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1. Despite the fact that insects are suffering a global decline, little is known about the extent to which species loss affects functional diversity. Thus, to understand the relationship between taxonomic and functional diversity metrics, we focused on saproxylic beetles, which perform vital functions in forest ecosystems. 2. Beetles were collected monthly for a year using flight interception traps placed in three protected areas of mature evergreen Mediterranean Quercus forests. Five morphological traits, one phenological trait, and one physiological surrogate trait were measured, and trophic guild identities were determined. Two taxonomic and six functional metrics were assessed to determine their relationships. 3. Taxonomic and functional diversity patterns, as well as trait patterns, differed depending on the protected areas, as they were influenced by habitat and microhabitat characteristics. Generally, a high number of species tended to support high values of functional richness, but did not ensure high values of functional redundancy, thus affecting the resilience of communities. 4. A 5% of species loss strongly affected the functional redundancy of communities, making them more vulnerable. Functional richness, however, was not affected, since it was associated with a high number of redundant species. Thus, the way species loss affected communities is highly dependent on the presence of rare or redundant species. 5. Taxonomic and functional diversity metrics should be used as complementary tools for conservation purposes. In this way, the comparative analysis of taxonomic and functional patterns may help to predict the vulnerability of saproxylic assemblages to changes or disturbances. [ABSTRACT FROM AUTHOR]

Aim High gamma diversity in tropical montane forests may be ascribed to high geographical turnover of community composition, resulting from population isolation that leads to speciation. We studied the evolutionary processes responsible for diversity and turnover in assemblages of tropical scarab beetles ( Scarabaeidae) by assessing DNA sequence variation at multiple hierarchical levels. Location A 300-km transect across six montane forests (900-1100 m) in Costa Rica. Methods Assemblages of Scarabaeidae (subfamilies Dynastinae, Rutelinae, Melolonthinae) including 118 morphospecies and > 500 individuals were sequenced for the cox1 gene to establish species limits with a mixed Yule-coalescent method. A species-level phylogenetic tree was constructed from cox1 and rrnL genes. Total diversity and turnover among assemblages were then assessed at three hierarchical levels: haplotypes, species and higher clades. Results DNA-based analyses showed high turnover among communities at all hierarchical levels. Turnover was highest at the haplotype level (community similarity 0.02-0.12) and decreased with each step of the hierarchy (species: 0.21-0.46; clades: 0.41-0.43). Both compositional and phylogenetic similarities of communities were geographically structured, but turnover was not correlated with distance among forests. When three major clades were investigated separately, communities of Dynastinae showed consistently higher alpha diversity, larger species ranges and lower turnover than Rutelinae and Melolonthinae. Main conclusions Scarab communities of montane forests show evidence of evolutionary persistence of communities in relative isolation, presumably tracking suitable habitats elevationally to accommodate climatic changes. Patterns of diversity on all hierarchical levels seem to be determined by restricted dispersal, and differences in Dynastinae could be explained by their greater dispersal ability. Community-wide DNA sequencing across multiple lineages and hierarchical levels reveals the evolutionary processes that led to high beta diversity in tropical montane forests through time. [ABSTRACT FROM AUTHOR]

. 1. Using a recently created database representing the joint effort of around 100 invertebrate taxonomists, this study uses the information on 52 arthropoda and 27 mollusca species that are endangered and critically endangered to examine to what extent invertebrate species are represented in existing Spanish protected areas. 2. As distribution information is available at a 100 km2 resolution, we consider different area thresholds to judge cells as being protected. 3. Approximately 19% of the area represented by the grid cells with observed occurrences rates as extant protected reserves, and 36% is included within the Natura 2000 network. 4. If having 50% of the cell area as a Natura 2000 reserve is considered as sufficient to have effective protection, almost 68% of species and 32% of probable populations (contiguous cell groups) would be represented. 5. However, 77% of species and 94% of probable populations are not represented in the current protected reserves if we establish that at least 95% of each cell area should belong to a reserve to provide effective protection. 6. Thus, existing conservation strategies, which are based primarily on the protection of certain areas and vertebrate species, may be insufficient to ensure the conservation of invertebrate species. [ABSTRACT FROM AUTHOR]

Aim Understanding the heterogeneous distribution of species on mountains is an important aim in ecology. Altitudinal gradients have enormous potential for improving our knowledge of trends in biodiversity and conservation. In this study, we investigated the variation in scarab beetle diversity (Dynastinae, Rutelinae and Melolonthinae) along an elevational tropical forest gradient. Location The Atlantic slope of the Guanacaste mountain range in Costa Rica. Methods Ultraviolet light traps placed in six forests situated from 100 to 1510 m were used. Changes in species composition and richness among elevations were investigated. Differences in the altitudinal patterns using different groups of species were examined: the whole assemblage, each separate subfamily and two different trophic habits (phytophagous or saproxylic). The effects of temperature, humidity and elevation on scarab distribution were tested using canonical correspondence analyses. The relationship between the community similarity of the studied forests and the altitudinal distance among them was also analysed. Results Species composition and richness changed along the gradient. The peak in species richness varied depending on the species group considered and in all cases occurred 500 or 800 m. Forests at these altitudes were also the richest in exclusive species. Species composition turnover among elevations appeared with a clear separation between lowland and highland fauna. The latter was lower in richness but also had exclusive species. Temperature, humidity and altitude affected species distribution, with altitude being the most important factor for all the subfamilies studied. Main conclusions Our results showed that species distribution fits a hump-shaped pattern. The peak of this pattern varied depending on the taxonomic group and mountain analysed, highlighting the importance of evolutionary processes as species distribution drivers. The fact that species richness peaked at elevations where human impact is currently important underlines the value of the development of conservation strategies for these areas. [ABSTRACT FROM AUTHOR]

Aim Landscape structure influences the distribution of animals, altering their movements and their ability to reach habitat patches. We analysed the spatial patterns of dung beetle species diversity in three differently structured natural landscapes in a Mediterranean protected area in the centre of the Iberian Peninsula. Location Cabañeros National Park, Central Spain. Methods Diversity components within (α) and among (β) the three main vegetation types in the reserve were compared by using a hierarchical nested design. These were forests, scrublands and grasslands embedded in three different landscapes, where each was the most dominant and structurally connected habitat. Results Species richness of grassland habitat did not vary across landscapes, but forest habitat showed lower species richness in the grassland-dominated landscape. Scrubland was the least species-rich habitat, but here again there was no significant variation across landscapes. However, in all cases, there was a significant influence of habitat context (configuration of habitat patches within landscape matrix) on similarity of species composition. These tended to be more similar to the dominant landscape matrix where they were embedded, rather than to the same habitat type in other landscapes. Additive partitioning of diversity showed higher than expected values of β in all landscapes, which indicated a structured response. Highest values of β in the grassland-dominated landscape suggest that this was the least connected landscape for dung beetles. Main conclusions Our results suggest that in homogeneous conditions of climate and trophic resources, landscape structure may well be more important than habitat type as a determinant of dung beetle distribution in the Mediterranean. [ABSTRACT FROM AUTHOR]

Aim The circum-Mediterranean region is one of the most complex regions of the Earth in terms of geography and natural history. The Old World species of the beetle subtribe Anisopliina (Scarabaeidae) feed almost exclusively on the pollen of grasses (Poaceae). Within this group, the ‘anisopliine clade’ forms a monophyletic group distributed mainly in the circum-Mediterranean region. Here, we reconstruct the biogeographical history of the anisopliine beetles in relation to the diversification of grasses, and compare this reconstruction with previous hypotheses concerning the evolution of the Mediterranean fauna and with palaeogeographical accounts of the history of this region. Location The Mediterranean region, including North Africa, the Western Mediterranean, Balkans–Anatolia, Middle East and Caucasus. Methods Dispersal–vicariance analysis (diva) was used to reconstruct ancestral distributions based on the morphological phylogeny and to infer the biogeographical processes that have shaped the observed distribution patterns. To account for phylogenetic uncertainty in the biogeographical reconstruction, we ran alternative ancestral distributions derived bydiva over a sample of trees obtained by bootstrapping the original data set, reflecting the relative confidence of the ancestral areas on the various clades in the phylogeny. Results The Eastern Mediterranean region and the Caucasus are inferred as the ancestral area of most of the anisopliine lineages. The Eastern Mediterranean region is also reconstructed as the source area of the majority of dispersal events, in particular towards North Africa and the Western Mediterranean. The Iberian Peninsula is inferred as part of the ancestral distribution of the anisopliine clade but also as the setting of several independent colonization events via both the North African platform ( Anthoplia) and a European dispersal route ( Anisoplia). Main conclusions Our results confirm the role played by the Eastern Mediterranean as an evolutionary cradle of diversity for Mediterranean lineages. This can be explained by a recent and intense orogenic activity that might have promoted isolation and allopatric speciation within lineages. Both the Anomalini fossil record and the close association of anisopliine beetles with grasses suggest that the anisopliine clade originated in the Late Tertiary and that its spatial and temporal evolution within the Mediterranean Basin coincided with that of its major food source, the Mediterranean Poaceae. [ABSTRACT FROM AUTHOR]

The Cetoniidae, the showiest of scarabs, comprises some 3900 species in 515 genera, distributed worldwide except for subpolar areas and some offshore New Zealand islands. Parsimony analysis of 76 larval and adult characters and 42 terminal taxa supports the monophyly of Cetoniidae (sensu Krikken, 1984 ), but not of the traditionally considered subfamilies Cetoniinae and Trichiinae (sensu Krikken, 1984 ). In the study taxon, larval characters are shown to be more informative than those of adults for deeper phylogeny. The evolution of some larval characters (head and legs) in relation to feeding habits is discussed on the basis of phylogenetic analysis. The results show an evolutionary shift from wood associations to a higher ecological plasticity that allows the larvae to feed on wide sources of organic matter (including compost, dung, gopher burrows, packrat middens, ant debris piles, etc.). [ABSTRACT FROM AUTHOR]

The subtribe Anisopliina (Scarabaeidae: Rutelinae: Anomalini) is associated with grasses, and its species are distributed in the Palaearctic, Oriental, Ethiopian, Nearctic and Neotropical biogeographical regions. Phylogenetic analysis of adult morphological characters was conducted to examine the monophyly and classification of the group, as well as to examine characters associated with grass pollinivory and graminivory. We review the biology, phylogeny and classification of the Anisopliina and provide an overview of each genus. The analysis of ninety-one morphological characters using parsimony does not support the monophyly of the subtribe Anisopliina. Instead, the results provide support for a group referred to here as the anisopliine clade, a circum-Mediterranean group, forming an internal clade within the well-supported tribe Anomalini. Sister group relationships are discussed, possibly being associated with a New World anomaline taxon. Character states associated with grass herbivory, including mouthpart and leg characters, are discussed based on the phylogenetic analysis. Within the Anomalini, an evolutionary shift from generalized leaf feeding to grass associations and grass pollen feeding is supported. [ABSTRACT FROM AUTHOR]

Thirty-eight characters derived from the larvae of Geotrupidae (Scarabaeoidea, Coleoptera) were analysed using parsimony and Bayesian inference. Trees were rooted with two Trogidae species and one species of Pleocomidae as outgroups. The monophyly of Geotrupidae (including Bolboceratinae) is supported by four autapomorphies: abdominal segments 3–7 with two dorsal annulets, chaetoparia and acanthoparia of the epipharynx not prominent, glossa and hypopharynx fused and without sclerome, trochanter and femur without fossorial setae. Bolboceratinae showed notable differences with Pleocomidae, being more related to Geotrupinae than to other groups.Odonteusspecies (Bolboceratinae s.str.) appear to constitute the closest sister group to Geotrupinae. Polyphyly of Bolboceratinae is implied by the following apomorphic characters observed in the‘ Odonteuslineage’: anterior and posterior epitormae of epipharynx developed, tormae of epipharynx fused, oncyli of hypopharynx developed, tarsal claws reduced or absent, plectrum and pars stridens of legs well developed and apex of antennal segment 2 with a unique sensorium. A‘ Bolbelasmuslineage’ is supported by the autapomorphic presence of various sensoria on the apex of the antennal segment, and the subtriangular labrum (exceptEucanthus). This group constituted byBolbelasmus,BolbocerosomaandEucanthusis the first evidence for a close relationship among genera, but more characters should be analysed to test the support for the clade. A preliminary classification at tribe level of Geotrupinae is suggested as follows: Chromogeotrupini (type genusChromogeotrupes), Lethrini (type genusLethrus), Taurocerastini (type genusTaurocerastes) and Geotrupini (type genusGeotrupes). Some ecological facts of Geotrupidae evolution could also be explained by the present results, such as those related to diet and nesting behaviour. Both coprophagy and male–female co-operation in nesting appear as derived traits. [ABSTRACT FROM AUTHOR]

The thermoregulation strategies of Scarabaeus sacer L. and Scarabaeus cicatricosus Lucas were studied in the Doñana National Park, Spain. In this area, both species coexist, showing the same habitat and food preferences. However, S. cicatricosus is active during warmer parts of the day compared to S. sacer. Both species thermoregulate their thoracic temperature but, whereas the abdomen of S. sacer is a passive thermal window, S. cicatricosus actively thermoregulates abdominal temperature by increasing heat transfer from the thorax to the abdomen at high Ta values. In the case of S. sacer, their endothermy indicates an adaptive capacity to thorax heat retention, as occurs mainly in winter-flying insects. This mechanism, possibly related to the aerodynamic flight posture in Scarabaeinae, could be an effective barrier to retard the rate of abdominal heat loss during flight. This endothermic strategy makes flight difficult at higher temperatures, although it allows flight during cooler periods of the day. On the other hand, S. cicatricosus showed a different adaptive behaviour to S. sacer. In this case, a significant decrease in abdominal heat loss at higher ambient temperatures would indicate a decrease in heat transfer from the thorax to the abdomen, as occurs in some desert and semiarid insects. This 'heat exchanger' mechanism observed in S. cicatricosus could be due to the irregular posture adopted during flight, with the posterior legs clearly extended and separate from the body. This behaviour increases turbulence and convective cooling, favouring exposure of the soft abdominal tergal cuticle and, subsequently, water loss. Thus, for S. cicatricosus, the well-adapted 'heat exchanger' permits flight during periods of the day when temperatures would possibly be lethal for those species with high endothermy. From an adaptive viewpoint, these mechanisms of thermoregulation may explain how both closely-related sympatric species respond in different ways to environmental temperature, favouring their coexistence. [ABSTRACT FROM AUTHOR]

Abstract. A study to assess the influence of abiotic (climatic conditions) and biotic factors (food resources, habitat preference and human activity) on endemism patterns of dung beetles in the Mediterranean region was conducted in the Iberian Peninsula and the Balearic Islands. The Thermicity Index (It), the Mediterraneity Index (Im3) and the Aridity Index (Ia) were used to assess the influence of abiotic factors. Relative rabbit density (DR), the proportion of landscape used historically for grazing by sheep and goats and the nature of the food resource were used to assess the influence of biotic factors. Relative endemism (EN) of dung beetle assemblages was positively and significantly related with all of the factors considered. However, the Aridity and Mediterraneity Indices are the best predictors of EN. The predicted endemism (EN = 0.017 Ia + 0.004 Im3 + 0.422) was highly positively and significantly related with the observed endemism. Dung beetle assemblages with the highest relative endemism were observed in the south-eastern part of the Iberian Peninsula. This distribution corresponded to the highest Aridity and Mediterraneity. In contrast, dung beetle assemblages with lower endemism were located in more humid and temperate areas. Assemblages of dung beetles with the highest endemism comprise many species adapted to aridity and the exploitation of dry dung pellets. Conservation of traditional grazing activity by pellet-dropping sheep and goats might benefit the maintenance of dung beetle biodiversity in Mediterranean ecosystems. [ABSTRACT FROM AUTHOR]