numbers and distribution of antennal sensilla were found.     

An unexpected clade of South American ground beetles (Coleoptera,Carabidae, Bembidion)

Phylogenetic relationships of the Antiperyphanes Complex of the genus Bembidion are inferred using DNA sequences from seven genes (two nuclear ribosomal, four nuclear protein coding, and one mitochondrial protein coding). Redefined subgenera within the complex are each well-supported as monophyletic. Most striking was the discovery that a small set of morphologically and ecologically heterogeneous species formed a clade, here called subgenus Nothonepha. This unexpected result was corroborated by the discovery of deep pits in the lateral body wall (in the mesepisternum) of all Nothonepha, a trait unique within Bembidion. These pits are filled with a waxy substance in ethanol-preserved specimens. In one newly discovered species (Bembidion tetrapholeon sp. n., described here), these pits are so deep that their projections into the body cavity from the two sides touch each other internally. These structures in Bembidion (Nothonepha) are compared to very similar mesepisternal pits which have convergently evolved in two other groups of carabid beetles. The function of these thoracic pits is unknown. Most members of subgenus Nothonepha have in addition similar but smaller pits in the abdomen. A revised classification is proposed for the Antiperyphanes Complex.     

Chromosomal diversity and evolution in the ground beetle genus Bembidion and related taxa (Coleoptera: Carabidae: Trechitae)

Chromosome numbers and sex chromosome systems of 154 previously unstudied Bembidion species are described. The genus is nearly uniform: males of 176 of 205 species are 2n=22+XY. Karyotypes are presented for 30 species. There is some variation among species in size of Y and size of autosomes. Within most species autosomes are subequal in size, and metacentric or submetacentric. Subterminal secondary constrictions and B chromosomes are reported from several species.The supertribe Trechitae (Zolini + Trechini + Pogonini + Bembidiini) is hypothesized to be primitively male 2n=22+X or 24+X, and the ancestral Bembidion stock 2n=22+XY. Conclusions are based on the most parsimonious hypothesis of ancestral state given an inferred phylogeny of the group, rather than the widespread-is-primitive arguments used previously. Evolution within Bembidion away from the presumably-primitive 2n=22+XY is discussed. Six lineages have lost Y chromosomes; seven have undergone changes in autosome number. It is not known why such changes are so scarce, nor what particular rearrangements led to the observed diversity. Nonetheless, the cytogenetic data can be used to infer a monophyletic origin of groups possessing derived chromosome numbers or sex chromosomes, and to help resolve species limits.     

Genus Bembidion Latreille (Coleoptera: Carabidae) in New Zealand: A revision

The cosmopolitan genus Bembidion is represented in New Zealand by 20 species, of which 19 are endemic; B. brullei appears to be a recent introduction. On phenetic characters the species fall into 7 subgenera, as follows: Zeplataphus n.subg.—maorinum Bates, dehiscens Broun, charile Bates, granuliferum n.sp., townsendi n.sp., tairuense Bates; Zeactedium Netolitzky—orbiferum Bates, musae Broun; Zeperyphodes n.subg.—callipeplum Bates; Zeperyphus n.subg.—actuarium Broun; Zemetal‐lina n.subg.—chalceipes Bates, solitarium n.sp., anchonoderum Bates, tekapoense Broun, wanakense n.sp., urewerense n.sp., hokitikense Bates, parviceps Bates; Ananotaphus Netolitzky—rotundicolle Bates; Notaphus Stephens—brullei Gemminger & Harold. The North Island population of maorinum is distinct from the typical South Island form in having reduced microscrulpture on the elytra, and is here separated as levatum n.ssp. An apparent geographic isolate of anchonoderum, represented by 2 females from Stewart Island, is provisionally recognised as stewartense n.ssp. The polymorphic complex within subg. Ananotaphus is here regarded as a single species, of which the North Island population is sufficiently distinct to warrant subspecific status as eustictum Bates; however, intergrades occur in the north‐west of the South Island. The following names fall into synonymy: latiusculum Broun (= maorinum); diaphanum Broun (= musae); nesophilum Broun (= callipeplum)’, tinctellum Broun (= chalceipes);antipodum Broun (= anchonoderum)’, tantillum Broun and probably attenuatum Broun (=hokitikense)’, clevedonense Broun and waikatoense Broun (= rotundicolle, ssp. eustictum)’, gameani Jeannel (= brullei). The relationships and aspects of the biology and ecology of the New Zealand Bembidion fauna are discussed.     

New species of the genus Psylliodes Latr. (Coleoptera, Chrysomelidae) from the Palaearctic Region

Six new species of the genus Psylliodes Latr. are described from the Palaearctic Region. P. concolor sp. n. from Georgia and P. diversicolor sp. n. from Turkey belong to the luteolus species-group. P. amurensis sp. n. and P. laxus sp. n. from the Russian Far East are included in the punctifrons species-group. P. submontanus sp. n. from the North Caucasus is provisionally included in the napi species-group, but it is similar to P. ozisiki Leon. et Arnold from the cupreus species-group. P. insularis sp. n. from the Canaries belongs to the chrysocephalus species-group.     

Karyotypes of Bembidion quadrimaculatum (L.) and Clivina fossor (L.) (Coleoptera: Carabidae)

Bembidion quadrimaculatum possesses 24 chromosomes: 2n male = 22 + XY, 2n female = 22 + XX; their structure is meta- and submetacentric and differences in length between them are slight. Achiasmatic meiosis has been identified in spermatogenesis. The diploid chromosome number in Clivina fossor is 44; 2n male = 42 + XY, 2n female = 42 + XX. The chromosome structure is meta-, submeta-, and subtelocentric and X is the longest element in the set. 1 to 2 chiasmata per bivalent occur in meiosis.     

A new species of the carabid genus <Emphasis Type="Italic">Harpalus</Emphasis> Latr. (Coleoptera,Carabidae) from Iran

Harpalus murzini sp. n. is described from northern Iran (Gilan Province). The new species belongs to the dimidiatus group and is closely related to H. caspius. A key to four species of the dimidiatus group is provided.     

The effects of habitat,density, gender and duration on overwintering success in Bembidion lampros (Coleoptera: Carabidae)

In temperate climates, winter mortality is a key factor influencing insect survival. In Bembidion lampros (Coleoptera: Carabidae), an abundant polyphagous predator in arable farmland, we investigated how habitat, population density, gender and duration of overwintering affect the winter mortality and body condition of survivors. In a field experiment, we introduced beetles at two population densities into isolators placed either in the interior of the field or in the grassy boundary. Half of the isolators were collected in mid‐March, and the remaining half were collected in mid‐April. Across all treatments, 44–92% of the beetles survived until spring. Survival was significantly higher in the field interior than in the boundary. The longevity of individuals starved after overwintering was higher in females than in males and was positively related to the length of time spent at the overwintering site. Population density was positively related to percentage survival but did not affect body condition. Our results suggest that biotic factors such as population density and predation, though little studied, may have a similar significance as abiotic factors. Even ubiquitous carabid species inhabiting agroecosystems have specific requirements for overwintering. Understanding these requirements is useful for developing techniques to support their overwintering success.     

Phylogeny of Bembidion and related ground beetles (Coleoptera: Carabidae: Trechinae: Bembidiini: Bembidiina)

The phylogeny of the large genus Bembidion and related genera is inferred from four nuclear protein-coding genes (CAD, wingless, arginine kinase, and topoisomerase I), ribosomal DNA (28S and 18S), and the mitochondrial gene cytochrome oxidase I (COI). 230 of the more than 1200 species of Bembidion are sampled, as well as 26 species of five related genera, and 14 outgroups. Nuclear copies (numts) of COI were found sparsely scattered through sampled species. The resulting phylogeny, based upon individual gene analyses and combined analyses using maximum likelihood and parsimony, is very well supported at most nodes. Additional analyses explored the evidence, and corroborate the phylogeny. Seven analyses, each with one of the seven genes removed from the combined matrix, were also conducted, and yielded maximum likelihood bootstrap trees sharing over 92% of their nodes with the original, well-resolved bootstrap trees based on the complete set of seven genes. All key nodes were present in all seven analyses missing a single gene, indicating that support for these nodes comes from at least two genes. In addition, the inferred maximum likelihood tree based on the combined matrix is well-behaved and self-predicting, in that simulated evolution of sequences on the inferred tree under the inferred model of evolution yields a matrix from which all but one of the model tree's clades are recovered with bootstrap value >50, suggesting that internal branches in the tree may be of a length to yield sequences sufficient to allow their inference. All likelihood analyses were conducted under both a proportion-invariable plus gamma site-to-site rate variation model, as well as a simpler gamma model. The choice of model did not have a major effect on inferred phylogenies or their bootstrap values. The inferred phylogeny shows that Bembidarenas is not closely related to Bembidiina, and Phrypeus is likely distant as well; the remaining genera of Bembidiina form a monophyletic group. Lionepha, formerly considered a subgenus of Bembidion, is shown to be outside of the clade of Asaphidion+Bembidion, and is separated as its own genus. B. (Phyla) obtusum is quite isolated within Bembidion, and there is some evidence that the remaining Bembidion form a clade. Within Bembidion, there are three large clades that are well-supported, the Bembidion, Odontium, and Ocydromus Series. The Bembidion Series contains Bembidion (s. str.), Notaphus, Furcacampa, Emphanes, Trepanedoris, Diplocampa, and related Holarctic species; all species from South America, Australia, New Zealand; and most species from southern Africa and Madagascar. All species in South America, except for members of Notaphus and Nothocys, form a clade, the Antiperyphanes Complex, which has independently radiated into body forms and niches occupied by multiple, independent Northern-Hemisphere forms. All species from New Zealand, including Zecillenus, and Australian species formerly placed in Ananotaphus together form a clade. Bembidion (s. str.) and Cyclolopha are in a clade with the Old World, Southern Hemisphere lineages Notaphocampa, Sloanephila, and Omotaphus. The large subgenus Notaphus appears to have originated in South America, with all Northern Hemisphere Notaphus arising from within a south-temperate grade. All major variation in frontal furrows on the head is contained within the Bembidion Series. The Odontium Series contains subgenera Hirmoplataphus and Hydriomicrus, which together are the sister clade of Odontium, Bracteon, Ochthedromus, Pseudoperyphus, and Microserrullula. The very large Ocydromus Series, dominant in the Holarctic region, includes the Ocydromus Complex, with many subgenera, including Hypsipezum and Leuchydrium; the phylogeny within this group is notably at odds with the current classification. Also included in the Ocydromus Series are Nepha and Bembidionetolitzkya, as well as the Princidium Complex, in which the intertidal B. (Cillenus) laterale falls. Outside these three series are a number of smaller groups, including the Plataphus Complex (containing Blepharoplataphus, Plataphus, the latter including Plataphodes); the Hydrium Complex (Metallina, Chlorodium, and Hydrium, which contains Eurytrachelus), whose sister group might be subgenus Andrewesa; Trechonepha and Liocosmius, which might be sisters; and B. (Melomalus) planatum, which is not close to Plataphus. There is some evidence that these groups plus the Ocydromus and Odontium Series form a clade. A few enigmatic groups were harder to place. The sister group of the pair Philochthus plus Philochthemphanes might be B. wickhami; Eupetedromus is well outside the three major series and not related to Notaphus; the high-elevation Asian group Hoquedela is a very isolated lineage. Notaphiellus is removed from synonymy with Nothocys, and placed in synonymy with Notaphus; Plataphodes is synonymized with Plataphus, as Plataphus is paraphyletic otherwise; Eurytrachelus is synonymized with Hydrium. A new subgenus, Lindrochthus, is described to house the distinctive B. wickhami. The implications of the inferred phylogeny for some morphological characters used in Bembidiina systematics are explored, and some of the most widely used (e.g., location of discal seta ed3 on the elytron, and shape of the shoulder) are shown to be notably homoplastic. For example, the location of elytral seta ed3 has undergone at least nine transitions between two states.     

First report of wing dimorphism in the genus Orthomus (Coleoptera: Carabidae)

Wing polymorphism has been reported for several carabid beetles. Traditionally, a great number of ecological and evolutionary studies have focused on this peculiarity, which has implications on dispersal power. Research based on Orthomus berytensis specimens from two sampling areas of Tenerife (Canary Islands, Spain) has shown that this species exhibits a wing dimorphism, instead of being brachypterous. This makes O. berytensis the first Orthomus wing dimorphic species to date. Statistical differences in macropterous percentage between both sexes and localities were found. Also, a sexual dimorphism in elytra length and width was found, both being higher in females.     

Korean species of the genus Coptodera Dejean (Coleoptera: Carabidae: Lebiinae)

A truncatipennes‐group species, Coptodera japonica Bates, is newly recorded from Korea. Here we provide a diagnosis and systematic accounts of C. japonica Bates and a key to the species of Korean Coptodera.     

Rediscovery of Bembidion (Lymnaeum) nigropiceum (Marsham) (= puritanum Hayward) in Massachusetts, with remarks on biology and habitat (Coleoptera, Carabidae, Bembidiini)

Bembidion (Lymnaeum) nigropiceum (Marsham) (=puritanum Hayward), a European species introduced into Massachusetts but presumed not to have become established, has been rediscovered during the Boston Harbor Islands All Taxa Biodiversity Inventory undertaken by the Museum of Comparative Zoology and the National Park Service. A summary is presented of treatment of this species in North America. Data on specimens collected are presented, along with observations on habitat and biology. Some speculations are presented about its highly specialized habitat in the gravel pushed up by high tide, which may act as a food-trapping sieve. A few words are included about future actions needed to resolve questions of distribution and behavior.     

C-bands on chromosomes of four beetle species (Coleoptera: Carabidae, Silphidae, Elateridae, Scarabaeidae)

The C-banding pattern of Bembidion geniculatum, Silpha atrata, Prosternon tesselatum, and Epicometis hirta are presented. All analysed species have pracentromeric C-bands on autosomes and chromosome X but the widest ones are visible in the karyotype of B. geniculatum. In S. atrata, P. tesselatum, and E. hirta sex chromosome y is heterochromatic, only B. geniculatum having the Y chromosome wholly euchromatin. The results indicate that on the chromosomes of the investigated species do not have a terminal and an intercalar segments of heterochromatin.     

Ground beetles of the Ukraine (Coleoptera, Carabidae)

A review of the ground beetles of the Ukrainian fauna is given. Almost 750 species from 117 genera of Carabidae are known to occur in the Ukraine. Approximately 450 species of ground beetles are registered in the Carpathian region. No less than 300 species of ground beetles are found in the forest zone. Approximately 400 species of Carabidae present in the forest-steppe zone are relatively similar in species composition to those in the forest territories. Some 450 species of Carabidae are inhabitants of the steppe zone. Representatives of many other regions of heterogeneous biotopes such as forest, semi desert, intrazonal, etc. can be found in the steppe areas. The fauna of Carabidae (ca. 100 species) of the lowlands of southern Ukraine (sandy biotopes), situated mostly in the Kherson region, is very peculiar. The fauna of the Crimean mountains contains about 300 species. Conservation measures for the Carabidae are discussed.     

Three species of the genus Laboulbenia (Laboulbeniales) parasitic on Chydaeus bedeli (Coleoptera, Carabidae) from Asia

Three species of the genus Laboulbenia were recorded on Chydaeus bedeli (Coleoptera, Carabidae, Harpalini) from high-altitude localities in Asia. Laboulbenia obtusa was obtained from tarsi of the midlegs of the male hosts; Laboulbenia acrogeniodontis was on the margins of the elytra of the male and female hosts; and Laboulbenia polyphaga was on the elytra (near the apex) and the pronotum (at the base) of the male and female hosts. Each of the three species is distinct in the shape of the perithecia and the structure of the appendages.     

Fossil carabids from Baltic amber -II - A new subgenus of Bembidion Latreille 1802 (Coleoptera: Carabidae: Bembidiini)

Bembidion (Archaeophilochthus) christelae a new subgenus and species belonging to the tribe Bembidiini (Coleoptera: Carabidae: Trechinae) preserved in a piece of Baltic amber (Eocene) is described. A comparison with recent fauna is made, and paleobiology of the species is hypothetized.     

Carabidae from Vietnam (Coleoptera)

Members of the Vietnamese carabid fauna are arranged and nomenclaturally revised. As the result, 178 species belonging to 74 genera of 12 subfamilies, Brachininae, Omophroninae, Carabinae, Scaritinae, Bembidiinae, Trechinae, Pterostichinae, Harpalinae, Callistinae, Oodinae, Odacanthinae, and Lebiinae, are recorded from Vietnam. Within these, four species, Abacetus leucocetes Bates, 1873, Ago-nom (Metacolpodes) buchanani Hope, 1831, Steno-lopus (Egadromd) difficilis (Hope, 1845), and Phe-ropsophus {Stenaptinus) beckeri Jedlicka, 1930 are newly recorded. A species checklist with taxonomie information, type localities and worldwide distribution are provided.