Phylogeny of minute carabid beetles and their relatives based upon DNA sequence data (Coleoptera, Carabidae, Trechitae)

The phylogeny of ground beetles of supertribe Trechitae is inferred using DNA sequences of genes that code for 28S ribosomal RNA, 18S ribosomal RNA, and wingless. Within the outgroups, austral psydrines are inferred to be monophyletic, and separate from the three genera of true Psydrina (Psydrus, Nomius, Laccocenus); the austral psydrines are formally removed from Psydrini and are treated herein as their own tribe, Moriomorphini Sloane. All three genes place Gehringia with Psydrina. Trechitae is inferred to be monophyletic, and sister to Patrobini.Within trechites, evidence is presented that Tasmanitachoides is not a tachyine, but is instead a member of Trechini. Perileptus is a member of subtribe Trechodina. Against Erwin's hypothesis of anillines as a polyphyletic lineage derived from the tachyine genus Paratachys, the anillines sampled are monophyletic, and not related to Paratachys. Zolini, Pogonini, Tachyina, and Xystosomina are all monophyletic, with the latter two being sister groups. The relationships of the subtribe Bembidiina were studied in greater detail. Phrypeus is only distantly related to Bembidion, and there is no evidence from sequence data that it belongs within Bembidiina. Three groups that have been recently considered to be outside of the large genus Bembidion are shown to be derived members of Bembidion, related to subgroups: Cillenus is related to the Ocydromus complex of Bembidion, Zecillenus is related to the New Zealand subgenus Zeplataphus, and Hydrium is close to subgenus Metallina. The relationships among major lineages of Trechitae are not, however, resolved with these data.     

The effects of forestry on carabid beetles (Coleoptera: Carabidae) in boreal forests

As compared to natural forests, managed boreal forests are younger, more homogeneous in terms of tree age and species composition, and consist of smaller fragments. Here we examine the effects of such characteristics caused by forestry on carabid beetles (Coleoptera, Carabidae) in the boreal region. The main results are the following. (1) Fragmentation of forests and the size of a fragment appear not to be crucial for the survival of the majority of forest carabids, as they tend to be distributed over various successional stages, but species requiring old-growth habitats suffer. (2) For carabids there appear to be no or very few edge specialist species, and forest-open land edges appear to be effective barriers for species associated with forest or open habitat. However, generalist species easily cross the edge, and edges of forest fragments may be invaded by species from the surrounding open habitat. (3) Habitat change following clear-cutting dramatically changes the composition of carabid assemblages: species restricted to mature forests disappear and open-habitat species invade, while habitat generalists survive at least in the short term. Carabid diversity can probably best be maintained if forest management mimics natural processes, maintains natural structures and includes the natural composition of vegetation and other structural elements (such as dead wood) within the stands, provided that these forest features can be maintained and recreated through forest management practices. At a larger scale, the whole spectrum of forest types and ages (especially old-growth forests), and different successional processes (especially fire) should be maintained. These require the development and use of innovative logging methods, and the planning, implementation, and assessment of landscape-scale ecological management strategies.     

The ground beetles (Coleoptera: Carabidae) of exposed riverine sediments in Scotland and northern England

A survey of invertebrates on exposed riverine sediments (ERS) in four catchments of rivers in Scotland and northern England was carried out in 1996 and 1997 using pitfall traps. The resulting 179 lists of ground beetle species were used in ordination and classification analyses to determine the different types of habitats and the environmental factors influencing species assemblage distribution. Types of ERS habitat differed between highland and lowland catchments; there were also different numbers of habitat types in each catchment. The distribution of ERS within catchments was related to geology, position in catchment, sediment composition and the amount of vegetation. Evidence of the positive nature of river engineering was identified and the role of river management is discussed. The numbers of nationally rare and scarce ground beetle species recorded from the ERS habitats in each catchment are reported and the factors affecting their distribution discussed.     

Impact of commercial forest management on geophilous carabid beetles (Coleoptera: Carabidae) in tall,wet Eucalyptus obliqua forest in southern Tasmania

Abstract The impact of silvicultural practices (clearfelling, slash burning and sowing) on carabid beetle communities in tall wet Eucalyptus obliqua forests in southern Tasmania was examined using traplines of pitfalls in a chronosequence of regenerating coupes. Total species richness (n= 18) was modest compared to other temperate forests and was not systematically changed by forest management, although it was considerably lower in 20 year old regrowth than in younger regrowth or old-growth controls, possibly due to habitat simplification. The Shannon-Weiner Index was at a minimum in intermediate aged regeneration. TWINSPAN analysis assisted recognition of beetle communities typical of broad stages in the forest succession, with a major dichotomy between most old-growth sites plus young sites and intermediate plus advanced regeneration sites. Vector fitting of environmental variables in an ordination of the sites by non-metric multidimensional scaling revealed a significant influence for the age of regeneration and litter depth, but not bare ground percentage or soil pH. Survival of carabids in the habitat mosaic created by commercial forestry activity is likely to depend on both reinvasion from edges and survival of individuals in the fire-protected refuges that exist within the coupes. Pioneer species were winged, small in size, and non-endemic whereas the opposite was true of the fauna in the older sites. Carabids in eucalypt forests have good potential as indicators but their seasonality in occurrence demands that sampling be extended over most of the year.     

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.     

Habitat choice of some field-inhabiting carabid beetles (Coleoptera: Carabidae) studied by recapture of marked individuals

ABSTRACT.

• 1 Habitat choice of some field-inhabiting carabid beetles was studied in the field, and included the adult-overwintering Bembidion lampros Herbst, Pterostichus cupreus L. and Agonum dorsale Pontoppidan, and the larval-overwintering Trechus secalis Paykull, Pterostichus melanarius Illiger, P.niger Schaller and Harpalus rufipes De Geer. These were compared to the forest-inhabiting and adult-overwintering species, Carabus nemoralis Müller.
• 2 Marked beetles were released in the centre of a circular enclosure which was placed on the edge between a cereal field and a wood. The direction of movement in this circle was compared with that in a control circle placed entirely in the cereal field.
• 3 In the field/wood circle, adult-overwinterers and the larval-overwin-terer H.rufipes choose to move into the cereal field. The majority of T.secalis. P.melanarius and C.nemoralis, however, moved into the wood, while P.niger exhibited no preference.
• 4 In the field circle, adult-overwinterers moved towards the more sun-exposed parts of the circle, whereas the movements of larval-overwin-terers were random.
• 5 Some species-specific characteristics considered important for habitat choice by these carabid beetles, and the relevance of these characteristics for their dispersal behaviour are discussed.

     

Movements of carabid beetles (Coleoptera: Carabidae) inhabiting cereal fields: a field tracing study

Summary The individual movements of four carabid species were investigated by tracing beetles in the field using a portable radar system (harmonic radar). The field-inhabiting a Pterostichus melanarius Illiger, P. niger Schaller and Harpalus rufipes De Geer were traced in cereal fields. The movements of P. melanarius Mnd P. niger were also tracked in a wood and then compared with the movements of the forest-species Carabus nemoralis Müller. P. melanarius and P. niger displayed directed movements in cereal fields, although P. niger moved faster than P. melanarius. The movement pattern of H. rufipes in the field, and C. nemoralis and P. malanarius in the wood can be described as a c correlated random walk. In cereal fields, P. melanarius burrowed actively into the soil, and P. niger searched out crevices. In contrast, none of these two carabids burrowed in the forest soil, whereas C. nemoralis did. No statistically significant difference was observed between the movements of hungry and satiated beetles, although starved beetles tended to disperse shorter distances per move and turned back more often (especially P. niger). The significance of the various movement patterns exhibited by the species studied for their reproduction and survival on arable land is discussed.     

转植酸酶基因玉米对步甲群落动态的影响

本研究以陷阱法研究转植酸酶基因玉米及其亲本玉米在整个生长季节对地表步甲类群的影响。每个玉米品系设置6块样地, 两个玉米品系交替排列, 每个样地设置2个采样点, 共设置24个样点, 整个玉米生育期取样8次。本研究共采集步甲标本8 012头, 隶属于11属23种, 其中黄斑青步甲Chlaenius micans个体数量分别占个体总数的87.54%, 为玉米田内的主要常见物种, 蠋步甲Dolichus halensis、 后斑青步甲Chlaenius posticalis和单齿蝼步甲Scarites terricola个体数量分别占除黄斑青步甲之外所有步甲个体总数的34.77%, 31.16%和6.21%, 这些物种组成当地玉米田内的常见物种。步甲物种多样性与常见物种个体数量随季节变化明显, 且转植酸酶基因玉米与其亲本玉米田内步甲物种多样性和常见物种个体数量的季节变化趋势相似度较高。重复测量方差分析（Repeated ANOVA）结果表明, 转植酸酶基因玉米对步甲物种丰富度、 物种多度、 香农威纳多样性指数和均匀度指数均没有显著影响; 转植酸酶基因玉米田内的后斑青步甲个体数量明显增加, 但其他常见物种没有显著变化。基于非度量多维度（NMDS）的群落结构分析表明转植酸酶基因玉米与亲本玉米田内的步甲群落结构非常相似。本文结果表明转植酸酶基因玉米的种植对步甲物种多样性及常见物种没有明显影响。     

Absence asymmetry: the evolution of monorchid beetles (Insecta: Coleoptera: Carabidae)

Asymmetrical monorchy, or the complete absence of one testis coupled with the presence of its bilateral counterpart, is reported for 174 species of the carabid beetle tribes Abacetini, Harpalini, and Platynini (Insecta: Coleoptera: Carabidae) based on a survey of over 820 species from throughout the family. This condition was not found in examined individuals of any other carabid beetle tribes, or of other adephagan beetle families. One monorchid taxon within Platynini exhibits symmetrical vasa deferentia at the beginning of the pupal stadium, suggesting that developmental arrest of the underdeveloped vas deferens takes place in pupation. The point at which development of the testis is interrupted is unknown. Complete absence of one organ of a bilateral pair--absence asymmetry--is rarely found in any animal clade and among insects is otherwise only known for testes in the minute-sized beetles of the family Ptiliidae, ovaries in Scarabaeinae dung beetles, and ovaries of some aphids. Based on current phylogenetic hypotheses for Carabidae, testis loss has occurred independently at least three times, and up to five origins are possible, given the variation within Abacetini. Clear phylogenetic evidence for multiple independent origins suggests an adaptive or functional cause for this asymmetry. A previously posited taxon-specific hypothesis wherein herbivory in the tribe Harpalini led to testis loss is rejected. Optimal visceral packing of the beetle abdomen is suggested as a general explanation. Specifically, based on the function of various organ systems, we hypothesize that interaction of internal organs and pressure to optimize organ size and space usage in each system led to the multiple origins and maintenance of the monorchid condition. Testes are the only redundant and symmetrically paired structures not thought to be developmentally linked to other symmetrical structures in the abdomen. Among all possible organs, they are the most likely--although the observed frequency is very small--to bypass constraints that maintain bilateral symmetry, resulting in absence asymmetry. However, based solely on our observations of gross morphology of internal organs, no function conclusively explains the ontogenetic loss of one testis in these taxa. Unlike the analogous absence asymmetry of organs in other animal groups, no dramatic body-form constraint--e.g., snakes and lung loss, ptiliid beetles' small body-size and relatively giant sperm--or adaptive scenario of improved locomotory performance--e.g., birds and ovary loss due to flight constraints-applies to these carabid beetles. We tentatively suggest that testis loss is driven wholly by an interaction among the internal organs of these beetles, possibly due to selective pressure to maximize the comparatively large accessory glands found in these taxa. However, as the ordering of these evolutionary events of testis loss and accessory gland size increase is not known, large accessory glands might have secondarily evolved to compensate for a decreased testicular output.     

Taxonomic review of the Pterostichini and Loxandrini fauna of New Caledonia (Coleoptera, Carabidae)

The generic-level taxa included in Pterostichini and Loxandrini from New Caledonia are reviewed and a key to genera and species provided. Two new genera are described, Paniestichus and Abacophrastus, with the following new species: Paniestichus subsolianus, Abacophrastus millei, Abacophrastus chapes, Abacophrastus carnifex, Abacophrastus hobbit, Abacophrastus megalops, Abacophrastus reflexus and Abacophrastus bellorum. Abacoleptus curtus new species, is described. Notonomus irideus and Notonomus savesi are moved to Prosopogmus. Four new species of Prosopogmus are described: Prosopogmus koghisensis, Prosopogmus lescheni, Prosopogmus fortis and Prosopogmus aoupiniensis. Homalosoma griseolum is moved to Sphodrosomus. Cerabilia is newly recorded from New Caledonia and the genus is moved from Platynini to Loxandrini and therefore is the first report of Loxandrini from New Caledonia. An apparent adventive from Australia, Darodilia, is newly reported from New Caledonia.     

A survey of ground beetles (Coleoptera: Carabidae) in Ahuriri Scenic Reserve,Banks Peninsula,and comparisons with a previous survey performed 30 years earlier

Carabidae (Coleoptera) are commonly used as indicators of ecosystem health due to their high diversity, conservation value and relative ease of identification. Carabids were monitored at Ahuriri Scenic Reserve, Port Hills, Canterbury, from August 2007 to September 2008 using pitfall traps. Carabid captures, species richness and distribution across the reserve were compared with those found in a similar study in 1977–1978. Monthly carabid captures were highly correlated with average temperature in both studies. There was little evidence of carabids responding to habitat factors, such as leaf litter depth, pH or ground cover. However, Megadromus antarcticus was positively associated with canopy openness and ground cover, whereas Holcaspis angustula was more abundant where canopy cover was extensive and there was a relatively high incidence of bare rock. Thirteen species of carabid were recorded in 1977–1978, of which six were absent, four had decreased, one was stable and two species increased in abundance from collections made in 2007–2008. The loss of one Banks Peninsula and five New Zealand endemic beetle species from this reserve is of concern, and future work should aim to gain more detailed information on the habitat requirements of these species to assist conservation management strategies.     

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.     

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.     

On five species of the tribes Abacetini and Pterostichini (Coleoptera,Carabidae)

Metabacetus willi sp. n. (type locality: Indonesia, Central Java Province, Purworejo Regency, Kaligesing District, cave Seplawan near Donorejo) and Rhytiferonia beroni sp. n. (type locality: Papua New Guinea, West Sepik Province, Bonforok bil, Tifalmin, 1600 m) are described. Two new combinations: Poecilus (Ancholeus) campania (Andrewes, 1937), comb. n. of Feronia campania Andrewes, 1937, Aristochroa poecilma (Andrewes, 1937), comb. n. of Feronia poecilma Andrewes, 1937, and a new synonymy: Pterostichus (Oreophilus) podgoricensis B. Guéorguiev, 2013, syn. n. of Pterostichus (Oreophilus) flavofemoratus pinguis (Dejean, 1828), are proposed, too.     

A new species of Lesticus Dejean, 1828 (Coleoptera,Carabidae) from the Finisterre Range,Papua New Guinea and a key to the genera of pterostichine-like Harpalinae of New Guinea

Lesticus finisterrae (Carabidae: Pterostichini) sp. n. (type locality: Finisterre Range, Papua New Guinea), is described and characters to differentiate it from other “Trigonotomi” species are given. A key to the genera of pterostichine-like Harpalinae of the island, including all genera of Morionini, Cratocerini, Drimostomatini, Abacetini, Loxandrini and Pterostichini, is provided. The genus Rhytisternus (Pterostichini) is for the first time reported from New Guinea, represented by the likely adventive species Rhytisternus laevis (Macleay). The previously unknown male of Stegazopteryx ivimkaensis Will (Drimostomatini) is described.     

A synopsis of the New Zealand Anillini (Coleoptera: Carabidae: Bembidiinae), with descriptions of new genera and species

Abstract

Pelodiaetodes n.gen. (type-species P. prominens n.sp.) and Hygranillus n.gen. (type-species H. kuscheli n.sp.) are described, and the following new synonymies are established: Nesamblyops oreobius (Broun, 1893) = N. coriaceus (Broun, 1908); Zeanillus pallidus (Broun, 1884) = Z. marginatus (Broun, 1914). Nine species of New Zealand Anillini are now recognised; they are distributed between five genera.     

Results from two sampling techniques for carabid beetles (Coleoptera: Carabidae) in temporarily flooded and terra firme rainforest of western Amazonia

Carabidae beetles (Coleoptera) were sampled by flight intercept traps (FITs) and hand sampling in the understory of a lowland rainforest in eastern Ecuador. Sampling occurred in June and July of 2011 and 2012 at 24 sites within temporarily flooded forest (FP) and non-flooded terra firme (TF) forest. A total of 674 Carabidae (excluding Cicindelinae) individuals, representing 18 tribes, 50 genera and 133 morphospecies, were collected. Significantly more individuals were collected by hand compared to FITs, particularly by hand in FP forests. After correction through sample-based rarefaction there are no significant differences in species richness between sampling methods and forest types. Ordination and dissimilarity analysis show FITs and hand sampling target different portions of the Carabidae community. Understanding sampling biases for various sampling methods and how it may be affected by forest type will result in more accurate and efficient surveys in tropical forests.     

Trends in detoxification enzymes and heavy metal accumulation in ground beetles (Coleoptera: Carabidae) inhabiting a gradient of pollution

Non-specfic carboxylesterase and glutathione S-transferase activity was measured in the ground beetle, Pterosthicus oblongopunctatus (Coleoptera: Carabidae), from five sites along a gradient of heavy metal pollution. A previous study determined that beetles from the two most polluted sites (site codes OLK2 and OLK3) were more susceptible to additional stressors compared with beetles from the reference site (Stone et al., Environ. Pollut. 113, 239-244 2001), suggesting the possibility of physiological impairment. Metal body burdens in ground beetles from five sites along the gradient ranged from 79 to 201 microg/g Zn, 0.174 to 8.66 microg/g Pb and 1.14 to 10.8 microg/g Cd, whereas Cu seemed to be efficiently regulated regardless of metal levels in the soil. Beetle mid- and hindguts were homogenized and the soluble fraction containing glutathione S-transferase (GST) and carboxylesterase (CaE) was assayed using kinetic analyses. Significantly higher levels of GST were found only in female beetles from the most polluted sites (OLK2 and OLK3; P=0.049, P<0.001, respectively) compared with the reference site (OLK7). In addition, OLK3 females had significantly higher levels of CaE compared with the reference beetles (P=0.01). Male beetles did not differ in enzyme activity along the metal gradient. Overall, obvious trends in detoxification enzymes were not detected in ground beetles in association with metal body burdens.     

Pselaphotumulus,a new genus of pselaphine endemic to New Zealand (Coleoptera: Staphylinidae: Pselaphinae: Pselaphitae)

ABSTRACT

Pselaphotumulus Owens and Carlton, gen. nov., is described as the sixth genus in the tribe Pselaphini known from New Zealand. Three new species are described: Pselaphotumulus aorerei, sp. nov., Pselaphotumulus dubius, sp. nov. and Pselaphotumulus unus, sp. nov.. Three species, Pselaphus cavelli (Broun 1893), Pselaphus oviceps (Broun 1917) and Pselaphus urquharti (Broun 1917) are transferred to Pselaphotumulus, nov. combs. Lectotypes from type series in the New Zealand Broun Collection (Natural History Museum, London) are designated for these three species. Habitus photographs, distributional maps and line drawings of diagnostic characters are provided for each species. A key to species is provided. Searches of museum collections have not yielded representatives outside of New Zealand, suggesting that this is the first endemic genus in the tribe Pselaphini described from the New Zealand’s main islands, specifically, the South Island. Pselaphotumulus species exhibit restricted distribution patterns that approximately coincide with the Pacific/Indo Australian Plate boundary.     

Activity patterns, diets and interspecific interactions of coexisting spring and autumn breeding carabids: Carabusyaconinus and Leptocarabus kumagaii (Coleoptera, Carabidae)

Abstract. 1. Activity patterns, diets and interspecific interactions of two coexisting carabid beetles, Carabus yaconinus B. and Leptocarabus kumagaii K. et K., were studied in a lowland habitat in Japan.
2. Larvae of C. yaconinus appeared from May to October and fed on earthworms. Those of L. kumagaii appeared from October to the next May and fed on dipteran and lepidopteran larvae.
3. In the adult stages, feeding habits and daily activities of the two species were similar. The beetles fed mainly on earthworms and other animal materials, but various plant materials were also utilized.
4. Direct competition for food items was observed both intra- and interspecifically. Interspecific interactions occurred only in early summer, however, because of the difference in breeding seasons (i.e. spring for C. yaconinus and autumn for L. kumagaii).
5. The difference in seasonal life history patterns between these carabids is a possible mechanism of coexistence.