Papers by Lars Vilhelmsen
Figure 2. Pro- and mesothorax of Trigonaloidea. A, Taeniogonalos gundlachii, mesopectus, posterio... more Figure 2. Pro- and mesothorax of Trigonaloidea. A, Taeniogonalos gundlachii, mesopectus, posterior view; B, T. gundlachii, mesopleuron, median view; C, Orthogonalys pulchella, pro- and mesothorax, median view; D, T. gundlachii, propleuron and procoxa, median view (anterior to the left); E, T. gundlachii, mesoscutum, anterior view. Abbreviations in Appendices 2 and 3.
Zootaxa, Oct 16, 2014
Orussus melanosoma Lee & Wei, sp. nov. from South Korea is described and illustrated. Phylogeneti... more Orussus melanosoma Lee & Wei, sp. nov. from South Korea is described and illustrated. Phylogenetic analyses place the new species basally in Orussus, together with other species from the Far East. A key to species of Orussidae from the eastern Palaearctic is provided.
Systematic Entomology, Nov 5, 2021
ABSTRACTWe present the first phylogenetic hypothesis for the cosmopolitan family Argidae based on... more ABSTRACTWe present the first phylogenetic hypothesis for the cosmopolitan family Argidae based on both molecular and morphological data. Furthermore, we present a biogeographic scenario based on a dated phylogeny and interpret the evolutionary history of the family. Information from sequences of eight genes is analysed to reconstruct the phylogeny of the Argidae based on maximum likelihood and Bayesian inference. Total evidence Bayesian analyses are also conducted, combining the molecular dataset with data from adult and larval morphology. For the historical biogeographic reconstruction, divergence times are estimated using node dating with uncorrelated relaxed‐clock analysis, and ancestral biogeographical distributions are estimated applying a Dispersal Extinction Cladogenesis model and a Bayesian binary model. Argidae s.s. is retrieved as monophyletic in all analyses and the clade Zenargidae + (Argidae + Pergidae) is better supported when combining molecular and morphological characters, and when excluding the saturated third codon position in the molecular analysis. Within Argidae, two large clades corresponding to the subfamilies Arginae and Sterictiphorinae sensu Benson are retrieved as monophyletic. The ancestral distribution of Arginae and Sterictiphorinae is estimated to be the Australian and Neotropical regions. Divergence of Argidae‐Pergidae and Arginae‐Sterictiphorinae is estimated to occur in the middle‐upper Jurassic before or during the east‐west Gondwana breakup. Diversification of Argidae may be associated with the radiation of angiosperms in the Early Cretaceous, especially for the Neotropical Sterictiphorinae after the separation of South America and Africa. Arginae were probably introduced to the northern hemisphere by dispersal to Africa and/or India and subsequent continental collision with Eurasia in the Cenozoic. The occurrence of Sterictiphorinae in the northern hemisphere is more difficult to explain. Maternal care and brood guarding behaviour evolved independently in Argidae and Pergidae, with a single origin in a subclade (Dielocerini) of Sterictiphorinae.
Arthropod Structure & Development, Oct 1, 2003
The structure of the ovipositor sheaths is examined in a number of parasitic wasps as well as a f... more The structure of the ovipositor sheaths is examined in a number of parasitic wasps as well as a few non-parasitic taxa. Parasitic wasps in the families Aulacidae, Braconidae, Ichneumonidae, Megalyridae, and Stephanidae with a very elongate, external ovipositor have the lateral wall of the sheath finely transversely subdivided by narrow furrows. This makes the sheath highly flexible, allowing it to support the ovipositor proper during ovipositing. The taxa having such a flexible sheath all drill into wood to lay their eggs. Support at the tip of the ovipositor is crucial in the initial stages of oviposition. The flexible ovipositor sheath is possibly a very ancient trait, arising prior to the radiation of extant Apocrita. This is corroborated by their presence in fossil Hymenoptera from the Upper Jurassic. The occurrence of short and/or concealed ovipositors with rigid sheaths in many extant apocritan taxa must therefore be assumed to be the result of multiple reversals.
Labriocimbex sinicus Yan & Wei gen. et sp. nov. of Cimbicidae is described. The new genus is simi... more Labriocimbex sinicus Yan & Wei gen. et sp. nov. of Cimbicidae is described. The new genus is similar to Praia Andre and Trichiosoma Leach. A key to extant Holarctic genera of Cimbicinae is provided. To identify the phylogenetic placement of Cimbicidae, the mitochondrial genome of L. sinicus was annotated and characterized using highthroughput sequencing data. The complete mitochondrial genome of L. sinicus was obtained with a length of 15,405 bp (GenBank: MH136623 ; SRA: SRR8270383) and a typical set of 37 genes (22 tRNAs, 13 PCGs, and two rRNAs). The results demonstrated that all PCGs were initiated by ATN codon, and ended with TAA or T stop codons. The study reveals that all tRNA genes have a typical cloverleaf secondary structure, except for trnS1. Remarkably, the secondary structures of the rrnS and rrnL of L. sinicus were much different from those of Corynis lateralis. Phylogenetic analyses verified the monophyly and positions of the three Cimbicidae species within the superfamily Tenthredinoidea and demonstrated a relationship as (Tenthredinidae + Cimbicidae) + (Argidae + Pergidae) with strong nodal supports. Furthermore, we found that the generic relationships of Cimbicidae revealed by the phylogenetic analyses based on COI genes agree quite closely with the systematic arrangement of the genera based on the morphological characters. Phylogenetic tree based on two methods shows that L. sinicus is the sister group of Praia with high support values. We suggest that Labriocimbex belongs to the tribe Trichiosomini of Cimbicinae based on adult morphology and molecular data. Besides, we suggest to promote the subgenus Asitrichiosoma to be a valid genus.
European journal of taxonomy, May 25, 2021
Many early taxonomic works on North American bees were published by Europeans using specimens col... more Many early taxonomic works on North American bees were published by Europeans using specimens collected in the New World, some with type locations so imprecise that uncertainty on the nomenclatural status remains to this day. Two examples come from Fabricius (1745-1808) who described Andrena virescens Fabricius, 1775 and Apis viridula Fabricius, 1793 from "America" and "Boreal America", respectively. The former species of Agapostemon Guérin-Méneville, 1844 occurs across most of the United States and southern Canada, the latter presumed an endemic to Cuba. The type materials of these two taxa have never been compared to each other, though a morphology-based phylogenetic analysis placed both in distinct species groups. Here we synonymize Apis viridula under Ag. virescens, thereby making Ag. femoralis (Guérin-Méneville, 1844) available as the name for the Cuban species. A lectotype for Ag. femoralis (the type species for the genus Agapostemon) is hereby designated to stabilize this taxonomy. We also synonymize Ag. obscuratus Cresson, 1869 under Ag. femoralis, suggesting that it represents a dark colour polymorphism. As Ag. cubensis Roberts, 1972 is a junior secondary homonym of Ag. cubensis (Spinola, 1851), we offer Ag. robertsi as a replacement name for the former.
Insect Systematics & Evolution, 2010
Th ree new species of Ophrynon Middlekauff , 1983, O. dominiqueae , O. martini and O. patricki , ... more Th ree new species of Ophrynon Middlekauff , 1983, O. dominiqueae , O. martini and O. patricki , are added to the only species known previously, O. levigatus Middlekauff , 1983. All species are described, illustrated and keyed. Cladistic analyses under diff erent weighting conditions retrieved Ophrynon as monophyletic, but the topologies produced for the higher ophrynopine taxa are quite variable. Th e possible closest relatives of Ophrynon are Argentophrynopus Vilhelmsen & D.R. Smith, 2002 and Kulcania Benson, 1935. Th e internal phylogeny of Ophrynon is O. martini + (O. levigatus + (O. dominiqueae + O. patricki)). Th e distribution of Ophrynon is restricted to central and southern California. Supposedly, the species belong to the californo-eremial distribution type and are the result of a comparatively recent speciation process.
Journal of Natural History, Nov 14, 2013
ABSTRACT The variation in ovipositor sculpture and additional features of the ovipositor apparatu... more ABSTRACT The variation in ovipositor sculpture and additional features of the ovipositor apparatus in Tenthredinoidea is explored. Two outgroups and 85 species of tenthredinoids were examined, the emphasis of the taxon sampling being on Nematinae. Twenty-four morphological characters where scored and the data set was analysed under different weighting conditions. The topologies derived from analysing the character set in TNT are poorly resolved and highly unstable when analytical settings are changed. Only a few clades, the Tenthredinoidea s. str., the Diprionidae, the Cimbicidae and the Argidae + Pergidae are consistently retrieved. The functional implications of the variation in ovipositor morphology within Tenthredinidae is discussed in relation to the qualities of the host plant, e.g. the hardness and the depth of penetration in plant tissue.
Invertebrate Systematics, 2015
Zoologica Scripta, 1999
The occipital region in representatives of all the`symphytan' families and a few apocritans is in... more The occipital region in representatives of all the`symphytan' families and a few apocritans is investigated. A new character, the occurrence of occipital sulci, is described. The absence of occipital sulci and corresponding internal ridges is a putative synapomorphy for the Xiphydriidae, Orussoidea, and Apocrita. Independent loss has occurred in the cimbicid genus Zaraea. The evolution of sclerotizations between the occipital and oral foramina in basal Hymenoptera is discussed. A postoccipital bridge, an internal structure formed by the fusion of the apodemes for the profurco-postoccipital muscles, forming a ventral connection between the tentorial arms, is present in Cimbex, Cephoidea, and Syntexis. An external sclerotization, the hypostomal bridge, is present in Corynis, Cephoidea, Siricoidea', Orussoidea, and Apocrita. In Syntexis, the postoccipital and hypostomal bridges are continuous. It is difficult to decide whether a hypostomal bridge is also present in the Pamphilioidea, or the external sclerotization in this superfamily has evolved independently. In the Siricidae, Orussidae, and some Apocrita, a postgenal bridge largely replaces the hypostomal bridge. The postgenal bridge has evolved at least twice independently in the Hymenoptera.
Zoologica Scripta, Oct 1, 2000
The skeleto-musculature of the ovipositor apparatus and the external sculpture of the 1st and 2nd... more The skeleto-musculature of the ovipositor apparatus and the external sculpture of the 1st and 2nd valvulae was studied in representatives from all 'symphytan' families. Nineteen informative characters were coded and scored. The distribution of character states are discussed with reference to recent cladistic treatments of the Hymenoptera. Putative autapomorphies of the Hymenoptera are the presence of cordate apodemes on T9 and basal articulations and associated musculature between the 2nd valvifers and the 2nd valvulae. It is a ground plan feature of the order to have the gonocoxites of abdominal segment 8 fused with the gonangula. The configuration of the musculature of the ovipositor apparatus did not display much variation among the taxa examined, except within the Pamphilioidea. There is considerable variation in the external ovipositor sculpture within the Tenthredinoidea. Putative synapomorphies for the tenthredinoid families except the Blasticotomidae are the presence of alternating strongly and weakly sclerotized zones on the first and/or second valvulae and the presence of serrulae on the sawteeth. The presence of transverse rows of large ctenidia on the 1st valvulae is an autapomorphy of the Diprionidae. Fusion of the 2nd valvifers and the 3rd valvulae is a synapomorphy for the Argidae + Pergidae. The ovipositor apparatus of the Pamphilioidea is highly derived, putative autapomorphies being the close association between T9 and the first valvifers, the reduction of the distal parts of the 1st valvulae, and the fusion of the 2nd valvulae for their entire length. The changes in the ovipositor apparatus of Pamphilioidea are associated with a decrease in the amount of work it has to perform during ovipositing, as the eggs are placed predominantly externally on the substrate. The ovipositor apparatus of the 'Siricoidea' is enlarged and modified for ovipositing into wood. Putative synapomorphies of the 'Siricoidea' + Orussidae + Apocrita are the presence of sawteeth only distally on the ovipositor and elongation of the cordate apodemes of T9.
Insect Systematics & Evolution, 2002
The genera Argentophrynopus Vilhelmsen & Smith, gen. n., Guiglia Benson, Kulcania Benson,... more The genera Argentophrynopus Vilhelmsen & Smith, gen. n., Guiglia Benson, Kulcania Benson, Ophrella Middlekauff, Ophrynon Middlekauff, Ophrynopus Konow, and Stirocorsia Konow of the wasp family Orussidae are revised. This group of genera of Orussidae occurs from southern United States to Chile and in Japan, southeastern Asia, and Australia. In total, 27 species are recognized, including Argentophrynopus enigmus Vilhelmsen & Smith, sp. n., A. gauldi Vilhelmsen & Smith, sp. n., Guiglia rubicunda Schmidt, sp. n., Ophrynopus carinatus Vilhelmsen & Smith, sp. n., and Ophrynopus hansoni Vilhelmsen & Smith, sp. n. Guiglia queenslandensis coronata Rayment is considered a new junior synonym of G. sericatus (Mocsáry); G. aureola Benson is considered a species inquirenda. Kulcania tomentosa (Middlekauff), comb. n., is transferred from Ophrynopus. Ophrynopus philippinensis Guiglia is considered a new junior synonym of Stirocorsia kohli Konow; Ophrynella rossi Yasumatsu, syn. n., and Oryssus trifasciatus Cameron, rev. stat., are both considered junior synonyms of Stirocorsia maculipennis (Smith). Keys and distribution maps of all recognized species are provided, as well as descriptions of all genera.
Zoologischer Anzeiger, 2000
Abstract The skeleto-musculature of the cervical region and prothorax has been studied in represe... more Abstract The skeleto-musculature of the cervical region and prothorax has been studied in representatives of all'symphytan'families. Thirty informative characters have been coded and scored. The distribution of character states are discussed with reference to recent cladistic ...
Insect Systematics & Evolution, 2002
The genera Argentophrynopus Vilhelmsen & Smith, gen. n., Guiglia Benson, Kulcania Benson,... more The genera Argentophrynopus Vilhelmsen & Smith, gen. n., Guiglia Benson, Kulcania Benson, Ophrella Middlekauff, Ophrynon Middlekauff, Ophrynopus Konow, and Stirocorsia Konow of the wasp family Orussidae are revised. This group of genera of Orussidae occurs from southern United States to Chile and in Japan, southeastern Asia, and Australia. In total, 27 species are recognized, including Argentophrynopus enigmus Vilhelmsen & Smith, sp. n., A. gauldi Vilhelmsen & Smith, sp. n., Guiglia rubicunda Schmidt, sp. n., Ophrynopus carinatus Vilhelmsen & Smith, sp. n., and Ophrynopus hansoni Vilhelmsen & Smith, sp. n. Guiglia queenslandensis coronata Rayment is considered a new junior synonym of G. sericatus (Mocsáry); G. aureola Benson is considered a species inquirenda. Kulcania tomentosa (Middlekauff), comb. n., is transferred from Ophrynopus. Ophrynopus philippinensis Guiglia is considered a new junior synonym of Stirocorsia kohli Konow; Ophrynella rossi Yasumatsu, syn. n., and Oryssus trifasciatus Cameron, rev. stat., are both considered junior synonyms of Stirocorsia maculipennis (Smith). Keys and distribution maps of all recognized species are provided, as well as descriptions of all genera.
Organisms Diversity & Evolution, Nov 29, 2007
Arthropod Structure & Development, 2011
We survey morphological features of larval and adult wasps that undergo their entire larval devel... more We survey morphological features of larval and adult wasps that undergo their entire larval development inside wood and interpret them in view of the lifestyle. The evolution of some of the characters is explored by mapping them on a recently published phylogeny of Hymenoptera. Based on this phylogeny, it is reasonable to assume that wood-living wasps evolved from a xylophagous/mycetophagous stage as displayed by woodwasps to a carnivorous/parasitoid lifestyle, preying on woodboring insect larvae. The latter mode of life is probably ancestral to the Apocrita which comprise the majority of the order; they share this lifestyle with their sister group, the Orussidae. However, most apocritan wasps have radiated into other habitats, the Orussidae and Stephanidae apparently being the only taxa that have retained the ancestral lifestyle of carnivorous wasps. Other apocritan lineages associated with wood (e.g., Aulacidae, Megalyridae, basal Cynipoidea and some Ichneumonoidea and Chalcidoidea) possibly entered this habitat secondarily and independently acquired morphological traits associated with it. The woody habitat was occupied by Hymenoptera during a crucial stage in their evolution where the transition from the phytophagous to carnivorous lifestyle took place. The anatomy of both larva and adults was extensively transformed in the process.
Systematic Biology, Aug 12, 2015
Directional evolution has played an important role in shaping the morphological, ecological, and ... more Directional evolution has played an important role in shaping the morphological, ecological, and molecular diversity of life. However, standard substitution models assume stationarity of the evolutionary process over the time scale examined, thus impeding the study of directionality. Here we explore a simple, nonstationary model of evolution for discrete data, which assumes that the state frequencies at the root differ from the equilibrium frequencies of the homogeneous evolutionary process along the rest of the tree (i.e., the process is nonstationary, nonreversible, but homogeneous). Within this framework, we develop a Bayesian approach for testing directional versus stationary evolution using a reversible-jump algorithm. Simulations show that when only data from extant taxa are available, the success in inferring directionality is strongly dependent on the evolutionary rate, the shape of the tree, the relative branch lengths, and the number of taxa. Given suitable evolutionary rates (0.1-0.5 expected substitutions between root and tips), accounting for directionality improves tree inference and often allows correct rooting of the tree without the use of an outgroup. As an empirical test, we apply our method to study directional evolution in hymenopteran morphology. We focus on three character systems: wing veins, muscles, and sclerites. We find strong support for a trend toward loss of wing veins and muscles, while stationarity cannot be ruled out for sclerites. Adding fossil and time information in a total-evidence dating approach, we show that accounting for directionality results in more precise estimates not only of the ancestral state at the root of the tree, but also of the divergence times. Our model relaxes the assumption of stationarity and reversibility by adding a minimum of additional parameters, and is thus well suited to studying the nature of the evolutionary process in data sets of limited size, such as morphology and ecology.
Zoologica Scripta, Apr 1, 1996
The skeletal and muscular morphology of the preoral cavity, including the labrum, hypopharynx and... more The skeletal and muscular morphology of the preoral cavity, including the labrum, hypopharynx and labium, was examined in the imago in representatives of all the 'symphytan'families as well as the apocritan families Stephanidae, Megalyridae and ...
Journal of Zoological Systematics and Evolutionary Research, Apr 27, 2009
Abstract A cladistic analysis of the lower Hymenoptera, including all the 'symphytan&... more Abstract A cladistic analysis of the lower Hymenoptera, including all the 'symphytan'families and the apocritan families Stephanidae, Megalyridae, Trigonalyidae, Ibaliidae, Vespidae and Gasteruptiidae, has been undertaken. A total of 98 characters were scored for 21 taxa ...
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Papers by Lars Vilhelmsen