Novelty and emergent patterns in sperm: Morphological diversity and evolution of spermatozoa and sperm conjugation in ground beetles (Coleoptera: Carabidae)

The beetle family Carabidae, with about 40,000 species, exhibits enough diversity in sperm structure and behavior to be an excellent model system for studying patterns and processes of evolution. We explore their potential, documenting sperm form in 177 species of ground beetles using light microscopy and collecting data on one qualitative and seven quantitative phenotypic traits. Our sampling captures 61% of the tribal-level diversity of ground beetles. These data highlight the notable morphological diversity of sperm in ground beetles and suggest that sperm in the group have dynamic evolutionary histories with much morphological innovation and convergence. Sperm vary among species in total length (48–3,400 μm), head length (0.5–270 μm), and head width (0.2–6.3 μm). Most ground beetles make sperm with heads that are indistinct from the flagella at the gross morphological level. However, some or all Omophron, Trachypachus, and Dyschiriini make broad-headed sperm that show morphological differences between species. Most ground beetles package their sperm into groups of sperm, termed conjugates, and ground beetles show variation in conjugate form and in the number and arrangement of sperm in a conjugate. Most ground beetles make sperm conjugates by embedding their sperm in a hyaline rod or spermatostyle. The spermatostyle is remarkably variable among species and varies in length from 17 to 41,000 μm. Several unrelated groups of ground beetles make only singleton sperm, including Nebriinae, Cicindelinae, many Trechinae, and the tribe Paussini. In order to study patterns in sperm evolution, we combine these data with a low-resolution phylogeny of ground beetles. Results from modern comparative analyses suggest the following: (a) sperm differ from conjugates in some aspect of their underlying evolutionary process, (b) sperm have influenced conjugate evolution and vice versa, and (c) conjugation with a spermatostyle likely evolved early within the history of Carabidae and it has been lost independently at least three times.     

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.     

Effects of habitat complexity on forest beetle diversity: do functional groups respond consistently?

We examined the responses of a beetle assemblage to habitat complexity differences within a single habitat type, Sydney sandstone ridgetop woodland, using pitfall and flight‐intercept trapping. Six habitat characters (tree canopy cover, shrub canopy cover, ground herb cover, soil moisture, amount of leaf litter, and amount of logs, rocks and debris) were scored between 0 and 3 using ordinal scales to reflect habitat complexity at survey sites. Pitfall trapped beetles were more species rich and of different composition in high complexity sites, compared with low complexity sites. Species from the Staphylinidae (Aleocharinae sp. 1 and sp. 2), Carabidae (Pamborus alternans Latreille), Corticariidae (Cartodere Thomson sp. 1) and Anobiidae (Mysticephala Ford sp. 1) were most clearly responsible for the compositional differences, preferring high complexity habitat. Affinities between general functional groupings of pitfall‐trapped beetles and habitat variables were not clear at a low taxonomic resolution (family level). The composition and species richness of flight‐intercept‐trapped beetles were similar in high and low complexity sites. Our study demonstrates that discrete responses of the various functional groups of beetles are strongly associated with their feeding habits, indicated by differing habitat components from within overall composite habitat complexity measures. Although habitat preferences by beetle species may often reflect their foraging habits, clarification of the causal mechanisms underpinning the relationships between habitat complexity and beetles are critical for the development of general principles linking habitat, functional roles and diversity.     

The influence of habitat type on the population dynamics of ground beetles (Coleoptera: Carabidae) in marshland

This study presents qualitative and quantitative research on communities of ground beetles (Carabidae) occurring in 2001 and 2003 in three types of habitats located near Bydgoszcz city, Poland (UTM CD19), on the Vistula river floodplain: elm and ash riparian forest (RF), poplar monoculture (P), and unintensively-farmed meadows (M). During both studied years 57 species of ground beetles were found in riparian forests (RF), 56 species in plot P and 47 in plot M. In 2001 from all sample plots 3845 beetles were collected; in 2003- 9589. In both years most specimens came from plot RF (3123 in 2001 and 6224 in 2003). From plot P 2656 specimens of ground beetles were collected and from plot M 1430 specimens. Statistically significant differences were found in the abundance of Carabidae, among sampling areas and the years of study.     

A new species of <Emphasis Type="Italic">Laboulbenia</Emphasis> parasitic on tiger beetles

Laboulbenia phaeoxanthae sp. nov. (Ascomycota: Laboulbeniales) parasitic on the tiger beetle Phaeoxantha aequinoctialis bifasciata (Coleoptera: Carabidae: Cicindelinae) from Brazil is described. The new species seems to be closely related to Laboulbenia pheropsophi, but can be distinguished from the latter by the much more slender and elongate habit, and by the different structure of the appendages. The new species is only the second Laboulbeniales species reported on tiger beetles. The widespread Laboulbenia variabilis was previously recorded on Tetracha fulgida from Ecuador.     

Effect of beetle size on predation of cabbage root fly eggs by ground beetles

Laboratory studies were done to determine the numbers of cabbage root fly (Delia radicum L.) (Diptera: Anthomyiidae) eggs eaten by sixty species of ground beetles (Coleoptera: Carabidae) collected from a range of habitats. For 35 species that were between 2.7 mm and 10 mm long, there was a linear relationship between the numbers of eggs eaten and beetle length. For each 1 mm increase in length above 2.7 mm, an additional 18 eggs were eaten/beetle/day. Eight further species within the above size range did not eat any fly eggs, as these species are known to be either phytophagous or to feed on moving prey. Large numbers of eggs were eaten by beetles in 13 of the 25 genera tested. Based on size, maximum numbers of eggs were eaten by 7 species of Agonum Bonelli, 5 species of Amara Bonelli and seven species of Bembidion Latreille. Only one of the 6 species of Pterostichus Bonelli tested, P. strenuus (Panz.), ate the expected numbers of eggs, the other species being too large to feed from such small prey items as cabbage root fly eggs. The numbers of beetles larger than 10 mm that ate eggs was highly variable and so 10 mm was considered the upper size limit of carabid predators of cabbage root fly eggs.     

Parachordodes tegonotus n. sp. (Gordioidea: Nematomorpha), a hairworm parasite of ground beetles (Carabidae: Coleoptera), with a summary of gordiid parasites of carabids

A new species of hairworm, Parachordodes tegonotus n. sp. (Gordioidea: Nematomorpha) is described from three species of ground beetles (Carabidae: Coleoptera) from the state of Oregon. This is the first record of Parachordodes Camerano parasitising carabid beetles in North America. Diagnostic characters for the new species include size, colour, the nature and arrangement of the areoles, and the character, shape and extant of ornamentation on the ventral surface of the male tail. Encysted hairworm larvae found in the internal tissues of mayfly and caddisfly larvae at the type-locality were presumed to be those of P. tegonotus, indicating an indirect life-cycle involving paratenic hosts. A worldwide host list shows that some 70 species of ground beetles have been documented as developmental hosts to hairworms belonging to at least five genera, namely Gordius, Parachordodes, Dacochordodes, Gordionus and Paragordionus.     

小兴安岭森林地表鞘翅目成虫对不同干扰梯度的响应

为探讨小兴安岭凉水自然保护区森林生态系统中地表鞘翅目成虫群落对不同人为干扰梯度的响应,于2015年7月、8月和10月分别对轻度干扰[原始阔叶红松林(KY)和谷地云冷杉林(YL)]、中度干扰[阔叶红松择伐林(ZF)和次生白桦林(BH)]及重度干扰[落叶松人工林(RL)和红松人工林(RHS)]的6个林型进行取样调查。结果表明:(1)整个采样周期共捕获地表鞘翅目成虫879只,隶属9科44物种;其中轻度干扰生境共捕获6科29种251只(KY捕获5科21种150只,YL捕获4科20种101只),中度干扰生境捕获6科27种276只(ZF捕获3科20种144只,BH捕获6科23种132只),重度干扰生境捕获6科29种352只(RL捕获4科22种232只,RHS捕获5科17种120只)。(2)7月和8月步甲科和葬甲科占据数量优势,10月步甲科和葬甲科成虫数量明显减少而隐翅虫科数量占优势;不同林型及不同干扰梯度地表鞘翅目成虫物种总数和总个体数于7、8、10月均表现为下降趋势,且群落多样性也呈不同程度下降。(3)林型和月份对地表鞘翅目成虫群落结构具有显著影响,干扰梯度对群落结构无显著影响;6个林型之间地表鞘翅目成虫个体数量具有显著差异,但在物种组成上无显著差异;不同干扰梯度间地表鞘翅目成虫个体数、物种数无显著差异,且随干扰梯度变化没有明显的梯度性规律;相似性系数和聚类分析表明,属于同一干扰梯度的两个林型没有表现出高度的相似性。本研究表明干扰梯度不是决定各林型间地表鞘翅目成虫群落多样性存在差异的主要原因,凉水森林生态系统地表鞘翅目成虫对不同干扰梯度的响应不符合中度干扰假说,林型和时间则是影响地表鞘翅目成虫群落组成的显著因素,本实验为地表生物多样性保护和森林生态系统管理提供数据支撑。     

Use of simple remote sensing tools to expedite surveys for rare tiger beetles (Insecta: Coleoptera: Cicindelidae)

Tiger beetles (Insecta: Coleoptera: Cicindelidae) often occupy small patches of suitable habitat in otherwise unsuitable landscapes. Such patches are easily overlooked, which may lead to underestimates of both the number of occurrences and the overall population size. In this study, simple World Wide Web-based tools (Google Earth and Microsoft Terraserver) were used to search high-resolution satellite imagery for patches of suitable habitat for globally and regionally rare tiger beetles on a 3,278 ha wildlife refuge in Maryland, USA. This tract is largely forested but contains scattered small open areas of sand and clay soils that are potential habitat for tiger beetles of conservation concern. Visual inspection of remotely sensed imagery resulted in the identification of 19 potential habitat patches, 15 of which yielded tiger beetle populations when surveyed on the ground. The number of species of tiger beetles recorded from this tract was increased from 3 to 8 and two new sites were discovered for the state sensitive species Cicindela scutellaris rugifrons Dejean. In addition, a small population of C. splendida Hentz was discovered, a species last reported from Maryland in 1948. The technique described here shows great promise for locating small patches of potential tiger beetle habitat in otherwise unsuitable landscapes.     

Effects of forest fragmentation on species richness and composition of ground beetles (Coleoptera: Carabidae and Brachinidae) in urban landscapes

To clarify the effects of forest fragmentation in urban landscapes on the abundance, species richness, dominance, and species composition of ground beetles (Coleoptera: Carabidae and Brachinidae), we compared the beetles collected in 12 pitfall traps from April to July and from September to November between three continuous suburban forests and eight isolated urban forests (0.06–1.02 ha), most of which were in the precincts of shrines and temples in Hanshin District, Honshu, Japan. A total of 28 species and 4178 individuals of ground beetles were collected. Segregation of urban forests from continuous suburban forests has changed the species composition and resulted in the loss of some large‐sized forest species and the addition of some non‐forest species. Simpson's index of dominance (λ) also increased in the urban forests. The richness of forest species markedly decreased with the reduction in forest area but not with the distance from continuous forests, although the species richness of non‐forest species did not change with them. Also, species composition changed only with forest area. These findings indicate that continuous forests do not necessary serve as a “mainland” for urban forest species and that every urban habitat, however small in size, acts as a temporary reservoir of species. In comparison with populations of small‐sized species, populations of large‐sized forest species appeared to decline more readily during forest fragmentation.     

What do we know about winter active ground beetles (Coleoptera,Carabidae) in Central and Northern Europe?

This paper summarizes the current knowledge on winter active Carabidae in Central and Northern Europe. In total 73 winter active species are listed, based on literature and own observations. Ground beetles are among the three most numerous Coleoptera families active during the autumn to spring period. The winter community of Carabidae is composed both of larvae (mainly autumn breeding species) and adults, as well as of epigeic species and those inhabiting tree trunks. Supranivean fauna is characterized by lower species diversity than the subnivean fauna. The activity of ground beetles decreases in late autumn, is lowest during mid-winter and increases in early spring. Carabidae are noted as an important food source in the diet of insectivorous mammals. They are also predators, hunting small winter active invertebrates.     

Low intensity surface fire instigates movement by adults of Calosoma frigidum (Coleoptera, Carabidae)

The genus Calosoma (Coleoptera: Carabidae) is a group of large, sometimes ornate beetles, which often voraciously attack caterpillars. Many studies have reported Calosoma beetles being highly conspicuous during defoliator outbreaks. Based on observations of individual beetle behavior, patterns of activity density and phenology we provide a hypothesis on how environmental cues may synchronize Calosoma activity with periods of high defoliation. We have observed that adults of Calosoma frigidum construct underground burrows similar to those reported to be created by larvae for pupation. We propose that small increases in soil surface temperature caused either by defoliation events or decreased albedo of blackened, burned soil causes beetles to leave their underground burrows and begin foraging. Indirect support for this hypothesis comes from high levels of adult Calosoma frigidum collected in relatively small patches of burned forest (<200m(2)) relative to the surrounding mosaic of unburned forest shortly after a prescribed surface burn.     

Remarkable discovery in a cave of south west Morocco: Siagona taggadertensis n.sp. (Carabidae: Siagoninae)

Five species of the genus Siagona Latreille 1804 (Carabidae: Siagoninae) are known from Morocco. Siagona taggadertensis n. sp., a remarkable species of Carabidae from a Moroccan cave is described. This insect is spectacular by its large size (one of the largest species of the genus), its external morphology, which isolates it from all the north african representant of the genus, and for the conditions of its discovery: its only known from the remains of six specimens, all of them found in the cave of Taggadert, western Atlas. Hypotheses concerning the ecology of this species are discussed.     

Pre-Mating Reproductive Isolation in Tiger Beetles (Carabidae: Cicindelinae): an Examination of the Role of Visual and Morphological Feedback

The biological species concept depends on mechanisms isolating reproduction to taxonomically define a species and to explain the disadvantages, and subsequent lack, of hybridization. In co-occurring tiger beetles (Carabidae: Cicindelinae), these between-species barriers are not readily apparent. To understand the characteristics potentially responsible for this isolation, we conducted behavioral and morphological experiments with two pairs of tiger beetle species. Choice trials showed males use visual cues to trigger the pursuit of potential mates and prefer females who appear similar in color. Geometric morphometric techniques allowed us to quantify shape differences in the mandibles of all four species. Previously, it was proposed that the mandible-coupling sulcus pairing acted as a lock-and-key mechanism to provide feedback necessary for species recognition. Our results, however, suggest that although there are significant differences in shape, these differences do not provide any feedback to the male or enhance his ability to retain his grasp on the female. Taken together, these findings suggest that vision, rather than the traditional mandibular lock-and-key hypothesis, may play a key role in reproductive isolation for co-occurring tiger beetle species.     

Invasions by ladybugs,ladybirds, and other predatory beetles

Species of predatory Coleoptera have become abundant in new geographic regions recently, raising concerns for invaded ecosystems. We address this topic by focusing on invasive alien ladybird beetles (Coccinellidae; known also as ladybugs). Humans appear directly or indirectly responsible for all or most ladybird invasions. Factors hypothesized to have promoted ladybird invasions include genetic diversity (e.g., for polymorphism), phenotypic plasticity, adaptation and genetic shift, generalized diet and habitat preferences, flexible life history and reproduction, large body size, and release from enemies. Factors such as climate, habitat and prey availability, and biotic resistance may sometimes prevent or slow ladybird invasions. Indigenous species (e.g., herbivores) may suffer from invasions, and biological control programs may be affected. Species of indigenous ladybirds throughout the world are reported to have declined in abundance following ladybird invasions, with increased competition and/or intraguild predation most often hypothesized or inferred. Similar recent studies especially of ground beetles (Carabidae) also make clear the potential of invasive alien predatory Coleoptera to disrupt invaded natural and agricultural ecosystems.     

Static antennae act as locomotory guides that compensate for visual motion blur in a diurnal,keen-eyed predator

High visual acuity allows parallel processing of distant environmental features, but only when photons are abundant enough. Diurnal tiger beetles (Carabidae: Cicindelinae) have acute vision for insects and visually pursue prey in open, flat habitats. Their fast running speed causes motion blur that degrades visual contrast, forces stop-and-go pursuit and potentially impairs obstacle detection. We demonstrate here that vision is insufficient for obstacle detection during running, and show instead that antennal touch is both necessary and sufficient for obstacle detection. While running, tiger beetle vision appears to be photon-limited in a way reminiscent of animals in low-light habitats. Such animals often acquire wide-field spatial information through mechanosensation mediated by longer, more mobile appendages. We show that a nocturnal tiger beetle species waves its antennae in elliptical patterns typical of poorly sighted insects. While antennae of diurnal species are also used for mechanosensation, they are rigidly held forward with the tips close to the substrate. This enables timely detection of path obstructions followed by an increase in body pitch to avoid collision. Our results demonstrate adaptive mechanosensory augmentation of blurred visual information during fast locomotion, and suggest that future studies may reveal non-visual sensory compensation in other fast-moving animals.     

Selection of breeding territory by little bustard (<Emphasis Type="Italic">Tetrax tetrax</Emphasis>) males in Central Spain: the role of arthropod availability

The influence of arthropod availability on the formation of male little bustard breeding territories in central Spain was studied in two consecutive years. Arthropods (68,217 individuals) were sampled using pitfall trapping and male territories estimated by means of kernel functions. Hymenopterans (ants), beetles, mites and spiders were the main taxa found in the samples, while the highest contributions in terms of biomass came from hymenopterans, beetles and orthopterans. The only group that exhibited significant biomass differences between territory and non-territory locations was that of the beetles; this pattern was consistent between years. Carabidae was the main beetle family in terms of biomass, followed by other large beetles such as Tenebrionidae, Cleridae, Scarabeidae and Elateridae. The biomass of Carabidae that were larger than 15 mm was significantly higher within territories than outside them. The biomass of Cleridae was greater in fallows, while Elateridae were more abundant in arable fields. The total biomass of beetles that were larger than 15 mm was higher in arable fields than in fallows. Family richness differed between years, while diversity showed differences between agricultural fields (it was highest in fallows), although these variables did not differ between territory and non-territory locations. Results indicate that large beetles (particularly carabids) seem to be a valuable food resource that is defended by little bustard males during the highly energy-demanding breeding season, which is discussed here in relation to habitat selection, the mating system and its implications for the conservation of the species.     

Body size changes in ground beetle assemblages – a reanalysis of Braun et al. (2004)'s data

Abstract.  1. Data in Braun et al .'s (2004) recent paper on size trends in ground beetles (Carabidae) are re-analysed, after re-classifying the species according to feeding categories.
2. When ground beetles are classified as predatory species, mixed feeders or herbivores, and evaluated separately, the size trends indicate group-specific differences. Most of these do not support the efficiency-specialisation hypothesis.
3. Taxonomic groups rarely fit into one ecological category and ecological analyses should explicitly consider this hurdle.     

Communities of ground‐dwelling arthropods in conventional and transgenic maize: background data for the post‐market environmental monitoring

To verify the validity of concerns about environmental safety of maize expressing insecticidal Cry toxins (referred to as Bt maize), we compared communities of ground beetles (Carabidae), rove beetles (Staphylinidae) and spiders (Araneae) in plots planted either with Bt maize cultivar YieldGard^® or with the non‐transgenic parental cultivar Monumental. Each cultivar was grown on 5 plots of 0.5 ha for three consecutive years. To increase the field load of Cry toxin, the fully grown maize of the first study year was shredded to small pieces that were ploughed into the soil. Arthropods were collected in pitfall traps and determined to the species level. The abundance and species richness of all studied groups greatly varied over the season and between the seasons but without statistically significant differences between the Bt and non‐Bt plots. A single spider species and three ground beetle species dominated in the catches every year, whereas a set of 1–4 most abundant rove beetle species changed every year. Frequently occurring species were typical for most of Europe. The total counts of ground beetles, rove beetles and spiders collected once or twice per season are proposed to serve as bioindicators in the post‐market environmental monitoring (PMEM).     

Dry season ecology of riverine tiger beetles in Kruger National Park, South Africa

The life cycles of many African species of tiger beetles (Insecta: Coleoptera: Cicindelidae) exhibit pronounced seasonality, with adult emergence and reproductive activity closely associated with seasonal rains. Anecdotal reports have suggested that adults of certain riverine tiger beetle species may be active during the dry season near perennial water sources. To test this hypothesis, fifteen sites along three perennial rivers in Kruger National Park, South Africa, were surveyed for tiger beetles in early September, 2006. Thirteen sites yielded adult beetles, with six species represented [ Chaetodera regalis (Dejean), Cylindera disjuncta (Dejean), Habrodera nilotica (Dejean), Lophyra neglecta intermediola (Horn), Lophyridia fimbriata imperatrix (Srnka), and Myriochile melancholica (Fabricius)]. Microhabitat associations of adults of all six species and ovipositional/larval substrates of five of the six species are reported, along with observations on predatory and reproductive behaviours and the possible presence of an anti-predator chemical defence in C. regalis and L. fimbriata imperatrix .