Annotated checklist of the Hydraenidae (Coleoptera: Staphylinoidea) recorded from Iran

According to our analysis of literature sources on the family Hydraenidae (Coleoptera: Staphylinoidea) of Iran, it is represented by 67 species and three genera: Hydraena Kugelann, 1794 (21 species), Limnebius Leach, 1815 (six species) and Ochthebius Leach, 1815 (40 species). Twenty-two species are supposed to be endemic to Iran. Here, for the first time, we summarize species distribution data in Iranian provinces and present them in a checklist of Iranian Hydraenidae. We also present two additional species lists: one with incorrect records (one species and one subspecies) and the other with unidentified species.     

Comparative morphology of the head of selected sporophagous and non‐sporophagous aleocharinae (Coleoptera: Staphylinidae): Musculature and hypopharynx‐prementum complex

To investigate whether specialization to spore‐ (or pollen‐) feeding in advanced Aleocharinae is mirrored by their head anatomy, we compiled and compared synchrotron X‐ray micro‐tomography datasets for 11 Aleocharinae in conjunction with previous data for two aleocharine and six outgroup species (two nonstaphylinids, four staphylinids). We describe the presence/absence of head muscles and investigate the variability of points of origin by character mapping analyses. Monophyly of Aleocharinae is supported by the absence of M. 48 (M. tentoriobuccalis anterior), and by changes in the origins of Mm. 1, 2, 17, 18, 28, 29, 30. Within Aleocharinae the origins of the labial muscles (Mm. 28–30) have shifted posteriorly to the gula, which might enhance the movement posterad of the hypopharynx and partly compensate for the loss of M. 48. We also analyzed the general organization of the hypopharynx‐prementum complex and the fine structure of the mandibles through SEM studies. In the absence of grinding mandibular molae like those of most mycophagous Coleoptera, seven aleocharine species studied have evolved “pseudomolae” at the ventral side of the mandibles that replace true molae as secondary grinding surfaces. In these species, the hypopharynx is elevated and displaced anteriorly, bearing a bowl‐like depression on its surface that functions as a mortar where spores are ground between the hypopharynx and the mandibles. Two of these species are not yet known to feed on spores or pollen. Another species (Oxypoda alternans) is thought to feed on fungus material but bears no pseudomolae on its mandibles. J. Morphol. 271:910–931, 2010. © 2010 Wiley‐Liss, Inc.     

Head morphology of selected Staphylinoidea (Coleoptera: Staphyliniformia) with an evaluation of possible groundplan features in Staphylinidae

In insects, anatomical features of the head have been found to provide important information for phylogenetic and comparative evolutionary studies. We analyzed the internal head morphology of three (omaliine, tachyporine, oxyteline group) out of the four subfamily groups of the beetle family Staphylinidae plus two non‐staphylinid outgroups (i.e., Agyrtidae and Leiodidae). Synchrotron X‐ray micro‐tomography was used to obtain comparative head anatomical datasets of eight species to describe (i) the presence/absence of muscles inside the head capsule and (ii) the variability in their points of origin. Nineteen of these muscles were phylogenetically informative (nine with respect to presence/absence and eleven with respect to the origin; one muscle had an influence on both analyses) and were used in character mapping analyses to reconstruct groundplan patterns of the head musculature in Staphylinidae and their subgroups. Three muscles (Mm. 7, 9, 50) were identified as possibly autapomorphic for Staphyliniformia. The taxon (Agyrtidae + Leiodidae) is supported by the absence of M. 9. The monophyly of the tachyporine group is supported by a common origin of M. 4. Aleocharinae, a subfamily within the tachyporine group, is supported by the absence of M. 42 and possibly by numerous points of origin of various muscles (Mm. 1, 17, 28, 29, 30). Our analysis of the general organization of the hypopharynx‐prementum‐complex has revealed that this complex is organized in a similar way in the investigated staphylinoids, i.e., with the prementum lying anteriorly to and being in line with the hypopharynx and the mentum. We have found deviating conditions in the investigated species of the Aleocharinae, in which the prementum can be largely retracted posteriorly. Consequently, it is sandwiched between the ventral mentum and the dorsal hypopharyngeal region. The hypopharyngeal region is thus lifted dorsad to a large extent, approaching the cibarial roof. This situation is paralleled by a loss of the hypopharyngeal retractor (M. 42) and a shift of origin of premental retractors (Mm. 28–30) posteriorly toward the gula. J. Morphol. 270:1503–1523, 2009. © 2009 Wiley‐Liss, Inc.     

Comparative trophic morphology in eight species of damselfishes (Pomacentridae)

Damselfishes show significant biodiversity in the coral reefs. To better understand such diversity, an ecomorphological approach was investigated in the trophic morphology of eight species of Pomacentridae (Chromis acares, C. margaritifer, Dascyllus aruanus, D. flavicaudus, Pomacentrus pavo, Plectroglyphidodon johnstonianus, Pl. lacrymatus and Stegastes nigricans) belonging to different trophic guilds (zooplankton, algal, coral polyp feeders and omnivores). Geometric morphometrics were used to quantify size and shape variations in four skeletal units: (1) neurocranium, (2) suspensorium and opercle, (3) mandible and (4) premaxilla. This method allowed us to reveal shape and size differences correlated to functional diversity both within and between trophic guilds. Among zooplanktivores, C. margaritifer, D. aruanus and D. flavicaudus have a high and long supraoccipital crest, short mandibles forming a small mouth and high suspensoria and opercles. These three species can be considered to be suction feeders. In the same guild, C. acares shows opposite characteristics (long and thin mandibles, lengthened neurocranium and suspensorium) and can be considered as a ram feeder. Among herbivores and corallivores, the two species of Plectroglyphidodon and S. nigricans can be considered as grazers. Differences in skeletal shape are mainly related to improving the robustness of some skeletal parts (broad hyomandibular, short and high mandibles). The shapes of P. pavo, which feeds largely on algae, strongly differ from that of the other three grazers exhibiting similar morphological characteristics to C. acares (e.g., long and shallow suspensorium, lengthened neurocranium). This highlights likely differences concerning cutting or scraping method. Finally, no strong correlations exist between size and shapes in the eight studied species. Size difference among species having a very similar shape could be viewed as a factor optimizing resource partitioning.     

Pretarsal structures in Leiodidae and Agyrtidae (Coleoptera,Staphylinoidea)

We analysed pretarsal characters of 87 species of Leiodidae (including 10 cholevines and representatives of all tribes and ca. 60% of the genera of non‐cholevines), five species of Agyrtidae, and nine representatives of outgroup taxa (Hydraenidae, Staphylinidae, Hydrophilidae, and Histeridae) using scanning electron microscopy. We focused our observations on the architecture of the empodium (including the sclerites and associated setae), the shape and composition of the medial projection of the distal margin of the terminal tarsomere, and the armature of the claws, which were considered a promising source of information for delimiting supraspecific taxa in our previous study. We identified several diagnostic features and recognize potential synapomorphies at the tribal, subtribal and generic levels. The internal systematic arrangement and/or even the monophyletic status of most of the subfamilies of Leiodidae (Camiarinae, Catopocerinae, Leiodinae, and Platypsyllinae) are challenged. We identified potential synapomorphies for Camiarinae (Camiarini and Agyrtodini) and Leiodinae. The non‐monophyly of Cholevinae is possible because part of the tribe (Anemadini, Eucatopini, and Oritocatopini) shares potentially apomorphic features with Leiodinae (e.g., a triangular medial projection with a diagonal row of conical spines), whereas another part (Leptodirini and Ptomaphagini) shares a potentially apomorphic feature with Coloninae and Platypsyllinae (a typical medial projection with two distinct triangular projections).     

天牛科5亚科昆虫成虫口器形态比较

观察比较了天牛科5亚科昆虫成虫口器的形态特征。结果表明:沟胫天牛亚科上颚切齿钝,花天牛亚科和天牛亚科中访花天牛上颚内缘具毛刷,沟胫天牛亚科和锯天牛亚科天牛不具毛刷;锯天牛亚科下颚内颚叶退化,沟胫天牛亚科、花天牛亚科、天牛亚科中内颚叶、外颚叶均发达,沟胫天牛亚科下颚须第4节纺锤状,锯天牛亚科、花天牛亚科棒状,天牛亚科形态多样;花天牛亚科和天牛亚科中访花天牛下唇唇舌发达,分2叶且膜质,其他亚科中唇舌均有骨化。锯天牛亚科天牛内颚叶退化,与其成虫阶段不取食有关,花天牛亚科和天牛亚科中访花天牛唇舌发达且成膜质,与访花习性有关。     

Comparative morphology of the tentorium and hypopharyngeal–premental sclerites in sporophagous and non‐sporophagous adult Aleocharinae (Coleoptera: Staphylinidae)

We elucidate the configuration of the tentorium and the sclerites of the hypopharynx–prementum complex in selected spore‐ (pollen‐) and non‐spore‐feeding Aleocharinae (Staphylinidae) by presenting the first comparative 3D reconstructions of these structures for 19 staphylinoid beetle species (six outgroups, 13 Aleocharinae). General organization of the tentorium follows the groundplan previously proposed for adult Staphylinidae, although some taxa have reduced or lost the dorsal (all Aleocharinae studied, Agathidium mandibulare [Leiodidae]) or anterior (Omalium rivulare [Omaliinae], Anotylus sculpturatus [Oxytelinae]) tentorial arms. All species investigated have premental and hypopharyngeal sclerites that are partly homologizable across taxa. We clarified that Musculus praementopalpalis externus originates from the premental sclerite, resolving its unclear origin reported in our previous publications. Eight of 13 investigated Aleocharinae species are spore/pollen feeders, six obligatorily. Three of these six (Eumicrota, Gyrophaena fasciata, G. gentilis) have grinding pseudomolae and a fully developed hypopharyngeal suspensorium with posterior bridge and anterior elongations; the remaining three (Oxypoda, Pagla, Polylobus) lack pseudomolae and suspensorial bridge, but have the suspensorium elongated anteriorly. The dorsolateral side of the hypopharyngeal sclerite interacts with the pseudomola. Obligate sporophagy/pollinivory apparently arose at least three times in Aleocharinae, not always involving the pseudomola–hypopharynx grinding mechanism.     

Factors affecting the structure of Coleoptera assemblages on bracket fungi (Basidiomycota) in a Brazilian forest

Insect–fungal interactions are an important but understudied aspect of tropical forest ecology. Here we present the first large‐scale study of insect communities feeding on the reproductive structures of macrofungi (basidiomes) in the Neotropics. This trophic interaction is not well characterized in most ecosystems; however, beetle consumption of basidiomes is thought to be affected by fungal factors, via mechanisms analogous to those observed in plant–herbivore interactions and in some interactions with fungi as hosts in the Holarctic region. We investigated how the composition of beetle assemblages varies as a function of fungal taxonomic distance, basidiome consistency, and hyphal systems. We collected 367 basidiomes belonging to the orders Polyporales and Hymenochaetales in the subtropical Araucaria angustifolia forest region of southern Brazil, along with any fauna present or without it. Basidiomes were maintained individually in the laboratory in plastic containers for up to three months to allow beetles to develop to adulthood, at which point the beetles were collected. We found that 207 basidiome specimens representing 40 species were associated with beetles. We recorded 447 occurrences of Coleoptera, representing 90 morphospecies from 20 families. We found that assemblages of fungivorous Coleoptera were more similar among more closely related fungi. Furthermore, the beetle assemblages varied as a function of basidiome toughness, which is influenced by sporocarp consistency and hyphal system type. The associations between beetles and basidiomes resemble those reported previously in temperate zones, suggesting continuity in the structure of such associations across a wide latitudinal range.     

捕食性颏花金龟口器形态的适应进化及功能研究(鞘翅目:金龟科:花金龟亚科)

颏花金龟为捕食性,是花金龟亚科12族中食性最为特殊的类群,以其下唇的前颏极度扩大而得名。依据捕食对象及对应形态变异类型的不同,其可分为蚁食性和蚜食性两类。由于颏花金龟的食性异于其他种类花金龟的植食性,栖息环境也有极大的不同,故其口器适应进化出诸多形态类型。通过比较颏花金龟、花金龟亚科其他种类、其他植食性金龟(丽金龟和犀金龟两个亚科为代表)以及典型肉食性甲虫口器的形态,对颏花金龟口器特殊结构及功能进行了分析,并初步探索了取食机制及颏花金龟的生存策略。     

Implication of the feeding limb morphology for herbivorous feeding in some freshwater cyclopoid copepods

1. The mouthparts of five species of adult planktonic cyclopoid copepods (Cyclops vicinus, C. abyssorum, Acanthocyclops robustus, Mesocyclops leuckarti and Thermocyclopscrassus), in particular the distance between setae and setules of the maxilliped (which can indicate the ability to retain small particles), were compared using electron and light microscopy. 2. The mesh‐sizes of the food‐collection grid formed by these setae and setules ranged between 4.6 and 13.2 μm; the area covered by the grid ranged between 6000 and 32 000 μm². 3. Mesh‐size was not simply correlated with body size. Cyclops abyssorum and M. leuckarti have the coarsest meshes and T. crassus the finest, while C. vicinus and A. robustus were intermediate. 4. The results suggest that cyclopoid copepods are able to retain particles in the size range of nanoplankton and that differences in mesh‐sizes between species may explain differences in the ability to subsist and reproduce on a diet of small algae.     

A phylogeny of Xanthopygina (Insecta,Coleoptera) reveals major lineages and the origin of myrmecophily

Xanthopygina is a group of colourful, neotropical rove beetles (Staphylinidae) comprising 28 genera and more than 350 species. While many genera are found on rotting fruits, carrion, dung and mushrooms, several taxa have evolved associations with social insects. Previous phylogenetic analyses have used only a subset of genera and were based solely on molecular data. In this paper, we performed Bayesian and maximum likelihood phylogenetic analyses of all known genera, including three potentially new genera, based on molecular (4,797 bp) and morphological (91) characters. Our results reaffirmed the monophyly of the subtribe and divided it into eight major lineages: the Elmas, Gastrisus, Isanopus, Ocyolinus, Plociopterus, Smilax, Trigonopselaphus and Xanthopygus groups. We hypothesized that myrmecophily evolved once in the common ancestor of the Smilax group, and that sphecophily possibly evolved twice, in the Trigonopselaphus and Xanthopygus groups.     

Pollen grains adhere to the moist mouthparts in the flower visiting beetle Cetonia aurata (Scarabaeidae, Coleoptera)

Flower visiting beetles possess numerous structural adaptations of their mouthparts to adhere and ingest pollen grains. Using a Cryo-SEM approach the examination of the mouthparts in rapidly frozen Cetonia aurata (Scarabaeidae) indicates a previously unknown technique of pollen uptake in Coleoptera. Cryo-SEM micrographs of the mouthparts reveal a fluid covering the bristles on the buccal surface. In this way the bristles of the galeae form a wet brush which represents the primary organ of pollen uptake. The fluid improves adhesion of pollen to bristles lacking any specialized adhering surface or highly sculptured cuticle as present in other pollen feeding Coleoptera. The well developed mola region of the otherwise non-biting mandibles of C. aurata indicates that these beetles open pollen grains mechanically before ingestion. Examination of gut content demonstrated that crushed and intact pollen occur in all regions. The Cryo-SEM method represents a new approach to study functional morphology including the interaction of microstructures and fluids on cuticular surfaces of insects. Handling editor: Gimme Walter     

On the evolution of adult head structures and the phylogeny of Hydraenidae (Coleoptera, Staphyliniformia)

External and internal head structures of adults of Orchymontiinae, Prosthetopinae, Hydraeninae and Ochthebiinae were studied and those of Ochthebius semisericeus and Limnebius truncatellus are described in detail. The results are evaluated with respect to their relevance for a reconstruction of hydraenid phylogeny and also compared with structural features found in adults of other staphyliniform families. The monophyly of Hydraenidae is supported by the presence of a plate‐like, trilobed premento‐hypopharyngeal extension, an unusual origin of m. tentoriohypopharyngalis, dorsal tentorial arms firmly fused with the head capsule, modified basal antennomeres, and palpigers connected by a transverse sclerotized bar. Orchymontiinae are monophyletic and the basal sister group of the remaining Hydraenidae. The presence of a ventral transverse genal bulge and of a pubescent antennal club with more than two antennomeres (reversal in some prosthetopines: e.g. Mesoceration abstrictum) are possible apomorphies of Hydraenidae excluding Orchymontiinae. Prosthetopinae are probably monophyletic and the sister group of Ochthebiinae + Hydraeninae. The latter clade is characterized by a distinct cupula formed by antennomere VI, a loose five‐segmented pubescent antennal club, and a modified antennal musculature. The presence of an unusual tentorio‐pharyngeal dilator is a shared derived feature of Ochthebiinae and the genus Davidraena. The monophyly of Ochthebiinae was confirmed and Ochtheosus is the sister group of the remaining ochthebiine genera, which are characterized by a perforated wall‐like structure formed by the posterior tentorial arms. The absence of this tentorial modification and the fimbriate galea are plesiomorphies retained in Ochtheosus. Calobius differs strongly from other subgenera of Ochthebius and a generic status may be appropriate. The monophyly of Hydraeninae is not supported. Hydraena was confirmed as a clade and Laeliaena and Limnebius are sister groups. The latter genus is characterized by several autapomorphies. The basal position of Orchymontiinae and Prosthetopinae suggests a Gondwanan origin of Hydraenidae and a primary preference for life in running water. Important evolutionary changes of head structures are complex transformations of the antennae and related structures. Yet, the use of the antennae as accessory breathing organs is not a groundplan feature of the family. The results of this study strengthen the case of staphylinoid affinities of Hydraenidae.     

Functional correlation between habitat use and leg morphology in birds (Aves)

Many of the morphological features of animals are considered to be adaptations to the habitat that the animals utilize. The habitats utilized by birds vary, perhaps more than for any other group of vertebrates. Here, we study possible adaptations in the morphology of the skeletal elements of the hind limbs to the habitat of birds. Measurements of the lengths of the femur, tibiotarsus and tarsometatarsus of 323 bird species from 74 families are used together with body mass data, taken from the literature. The species are separated into six habitat groups on the basis of literature data on leg use. A discriminant analysis of the groups based on leg morphology shows that swimming birds, wading birds and ground living species are more easily identified than other birds. Furthermore, functional predictions are made for each group based on ecological and mechanical considerations. The groups were tested for deviation from the norm for all birds for three indices of size- and leg-length-independent measures of the bones and for a size-independent-index of leg length. Several of the groups deviate significantly from the norm for one or more of the indices used, suggesting habitat-related adaptations in the leg morphology of birds. The results indicate that stability is an important factor affecting the leg morphology of primarily long-legged birds. The femur seems to be more important than previously thought because several of the groups have high femur indices, suggesting a positive selection pressure on this bone. On a general basis, the results suggest that the effect of leg length should be taken into consideration when discussing adaptations of mass-independent lengths of the long bones of the legs of birds.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79, 461–484.     

Larval morphology of Panorpodes kuandianensis (Insecta,Mecoptera, Panorpodidae) and its evolutionary implications

Larval characters play a significant role in evolutionary and systematic studies of holometabolous insects. However, Panorpodidae, a derived family of Mecoptera, are largely unknown in their immature stages to date. Here, the first instar larva of the short-faced scorpionfly Panorpodes kuandianensis Zhong, Zhang & Hua, 2011 is described and illustrated using light and scanning electron microscopy. The larva of Panorpodes is remarkable for the absence of compound eyes on the head and the presence of seven small unpaired proleg-like processes along the midventral line on abdominal segments II–VIII. The homology of these unpaired appendage-like processes, their ecological adaptation, and the evolutionary implications of some larval characters of Panorpodidae are discussed.