Phylogenetic studies of the subgenus Nialoe (s. lat.) (Coleoptera, Carabidae, Genus Pterostichus), part 2: The Asymmetricus species group

Pterostichus (Nialoe) asymmetricus Bates and its allies are revised based on the membranous parts of their genitalia. Four new taxa, P. (N.) ovaliphallus sp. nov., P. (N.) fujimurai ibukiyamanus ssp. nov., P. (N.) shotaroi kiimontanus ssp. nov., and P. (N.) basilobatus sp. nov. are described. A cladistic analysis based on 31 morphological characters reveals that Daisenilaoe (s. str.) Nakane and Straneo is included as one clade within Nialoe (s. str.). The speciation and dispersal process of this species group are also discussed.     

Notes on two species of the subgenus Lyrothorax Chaudoir (Coleoptera: Carabidae; genus Pterostichus), Pterostichus amagisanus Tanaka and Ishida and Pterostichus fujitai Tanaka and Ishida

Two species of the subgenus Lyrothorax Chaudoir (Coleoptera: Carabidae; genus Pterostichus ), Pterostichus amagisanus Tanaka and Ishida and Pterostichus fujitai Tanaka and Ishida, were revised based on the male endophallus (inner sac everted from aedeagus). P. amagisanus was newly recorded based on a single male from Kyushu, southwest Japan, far from its known distribution (Honshu; the Fuji-Hakone-Izu volcano area), although additional materials are necessary to confirm this record. Despite a highly disjunct distribution, no conspicuous difference was recognized in either external or genital characters between the materials from Honshu and Kyushu. The nominal species P. fujitai was separated into two species, P. fujitai (Honshu) and Pterostichus eoyoritomus sp. nov. (Shikoku; type locality: Mount Jingayama); these two species have some significant differences in the endophallic structures. Character states in male genitalia suggest a sister relationship between P. eoyoritomus sp. nov. and Pterostichus yoritomus Bates.     

Structure of the male reproductive accessory glands of Pterostichus nigrita (Coleoptera: Carabidae), their role in spermatophore formation

Accessory gland secretions of male insects have many important functions including the formation of spermatophores. We used light and electron microscopy to investigate the structure of the accessory glands and posterior vasa deferentia of the carabid beetle Pterostichus nigrita to try to determine where spermatophore material is produced. Each accessory gland and posterior vas deferens had an outer layer of longitudinal muscle, beneath which was a layer of connective tissue and a thin band of circular muscle, all of which surrounded a layer of epithelial cells lining the lumen of the ducts. Based on the ultrastructure of the epithelial cells, and their secretory products, we identified two epithelial cell types in each region (distal and proximal) of the accessory glands and four types in the posterior vas deferens. Most secretory products, which stained positively for proteins and some mucins, were released into the lumen of the ducts by apocrine secretion. The accessory glands produced one type of secretory product whereas in posterior vasa deferentia, four types of secretory products were found layered in the lumen. Our results suggest that most of the structural material used to construct a spermatophore is produced by the cells of the posterior vasa deferentia.     

Geographic variability of the life cycle in Pterostichus melanarius (Coleoptera, Carabidae)

In Pterostichus melanarius, the seasonal dynamics of activity is characterized by two peaks (in mid-June-early July and in late July-mid-August). In southwestern Moldova, these peaks were registered twenty days later than in Moscow Province. In both regions, larvae and adults hibernate. Both hibernated immature adults and adults that had already bred before wintering start reproduction simultaneously. In hibernated immature individuals, the average number of eggs per female constituted 1.5 times that in the postgenerative females. In the steppe zone, the total number of eggs laid by immature females was closed to that laid by previously bred females, while in the southern taiga zone, hibernated immature females laid a greater number of eggs. In the steppe zone, females of a new generation lay a somewhat lower number of eggs in comparison with hibernated immature females, whereas the total number of laid eggs is significantly higher. In the southern taiga zone, both the average number of laid eggs and their total number are equal in females of a new generation and hibernated immature females. In Moldova, the majority of individuals of a new generation breed in the current season, whereas in Moscow Prov., about half the individuals hibernate without breeding. Over the largest part of the range, the life cycle of P. melanarius combines the one-year development with hibernating larvae with two-year development with the hibernating immature and postgenerative adults. The two subpopulation groups, wintering at different phases of the ontogenesis, are totally isolated at the northern border of the range. Within each subpopulation., individuals are characterized by the biennial life cycle; as a result, the polyvariant character of the life cycle turns into the monovariant one. We suggest that such a life cycle should be designated as the compensatory monovariant cycle, and populations, in which it is realized, as byside populations (from the English “side by side”).     

Two new species of Pterostichus Bonelli subgenus Pseudoferonina Ball (Coleoptera, Carabidae, Pterostichini) from the mountains of central Idaho, U.S.A

Two new species of Pterostichus Bonelli subgenus Pseudoferonina Ball, are described from the mountains of central Idaho: Pterostichus bousqueti Bergdahl [type locality = small tributaries of South Fork of Payette River watershed, ca. 1170 m (3840 ft), 44.0675°; -115.6822°, near Lowman, Salmon River Mountains, Boise County, Idaho, U.S.A.] and Pterostichus lolo Bergdahl [type locality = Cottonwood/Orogrande Creek, ca. 870 m (2850 ft), 46.5528°; -115.5522°, North Fork of Clearwater River watershed, Clearwater Mountains, near Bungalow, Clearwater County, Idaho, U.S.A.]. Males of Pterostichus bousqueti and Pterostichus lolo are easily distinguished from each other and the seven previously described Pseudoferonina species by the form of the median lobe of the aedeagus, and from most individuals of the other species of Pseudoferonina in Idaho by features of pronotal shape and macrosculpture. Both species appear to be obligate ripicolous hygrophiles, restricted in distribution primarily to the margins of small montane streams in forested areas. Widespread intensive stream surveys for Pseudoferonina over many years indicate the geographic ranges of both species are highly localized, and additional undescribed species may occur in Idaho.     

A key to species of subgenus Lithochlaenius (Coleoptera, Carabidae, Chlaeniini, Chlaenius), with descriptions of three new species

Three new species of genus Chlaenius Bonelli subgenus Lithochlaenius Kryzhanovskij are described from China: Chlaenius chuanqianensis Liu & Liang, sp. n. (type locality: Xishui, Guizhou Province), Chlaenius linwensini Liu & Liang, sp. n. (type locality: Fujian Province), and Chlaenius propeagilis Liu & Kavanaugh, sp. n. (type locality: Gaoligongshan, Yunnan Province). Seven species of the subgenus are redescribed: Chlaenius agiloides Jedli?ka, Chlaenius formosensis Lorenz, Chlaenius agilis Chaudoir, Chlaenius leishanensis Kirschenhofer, Chlaenius noguchii Bates, Chlaenius rambouseki Lutshnik, and Chlaenius wrasei Kirschenhofer. Additional taxonomic changes include the following: Chlaenius formosanus Jedli?ka is treated as a junior synonym of Chlaenius rambouseki Lutshnik and Chlaenius anchomenoides Bates, syn. n. and Chlaenius nuristanus Jedli?ka as junior synonyms of Chlaenius agilis Chaudoir, syn. n.Chlaenius latroLaFerté-Sénectère is considered a nomen nudum stat. n. and unavailable, leaving Chlaenius agilisChaudoir as the next available name. Chlaenius nuristanusaberration rubridipesJedli?ka is also an unavailable name. Chlaenius formosensisLorenz (=Chlaenius formosanusHabu) is returned to species status stat. n. A key to adults of the 10 known species of subgenus Lithochlaenius is provided.     

Phylogenetic relationships of Dalyat mirabilis Mateu, 2002, with a revised molecular phylogeny of ground beetles (Coleoptera, Carabidae)

Dalyat mirabilis Mateu 2002 (Coleoptera: Carabidae) is a cave species recently described from SE Spain, which, based on morphological analyses, has been related to the Promecognathinae (with one genus in western North America and four genera in South Africa). In this paper, we investigated the phylogenetic relationships of the main lineages of family Carabidae, and the placement of Dalyat among them, with the 18S rRNA full sequence and a fragment of wingless with the use of parsimony, a fast maximum likelihood algorithm (implemented in the program phyml ), and Bayesian posterior probabilities. Although with wingless alone the relationships of Dalyat were not robustly supported, both with 18S rRNA and in a combined analysis there was a strong support for a sister relationship between Promecognathus and Dalyat with the three methods used. Using a molecular-clock approach the two lineages were estimated to have diverged at a similar (or slightly earlier) age than the origin of Harpalinae, known to have radiated in the Cretaceous. This is compatible with a vicariant origin of the lineage leading to Dalyat because of the isolation of the Iberian plate from Pangea in the late Jurassic to early Cretaceous. Other robust relationships within the Carabidae are the monophyly of Harpalinae (including Morionini, Peleciini and Pseudomorphini), its sister relationship with Brachininae, and the inclusion of these two subfamilies together with Scaritini and the austral Psydrinae in a strongly supported clade (the 'higher' Carabidae).     

Morphological association between spermatophores and male genitalia in carabid beetles of the tribe Pterostichini (Coleoptera: Carabidae)

A morphological association between genitalia and ejaculates could provide insight into the function and evolution of genitalia. In this study, the morphologies of the ejaculates and male genitalia of 15 species of Pterostichini and two species of Platynini (Coleoptera: Carabidae) are described. All the species examined formed a spermatophore, the morphology of which could be classified into three types based on its relative volume in the female vaginal cavity and the presence or absence of a pluglike conformation. Male genital morphology could be divided into two types by the direction of the endophallus and gonopore. Species with a strongly bent endophallus invariably formed a pluglike spermatophore. The results suggest that the peculiar shape of endophallus found in some species of Pterostichini may function in forming the pluglike structure of the spermatophore.     

Cladistic analysis, phylogeny and biogeography of the Hawaiian Platynini (Coleoptera: Carabidae)

The 128 known native Hawaiian species of the tribe Platynini are analysed cladistically. Cladistic analysis is based on 206 unit-coded morphological characters, and also includes forty-one outgroup taxa from around the Pacific Rim. Strict consensus of the multiple equally parsimonious cladograms supports the monophyly of the entire species swarm. The closest outgroup appears to be the south-east Asian-Pacific genus Lorostema Motschulsky, whose species are distributed from India and Sri Lanka to Tahiti, supporting derivation of the Hawaiian platynines from a source in the western or south-western Pacific. The biogeographic relationships of the Hawaiian taxa are analysed using tree mapping, wherein items of error are minimized. The area cladogram found to be most congruent with the phylogenetic relationships, and most defensible based on underlying character data is {Kauai[Oahu(Hawaii{Lanai[East Maui(West Maui + Molokai)]})]}. This progressive vicariant pattern incorporates progressive colonization from Kauai, and vicariance of the former Maui Nui into the present islands of Molokai, Lanai, West Maui and East Maui. The evolution of flightlessness, tarsal structure, pronotal setation and bursal asymmetry are evaluated in the context of the cladogram. Brachyptery is a derived condition for which reversal is not mandated by the cladogram, although repeated evolution of reduced flight wings is required. Tarsal structure supports Sharp's (1903) recognition of Division 1 as a monophyletic assemblage, but exposes his Division 2 as a paraphyletic group requiring removal of the genus Colpocaccus Sharp. Pronotal setation is exceedingly homoplastic, and is not useful for delimiting natural groups. Left-right asymmetry of the bursa copulatrix reversed twice independently, resulting in mirror-image bursal configurations in B. rupicola and Prodisenochus terebratus of East Maui. The amount of character divergence is greater among species comprising Division 1 than among species of its sister group, the redefined Division 2. Based on superior fit of Division 1 relationships to the general biogeographic pattern, a greater speciation rate coupled with more extensive extinction is rejected as the cause for this greater divergence. Intrinsic differentiation in the processes underlying cuticular evolution appears to be more consistent with the observed biogeographic and morphological patterns.     

Phylogenetic studies of Meloidae (Coleoptera), with emphasis on the evolution of phoresy

One of the most recent classifications of Meloidae is based on the assumption that phoretic first‐instar larvae evolved twice in the family, once in Meloinae and again in Nemognathinae. Within Meloinae, this scheme places all presumed phoretic taxa in Meloini regardless of other characteristics. This paper challenges this classification with a cladistic analysis of all meloid genera whose first‐instar larvae were available for study. It concludes that phoresy evolved several times in Meloinae alone and that Meloini, when defined to include all phoretic genera, is polyphyletic. Cladistic support also is presented for four subfamilies of Meloidae and for several of the traditional tribes recognized in recent classifications.     

Chromosomes and identification of the sibling species Pterostichus nigrita (Paykull) and P. rhaeticus Heer (Coleoptera: Carabidae)

The karyotypes of Pterostichus nigrita (Paykull) and P. rhaeticus Heer are described. Both species have eighteen pairs of autosomes, sex chromosomes which are XO (♂) and XX (♀), plus a variable number of totally heterochromatic B chromosomes. Males may be identified by the form of the inflated endophallus, but the shape of the right paramere is not a reliable character. Females may be identified by the form of the eighth abdominal sternite.     

Comparative morphology of the male genitalia of Aphididae (Insecta,Hemiptera): part 1

The present study provides new data concerning the morphology of the male genitalia of Aphididae and unifies their nomenclature. The structure of the male genitalia of 31 species from 26 genera of Aphididae was studied with light and scanning electron microscopy. In the studied species, the genitalia of males consist of a phallus composed of the sclerotized basal part with its articulation and a membranous apical part—an aedeagus. Laterally of the phallus, there is a pair of setose parameres. The shape of the aedeagus, the shape and length of the sclerotized basal part and its articulation as well as the variability of parameres in their form and the number of setae are recognized as important systematic signs of the genitalia. These characters are considered in conjunction with the phylogenetic relationships among the studied taxa.     

A New Species of the Oriental Ground-Beetle Genus Dioryche MacLeay (Coleoptera,Carabidae) from Oman

Entomological Review - Dioryche MacLeay, 1825 is a moderately diverse selenophorine genus distributed in the Oriental Region. In this contribution, D. orszuliki sp. n. is described from the extreme...     

Phylogeny of the Genus Cladonia s.lat. (Cladoniaceae, Ascomycetes) Inferred from Molecular, Morphological, and Chemical Data

Phylogenetic relationships within the genus Cladonia , including Cladina (Cladoniaceae, Lecanoromycetes), were reconstructed based upon simultaneous analyses of DNA sequences and morphological and chemical data. We used sequences from the internal transcribed spacer 1 (ITS1), the 5.8 rDNA gene, and the internal transcribed spacer 2 (ITS2) of the nuclear rDNA gene cluster, and partial sequences from the protein-coding β-tubulin gene. The analyses included 235 specimens of 168 taxa representing all currently recognized sections of Cladonia and Cladina and the outgroup genera Cladia, Pycnothelia , and Ramalea . Analyses were performed using optimization alignment with three different parameter values. The results of all analyses support the inclusion of Cladina in Cladonia . The current sectional division of Cladonia was not supported, and a new provisional classification for the genus is proposed.     

兜兰属宽瓣亚属（广义）的被充研究

对新种小囊兜兰Paphiopedilum globulosum与新变种扁球兜兰P.micranthum var.oblatum作了描述与绘图。两者均为采自云南东南部的栽培植物,与硬叶兜兰P.micranthum近似。但小囊兜兰具有宽得多的叶（宽2．8－3．4cm),先端多少呈钩状、宽1．1－1．2cm的辱瓣,以及与唇瓣囊口几乎同样大小的退化雄蕊,而扁球兜兰则叶宽2．2－3cm,唇瓣扁球形,囊深不超过1cm,可区别于硬叶兜兰。上述种类均属于宽瓣亚属（广义）。该亚属有16个种产中国、越南或两地共有。本文提供了分种检索表。     

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.     

The Influence of Mucus Production by the Slug,Deroceras reticulatum,on Predation by Pterostichus madidus and Nebria brevicollis (Coleoptera: Carabidae)

Slugs are important pests in many agricultural crops and potential biological control agents are being studied as an alternative to molluscicides. Carabid beetles may be able to reduce slug populations, but the defence mucus of slugs may deter some predator attacks. Interactions between the carabids Pterostichus madidus (Fabricius) and Nebria brevicollis (Fabricius) with healthy and 'stressed' (unable to produce defence mucus) Deroceras reticulatum (Müller) were investigated in laboratory conditions. Both beetle species consumed significantly more stressed slugs than controls. Defence mucus production by control slugs hampered beetle attacks. These generalist beetle species did not direct their attacks at vulnerable parts of the prey as equal numbers of contacts were made on the slug head, mantle and tail. Blowfly ( Calliphora sp.) larvae are readily consumed by P. madidus and N. brevicollis . Calliphora larvae coated in slug defence mucus were less acceptable to both beetle species compared with control larvae. Results indicate that these generalist beetle species are unable to overcome the defence mucus production of healthy slugs. Slugs sub-lethally poisoned by molluscicides may be a more suitable prey item due to a reduction in defence mucus production.     

A New Species of the Ground-Beetle Genus Cymindis Latreille, 1805 (Coleoptera,Carabidae: Lebiini) from Sichuan Province,China

Entomological Review - A new species of the genus Cymindis Latreille, 1805, C. (Paracymindis) batangensis Kabak et Liang, sp. n., related to C. wrasei Kabak, 2006, is described from the Lanashan...     

Catalogue and Check-list of the Carabid Subgenus Cribramara Kryzhanovskij, 1964 (Genus Amara Bonelli, 1810) (Coleoptera,Carabidae: Zabrini)

Entomological Review - New and specified distributional data on eight hitherto known species of the subgenus Cribramara Kryzhanovskij, 1964 (genus Amara Bonelli, 1810) are provided and mapped. The...     

Dorsipes caspius n. sp. (Acari: Podapolipidae), a subelytral parasite of Pterostichus caspius (Menetries) (Coleoptera: Carabidae) with notes on host range of the genus and the distribution of the platysmae group

The ectoparasitic mite Dorsipes caspius n. sp. (Heterostigmata: Podapolipidae) belonging to the platysmae species group collected from the beneath elytra of Pterostichus (Lyrothorax) caspius (Menetries) (Coleoptera: Carabidae) in northern Iran, is described. This is the first record of species of the platysmae group from the Middle East. Keys to adult and larval female stages of world species of the platysmae group are provided. The host range of all species of the genus and the distribution of all representatives of the group are discussed.