Early burst in body size evolution is uncoupled from species diversification in diving beetles (Dytiscidae)

Changes in morphology are often thought to be linked to changes in species diversification, which is expected to leave a signal of early burst (EB) in phenotypic traits. However, such signal is rarely recovered in empirical phylogenies, even for groups with well‐known adaptive radiation. Using a comprehensive phylogenetic approach in Dytiscidae, which harbours ~4,300 species with as much as 50‐fold variation in body size among them, we ask whether pattern of species diversification correlates with morphological evolution. Additionally, we test whether the large variation in body size is linked to habitat preference and whether the latter influences species turnover. We found, in sharp contrast to most animal groups, that Dytiscidae body size evolution follows an early‐burst model with subsequent high phylogenetic conservatism. However, we found no evidence for associated shifts in species diversification, which point to an uncoupled evolution of morphology and species diversification. We recovered the ancestral habitat of Dytiscidae as lentic (standing water), with many transitions to lotic habitat (running water) that are concomitant to a decrease in body size. Finally, we found no evidence for difference in net diversification rates between habitats nor difference in turnover in lentic and lotic species. This result, together with recent findings in dragonflies, contrasts with some theoretical expectations of the habitat stability hypothesis. Thus, a thorough reassessment of the impact of dispersal, gene flow and range size on the speciation process is needed to fully encompass the evolutionary consequences of the lentic–lotic divide for freshwater fauna.     

Thermal tolerance and geographical range size in the Agabus brunneus group of European diving beetles (Coleoptera: Dytiscidae)

Aim Within clades, most taxa are rare, whilst few are common, a general pattern for which the causes remain poorly understood. Here we investigate the relationship between thermal performance (tolerance and acclimation ability) and the size of a species’ geographical range for an assemblage of four ecologically similar European diving beetles (the Agabus brunneus group) to examine whether thermal physiology relates to latitudinal range extent, and whether Brown’s hypothesis and the environmental variability hypothesis apply to these taxa. Location Europe. Methods In order to determine the species tolerances to either low or high temperatures we measured the lethal thermal limits of adults, previously acclimated at one of two temperatures, by means of thermal ramping experiments (± 1°C min^?1). These measures of upper and lower thermal tolerances (UTT and LTT respectively) were then used to estimate each species’ thermal tolerance range, as total thermal tolerance polygons and marginal UTT and LTT thermal polygons. Results Overall, widespread species have higher UTTs and lower LTTs than restricted ones. Mean upper lethal limits of the Agabus brunneus group (43 to 46°C), are similar to those of insects living at similar latitudes, whilst mean lower lethal limits (?6 to ?9°C) are relatively high, suggesting that this group is not particularly cold‐hardy compared with other mid‐temperate‐latitude insects. Widespread species possess the largest thermal tolerance ranges and have a relatively symmetrical tolerance to both high and low temperatures, when compared with range‐restricted relatives. Over the temperature range employed, adults did not acclimate to either high or low temperatures, contrasting with many insect groups, and suggesting that physiological plasticity has a limited role in shaping distribution. Main conclusions Absolute thermal niche appears to be a good predictor of latitudinal range, supporting both Brown’s hypothesis and the environmental variability hypothesis. Restricted‐range species may be more susceptible to the direct effect of climate change than widespread species, notwithstanding the possibility that even ‘thermally‐hardy’, widespread species may be influenced by the indirect effects of climate change such as reduction in habitat availability in Mediterranean areas.     

Dispersal ability rather than ecological tolerance drives differences in range size between lentic and lotic water beetles (Coleoptera: Hydrophilidae)

Aim In aquatic ecosystems, standing (lentic) and running (lotic) waters differ fundamentally in their stability and persistence, shaping the comparative population genetic structure, geographical range size and speciation rates of lentic versus lotic lineages. While the drivers of this pattern remain incompletely understood, the suite of traits making up the ability of a species to establish new populations is instrumental in determining such differences. Here we explore the degree to which the association between habitat type and geographical range size results from differences in dispersal ability or fundamental niche breadth in the members of the Enochrus bicolor complex, an aquatic beetle clade with species across the lentic–lotic divide. Location Western Mediterranean, with a special focus on North Africa, the Iberian Peninsula and Sicily. Methods DNA sequences for four loci were obtained from species of the E. bicolor complex and analysed using phylogenetic inference. Dispersal and establishment abilities were assessed in lentic–lotic species pairs of the complex, using flight wing morphometrics and thermal tolerance ranges as surrogates, respectively. Results There were clear differences in range size between the lotic and lentic taxa of the complex, which appears to have had a lotic origin with two transitions to standing waters. Only small differences were observed in temperature tolerance and acclimation ability between the two lotic–lentic sister species studied. By contrast, wing morphometrics revealed clear, consistent differences between lotic and lentic Enochrus species pairs, the latter having a higher dispersal capacity. Main conclusions We hypothesize that there have been two habitat shifts from lotic to lentic waters, which have allowed marked expansions in geographical range size in western Mediterranean species of the E. bicolor complex. Differences in dispersal rather than in establishment ability appear to underlie differences in geographical range extent, as transitions to lentic waters were associated with changes in wing morphology, but not in thermal tolerance range. In this lineage of water beetles, selection for dispersal in geologically short‐lived lentic systems has driven the evolution of larger range sizes in lentic taxa compared with those of their lotic relatives.     

Bergmann's rule in the ant lion Myrmeleon immaculatus DeGeer (Neuroptera: Myrmeleontidae): geographic variation in body size and heterozygosity

Aim Geographic variation in body size and heterozygosity were surveyed for discrete populations of the ant lion, Myrmeleon immaculatus DeGeer, collected from the central and northeastern United States. Location Collection sites were located in the central and eastern United States ranging from western Oklahoma to northern New York. Methods We collected 872 M. immaculatus larvae from thirty-four collecting sites. At each site, we randomly sampled ant lion pits and collected between fifteen and fifty-two larvae in total. Larvae were preserved in 95% ETOH for morphological analysis and frozen in a ?80°C freezer for protein electrophoresis. We measured the body size of eighty-five preserved adult M. immaculatus obtained from museum collections using head width as an indicator of body size. Five enzymes [GPI (glucose phosphate isomerase), MDH (malate dehydrogenase), PEP (peptidase), DIA (diaphorase) and SOD (superoxide dismutase)] were used in the heterozygosity analyses. Results Larval and adult body size increased with latitude, but decreased with elevation. Average heterozygosity, measured at five polymorphic loci, also increased significantly with latitude. Minimum temperature variance was the best predictor of body size, whereas precipitation and maximum temperature were the best predictors of heterozygosity. Populations were genetically differentiated from one another and showed a pattern of isolation by distance, as measured by Wright's F_st values and Nei's genetic distances. Main conclusions Sampling artifacts, heat conservation, character displacement, cell-size variation, density-dependent mortality, and differential dispersal probably cannot account for latitudinal variation in ant lion body size. Our results implicate the importance of diurnal photoperiod, which varies with latitude, but not with elevation. Because photoperiod often controls growth, diapause, and metamorphosis, it may be an important determinant of latitudinal clines in body size and life history of insects.     

Evolution of subterranean diving beetles (Coleoptera: Dytiscidae: Hydroporini, Bidessini) in the arid zone of Australia

Calcrete aquifers in arid inland Australia have recently been found to contain the world's most diverse assemblage of subterranean diving beetles (Coleoptera: Dytiscidae). In this study we test whether the adaptive shift hypothesis (ASH) or the climatic relict hypothesis (CRH) is the most likely mode of evolution for the Australian subterranean diving beetles by using a phylogeny based on two sequenced fragments of mitochondrial genes (CO1 and 16S-tRNA-ND1) and linearized using a relaxed molecular clock method. Most individual calcrete aquifers contain an assemblage of diving beetle species of distantly related lineages and/or a single pair of sister species that significantly differ in size and morphology. Evolutionary transitions from surface to subterranean life took place in a relatively small time frame between nine and four million years ago. Most of the variation in divergence times of the sympatric sister species is explained by the variation in latitude of the localities, which correlates with the onset of aridity from the north to the south and with an aridity maximum in the Early Pliocene (five mya). We conclude that individual calcrete aquifers were colonized by several distantly related diving beetle lineages. Several lines of evidence from molecular clock analyses support the CRH, indicating that all evolutionary transitions took place during the Late Miocene and Early Pliocene as a result of aridification.     

Energy and interspecific body size patterns of amphibian faunas in Europe and North America: anurans follow Bergmann's rule, urodeles its converse

Aim To describe broad‐scale geographical patterns of body size for European and North American amphibian faunas and to explore possible processes underlying these patterns. Specifically, we propose a heat balance hypothesis, as both heat conservation and heat gain determine the heat balance of ectotherms, and test it along with five other hypotheses that have a possible influence on body size gradients: size dependence, migration ability, primary productivity, seasonality and water availability. Location Western Europe and North America north of Mexico. Methods We processed distribution maps for native amphibian species to estimate the mean body size in 110 × 110 km cells and calculated eight environmental predictors to explore the relationship between environmental gradients and the observed patterns. We used least squares regression modelling and model selection approaches based on information theory to evaluate the relative support for each hypothesis. Results We found consistent body size gradients and similar relationships to environmental variables within each amphibian group in Europe and North America. Annual potential evapotranspiration, a measure of environmental energy, was the strongest predictor of mean body size in both regions. However, the contrasting responses to ambient energy in each group resulted in opposite geographical patterns, i.e. anurans increased in size from high‐ to low‐energy areas in both continents and urodeles showed the opposite pattern. Main conclusions Our results support the heat balance hypothesis, suggesting that the thermoregulatory abilities of anurans would allow them to reach larger sizes in colder climates by optimizing the trade‐off between heating and cooling rates, whereas a lack of such strategies among urodele faunas would explain why these organisms tend to be smaller in cooler areas. These findings may also have implications for the role of climate warming on the global decline of amphibians.     

The relationship between the ventral nerve cord,body size and phylogeny in ground beetles (Coleoptera: Carabidae)

The ventral nerve cord in the family Carabidae (considered in the widest sense! exhibits variations in the degree of fusion of thoracic and abdominal ganglia. There are usually three discrete thoracic ganglia and between one and seven discrete abdominal ganglia, the number differing between tribes. Of the 44 tribes and 177 species examined, 38 tribes contained species showing no differences in the degree of ventral nerve cord consolidation. However, the remaining six tribes showed variations in the degree of ventral nerve cord consolidation between genera (Lebiini, Cychrini, Nebriini, Scaritini, Licinini and Brachinini), whilst one genus showed variations between species (Leistus, Nebriini). No variation in ventral nerve cord consolidation was observed in conspecifics. The degree of ventral nerve cord consolidation is inversely proportional to overall body length. With respect to phylogeny, the degree of consolidation of the nerve cord docs not consistently support the traditional Carabinae-Harpalinae subfamily division. However, the Harpalinae always have four or less discrete abdominal ganglia (with the sole exception of the Broscinij, whilst the Carabinae exhibit almost the whole range of variations. Thus the Harpalinae (or the major pari of it) may be a monophyletic group, but this is not true of the Carabinae. Trends in the degree of ventral nerve cord consolidation for the various tribes were noted, and phylogenetic implications were evaluated wherever possible.     

Characterization and localization of Rickettsia sp. in water beetles of genus Deronectes (Coleoptera: Dytiscidae)

In the present study, Rickettsia sp. was detected in four water beetles of the genus Deronectes ( Dytiscidae ) for the first time. Rickettsiae were found in 100% of examined specimens of Deronectes platynotus (45/45), 39.4% of Deronectes aubei (28/71), 40% of Deronectes delarouzei (2/5) and 33.3% of Deronectes semirufus (1/3). Analysis of 16S rRNA gene sequences revealed a phylogenetic relationship with rickettsial isolates of Limonia chorea ( Diptera ), tentatively classified as members of the basal ancestral group. Phylogenetic analysis of the gltA (citrate synthase) gene sequences showed that Deronectes symbionts were closest to bacterial symbionts from spiders. Ultrastructural examinations revealed typical morphological features and intracellular arrangements of rickettsiae. The distribution, transmission and localization of Rickettsia sp. in D. platynotus were studied using a diagnostic PCR assay and FISH. Eggs from infected females of D. platynotus were all Rickettsia -positive, indicative of a vertical transmission.     

Phylogeography of the Namib Desert dung beetles Scarabaeus (Pachysoma) MacLeay (Coleoptera: Scarabaeidae)

Aim Namib biogeography in many instances remains reliant on advanced and detailed systematic studies. This study attempts to combine molecular phylogenetic data, geology and palaeo‐climatic data to (i) resolve the relationships of the 13 morphological species of Scarabaeus (Pachysoma) and (ii) relate their evolution to past climatic and geological events. Location South Africa and Namibia. Methods Sequencing of a 1197 bp segment of the mitochondrial cytochrome oxidase I (COI) gene of the 13 species within Scarabaeus (Pachysoma) was undertaken. Analyses performed included Parsimony and Maximum Likelihood as well as imposing a molecular clock. Results The molecular phylogeny showed strong support for 11 of the 13 morphological species. The remaining two species, S. (P.) glentoni and S. (P.) hippocrates, formed a complex and could not be assigned specific status on the basis of the COI gene phylogeny. Strong support for the three species formerly classified within the genus Neopachysoma was consistently obtained. The subgenus appears to have arisen c. 2.9 Ma. Species within the subgenus arose at different times, with the common ancestor to Neopachysoma and the hippocrates complex having evolved 2.65 and 2.4 Ma, respectively. Scarabaeus (P.) denticollis, S. (P.) rotundigenus, S. (P.) rodriguesi and S. (P.) schinzi are some of the youngest species, having diverged between 2 million and 600,000 years ago. Main conclusions Scarabaeus (Pachysoma) is a derived monophyletic clade within the Scarabaeini. The subgenus appears to be young in comparison with the age of the Namib Desert, which dates back to the Miocene (c. 15 Ma). The psammophilous taxa are shown to disperse with their substratum and habitat, barchan dunes. Clear south/north evolutionary gradients can be seen within the species of this subgenus, which are consistent with the unidirectional wind regime. Species with a suite of mostly plesiomorphic characters have a southerly distribution while their derived psammophilous relatives have central to northern Namib distributions. Major rivers such as the Orange, Buffels and Holgat appear to be gene barriers to certain species as well as areas of origin of speciation events.     

Molecular phylogenetics of Atlantic Shield Platynectes diving beetles (Coleoptera: Dytiscidae): a first glance at the evolution of the genus in the Amazon Basin

Neotropical diving beetles of the genus Platynectes are distributed across Central America, the Andes and different Precambrian shields in the Amazon Basin. Species from the northern Guiana Shield form a monophyletic clade, yet the phylogenetic relationships of the eastern Atlantic Shield species remain unknown. Here, we augmented an existing molecular dataset with a species from the Atlantic Shield that was not previously sampled. We reconstructed the phylogenetic relationships and estimated divergence times to understand the evolution of lineages dwelling in this region. The newly sampled specimens from the Atlantic Shield are recovered as sister taxa to Guiana Shield species. The dating analyses suggest a split between these two lineages in the late Oligocene to mid-Miocene, contemporary with the Miocenic geological remodeling of the Amazon Basin. Additional sampling in the Atlantic and Central Brazilian Shields will be determinant to test the monophyly of Platynectes species distributed in these ancient shields, and to fully understand the biogeographical history of diving beetles in the Amazon Basin.     

Fossil water‐penny beetles (Coleoptera: Psephenidae: Eubrianacinae) from the Eocene of Europe,with remarks on their phylogenetic position and biogeography

Abstract: A species of water‐penny beetles is recorded from larval instars from the Middle Eocene Messel pit fossil site in Germany. This species clearly belongs to the psephenid subfamily Eubrianacinae, but its precise systematic affinities remain unclear. It is the second fossil species of this taxon recorded from Europe. The holotype of the first fossil species, Eubrianax vandeli Bertrand and Laurentiaux, 1963, is lost. The high number of fossil specimens from Messel allowed discussion of stratigraphic and spatial occurrence of the eubrianacine species in the Messel pit, but its larval ontogeny could not be unravelled. Because of the high number of fossils and their wide occurrence, it is inferred that the species from Messel gen. sp. 1 was an autochthonous faunal element of the Eocene Lake Messel, which might indicate that some parts of the former Lake Messel had a shore area with stones. The analysis of the phylogenetic position of both Eocene eubrianacine species showed that their phylogenetic placement cannot be resolved because preservational influences limit the evaluation of characters. The historical biogeography of Psephenidae and Eubrianacinae is analysed and discussed. The fossil record shows that psephenid beetles have fossil members occurring outside their current distribution range, so interpretations of their biogeography based only on extant members can be misleading.     

基于线粒体基因组序列的鞘翅目肉食亚目水生类群系统发育分析

【目的】明确肉食亚目(Adephaga)水生类群线粒体基因组的基本特征,并基于线粒体基因组序列分析肉食亚目水生类群的系统发育关系。【方法】基于Illumina HiSeq X Ten测序技术测定了圆鞘隐盾豉甲Dineutus mellyi和齿缘龙虱Eretes sticticus的线粒体全基因组序列,对其进行了基因注释,并对其tRNA基因二级结构进行了预测分析。加上已公布的鞘翅目(Coleoptera)肉食亚目水生类群17个种的线粒体基因组序列,对该类群共19个种线粒体的蛋白质编码基因(protein-coding genes, PCGs)开展了比较基因组学分析,包括AT含量、密码子偏好性、选择压力等。基于13个PCGs的氨基酸序列和核苷酸序列,利用最大似然法(ML)和贝叶斯法(BI)分别构建鞘翅目肉食亚目水生类群的系统发育关系,并通过FcLM分析进一步评估伪龙虱科(Noteridae)和瀑甲科(Meruidae)的系统发育位置。【结果】圆鞘隐盾豉甲和齿缘龙虱的线粒体基因组全长分别为16 123 bp(GenBank登录号: MN781126)和16 196 bp(GenBank登录号: MN781132),都包含13个PCGs、22个tRNA基因、2个rRNA基因和1个D-loop区(控制区)。19个肉食亚目水生类群线粒体基因组PCGs的碱基组成都呈现A+T偏好性,在密码子使用上也都偏向于使用富含A+T的密码子;在进化过程中13个PCGs的进化模式相同,都受到纯化选择。基于线粒体基因组13个PCGs的氨基酸序列的肉食亚目水生类群的系统发育关系为(豉甲科Gyrinidae+(沼梭甲科Haliplidae+((壁甲科Aspidytidae+(两栖甲科Amphizoidae+龙虱科Dytiscidae))+(水甲科Hygrobiidae+(瀑甲科Meruidae+伪龙虱科Noteridae)))))。【结论】研究结果表明,豉甲科是肉食亚目水生类群的基部类群,接下来是沼梭甲科和龙虱总科;伪龙虱科和瀑甲科互为姐妹群,并一起作为龙虱总科内部的一个分支;两栖甲科与龙虱科具有更近的亲缘关系。     

A mark and recapture study of water beetles (Coleoptera: Dytiscidae) in a group of semi-permanent and temporary ponds

A year mark and recapture study of aquatic Dytiscidae (Coleoptera) in a group of seven semi-permanent and temporary ponds in Cheshire (UK) is described. Pond water levels showed strong seasonal fluctuations with most ponds drying and refilling on several occasions over the study period. Sampling and labelling of aquatic Dytiscidae was carried out at monthly intervals. Beetles were labelled by fixing numbered waterproof paper labels to the elytra with cyanoacrylate `Superglue'. A total of 780 beetles were marked and released of which 11.5% were subsequently recaptured over varying time intervals of one month to eighteen months. The six species marked (Agabus bipustulatus (Linnaeus, 1767), Acilius sulcatus (Linnaeus, 1758), Colymbetes fuscus (Linnaeus, 1758), Dytiscus marginalis (Linnaeus, 1758), Ilybius ater (De Geer, 1774) and Ilybius fuliginosus (Fabricius, 1792)) are all known to be highly flight mobile. Colymbetes fuscus did not move between ponds as a result of ponds drying up in the summer. The behaviour of Dytiscus marginalis and Acilius sulcatus was found to be in broad agreement with previous accounts of dispersal from over-wintering ponds into temporary ponds in the spring with movement back to permanent ponds in the summer as the temporary ponds dry up. In contrast, Agabus bipustulatus did not migrate when ponds dried up in the summer but remained in terrestrial vegetation in damp pond basins for several months after open water had disappeared, only migrating if the basin dried out completely. Both species of Ilybius were captured and marked in very low numbers and no recaptures were made. As more ponds became temporary over the years of the study Agabus bipustulatus persisted while the other five species declined or were lost altogether.     

The influence of matrix and edges on species richness patterns of ground beetles (Coleoptera: Carabidae) in habitat islands

Aim The aim of this study was to analyse whether, and how, the inclusion of habitat specialists and edge‐preferring species modifies the species–area relationship predictions of the island biogeography theory for an insect group (ground beetles, Coloptera: Carabidae) living in natural fragments. Species–habitat island area relationships applied to terrestrial habitat islands can be distorted by the indiscriminate inclusion of all species occurring in the fragments. Matrices surrounding terrestrial habitat fragments can provide colonists that do not necessarily distinguish the fragment from the matrix and can survive and reproduce there. Edge‐preferring species can further distort the expected relationship, as smaller fragments have larger edge:core ratios. Location Nineteen forest fragments were studied in the Bereg Plain, Hungary, and SW Ukraine. This area contains natural forest patches, mainly of oak and hornbeam, and supports a mountain entomofauna. Methods Ground beetles (Carabidae) present in the 19 forest patches were categorized into generalists, forest specialists and edge‐preferring species. We analysed the relationship between species richness and fragment area using species richness in the different categories. Results The assemblages contained a high share of generalist species (species that occur also in the surrounding matrix). Forest patch size and the number of generalist species showed a marginally significant negative relationship, indicating that generalist species were more important in smaller patches. Forest specialist species richness was correlated positively with patch area. Edge‐preferring species were shown to influence the species–area relationship: the number of edge‐preferring species increased with the edge:area ratio. Main conclusions Both generalist and edge‐preferring species can considerably distort the species–area relationship. Island biogeography theory can be applied to habitat islands only if the habitat islands are defined correctly from the viewpoint of the target species.     

Broad‐scale geographic patterns in body size and hind wing development of western Palaearctic carabid beetles(Coleoptera: Carabidae)

Research into large‐scale ecological rules has a long tradition but has received increasing attention over the last two decades. Whereas environmental, especially climatic, influences on the geographic distribution of species traits such as body size are well understood in mammals and birds, our knowledge of the determinants and mechanisms which shape spatial patterns in invertebrate traits is still limited. This study analyzes macroecological patterns in two traits of the highly diverse invertebrate taxon of carabid beetles: body size and hind wing development, the latter being directly linked to species’ dispersal abilities. We tested for potential impacts of environmental variables (spatial, areal, topographic and climate‐related) representing both contemporary conditions and historical processes on large‐scale patterns in the two traits. Regression models revealed hump‐shaped relationships with latitude for both traits in the categories 1) all species, 2) widespread and 3) endemic (restricted‐range) species: body size and the proportion of flightless species increased from northern towards southern Europe and then decreased towards North Africa. The shared and independent influence of environmental factors was analyzed by variation partitioning. While contemporary environmental productivity and stability (represented by measures of ambient energy and water energy balance) had strong positive relationships with carabid body size, patterns in hind wing development were most notably influenced by topography (elevation range). Regions with high elevation range and low historical climate variability (since the last ice age), which likely offer long‐term stable habitats (i.e. glacial refugia), coincide with regions with high proportions of flightless species. Thus geographic patterns in carabid traits tend to be formed not only by recent climate but also by dispersal and historical climate and processes (i.e. glaciations and postglacial colonization).     

Desiccation resistance and water balance in southern African keratin beetles (Coleoptera, Trogidae): the influence of body size and habitat

Desiccation resistance and water balance were examined in the adults of seven trogid species, which differed both in body size and in the habitats from which they were collected. Body water contents (51–58% fresh mass) and desiccation rates at 27 °C (0.00026–0.00093 g h⁻¹) in these species were very similar to those of unrelated, similar-sized beetles from arid habitats. The keratin beetles differed markedly from many other adult Coleoptera by virtue of their very high haemolymph osmolality and inability to regulate haemolymph osmolality, and to catabolise lipids for water production, during desiccation. Like most other insects, the xeric trogid species had lower rates of water loss and longer survival times than trogids from mesic areas. This was due both to lower rates of water loss and to the larger body size of species from the more arid areas. Because absolute body water content was higher in large beetles than in small ones, larger body size conferred higher desiccation resistance on the very large Kalahari desert species. This suggests that there may be strong selection for large body size in such insects from arid areas. Most ecological and ecophysiological investigations of geographical variation in body size, and the species-body size distribution, have focused on temperature and metabolic rate as explanatory variables. This study suggests that attention should also be given to desiccation resistance. Accepted: 29 September 1997