Phylogenetic analysis of phenotypic characters of Tunicata supports basal Appendicularia and monophyletic Ascidiacea.

With approximately 3000 marine species, Tunicata represents the most disparate subtaxon of Chordata. Molecular phylogenetic studies support Tunicata as sister taxon to Craniota, rendering it pivotal to understanding craniate evolution. Although successively more molecular data have become available to resolve internal tunicate phylogenetic relationships, phenotypic data have not been utilized consistently. Herein these shortcomings are addressed by cladistically analyzing 117 phenotypic characters for 49 tunicate species comprising all higher tunicate taxa, and five craniate and cephalochordate outgroup species. In addition, a combined analysis of the phenotypic characters with 18S rDNA-sequence data is performed in 32 OTUs. The analysis of the combined data is congruent with published molecular analyses. Successively up-weighting phenotypic characters indicates that phenotypic data contribute disproportionally more to the resulting phylogenetic hypothesis. The strict consensus tree from the analysis of the phenotypic characters as well as the single most parsimonious tree found in the analysis of the combined dataset recover monophyletic Appendicularia as sister taxon to the remaining tunicate taxa. Thus, both datasets support the hypothesis that the last common ancestor of Tunicata was free-living and that ascidian sessility is a derived trait within Tunicata. "Thaliacea" is found to be paraphyletic with Pyrosomatida as sister taxon to monophyletic Ascidiacea and the relationship between Doliolida and Salpida is unresolved in the analysis of morphological characters; however, the analysis of the combined data reconstructs Thaliacea as monophyletic nested within paraphyletic "Ascidiacea". Therefore, both datasets differ in the interpretation of the evolution of the complex holoplanktonic life history of thaliacean taxa. According to the phenotypic data, this evolution occurred in the plankton, whereas from the combined dataset a secondary transition into the plankton from a sessile ascidian is inferred. Besides these major differences, both analyses are in accord on many phylogenetic groupings, although both phylogenetic reconstructions invoke a high degree of homoplasy. In conclusion, this study represents the first serious attempt to utilize the potential phylogenetic information present in phenotypic characters to elucidate the inter-relationships of this diverse marine taxon in a consistent cladistic framework.

The thorax of the cave cricket Troglophilus neglectus: anatomical adaptations in an ancient wingless insect lineage (Orthoptera: Rhaphidophoridae).

BACKGROUND: Secondary winglessness is a common phenomenon found among neopteran insects. With an estimated age of at least 140 million years, the cave crickets (Rhaphidophoridae) form the oldest exclusively wingless lineage within the long-horned grasshoppers (Ensifera). With respect to their morphology, cave crickets are generally considered to represent a `primitive' group of Ensifera, for which no apomorphic character has been reported so far. RESULTS: We present the first detailed investigation and description of the thoracic skeletal and muscular anatomy of the East Mediterranean cave cricket Troglophilus neglectus (Ensifera: Rhaphidophoridae). T. neglectus possesses sternopleural muscles that are not yet reported from other neopteran insects. Cave crickets in general exhibit some unique features with respect to their thoracic skeletal anatomy: an externally reduced prospinasternum, a narrow median sclerite situated between the meso- and metathorax, a star-shaped prospina, and a triramous metafurca. The thoracic muscle equipment of T. neglectus compared to that of the bush cricket Conocephalus maculatus (Ensifera: Tettigoniidae) and the house cricket Acheta domesticus (Ensifera: Gryllidae) reveals a number of potentially synapomorphic characters between these lineages. CONCLUSIONS: Based on the observed morphology we favor a closer relationship of Rhaphidophoridae to Tettigoniidae rather than to Gryllidae. In addition, the comparison of the thoracic morphology of T. neglectus to that of other wingless Polyneoptera allows reliable conclusions about anatomical adaptations correlated with secondary winglessness. The anatomy in apterous Ensifera, viz. the reduction of discrete direct and indirect flight muscles as well as the strengthening of specific leg muscles, largely resembles the condition found in wingless stick insects (Euphasmatodea), but is strikingly different from that of other related wingless insects, e.g. heel walkers (Mantophasmatodea), ice crawlers (Grylloblattodea), and certain grasshoppers (Caelifera). The composition of direct flight muscles largely follows similar patterns in winged respectively wingless species within major polyneopteran lineages, but it is highly heterogeneous between those lineages.

The thoracic morphology of the wingless dune cricket Comicus calcaris (Orthoptera: Schizodactylidae): Novel apomorphic characters for the group and adaptations to sand desert environments.

Schizodactylidae, splay-footed or dune crickets, represents a distinct lineage among the highly diverse orthopteran subgroup Ensifera (crickets, katydids and allies). Only two extant genera belong to the Schizodactylidae: the winged Eurasian genus Schizodactylus, whose ecology and morphology is well documented, and the wingless South African Comicus, for which hardly any studies providing morphological descriptions have been conducted since its taxonomic description in 1888. Based on the first in-depth study of the skeletomuscular system of the thorax of Comicus calcaris Irish 1986, we provide information on some unique characteristics of this character complex in Schizodactylidae. They include a rigid connection of prospinasternite and mesosternum, a T-shaped mesospina, and a fused meso- and metasternum. Although Schizodactylidae is mainly characterized by group-specific anatomical traits of the thorax, its bifurcated profuca supports a closer relationship to the tettigonioid ensiferans, like katydids, wetas, and hump-winged crickets. Some specific features of the thoracic musculature of Comicus seem to be correlated to the skeletal morphology, e.g., due to the rigid connection of the tergites and pleurites in the pterothorax not a single direct flight muscle is developed. We show that many of the thoracic adaptations in these insects are directly related to their psammophilous way of life. These include a characteristic setation of thoracic sclerites that prevent sand grains from intrusion into vulnerable membranous areas, the striking decrease in size of the thoracic spiracles that reduces the respirational water loss, and a general trend towards a fusion of sclerites in the thorax.

Extreme convergence in egg-laying strategy across insect orders.

The eggs of stick and leaf insects (Phasmatodea) bear strong resemblance to plant seeds and are commonly dispersed by females dropping them to the litter. Here we report a novel egg-deposition mode for Phasmatodea performed by an undescribed Vietnamese species of the enigmatic subfamily Korinninae that produces a complex egg case (ootheca), containing numerous eggs in a highly ordered arrangement. This novel egg-deposition mode is most reminiscent of egg cases produced by members of unrelated insect orders, e.g. by praying mantises (Mantodea) and tortoise beetles (Coleoptera: Cassidinae). Ootheca production constitutes a striking convergence and major transition in reproductive strategy among stick insects, viz. a shift from dispersal of individual eggs to elaborate egg concentration. Adaptive advantages of ootheca formation on arboreal substrate are likely related to protection against parasitoids and desiccation and to allocation of specific host plants. Our phylogenetic analysis of nuclear (28S, H3) and mitochondrial (COI, COII) genes recovered Korinninae as a subordinate taxon among the species-rich Necrosciinae with Asceles as sister taxon, thus suggesting that placement of single eggs on leaves by host plant specialists might be the evolutionary precursor of ootheca formation within stick insects.