DNA barcoding of Austrian snow scorpionflies (Mecoptera, Boreidae) reveals potential cryptic diversity in Boreus westwoodi.

BACKGROUND: Snow scorpionflies (genus Boreus) belong to a family of Mecoptera, Boreidae, that has been vastly neglected by entomological researchers due to their shift in seasonality to the winter months. Their activity during this time is regarded as a strategy for predator avoidance and regular sightings on snow fields suggest that this also facilitates dispersal. However, many aspects about snow scorpionflies, especially systematics, taxonomy, distribution of species, phylogenetics and phylogeography have remained fairly unexplored until today. In this study, we fill some of these gaps by generating a reference DNA barcode database for Austrian snow scorpionflies in the frame of the Austrian Barcode of Life initiative and by characterising morphological diversity in the study region. METHODS: Initial species assignment of all 67 specimens was based on male morphological characters previously reported to differ between Boreus species and, for females, the shape of the ovipositor. DNA barcoding of the mitochondrial cytochrome c oxidase subunit 1 (COI) gene was carried out for all 67 samples and served as a basis for BIN assignment, genetic distance calculations, as well as alternative species delimitation analyses (ABGD, GMYC, bGMYC, bPTP) and a statistical parsimony network to infer phylogenetic relationships among individual samples/sampling sites. RESULTS: Morphological investigations suggested the presence of both Boreus hyemalis and Boreus westwoodi in Austria. DNA barcoding also separated the two species, but resulted in several divergent clades, the paraphyly of B. westwoodi in Austria, and high levels of phylogeographic structure on a small geographic scale. Even though the different molecular species delimitation methods disagreed on the exact number of species, they unequivocally suggested the presence of more than the traditionally recognized two Boreus species in Austria, thus indicating potential cryptic species within the genus Boreus in general and especially in B. westwoodi.

Morphological adaptations to silk production by adult females in the pollen wasp genus Quartinia (Masarinae, Vespidae) - a keystone character for ground nesting in dry sand habitats.

While many pollen wasps nest in hard clayey soil or in rigid sand or use these kinds of substrates to build aerial earthen cells, all representatives of the genus Quartinia, in which nesting behavior has been studied so far, construct their nests in habitats with loose sand. The walls of the burrow are stabilized by a silky excretion that is applied on their inner surfaces during nest construction. The lining-behavior of Quartinia females is described. Representatives of the genus Quartinia and other pollen wasp taxa have been studied comparatively using histology, microcomputed tomography and scanning electron microscopy for identifying the structures correlated with lining-behavior. Thus, we can document a change in the morphology of the maxillary gland for females of the genus Quartinia, as well as the existence of a process of the galea which likely serves the production of the silky threads. The fact that these modifications are missing in males is corroborating their function in brood care. Two possible ways of silk thread production are discussed. The newly discovered structures are key adaptations of the genus Quartinia that enabled the inhabitation of new habitats and thus probably facilitated the species diversification of this genus.

Small, but oh my! Head morphology of adult Aleuropteryx spp. and effects of miniaturization (Insecta: Neuroptera: Coniopterygidae).

We present the first morphological study of the internal head structures of adults of the coniopterygid genus Aleuropteryx, which belong to the smallest known lacewings. The head is ventrally closed with a gula, which is unique in adult Neuroptera and otherwise developed in Megaloptera, the sister group of Neuroptera. The dorsal tentorial arms are directed posteriorly and fused, forming an arch that fulfills functions otherwise taken by the tentorial bridge. A newly found maxillary gland is present in both sexes. Several structural modifications correlated with miniaturization are recognized: a relative increase in the size of the brain, a reduction in the number of ommatidia and diameter of the facets, a countersunken cone-shaped ocular ridge, and a simplification of the tracheal system. The structure of the head differs strikingly from that of the previously studied species Coniopteryx pygmaea, indicating a greater variability in the family Coniopterygidae, which might be another effect of miniaturization.

Head anatomy of adult Coniopteryx pygmaea: Effects of miniaturization and the systematic position of Coniopterygidae (Insecta: Neuroptera).

External and internal head structures of adult Coniopteryx pygmaea Enderlein, 1906, one of the smallest known lacewings, are described in detail for the first time. Possible effects of miniaturization and two hypotheses on the phylogenetic position of Coniopterygidae are evaluated and compared with data from literature. Several convergent modifications in C. pygmaea and other miniaturized insect species are outlined, e.g., a relative increase in the size of the brain, simplification of the tracheal system with respect to the number of tracheae, and reduction of the number of ommatidia and diameter of the facets. Further, the ocular ridge is bell-shaped and countersunk into the head capsule. The cuticle is weakly sclerotized and equipped with wax glands which are unique in Neuroptera. The total number of muscles is not affected by miniaturization. The phylogenetic analysis yields Coniopterygidae as sistergroup to the dilarid clade based on one larval character, the shape of the stylets. The enforced basal position of Coniopterygidae is supported by one disputable synapomorphy of the remaining Neuroptera, the presence of paraglossae in adults.

Head anatomy of adult Sisyra terminalis (Insecta: Neuroptera: Sisyridae)--functional adaptations and phylogenetic implications.

The external and internal head anatomy of Sisyra terminalis is described in detail and compared with data from literature. A salivary pump consisting of a peculiar reservoir and a hitherto unknown muscle, M. ductus salivarii, is newly described for Neuroptera. The upward folded paraglossae form a secondary prolongation of the salivary system. These structures are discussed as functional adaptations for feeding on aphids and desiccated honeydew. In a phylogenetic analysis the basal position of the Sisyridae within Neuroptera is retrieved. The following new synapomorphies are postulated: (1) for Neuropterida, the presence of a M. submentomentalis and prepharyngeal ventral transverse muscles, and the absence of a M. submentopraementalis; (2) for Neuroptera and Sialidae, the presence of a mandibular gland; (3) for Neuroptera, the presence of four scapopedicellar muscles; (4) for Neuroptera exclusive Nevrorthidae and Sisyridae, the weakening of dorsal tentorial arms, the presence of a M. tentoriomandibularis medialis superior and the shifted origin of M. tentoriocardinalis.

The function and phylogenetic implications of the tentorium in adult Neuroptera (Insecta).

Despite several recent analyses on the phylogeny of Neuroptera some questions still remain to be answered. In the present analysis we address these questions by exploring a hitherto unexplored character complex: the tentorium, the internal cuticular support structure of the insect head. We described in detail the tentoria of representatives of all extant neuropteran families and the muscles originating on the tentorium using 3D microCT images and analyzed differences in combination with a large published matrix based on larval characters. We find that the tentorium and associated musculature are a source of phylogenetically informative characters. The addition of the tentorial characters to the larval matrix causes a basad shift of the Sisyridae and clearly supports a clade of all Neuroptera except Sisyridae and Nevrorthidae. A sister group relationship of Coniopterygidae and the dilarid clade is further corroborated. A general trend toward a reduction of the dorsal tentorial arms and the development of laminatentoria is observed. In addition to the phylogenetic analysis, a correlation among the feeding habits, the development of the maxillary muscles, and the laminatentoria is demonstrated.