Functional and Taxonomic Beta Diversity of Saproxylic Beetles in Mediterranean Forests: On What Factors Do They Depend?

Understanding how biodiversity is distributed across geographical and environmental gradients is a main goal of diversity sciences. However, since ecosystem processes are linked to variation in functional traits of the biota, examining functional beta diversity is particularly important. Our objective was to analyze the taxonomic and functional beta diversity patterns of saproxylic beetle assemblages in evergreen Quercus forest of Spain. We tested whether environmental or geographical distance had a greater influence on taxonomic and functional beta diversity, and if both measures of beta diversity were affected by the same environmental variables. We used 45 flight interception traps distributed in three protected areas over a 12-mo period to sample saproxylic beetles. We measured 13 environmental variables around each trap and the geographical distance between traps. For functional composition, we used 12 functional traits from four functional groups (morphological, phenological, trophic, and a surrogate of physiological). Our results showed that environmental differences between areas influenced the taxonomic and functional beta diversity components (replacement and loss/gain) but in different ways. While replacement components (higher for taxonomic composition) increased with environmental distance, the loss or gain components (higher for functional composition) remained constant, indicating that species replacement mostly involved functionally redundant species. Besides, environmental variables influencing both taxonomic and functional composition were strongly dependent on each area. In conclusion, in well-preserved Mediterranean forests, environmental filtering determines the taxonomic and functional composition of saproxylic beetle assemblages, by favoring species replacement but filtering species traits.

Influence of Traditional Management and Environmental Variables on Mediterranean Saproxylic Beetle Assemblages.

Mediterranean oak forests of the Iberian Peninsula host a great diversity of saproxylic beetles. For centuries, humans have carried out traditional management practices in this area, at both habitat and tree level, causing changes in forest structure. The aim of this study was to evaluate the anthropic effect of these traditional practices on saproxylic beetle diversity by measuring a set of environmental variables related to forest structure at both plot and tree level. Fauna was collected using window traps over a period of 12 mo. Multiple regression procedures showed which variables significantly affected the diversity of the studied assemblage. Our results demonstrated that the different metrics used to assess the diversity of assemblages responded variably depending on the management strategies applied and the level at which they were carried out. Certain management practices that disrupted the landscape from its natural state, such as the introduction of livestock or the local removal of particular trees, maximized species richness but, nevertheless, had a negative effect on the rest of diversity metrics analyzed. However, other practices such as pollarding, which involves the suppression of the main branch of the tree, had a positive effect on all diversity metrics evaluated as it promoted the formation of potential microhabitats for saproxylic fauna. We concluded that not all types and degrees of traditional forest management favor saproxylic beetle diversity and that different diversity metrics should be taken into consideration in future strategies for the protection and conservation of this fauna.

Saproxylic Beetle Assemblage Selection as Determining Factor of Species Distributional Patterns: Implications for Conservation.

The knowledge of the distributional patterns of saproxylic beetles is essential for conservation biology due to the relevance of this fauna in the maintenance of ecological processes and the endangerment of species. The complex community of saproxylic beetles is shaped by different assemblages that are composed of species linked by the microhabitats they use. We evaluate how different the species distribution patterns that are obtained can be, depending on the analyzed assemblage and to what extent these can affect conservation decisions. Beetles were sampled using hollow emergence and window traps in three protected areas of the Iberian Peninsula. Species richness, composition, and diversity turnover were analyzed for each sampling method and showed high variation depending on the analyzed assemblage. Beta diversity was clearly higher among forests for the assemblage captured using window traps. This method collects flying insects from different tree microhabitats and its captures are influenced by the forest structuring. Within forests, the assemblages captured by hollow emergence traps, which collect the fauna linked to tree hollows, showed the largest turnover of species, as they are influenced by the characteristics of each cavity. Moreover, the selection of the forest showing the highest species richness strongly depended on the studied assemblage. This study demonstrates that differences in the studied assemblages (group of species co-occurring in the same habitat) can also lead to significant differences in the identified patterns of species distribution and diversity turnover. This fact will be necessary to take into consideration when making decisions about conservation and management.

Facilitation among saproxylic insects inhabiting tree hollows in a Mediterranean forest: the case of cetonids (Coleoptera: Cetoniidae) and syrphids (Diptera: Syrphidae).

Tree hollows offer an ideal niche for saproxylic insects in mature Mediterranean forests, where Diptera and Coleoptera are the richest groups. Co-occurrence is frequently observed among many species of both groups in these microhabitats, and some of these species have been considered to facilitate the presence of other species by acting as ecosystem engineers. One of the systems that is found in Mediterranean tree hollows is formed by cetonid (Coleoptera: Cetoniidae) and syrphid (Diptera: Syrphidae) larvae. Here, cetonid larvae feed on wood and litter and produce a substrate that is easier to decompose. To assess the possible role of these larvae as facilitating agents for the saproxylic guild, we studied whether the presence of saprophagous Syrphidae inside tree hollows is associated with the activity of cetonid larvae. Furthermore, in laboratory conditions, we tested whether cetonid larvae activity can improve the development and fitness of the saprophagous syrphid species. Our results show that "cetonid activity" was the variable that best explained the presence of saprophagous syrphid species in natural conditions. Myathropa florea (L., 1758) was one of the species most influenced by this activity. The laboratory experiment gave similar results, demonstrating that an enriched substrate with Cetonia aurataeformis Curti, 1913 larval feces improves syrphid larval growth rate and fitness of adults (measured as longer wing length) of M. florea.

Molecules, wing pattern and distribution: an approach to species delimitation in the "loxurina group" (Lepidoptera: Lycaenidae: Penaincisalia).

The wide range of morphological variations in the "loxurina group" makes taxa identification difficult, and despite several reviews, serious taxonomical confusion remains. We make use of DNA data in conjunction with morphological appearance and available information on species distribution to delimit the boundaries of the "loxurina" group species previously established based on morphology. A fragment of 635 base pairs within the mtDNA gene cytochrome oxidase I (COI) was analysed for seven species of the "loxurina group". Phylogenetic relationships among the included taxa were inferred using maximum parsimony and maximum likelihood methods. Penaincisalia sigsiga (Bálint et al), P. cillutincarae (Draudt), P. atymna (Hewitson) and P. loxurina (C. Felder & R. Felder) were easily delimited as the morphological, geographic and molecular data were congruent. Penaincisalia ludovica (Bálint & Wojtusiak) and P. loxurina astillero (Johnson) represent the same entity and constitute a sub-species of P. loxurina. However, incongruence among morphological, genetic, and geographic data is shown in P. chachapoya (Bálint & Wojtusiak) and P. tegulina (Bálint et al). Our results highlight that an integrative approach is needed to clarify the taxonomy of these neotropical taxa, but more genetic and geographical studies are still required.

Molecules, wing pattern and distribution: an approach to species delimitation in the "loxurina group" (Lepidoptera: Lycaenidae: Penaincisalia)

The wide range of morphological variations in the "loxurina group" makes taxa identification difficult, and despite several reviews, serious taxonomical confusion remains. We make use of DNA data in conjunction with morphological appearance and available information on species distribution to delimit the boundaries of the "loxurina" group species previously established based on morphology. A fragment of 635 base pairs within the mtDNA gene cytochrome oxidase I (COI) was analysed for seven species of the "loxurina group". Phylogenetic relationships among the included taxa were inferred using maximum parsimony and maximum likelihood methods. Penaincisalia sigsiga (Bálint et al), P. cillutincarae (Draudt), P. atymna (Hewitson) and P. loxurina (C. Felder & R. Felder) were easily delimited as the morphological, geographic and molecular data were congruent. Penaincisalia ludovica (Bálint & Wojtusiak) and P. loxurina astillero (Johnson) represent the same entity and constitute a sub-species of P. loxurina. However, incongruence among morphological, genetic, and geographic data is shown in P. chachapoya (Bálint & Wojtusiak) and P. tegulina (Bálint et al). Our results highlight that an integrative approach is needed to clarify the taxonomy of these neotropical taxa, but more genetic and geographical studies are still required.

Taxonomy of Iberian Hoplia (Col., Scarabaeoidea, Hopliinae) based on mtDNA analysis.

The morphology of some Hoplia species (Scarabaeoidea: Hopliinae) is so variable that parapatric populations have often been considered different species or subspecies. In this study we analyze the nucleotide sequences of a fragment of mitochondrial gene cytochrome c oxidase subunit I (COI) of six species and two subspecies of Palaearctic Hoplia to reexamine the species limits. Based on the analysis of sequences from COI and morphological and ecological observations, we consider Hoplia freyi Baraud to be a junior synonym of Hoplia chlorophana Erichson and H. philanthus ramburi Heyden to be a junior synonym of H. philanthus philanthus (Fuessly). However, complete resolution of relationships among H. philanthus subspecies requires the addition of sequences from genes evolving faster than COI. Phylogenetic relationships among the species studied are discussed.