Agra, arboreal beetles of Neotropical forests: feisthameli group systematics (Carabidae)

Abstract. This paper is the sixth in a series designed to cover taxonomically all species groups of Agra , whose cumulative ranges extend from southernmost Texas to northernmost Argentina. The feisthameli, mixta, virgata and ohausi groups constitute the Feisthameli Section of the genus. The feisthameli group consists of seventeen species which have a composite range extending from Guatemala to Bolivia and southeastern Brazil.
Among this group, nine new species are described from the following type localities: chapada (BRAZIL, Chapada de GuimaráTes), phite (BOLIVIA, Provencia de Gutierrez), para (BRAZIL, Pará), rubra (BRAZIL, Para, SáTo Félix do Xingu), dation (BRAZIL, Minas Gerais, Teófilo Otoni), tuitis (PERU, San Martin, Tarapoto), memnon (BRAZIL, Espírito Santo, SáTo Mateus), lata (BOLIVIA, Santa Cruz), magdalena (COLOMBIA, Santander). A dot map illustrates the range of each taxon.
Distribution and relationships are discussed in general, but detailed cladistic analysis is deferred until taxonomy of other sections is completed. Species distributions correspond in part to pleistocene refuges suggested by other authors based on other classes of organisms; however, more species occur outside such centres than in them.     

Agra, arboreal beetles of Neotropical forests: mixta group, virgata group, and ohausi group systematics (Carabidae)

Abstract. This paper is the fifth in a series designed to cover taxono-mically all species-groups of the lebiine genus Agra , whose cumulative ranges extend from southernmost Texas to northernmost Argentina. The mixta, virgata, feisthameli and ohausi groups constitute Section Feisthameli of the genus, and a key to these groups is provided. The mixta group consists of two species which have a composite range extending from southeastern Brazil to middle Mexico; the virgata group consists of nine species which have a composite range extending from southeastern Brazil to Guatemala; and the ohausi group consists of four species which have a composite range restricted to southeastern Brazil. The speciose feisthameli group is covered in a separate paper.
Among the three groups in the present paper, eleven new species are described from the following type localities: perkinsorum (MEXICO, Nayarit, 17.6 km southwest of Compostela), vate (MEXICO, Veracruz, Los Tuxtlas, La Playa Escondida), nola (REPUBLIC OF PANAMA, Panama, Cerro Campana), cadabra (EQUADOR, Pichincha, 47 km south of Santo Domingo de los Colorados, Rio Palenque Station), chocha (WEST INDIES, Trinidad, Arima Valley), phainops (FRENCH GUIANA, Pariacabo), imaginis (BRAZIL, Goias, Jatahy), inca (PERU, Madre de Dios, Rio Tambopata Reserve, 30 km (air) southwest of Puerto Maldonado), itatiaya (BRAZIL, Rio de Janeiro, Parque Nacional Itatiaya), calamitas (BRAZIL, Bahia, Itamaraju), anthrax (BRAZIL, Espirito Santo, Parque Socretama). A dot map illustrates the range of each taxon.
Distribution and relationships are discussed in general, but detailed cladistic and biogeographic analysis is deferred until taxonomy of the four groups of Section Feisthameli is completed. Species distribution correspond to Pleistocene refuges suggested by other authors based on other groups and classes of organisms.     

Agra, arboreal beetles of Neotropical forests: pusilla group and piranha group systematics and notes on their ways of life (Coleoptera, Carabidae, Lebiini, Agrina)

Revisions of two new species groups of the genus Agra Fabricius are presented with the following species described as new: pusilla group - Agra cruciariasp. n. (Brazil), Agra gracesp. n. (Ecuador,Perú), Agra maxsp. n. (Brazil), Agra minasianussp. n. (Brazil),Agra notpusillasp. n. (Brazil), Agra pseudopusillasp. n. (Brazil); piranha group - Agra cesp. n. (Perú), Agra risserisp. n. (Bolivia,Brazil), Agra maiasp. n. (Bolivia), Agra piranhasp. n. (Ecuador); Agra tiputinisp. n. (Ecuador). Species of these two groups have adults that are the smallest in the entire genus, although this does not indicate they are closely related based on other attributes. All species are Amazonian in distribution.     

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.     

Ground beetles (Coleoptera: Carabidae) as bioindicators

One of the primary goals of research on bioindicators is to identifyspecies or other taxonomic units that would reliably indicate disturbances inthe environment, and reflect the responses of other species or the overallbiodiversity. However, there is no perfect bioindicator and selecting the mostsuitable one depends to a great extent on the goal of the survey. In this paperwe examine the suitability of carabids as bioindicators. Carabids are frequentlyused to indicate habitat alteration. They have been used in grasslands andboreal forests where species number and/or abundances have been noted to changealong a habitat disturbance gradient. A common trend is that large, poorlydispersing specialist species decrease with increased disturbance while smallgeneralist species with good dispersal ability increase. Some species are notaffected by moderate disturbance. There is, however, not enough research todetermine how suitable carabids are for biodiversity studies, or how well theyrepresent the response of other species. We conclude that carabids are usefulbioindicators, but as crucial understanding of their relationship with otherspecies is incomplete, they should be used with caution.     

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.     

Gap felling as a forest harvesting method in boreal forests: responses of carabid beetles (Coleoptera, Carabidae)

In order to study the ecological effects of logging combined with mechanical soil preparation, we sampled carabid beetles with pitfall trapping in nine spruce-dominated stands in central Finland in 1995–1998. Three of the stands were left intact as controls. Three 1600-m² openings per stand were logged in the winter 1995–1996 into six stands. In three of these stands, light soil preparation was applied. Logging affected the species assemblages, but soil preparation per se had no clear effects. Open-habitat species increased in abundance in the openings one year after logging, but catches of generalist species in the different treatments did not differ from each other. A forest species, Calathus micropterus , was least abundant in the prepared openings. The amounts of logging residue, exposed mineral soil and aggregated humus, as well as the abundance of red wood ants, significantly explained variation in carabid assemblages.     

Response of female beetles to LIDAR derived topographic variables in Eastern boreal mixedwood forests (Coleoptera, Carabidae)

Biodiversity monitoring is increasingly being bolstered with high resolution data derived from remote sensing such as LIDAR (Light Detection and Ranging). We derived a series of topographical variables, including slope, azimuth, ground curvature and flow accumulation from LIDAR images and compared these to captures of female carabids in pitfall traps in Eastern boreal mixedwood forests. We developed a series of species-specific logistic models predicting the proportion of females for eight dominant species, including Agonum retractum, Calathus ingratus, Platynus decentis, Pterostichus adstrictus, Pterostichus coracinus, Pterostichus pensylvanicus, Sphaeroderus nitidicollis and Synuchus impunctatus. We used these models to test three hypotheses related to how the modest topography in boreal forests could influence the availability of microhabitats and possibly potential sites for oviposition and larval development. In general, topographic features such as north facing slopes and high flow accumulation were important predictors of the proportion of females. Models derived from larger scale topography, such as hillsides or small watersheds on the order of ¼-1 ha were better predictors of the proportion of females than were models derived from finer scale topography such as hummocks and small depressions. We conclude that topography likely influences the distribution of carabids based on hydrological mechanisms rather than factors related to temperature. We further suggest based on the scale of responses that these hydrological mechanisms may be linked to the attenuation of past disturbances by wildfire and the propensity of unburned forest patches and fire skips.     

Proventricular trituration in adult carabid beetles (Coleoptera: Carabidae)

Adult carabids that ingest particles of food have a muscular proventriculus lined internally with an intima that bears sclerotized projections. In experiments where six species of carabid were allowed to feed upon Drosophila, the average size of cuticular remains found in post-proventricular regions of the gut was generally significantly smaller than those in the crop. This reduction in size of cuticular particles is likely to be due to proventricular trituration rather than prolonged exposure to digestive secretions. There is no evidence that material is triturated by the proventriculus and returned to the crop, the main seat of digestion in carabids, so proventricular trituration is unlikely to enhance digestion. By crushing large, potentially abrasive, cuticular particles, the proventriculus may serve to protect midgut epithelial tissue.     

A comprehensive molecular phylogeny of tiger beetles (Coleoptera,Carabidae, Cicindelinae)

Tiger beetles are a remarkable group that captivates amateur entomologists, taxonomists and evolutionary biologists alike. This diverse clade of beetles comprises about 2300 currently described species found across the globe. Despite the charisma and scientific interest of this lineage, remarkably few studies have examined its phylogenetic relationships with large taxon sampling. Prior phylogenetic studies have focused on relationships within cicindeline tribes or genera, and none of the studies have included sufficient taxon sampling to conclusively examine broad species patterns across the entire subfamily. Studies that have attempted to reconstruct higher‐level relationships of Cicindelinae have yielded conflicting results. Here, we present the first taxonomically comprehensive molecular phylogeny of Cicindelinae to date, with the goal of creating a framework for future studies focusing on this important insect lineage. We utilized all available published molecular data, generating a final concatenated dataset including 328 cicindeline species, with molecular data sampled from six protein‐coding gene fragments and three ribosomal gene fragments. Our maximum‐likelihood phylogenetic inferences recover Cicindelinae as sister to the wrinkled bark beetles of the subfamily Rhysodinae. This new phylogenetic hypothesis for Cicindelinae contradicts our current understanding of tiger beetle phylogenetic relationships, with several tribes, subtribes and genera being inferred as paraphyletic. Most notably, the tribe Manticorini is recovered nested within Platychilini including the genera Amblycheila Say, Omus Eschscholtz, Picnochile Motschulsky and Platychile Macleay. The tribe Megacephalini is recovered as paraphyletic due to the placement of the monophyletic subtribe Oxycheilina as sister to Cicindelini, whereas the monophyletic Megacephalina is inferred as sister to Oxycheilina, Cicindelini and Collyridini. The tribe Collyridini is paraphyletic with the subtribes Collyridina and Tricondylina in one clade, and Ctenostomina in a second one. The tribe Cicindelini is recovered as monophyletic although several genera are inferred as para‐ or polyphyletic. Our results provide a novel phylogenetic framework to revise the classification of tiger beetles and to encourage the generation of focused molecular datasets that will permit investigation of the evolutionary history of this lineage through space and time.     

Effects of pitfall trap size and sampling duration on collection of ground beetles (Coleoptera: Carabidae) in temperate forests

Pitfall trapping is a standard sampling method to compare the abundance or community structure of ground beetles. However, effects of sampling duration on biodiversity estimation of ground beetles according to different trap sizes have not been experimentally evaluated in temperate forests in Korea. Therefore, the objective of this study was to determine the interaction between trap sizes (SB, small‐sized bottle; MB, medium‐sized bottle; PC, plastic cup; PJ, plastic jar; PT, perforated type trap; FT, funnel‐type trap) and sampling duration (T1, 2 weeks × 2 sessions, 28 days; T2, 2 weeks × 4 sessions, 56 days; T3, 4 weeks × 2 sessions, 56 days) on estimation of ground beetle assemblages in Naejangsan National Park, a temperate forest in Korea. Funnel type and larger pitfall traps collected higher numbers of individuals and species than other trap sizes. Species composition of ground beetles was different by size of traps (SB, MB, and PC vs. PJ, PT, and FT). In particular, ground beetle composition in larger traps (PJ, PT, and FT) appeared to be influenced by environmental characteristics according to localities (e.g., soil characters and dominant tree species). These findings from our study support that pitfall trapping of ground beetles can be influenced by trap sizes per se as well as sampling durations and environmental characteristics. Thus, biodiversity monitoring in temperate forests should be conducted with long sampling duration (at least 28 days) using large‐sized traps (> 7.5 cm in diameter of trap mouthpart) considering expenses and study aims.     

The Mecyclothorax beetles (Coleoptera,Carabidae, Moriomorphini) of Tahiti,Society Islands

The 101 species of Mecyclothorax Sharp known to inhabit Tahiti Island, French Polynesia are taxonomically revised, including 28 species that are newly described: Mecyclothorax claridgeiae sp. n., Mecyclothorax jeanyvesi sp. n., Mecyclothorax poria sp. n., Mecyclothorax aano sp. n., Mecyclothorax papau sp. n., Mecyclothorax manina sp. n., Mecyclothorax everardi sp. n., Mecyclothorax ramagei sp. n., Mecyclothorax pitohitiensis sp. n., Mecyclothorax curtisi sp. n., Mecyclothorax hoeahiti sp. n., Mecyclothorax ninamu sp. n., Mecyclothorax kokone sp. n., Mecyclothorax paahonu sp. n., Mecyclothorax kayballae sp. n., Mecyclothorax ehu sp. n., Mecyclothorax papuhiti sp. n., Mecyclothorax tuea sp. n., Mecyclothorax taatitore sp. n., Mecyclothorax konemata sp. n., Mecyclothorax arboricola sp. n., Mecyclothorax rahimata sp. n., M. oaoa sp. n., Mecyclothorax maninapopoti sp. n., Mecyclothorax hunapopoti sp. n., Mecyclothorax fefemata sp. n., Mecyclothorax maninamata sp. n., and Mecyclothorax niho sp. n. Mecyclothorax muriauxioides Perrault, 1984 is newly synonymized with Mecyclothorax muriauxi Perrault, 1978. Lectotypes are designated for: Thriscothorax altiusculus Britton, 1938; Thriscothorax bryobius Britton, 1938; Mecyclothorax globosus Britton, 1948: and Mecyclothorax sabulicola Britton, 1948. Dichotomous identification keys augmented by dorsal habitus and male aedeagal photographs are provided to the various species-groups and all included species. The spermatophore of Mecyclothorax papau sp. n. is described, with the ampulla and collar found to correspond dimensionally to the length of the internal sac flagellar plate. Variation among characters of the female reproductive tract is presented for all newly described plus other representative species comprising the radiation. Taxa are assigned to species groups, modified from the classification of G.G. Perrault, based on derived character states polarized using the Australian outgroup taxon Mecyclothorax punctipennis (MacLeay). Much of the species-level diversity on this small Pacific island is partitioned allopatrically over very small distributional ranges. No species is shared between Tahiti Nui and Tahiti Iti, and nearly all species in Tahiti Nui are geographically restricted to one ridgelike massif of that volcano. Cladistically similar species are often distributed on different massifs suggesting that vicariance associated with erosional valley formation has facilitated speciation, however several instances in which sister species occupy sympatric distributions on the same ridge system demonstrate that speciation may also occur across extremely localized landscapes. Such localized differentiation is facilitated by the low vagility of these small-bodied, flightless predators whose fragmented populations can persist and diverge within spatially limited habitat patches. The intense philopatry of Tahitian Mecyclothorax spp. coupled with the highly dissected landscape has produced the geographically densest adaptive radiation on Earth. This radiation has occurred very rapidly, with species durations averaging 300,000 yr; a speciation rate similar to that observed in Hawaiian Oliarus planthoppers and Laupala crickets, and East African Rift lake cichlid fishes.     

The effects of green tree retention and subsequent prescribed burning on ground beetles (Coleoptera: Carabidae) in boreal pine-dominated forests

We studied how two methods to promote biodiversity in managed forests, i.e. green tree retention and prescribed fire, affect the assemblages of carabid beetles. Our experiment consisted of 24 study sites, each 3–5 ha in size, which had been prepared according to factorial design. Each of the eight treatment combinations determined by the two factors explored – tree retention level (0, 10, 50 m³/ha^?1 and uncut controls) and prescribed use of fire (yes/no) – was replicated three times. We sampled carabids using pitfall traps one year after the treatments. Significantly more individuals were caught in most of the burned sites, but this difference was partially reflective of the trap‐catches of Pterostichus adstrictus. The fire did not increase no. of P. adstrictus in the uncut sites as much as in the other sites. Species richness was significantly affected by both factors, being higher in the burned than in the unburned sites and in the harvested than in the unharvested sites. Many species were concentrated in the groups of retention trees in the burned sites, but only a few were in the unburned sites. The species turnover was greater in the burned than in the unburned sites, as indicated by the NMDS ordinations. Greater numbers of smaller sized species and proportion of brachypterous species were present in the burned sites. Fire‐favored species, and also the majority of other species that prefer open habitats were more abundantly caught in the burned sites than in the unburned sites. Dead wood or logging waste around the traps did not correlate with the occurrence of species. We conclude that carabids are well adapted to disturbances, and that frequent use of prescribed fire is essential for the maintenance of natural assemblages of carabid beetles in the boreal forest. Small retention tree groups can not maintain assemblages of uncut forest, but they can be important by providing food, shelter and breeding sites for many species, particularly in the burned sites.     

Histological structure of tripartite mushroom bodies in ground beetles (Insecta, Coleoptera: Carabidae)

Contrary to members of the suborder Polyphaga, ground beetles have been found to possess tripartite mushroom bodies, which are poorly developed in members of basal taxa and maximally elaborated in evolutionarily advanced groups. Nevertheless, they do not reach the developmental stage, which has been previously found in particular families of beetles. It has been pointed out that a new formation of the Kenyon cells occurs during at least the first months of adult life, and inactive neuroblasts are found even in one-year-old beetles. It has been suggested that there is a relation between the Kenyon cell number and development of the centers of Kenyon cell new-formation.     

Testing abundance-range size relationships in European carabid beetles (Coleoptera, Carabidae)

Four of the eight hypotheses proposed in the literature for explaining the relationship between abundance and range size (the sampling artifact, phylogenetic non-independence, range position and resource breadth hypotheses) were tested by using atlas data for carabid beetles (Coleoptera, Carabidae) from Belgium, Denmark and the Netherlands. A positive relationship between abundance and partial range size was found in all three countries, and the variation in abundance was lower for widespread species. Analysis of the data did not support three of the proposed hypotheses, but did support the resource breadth hypothesis (species having broader environmental tolerances and being able to use a wider range or resources will have higher local densities and be more widely distributed than more specialised species). Examination of species' characteristics revealed that widespread species are generally large bodied, generalists (species with wide niche breadths occurring in a variety of habitat types) and are little influenced by human-altered landscapes, while species with restricted distributions are smaller bodied, specialists (species with small niche breadths occurring in only one or two habitat types), and favour natural habitat. Landscape alteration may be an important factor influencing carabid abundance and range size in these three countries with a long history of human-induced environmental changes.     

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.