Lars Krogmann

Phylogenetic relationships among superfamilies of Hymenoptera

The first comprehensive analysis of higher‐level phylogeny of the order Hymenoptera is presented. The analysis includes representatives of all extant superfamilies, scored for 392 morphological characters, and sequence data for four loci (18S, 28S, COI and EF‐1α). Including three outgroup taxa, 111 terminals were analyzed. Relationships within symphytans (sawflies) and Apocrita are mostly resolved. Well supported relationships include: Xyeloidea is monophyletic, Cephoidea is the sister group of Siricoidea + [Xiphydrioidea + (Orussoidea + Apocrita)]; Anaxyelidae is included in the Siricoidea, and together they are the sister group of Xiphydrioidea + (Orussoidea + Apocrita); Orussoidea is the sister group of Apocrita, Apocrita is monophyletic; Evanioidea is monophyletic; Aculeata is the sister group of Evanioidea; Proctotrupomorpha is monophyletic; Ichneumonoidea is the sister group of Proctotrupomorpha; Platygastroidea is sister group to Cynipoidea, and together they are sister group to the remaining Proctotrupomorpha; Proctotrupoidea s. str. is monophyletic; Mymarommatoidea is the sister group of Chalcidoidea; Mymarommatoidea + Chalcidoidea + Diaprioidea is monophyletic. Weakly supported relationships include: Stephanoidea is the sister group of the remaining Apocrita; Diaprioidea is monophyletic; Ceraphronoidea is the sister group of Megalyroidea, which together form the sister group of [Trigonaloidea (Aculeata + Evanioidea)]. Aside from paraphyly of Vespoidea within Aculeata, all currently recognized superfamilies are supported as monophyletic. The diapriid subfamily Ismarinae is raised to family status, Ismaridae stat. nov.

New fossil insect order Permopsocida elucidates major radiation and evolution of suction feeding in hemimetabolous insects (Hexapoda: Acercaria)

With nearly 100,000 species, the Acercaria (lice, plant lices, thrips, bugs) including number of economically important species is one of the most successful insect lineages. However, its phylogeny and evolution of mouthparts among other issues remain debatable. Here new methods of preparation permitted the comprehensive anatomical description of insect inclusions from mid-Cretaceous Burmese amber in astonishing detail. These “missing links” fossils, attributed to a new order Permopsocida, provide crucial evidence for reconstructing the phylogenetic relationships in the Acercaria, supporting its monophyly, and questioning the position of Psocodea as sister group of holometabolans in the most recent phylogenomic study. Permopsocida resolves as sister group of Thripida + Hemiptera and represents an evolutionary link documenting the transition from chewing to piercing mouthparts in relation to suction feeding. Identification of gut contents as angiosperm pollen documents an ecological role of Permopsocida as early pollen feeders with relatively unspecialized mouthparts. This group existed for 185 million years, but has never been diverse and was superseded by new pollenivorous pollinators during the Cretaceous co-evolution of insects and flowers. The key innovation of suction feeding with piercing mouthparts is identified as main event that triggered the huge post-Carboniferous radiation of hemipterans, and facilitated the spreading of pathogenic vectors.

Phylogenetic implications of the mesosomal skeleton in Chalcidoidea (Hymenoptera, Apocrita) – tree searches in a jungle of homoplasy

Results from a comparative anatomical study of the mesosomal skeleton of Chalcidoidea are presented. External and internal features are described and illustrated for 39 chalcidoid taxa, representing 16 families and 29 subfamilies. This is the most comprehensive morphological study ever conducted for the superfamily. The mesosoma was dissected, macerated and investigated using scanning electron microscopy. The mesothorax and metathorax contributed most of the phylogenetically relevant information. The metafurca is highly variable within Chalcidoidea but seems to be relatively constant at the subfamily level. One hundred and fifty-four morphological characters were scored and analysed cladistically. Outgroup species were chosen from six apocritan superfamilies: Stephanoidea, Ceraphronoidea, Cynipoidea, Platygastroidea, Proctotrupoidea and Mymarommatoidea. Some previously suggested chalcidoid relationships were retrieved: (1) Pteromalidae: Pteromalinae + Miscogasterinae + Panstenoninae; (2) Perilampidae + Eucharitidae; (3) Chalcididae + Leucospidae + Eurytomidae; (4) Eulophidae: Eulophinae + Tetrastichinae + Entedoninae; and (5) Eupelmidae + Encyrtidae. Mymarommatoidea renders Chalcidoidea paraphyletic in our analyses; however, the taxon sample is too restricted to provide a robust hypothesis. Three previously unreported putative autapomorphies of Chalcidoidea were revealed: (1) presence of an exposed, triangular or diamond-shaped prosternum; (2) presence of a percurrent mesopleural sulcus anteriorly terminating in the acropleuron; and (3) presence of paired metapectal plates lateral to the metafurca.

Does early learning drive ecological divergence during speciation processes in parasitoid wasps

Central to the concept of ecological speciation is the evolution of ecotypes, i.e. groups of individuals occupying different ecological niches. However, the mechanisms behind the first step of separation, the switch of individuals into new niches, are unclear. One long-standing hypothesis, which was proposed for insects but never tested, is that early learning causes new ecological preferences, leading to a switch into a new niche within one generation. Here, we show that a host switch occurred within a parasitoid wasp, which is associated with the ability for early learning and the splitting into separate lineages during speciation. Lariophagus distinguendus consists of two genetically distinct lineages, most likely representing different species. One attacks drugstore beetle larvae (Stegobium paniceum (L.)), which were probably the ancestral host of both lineages. The drugstore beetle lineage has an innate host preference that cannot be altered by experience. In contrast, the second lineage is found on Sitophilus weevils as hosts and changes its preference by early learning. We conclude that a host switch has occurred in the ancestor of the second lineage, which must have been enabled by early learning. Because early learning is widespread in insects, it might have facilitated ecological divergence and associated speciation in this hyperdiverse group.

Biodiversity into your hands - A call for a virtual global natural history 'metacollection'

BackgroundMany scientific disciplines rely on correct taxon delineations and identifications. So does a great part of the general public as well as decision makers. Researchers, students and enthusiastic amateurs often feel frustrated because information about species remains scattered, difficult to access, or difficult to decipher. Together, this affects almost anyone who wishes to identify species or verify identifications. Many remedies have been proposed, but we argue that the role of natural history collections remains insufficiently appreciated. We suggest using state-of-the-art mass imaging technology and to join forces to create a global natural history metacollection on the internet, providing access to the morphology of tens of millions of specimens and making them available for automated digital image analysis.DiscussionRobotic high-resolution imaging technology and fast (high performance) computer-based image stitching make it now feasible to digitize entire collection drawers typically used for arthropod collections, or trays or containers used for other objects. Resolutions of 500 megapixels and much higher are already utilized to capture the contents of 40x50 cm collection drawers, providing amazing detail of specimens. Flanked by metadata entry, this helps to create access to tens of thousands of specimens in days. By setting priorities and combining the holdings of the most comprehensive collections for certain taxa, drawer digitizing offers the unique opportunity to create a global, virtual metacollection.The taxonomic and geographic coverage of such a collection could never be achieved by a single institution or individual. We argue that by joining forces, many new impulses will emerge for systematic biology, related fields and understanding of biodiversity in general.Digitizing drawers containing unidentified, little-curated specimens is a contribution towards the beginning of a new era of online curation. It also will help taxonomists and curators to discover and process the millions of “gems” of undescribed species hidden in museum accessions.SummaryOur proposal suggests creating virtual, high-resolution image resources that will, for the first time in history, provide access for expert scientists as well as students and the general public to the enormous wealth of the world’s natural history collections. We foresee that this will contribute to a better understanding, appreciation and increased use of biodiversity resources and the natural history collections serving this cause.

How to tackle the molecular species inventory for an industrialized nation-lessons from the first phase of the German Barcode of Life initiative GBOL (2012-2015).

Biodiversity loss is mainly driven by human activity. While concern grows over the fate of hot spots of biodiversity, contemporary species losses still prevail in industrialized nations. Therefore, strategies were formulated to halt or reverse the loss, driven by evidence for its value for ecosystem services. Maintenance of the latter through conservation depends on correctly identified species. To this aim, the German Federal Ministry of Education and Research is funding the GBOL project, a consortium of natural history collections, botanic gardens, and universities working on a barcode reference database for the countrys fauna and flora. Several noticeable findings could be useful for future campaigns: (i) validating taxon lists to serve as a taxonomic backbone is time-consuming, but without alternative; (ii) offering financial incentives to taxonomic experts, often citizen scientists, is indispensable; (iii) completion of the libraries for widespread species enables analyses of environmental samples, but the process may not hold pace with technological advancements; (iv) discoveries of new species are among the best stories for the media; (v) a commitment to common data standards and repositories is needed, as well as transboundary cooperation between nations; (vi) after validation, all data should be published online via the BOLD to make them searchable for external users and to allow cross-checking with data from other countries.

Cretevania bechlyi sp. nov., from Cretaceous Burmese amber (Hymenoptera: Evaniidae)

The fossil evaniid wasp Cretevania bechlyi sp. nov., is described based on a well preserved female specimen from Cretaceous Burmese amber. The new species is placed in the genus Cretevania Rasnitsyn, 1975 based on the elongation of the mid and hind trochantellus, the fore wing venation (e.g. first marginal cell triangular and broad, 2m-cu absent, second sub-marginal cell separated from first discal cell), the shape of the petiole (subcylindrical with distal extension) and other distinct morphological features. Cretevania bechlyi sp. nov. differs from all previously described species in having just 10 flagellomeres (11 in other members of the genus) and in the presence of notauli (absent in other species). The new species represents the first species of Cretevania from Burmese amber and significantly expands the known morphological diversity of Mesozoic Evaniidae.

Lower Cretaceous origin of long-distance mate finding behaviour in Hymenoptera (Insecta)

Flabellate antennae have evolved numerous times in males of several insect groups where they carry olfactory receptor neurons enabling the individuals to locate females over long distances. In Recent Hymenoptera, one of the largest species radiations among the Animalia, flabellate antennae are relatively uncommon. Here we describe Atefia rasnitsyni gen. et sp. nov. from the Lower Cretaceous Crato Formation of Brazil, representing the first fossil hymenopteran with a biflabellate antenna. The fossil indicates that long-distance mate finding behaviour in Hymenoptera was already well established in the Lower Cretaceous. The new fossil is taxonomically challenging and shares morphological characters with Recent Pergidae and Diprionidae. However, it lacks definite diagnostic features of either family and is here left unplaced as an early lineage of Tenthredinoidea. The occurrence and structural diversity of flabellate antennae in Recent Tenthredinoidea is discussed in a phylogenetic framework. http://zoobank.org/urn:lsid:zoobank.org:pub:D69866FA-831C-4B72-8E18-1CFA5B87D3DA

Insect Evolution in an Amberiferous and Stone Alphabet

This is an edited book with original scientific papers of the results of the 6th International Congress on Fossil Insects, Arthropods and Amber (FossilX3) held in Byblos, Lebanon in April, 2013. In the tradition of previous congresses, researchers from around the world gathered to discuss the latest developments and to build new co-operative endeavours. Recognizing that the future of our science is one of interdisciplinary collaboration, these meetings steadily grow in importance, and proceedings such as this reveal the latest hypotheses and conclusions, while inspiring others toward newer and greater goals.

Transcriptome sequence-based phylogeny of chalcidoid wasps (Hymenoptera: Chalcidoidea) reveals a history of rapid radiations, convergence, and evolutionary success

Chalcidoidea are a megadiverse group of mostly parasitoid wasps of major ecological and economical importance that are omnipresent in almost all extant terrestrial habitats. The timing and pattern of chalcidoid diversification is so far poorly understood and has left many important questions on the evolutionary history of Chalcidoidea unanswered. In this study, we infer the early divergence events within Chalcidoidea and address the question of whether or not ancestral chalcidoids were small egg parasitoids. We also trace the evolution of some key traits: jumping ability, development of enlarged hind femora, and associations with figs. Our phylogenetic inference is based on the analysis of 3,239 single-copy genes across 48 chalcidoid wasps and outgroups representatives. We applied an innovative a posteriori evaluation approach to molecular clock-dating based on nine carefully validated fossils, resulting in the first molecular clock-based estimation of deep Chalcidoidea divergence times. Our results suggest a late Jurassic origin of Chalcidoidea, with a first divergence of morphologically and biologically distinct groups in the early to mid Cretaceous, between 129 and 81 million years ago (mya). Diversification of most extant lineages happened rapidly after the Cretaceous in the early Paleogene, between 75 and 53 mya. The inferred Chalcidoidea tree suggests a transition from ancestral minute egg parasitoids to larger-bodied parasitoids of other host stages during the early history of chalcidoid evolution. The ability to jump evolved independently at least three times, namely in Eupelmidae, Encyrtidae, and Tanaostigmatidae. Furthermore, the large-bodied strongly sclerotized species with enlarged hind femora in Chalcididae and Leucospidae are not closely related. Finally, the close association of some chalcidoid wasps with figs, either as pollinators, or as inquilines/gallers or as parasitoids, likely evolved at least twice independently: in the Eocene, giving rise to fig pollinators, and in the Oligocene or Miocene, resulting in non-pollinating fig-wasps, including gallers and parasitoids. The origins of very speciose lineages (e.g., Mymaridae, Eulophidae, Pteromalinae) are evenly spread across the period of chalcidoid evolution from early Cretaceous to the late Eocene. Several shifts in biology and morphology (e.g., in host exploitation, body shape and size, life history), each followed by rapid radiations, have likely enabled the evolutionary success of Chalcidoidea.