Four new species of the Australian Pamborus Latreille (Coleoptera, Carabidae) carabid beetles

Abstract  Based on an examination of the type specimens of all the known species of the genus Pamborus Latrielle (Coleoptera, Carabidae) four new Australian species are recognised and are described as: P. euopacus sp. n., P. moorei sp. n., P. monteithi sp. n and P. cooloolensis sp. n. In addition, Pamborus viridis Gory, P. opacus Géhin and P. subtropicus Darlington are also redescribed based on the type material and lectotypes are designated for the first two species.     

Karyotypes of nine species of Cicindelini and cytotaxonomic notes on Cicindelinae (Coleoptera,Carabidae)

The male meioformula of seven species of Cicindelini from the Iberian Peninsula and one species from Canada is 9+XXXY. In addition, female data of Cylindera paludosa corroborate previous reports of males. The Palearctic species show a generalized karyotype which presents specific modifications, but it is not possible to establish if it is also present in species of other faunas.Present data indicate that multiple sex chromosomes is an ancestral condition for the subfamily Cicindelinae. It is postulated that the meioformula of the tribe Cicindelini is 9 or 10 plus XXY and that trends in karyotypic evolution are partially related to the geographic distribution of lineages. Numerical deviations observed in the tribe can be explained by changes occurring independently in autosomes and heterosomes, so that autosome-heterosome fusions do not seem to be a generalized event. A number of criteria may be developed for determining the polarity of change of karyotypic characters, thus making cytotaxonomic studies a valuable tool for understanding the systematics of Cicindelini.     

Parasitism of Neotropical Tiger Beetles (Coleoptera: Carabidae: Cicindelinae) by Anthrax (Diptera: Bombyliidae)

Bee flies (Bombyliidae) were recorded as parasitoids of larval tiger beetles at two rain forest localities (near São Paulo and Manaus) in Brazil. Anthrax gideon was reared from larvae of Oxycheila tristis. Up to 33 parasitoid larvae were found on a single tiger beetle host. Pupation of the bee fly took place in late August and the pupal stage lasted 14 days. The host digs horizontal burrows in contrast to the great majority of cicindelids, as does Pseudoxycheila tarsalis, the other known host of A. gideon. Two pupae of another undetermined Anthrax species were reared from larvae of Pentacomia ventralis in Central Amazonia. Pupation of this Anthrax sp. took place in October, the period of lowest host abundance.     

A reappraisal of the genus Megacaryon (Boraginaceae,Lithospermeae) based on molecular,morphological, and karyological evidence

The systematic position and phylogenetic relationships of Echium orientale, a rare endemic of the mountains of northern Turkey, were elucidated based on morphological, molecular, and karyological evidence.

Using nuclear and plastid DNA sequences, we found that this species is not included in Echium, but is rather at least as divergent from it as other related lineages, namely the South African Lobostemon and Echiostachys. Pollen characters revealed a strong affinity with Onosma, especially in the non-reticulate ornamentation of the tectum. Fruit morphology is unique in the genera of the Echium ‘alliance' (e.g. Echium, Pontechium, Lobostemon, and Echiostachys), because of the large size and the smooth, glossy nutlet surface as in most species of Onosma. Karyological observations show that this species is diploid with 2n = 12, and differing from both Lobostemon or Echium, where nearly all species have higher numbers (usually 2n = 14 and 2n = 16, respectively). Combined with a ‘megaherb' habit, the weak zygomorphism of the large flowers and a restricted range in the humid mountain forests of the Black Sea region, fruit, pollen and chromosome characters suggest an isolated position for E. orientale in the Echium alliance. We therefore propose its placement in the monotypic Megacaryon, a genus originally described by Boissier, but largely disregarded by later botanists. An original specimen is designated as generic lectotype.     

Molecular phylogeny and divergence time estimation of Cosmodela (Coleoptera: Carabidae: Cicindelinae) tiger beetle species from Southeast Asia

Cosmodela Rivalier, 1961, represents a genus of tiger beetles that currently contains 13 described species widely distributed across Southeast Asia. A phylogenetic analysis based on the mitochondrial DNA fragments 16S and COI and the nuclear marker wingless and a phylogeographic analysis using a COI fragment were carried out on two subspecies of Cosmodela aurulenta (C. a. juxtata and C. a. aurulenta). The results support the hypothesis that these two subspecies are significantly different to be considered as separate species that diverged during the Pleistocene. TAXONDNA analysis was used to investigate the capability of the COI region as a marker for discriminating both entities and to quantify intra‐ and intertaxa genetic variation. The minimum distance between C. aurulenta and C. juxtata was 2.7837%, and no overlap of intra‐ and intertaxa genetic divergence was observed. Both taxa, here considered as valid species, occur in sympatry in the Malay Peninsula, with C. aurulenta most probably originating from the area and C. juxtata a secondary colonizer that expanded southwards from the Asian mainland. Our data infer a continental origin of the Indonesian samples of C. aurulenta, and they most likely dispersed across the land bridges that emerged during glacial maxima to form Sundaland.     

Resolution of taxonomic problems in Australian Harpalini,Abacetini, Pterostichini,and Oodini (Coleoptera,Carabidae)

Taxonomic changes are made for several problematic Australian Carabidae in the tribes Harpalini, Abacetini, Pterostichini, and Oodini. Examination of types resulted in the synonymy of Veradia Castelnau, 1867 with Leconomerus Chaudoir, 1850; Nelidus Chaudoir, 1878, Feronista Moore, 1965, and Australomasoreus Baehr, 2007 with Cerabilia Castelnau, 1867; and newly combining Fouquetius variabilis Straneo, 1960 in the genus Pediomorphus Chaudoir, 1878; Australomasoreus monteithi Baehr, 2007 in the genus Cerabilia Castelnau, 1867; and Anatrichis lilliputana W.J. Macleay, 1888 in the genus Nanodiodes Bousquet, 1996. Cuneipectus Sloane, 1907 is placed in Pterostichini Bonelli, 1810, which is a senior synonym of Cuneipectini Sloane, 1907.     

Cytogenetic evidence for a complex of species within the taxon Anopheles maculatus (Diptera: Culicidae)

Two largely independent studies of chromosomes from natural populations of Anopheles maculatus provide evidence for several genetic species within the taxon. (1) Polytene chromosome variation shows four different rearrangements of arm 2 and three rearrangements of the X chromosome. There is strong evidence for three species. Two allopatric populations represent either dramatic geographic variation for two independent inversion systems within one of the genetic species, or represent two additional species. Their species status remains unresolved by this work. (2) Heterochromatic variation occurs in both X and Y chromosomes as revealed by Giemsa-banding of mitotic chromosomes from larval brains. The distribution and association of these various sex chromosomes give further evidence of a species complex. A preliminary correlation of these two kinds of chromosomal variation is given.     

Mating behavior and genitalic counterparts in tiger beetles (Carabidae: Cicindelinae)

Comparative studies on the structure of genitalia in Pseudoxychila tarsalis Bates and the copulating behavior in 5 species of Cicindela respectively complement similar findings by Freitag [1] on Cicindela spp. and Palmer [4] on P. tarsalis. These strengthen the hypothesis that in tiger beetles the flagellum fits into the spermatheca duct during copulation; that the main function of the flagellum, which is closed at the apex and not connected to the ejaculatory duct, is to open and prepare the lumen of the spermatheca duct for sperm movement from the bursa copulatrix to the spermatheca; and that copulation proceeds in 3 phases: phase 1 in which the lumen of the spermatheca duct is cleared by the flagellum, phase 2 in which the flagellum is withdrawn from the spermatheca duct, and phase 3 in which semen is transferred from the gonopore of the ejaculatory duct to the bursa copulatrix, usually with a spermatophore.     

Integration of conflict into integrative taxonomy: fitting hybridization in species delimitation of Mesocarabus (Coleoptera: Carabidae)

In species differentiation, characters may not diverge synchronously, and there are also processes that shuffle character states in lineages descendant from a common ancestor. Species are thus expected to show some degree of incongruence among characters; therefore, taxonomic delimitation can benefit from integrative approaches and objective strategies that account for character conflict. We illustrate the potential of exploiting conflict for species delimitation in a study case of ground beetles of the subgenus Carabus (Mesocarabus), where traditional taxonomy does not accurately delimit species. The molecular phylogenies of four mitochondrial and three nuclear genes, cladistic analysis of the aedeagus, ecological niche divergence and morphometry of pronotal shape in more than 500 specimens of Mesocarabus show that these characters are not fully congruent. For these data, a three‐step operational strategy is proposed for species delimitation by (i) delineating candidate species based on the integration of incongruence among conclusive lines of evidence, (ii) corroborating candidate species with inconclusive lines of evidence and (iii) refining a final species proposal based on an integrated characterization of candidate species based on the evolutionary analysis of incongruence. This procedure provided a general understanding of the reticulate process of hybridization and introgression acting on Mesocarabus and generated the hypothesis of seven Mesocarabus species, including two putative hybrid lineages. Our work emphasizes the importance of incorporating critical analyses of character and phylogenetic conflict to infer both the evolutionary history and species boundaries through an integrative taxonomic approach.     

Cryptic species of the subgenus Morphnosoma Lutshnik (Coleoptera: Carabidae; genus Pterostichus) from Japan

Pterostichus thunbergi Morawitz and its related species of Japan are revised based on the structure of membranous parts of their genitalia for the first time, under the name of the thunbergi species group. This species group is monophyletic and is placed in the subgenus Morphnosoma Lutshnik. The monotypic subgenus Moritapterus Berlov, erected for P. thunbergi, is synonymized with Morphnosoma. Comparative study of the endophallus revealed that P. thunbergi michinoku Nakane is a junior synonym of P. habui and that four species should be recognized in this group. Three of them are sympatric with P. thunbergi and one of them, Pterostichus (Morphnosoma) robustistylis, is described as a new species. Molecular phylogeny based on the COI gene (mitochondrial [mt]DNA) supported its independence as a distinct species. The resultant trees of mtDNA and comparative study of morphology revealed that P. thunbergi is probably non‐monophyletic. In the present study we suggest the presence of further cryptic species of Morphnosoma in Japan.     

A morphological, karyological and chemical study of the Apteranthes (Caralluma) europaea complex

The Apteranthes europaea complex (Apocynaceae–Asclepiadoideae–Ceropegieae–Stapeliinae) distributed from Morocco/southern Spain along the North African coast of the Mediterranean Sea up to Sinai, Negev and southern Jordan is studied with emphasis on stem and flower morphology by SEM, chromosome sizes and flavonoid compounds. Apteranthes europaea was found to be rich in rather uncommon flavone glycosides. Of these, luteolin 4'-neohesperidoside represents the major flavonoid of all samples; luteolin-3'- O -(6'- O -sinapoylglucoside)-4'- O -neohesperidoside and luteolin-3'- O -(6'- O -feruloylglucoside)-4'- O -neohesperidoside are reported here for the first time. Flavonol glycosides also occur, but in much smaller quantities. The different flower morphs, variation in stem and corolla epidermal structures, slight variations in the length of the 2 n  = 22 chromosomes and in quantitative flavonoid composition are taxonomically best reflected by subdividing the complex into a western var. europaea (Europe and Africa) and an eastern var. judaica (Sinai, Arabia).  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 149 , 419–432.     

Multiple cryptic genetic units in Hypothenemus hampei (Coleoptera: Scolytinae): evidence from microsatellite and mitochondrial DNA sequence data

Hypothenemus hampei is the most important insect pest of coffee and has spread to most coffee‐growing countries worldwide. There have been very few studies and none have addressed the population genetics of the beetle using microsatellite markers. In the present study, 683 individuals collected from 37 locations in 18 countries worldwide were screened at nine polymorphic microsatellite loci. Sixty‐five out the 683 and six additional individuals were analyzed on a 400‐bp fragment of the mitochondrial cytochrome oxidase I gene. Bayesian clustering analysis and phylogenetic approaches were used to infer the genetic structure of H. hampei over the sampling that encompassed almost all its range. Microsatellite markers made it possible to achieve sufficiently significant power for the delineation of five morphocryptic evolutionary units. Supported by 27 new COI haplotypes, an unexpected considerably high level of genetic differentiation and genetic divergence was revealed between five geographically delineated clusters. Both markers and approaches showed that the clusters included specimens from (1) Ethiopia, (2) Kenya and Uganda, (3) Brazil, (4) Central America excluding Jamaica, and (5) all samples from Asia, West Africa, and Jamaica. These findings clearly suggest the existence of a ‘species complex in H. hampei’. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 113–129.     

A quarter of a century succession of epigaeic beetle assemblages in remnant habitats in an urbanized matrix (Coleoptera, Carabidae)

We studied the long-term (23-24 years) species turnover and succession of epigaeic beetle assemblages (Coleoptera: Carabidae, incl. Cicindelinae) in three remnant habitats [cottonwood (Populus spp.) and oak (Quercus spp.) stands, and old fields] that are embedded within highly urbanized areas in central Minnesota. A total of 9,710 beetle individuals belonging to 98 species were caught in three sampling years: 1980, 1981 and 2005 in pitfall traps in identical locations within each habitat. Results indicate that there were 2-3 times greater trap catches in 2005 than in 1980 (cottonwood and oak stands, and old fields) and 1.4-1.7 times greater species diversity of beetles in 2005 than in the 1980-1981 suggesting increased habitat association by beetles over time. Although there were no significant differences in catches between 2005 and 1981 (only cottonwood stands and old fields), there was a trend where more beetles were caught in 2005. At the species-level, 10 times more of an open-habitat carabid species, Cyclotrachelus sodalis sodalis LeConte, was caught in 2005 than in 1980. However, trap catches of five other abundant carabid species [Pterostichus novus Straneo, Platynus decentis (Say), Platynus mutus (Say), Calathus gregarius (Say), and Poecilus lucublandus lucublandus (Say)] did not change indicating population stability of some beetle species. These remnant habitats were increasingly colonized by exotic carabid species as Carabus granulatus granulatus Linneaus, Clivina fossor (Linneaus) and Platynus melanarius (Illiger), that were trapped for the first time in 2005. Species composition of epigaeic beetles was quite distinct in 2005 from 1980 with 39 species reported for the first time in 2005, indicating a high turnover of assemblages. At the habitat-level, greatest species diversity was in cottonwood stands and lowest was in old fields, and all habitat types in 2005 diverged from those in 1980s, but not cottonwood stands in 1981. As our sampled areas are among some of the last remnants of the original oak savanna habitats in central Minnesota, we hypothesize that conservation of these sites may be critical to maintaining epigaeic beetle assemblages under increased urbanization pressure.     

Evaluation of relationships within the endemic Hawaiian Platynini (Coleoptera: Carabidae) based on molecular and morphological evidence

Relationships among 69 species of Hawaiian Platynini, a monophyletic beetle radiation, was investigated based on evidence from five data partitions, comprising mitochondrial and nuclear DNA sequences (cytochrome oxidase II, 624 bp; cytochrome b, 783 bp; 28S rDNA, 668 bp; wingless; 441 bp) and morphology (206 features of external and internal anatomy). Results from individual and combined data analyses generally support the monophyly of three putative divisions within Platynini in Hawaii: Division 0 (Colpocaccus species group), Division 1 (Blackburnia species group), and Division 2 (Metromenus species group). However, relationships within and among these three divisions differ from previous morphological hypotheses. An extensive series of sensitivity analyses was performed to assess robustness of recovered clades under a variety of weighted parsimony conditions. Sensitivity analyses support the monophyly of Divisions 0 and 1, but were equivocal for the monophyly of Division 2. A phylogeny based on combined data suggests at least four independent losses/reductions of platynine flight wings. The combined analysis provides corroboration for biogeographic hypotheses, including (1) colonization of Kauai by Hawaiian Platynini with subsequent dispersal and colonization along the island chain from Oahu to Maui Nui to Hawaii Island and (2) incongruent area relationships among Eastern Molokai, West Maui, and Haleakala for two species triplets.     

Phylogeny and systematics of the Acrapex apicestriata (Bethune-Baker, 1911) species complex (Lepidoptera,Noctuidae, Noctuinae,Apameini, Sesamiina) with the description of eight new species from the Afrotropics

Twelve morphologically similar species of Acrapex Hampson 1894, (Lepidoptera, Noctuidae, Noctuinae, Apameini, Sesamiina), from Western, Central and Eastern Africa are reviewed. Eight of these species are new to science and are described: Acrapex akunamatata n. sp. and A. incrassata n. sp. from Kenya; A. gracilis n. sp., A. iringa n. sp., A. lukumbura n. sp. and A. rungwe n. sp. from Tanzania; A. soyema n. sp. from Ethiopia; and A. zoutoi n. sp. from Benin. All 12 species belong to a species complex that we hereby define as the Acrapex apicestriata group. Host-plants for three of the new species are recorded: Setaria incrassata (Hochst.) Hack. for Acrapex incrassata; Cymbopogon pospishilii (K. Schum.) C.E. Hubb. for A. rungwe; and Andropogon perligulatus Stapf. for A. zoutoi. We also conducted molecular phylogenetic analyses (using maximum likelihood and Bayesian inference) on a six gene multimarker molecular dataset (four mitochondrial and two nuclear gene fragments; 4581 nucleotides in length) consisting of 15 Acrapex species (including seven species from the apicestriata group) and four outgroups species from the subtribe Sesamiina (from genera Busseola Thurau 1904, Sciomesa Tams & Bowden 1953, Pirateolea Moyal, Le Ru, Conlong, Cugala, Defabachew, Matama-Kauma, Pallangyo & Van den Berg 2010 and Sesamia Boisduval & Guenée 1852). Both maximum likelihood and Bayesian inference analyses yield a similar and well-supported topology, which supports the monophyly of the apicestriata group.     

Genome sizes of three species in the subtribe Carabina (Coleoptera: Carabidae)

Genome sizes of three species in the subtribe Carabina (Coleoptera: Carabidae: tribe Carabini) were estimated from nuclei of spermatozoa using a flow cytometry. The estimated haploid genome sizes were about 330 megabases (Mb) for Damaster blaptoides, about 280 Mb for Carabus uenoi and about 260 Mb for Carabus yamato.     

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.     

Evaluating evolutionary history in the face of high gene tree discordance in Australian Gehyra (Reptilia: Gekkonidae)

Species tree methods have provided improvements for estimating species relationships and the timing of diversification in recent radiations by allowing for gene tree discordance. Although gene tree discordance is often observed, most discordance is attributed to incomplete lineage sorting rather than other biological phenomena, and the causes of discordance are rarely investigated. We use species trees from multi-locus data to estimate the species relationships, evolutionary history and timing of diversification among Australian Gehyra—a group renowned for taxonomic uncertainty and showing a large degree of gene tree discordance. We find support for a recent Asian origin and two major clades: a tropically adapted clade and an arid adapted clade, with some exceptions, but no support for allopatric speciation driven by chromosomal rearrangement in the group. Bayesian concordance analysis revealed high gene tree discordance and comparisons of Robinson–Foulds distances showed that discordance between gene trees was significantly higher than that generated by topological uncertainty within each gene. Analysis of gene tree discordance and incomplete taxon sampling revealed that gene tree discordance was high whether terminal taxon or gene sampling was maximized, indicating discordance is due to biological processes, which may be important in contributing to gene tree discordance in many recently diversified organisms.     

Phylogeny of ground beetles subgenus Nialoe (s. lat.) Tanaka (Coleoptera: Carabidae; genus Pterostichus): A molecular phylogenetic approach

We constructed a phylogeny of the ground beetle subgenus Nialoe ( s. lat. ), genus Pterostichus (Coleoptera: Carabidae) based on two mitochondrial (cytochrome oxidase I and 16S ribosomal DNA) and one nuclear (28S ribosomal DNA) gene sequences. Thirty-three representative species of the group and three outgroup species were analyzed. The resultant trees (maximum parsimonious, maximum likelihood and Bayesian trees of the combined data of the three gene sequences) indicated that there are two large and three small lineages in the group, some of which were supported by a previous morphology-based phylogeny. In all the analyses, the small lineage composed of two Korean species is sister to the rest of the subgenus, but relationships of other four lineages differed among the analyses and remained unresolved. The implications of the present results are discussed in terms of taxonomy and biogeography of the group.