Exemplar Species Research Articles

A method for designing complex biosecurity surveillance systems: detecting non‐indigenous species of invertebrates on Barrow Island

AbstractAimWe developed a new method to design objective, risk‐based surveillance systems for non‐indigenous species of invertebrates, vertebrates and plants, which might be introduced to a natural area through an industrial project; here, we provide the invertebrate case study. The method addresses issues common to complex surveillance design problems: a statistical standard (e.g. power); information gaps; multiple targets of unclear identity; a large surveillance area of heterogeneous risk of invasion; integrating multiple sources of surveillance data; optimizing for cost.LocationBarrow Island, Western Australia.MethodsWe mapped the surveillance area for risk to target surveillance activities. An expert group identified a set of exemplar species and identified and characterized a set of detection methods for each, such that all potential invaders would be detected. We devised multi‐element surveillance systems to detect each exemplar to the design power (0.8), then integrated them to a single system that was optimized for cost.ResultsThe surveillance system was deployed on the island to specification over 1 year, then reviewed for redesign in a second period.Main conclusionsThe new method provided practical, risk‐based surveillance system designs that met application requirements and overcame complex issues common to many surveillance applications. A review of experiences from surveillance in the first year led to practical improvements and design efficiencies.

Treatise on the Isoptera of the World

A comprehensive compendium on the taxonomy and biology of the 3106 living and fossil species of the worlds termites is presented, along with reviews of Isoptera morphology and evolution, identification keys, the history of termite systematics, and summary of the worlds 363 significant pest species. A complete bibliography is provided of nearly 5000 references covering virtually all aspects of termite taxonomy and biology through December 2011.The morphology of Isoptera is thoroughly reviewed and illustrated with original scanning electron micrographs and photomicrographs, covering the cuticular anatomy and those internal organs that are taxonomically and phylogenetically significant, including several new character systems. Terminology is presented for the systems of tibial spines and spurs so as to establish homologs. Keys are presented to the nine living families of termites, and the world subfamilies and genera of Archotermopsidae, Hodotermitidae, Kalotermitidae, Mastotermitidae, Rhinotermitidae, Serritermitidae, Stolotermitidae, and Stylotermitidae. A key to subfamilies of the Termitidae is included. A detailed morphological diagnosis for each family and subfamily is provided, along with images of exemplar species. The history of isopteran research in taxonomy, systematics, morphology, paleontology, and biology is reviewed from 1758 to the present, with emphasis on transformative workers such as Holmgren, Silvestri, Emerson, Roonwal, Noirot, and Sands. Evolution of the Isoptera is reviewed, including the diversity and natural history of genera and species in all Zoogeographic regions, major patterns in social biology, the phylogeny of Recent and fossil genera and families, and 135 million years of fossils preserved as compressions, mineralized replicas, and in amber. The definitive sister group to the Isoptera is the monogeneric family of wood roaches, Cryptocercidae (Cryptocercus), so the taxonomic ranks of the two groups are now Infraorder Isoptera and Infraorder Cryptocercoidea within Order Blattaria (roaches and termites).The compendium summarizes the taxonomic history, nomenclature, distribution, type locality, and repository, and all significant aspects of natural history and biology for each species of the world, exclusive of pest control and colony inquilines (termitophiles). The classification of Recent and fossil lower termites (all those exclusive of family Termitidae) used in the compendium is from Engel et al. (2009), which is based on morphology and largely congruent with molecular studies. Rhinotermitidae s.s. (exclusive of Stylotermitidae) may be paraphyletic with respect to Termitidae, although the six traditional subfamilies of the former are used here. A separate section summarizes the nomenclatural changes made in the compendium, including new synonymies, new combinations, status novus, lectotype selection etc. A detailed list is provided of museums and other institutional collections that house type specimens. An index is included. The Treatise is intended to provide an authoritative foundation for taxonomic work on the Isoptera, present and future.

Tests of the Link between Functional Innovation and Positive Selection at Phytochrome A: The Phylogenetic Distribution of Far-Red High-Irradiance Responses in Seedling Development

We surveyed the phylogenetic distribution of far-red high-irradiance responses (FR-HIRs), which are important for early development of seedlings in shaded habitats, in exemplar species across seed plants to test whether the responses are unique to angiosperms and may have helped early angiosperms colonize the understory of Mesozoic forests dominated by ferns and gymnosperms. We show that dark-grown seedlings in most seed plants adopt an etiolated morphology and that being green in the dark is a derived condition in some conifers and gnetophytes. We also show that FR-HIR-induced seedling de-etiolation, mediated by phytochrome, is widely distributed in seed plants. Together, these data show that despite the ability of gymnosperms to green in the dark, most of them do not do so but link seedling development to light signals, as do angiosperms. Our data refute an obvious link between the origin of FR-HIRs and the signal of positive selection at the PHYA locus early in the history of angiosperms.

Evolutionary morphology of the hemolymph vascular system in scorpions: A character analysis

Phylogenetically informative characters from the internal anatomy of scorpions were first reported more than 150 years ago, but the subject received little attention after the mid-1920s. Recent investigations, using traditional dissection, illustration and histological sectioning, microscopy, and innovative new methods for investigating complex soft tissue anatomy identified a new wealth of variation. Additionally, these investigations confirmed the phylogenetic significance of previously identified structures. Building on earlier work we present a more detailed description of the hemolymph vascular system (HVS) in scorpions, based on comparison of the vascular structures of the heart and the branching pattern of the prosomal arteries among 45 exemplar species representing most of the major scorpion lineages. Using corrosion-casting, MicroCT in combination with computer-aided 3D-reconstruction, and scanning electron microscopy, we conceptualize a series of phylogenetically informative characters for the anterior aorta system and characters of the heart and associated structures (e.g. arrangement of the ostia) of scorpions. Furthermore, we optimize the possible evolution of these new characters on a previous hypothesis of scorpion phylogeny, and discuss alternative character state transformations, their evolutionary consequences, and possible underlying evolutionary mechanisms acting on the HVS.

Phylogeny of mosquitoes of tribe Culicini (Diptera: Culicidae) based on morphological diversity

Harbach, R. E., Kitching, I. J., Culverwell, C. L., Dubois, J. & Linton, Y.‐M. (2012). Phylogeny of mosquitoes of tribe Culicini (Diptera: Culicidae) based on morphological diversity. —Zoologica Scripta, 41, 499–514.Relationships among taxa of the mosquito tribe Culicini are explored using 169 morphological characters from 86 exemplar species representing the four genera and 26 subgenera of Culicini, most species groups and subgroups of subgenus Culex and an outgroup of five species from five other tribes. We analysed the data set, with multistate characters treated as unordered, under implied weights with values of K ranging from 1 to 20, implemented by TNT. Each analysis, except K = 4, produced a single most parsimonious (fittest) cladogram (MPC). The topology of the ingroup was identical for K = 6–11, whereas the MPCs for K = 14–20 differed only in the position of a single species, which occupied the same position in the K = 16 and K = 6–11 topologies. The K = 9 and K = 16 trees were given further consideration. In both these cladograms, Lutzia is sister to a clade comprising genera Culex, Deinocerites and Galindomyia. The two topologies have 13 clades in common, but their arrangements differ primarily because of Culex (Culex) duttoni acting as a ‘rogue’ taxon. We evaluated the effect of removing this species from the K = 9 and K = 16 analyses and chose the refined K = 9 topology as the best hypothesis of relationships within Culicini. Genus Culex is not monophyletic because it includes Deinocerites and Galindomyia as derived members of the New World subgenera. With the exception of subgenera Culex, Eumelanomyia and Neoculex, there is strong support for the monophyly of all genera and subgenera. Subgenus Culex would be monophyletic were four other subgenera included and three other taxa (the Afrotropical Cx. duttoni, Neotropical Cx. apicinus and the Australasian Atriceps Group) excluded.

Phylogenetics and taxonomy of Ventrops - the largest genus of Afrotropical Rhinophoridae (Diptera)

Three new species of the Afrotropical genus Ventrops Crosskey, 1977 (V. aethiopicus, sp. nov., V. freidbergi, sp. nov. and V. stuckenbergi, sp. nov.) are described, and all seven known species are included in a morphology-based phylogenetic analysis both to provide a rigorous argumentation for the generic affiliation of the newly described species, and to provide a first explicit hypothesis of phylogenetic relationships between the included species. Also, by including exemplar species from several genera of the Rhinophoridae as outgroups, this analysis is the first explicitly phylogenetic definition of the genus Ventrops within a broader systematic context. The seven species of Ventrops are arranged in three species groups (milichioides-group, hannemariae-group and aethiopicus-group).

The evolution of the tape measure protein: units, duplications and losses.

BackgroundA large family of viruses that infect bacteria, called phages, is characterized by long tails used to inject DNA into their victims' cells. The tape measure protein got its name because the length of the corresponding gene is proportional to the length of the phage's tail: a fact shown by actually copying or splicing out parts of DNA in exemplar species. A natural question is whether there exist units for these tape measures, and if different tape measures have different units and lengths. Such units would allow us to retrace the evolution of tape measure proteins using their duplication/loss history. The vast number of sequenced phages genomes allows us to attack this problem with a comparative genomics approach.ResultsHere we describe a subset of phages whose tape measure proteins contain variable numbers of an 11 amino acids sequence repeat, aligned with sequence similarity, structural properties, and simple arithmetics. This subset provides a unique opportunity for the combinatorial study of phage evolution, without the added uncertainties of multiple alignments, which are trivial in this case, or of protein functions, that are well established. We give a heuristic that reconstructs the duplication history of these sequences, using divergent strains to discriminate between mutations that occurred before and after speciation, or lineage divergence. The heuristic is based on an efficient algorithm that gives an exhaustive enumeration of all possible parsimonious reconstructions of the duplication/speciation history of a single nucleotide. Finally, we present a method that allows, when possible, to discriminate between duplication and loss events.ConclusionsEstablishing the evolutionary history of viruses is difficult, in part due to extensive recombinations and gene transfers, and high mutation rates that often erase detectable similarity between homologous genes. In this paper, we introduce new tools to address this problem.

A comparative survey of proboscis morphology and associated structures in fruit-piercing, tear-feeding, and blood-feeding moths in Calpinae (Lepidoptera: Erebidae)

Functional feeding categories for adult species of Calpinae are described. Structures associated with the proboscis were examined using exemplar species in fruit-piercing, blood-feeding, and tear-feeding species using both light microscopy and SEM methods. At least three genera currently placed in Calpini, and several others in related groups lack specialized piercing structures. The proboscis of the tear-feeding species, Hemiceratoides hieroglyphica, is equipped with specialized cuticular hooks not yet observed in other tear-feeding species. Tearing hooks moveable by blood pressure are restricted to Calpini species, and little additional variation within this taxon exists, suggesting proboscis morphology may not be strongly correlated with feeding behavior (e.g., fruit piercing vs. blood feeding). A glossary of terms and character codings for proboscis structures is provided, and morphologies for all included calpine taxa are described. We discuss the taxonomic significance of proboscis morphology in Calpinae and the evolutionary implications of their associated feeding behaviors. This survey indicates morphology provides powerful prediction, but not proof of lepidopteran adult food habits.

Consolida and Aconitella are an annual clade of Delphinium (Ranunculaceae) that diversified in the Mediterranean basin and the Irano‐Turanian region

AbstractThe Mediterranean and Irano‐Turanian biogeographic regions are biodiversity hotspots for mesophytic and xerophytic species, including many Delphinieae. This phylogenetically poorly understood tribe of Ranunculaceae consists of Consolida and Aconitella, with together ca. 52 species, and Delphinium and Aconitum, each with ca. 300 species. To infer the phylogeny of Consolida and Aconitella, we analyzed nuclear and chloroplast DNA sequences from 39 of their species and subspecies (44 taxa) plus a set of 30 exemplar species of Delphinium and Aconitum. We used a Bayesian relaxed clock model to estimate divergence times and a maximum likelihood approach to reconstruct ancestral areas. Aconitella forms a clade embedded in Consolida, and the latter is embedded in Delphinium. Consolida s.l. (including Aconitella) comprises two clades in the Irano‐ Turanian region and three in the Mediterranean basin. The latter clades’ inferred crown ages of 5.1, 4.4, and 2.8 Ma suggests that the repeated drying‐up of the Mediterranean, concomitant with and following the Messinian salinity crisis (5.96–5.33 Ma), may have facilitated their westward expansion. While there is clear geographic structure towards the tips of the Consolida s.l. tree, a likely ancestral area could not be inferred. However, the initial diversification of Consolida s.l., which occurred ca. 17 Ma ago, falls in a period when the climate in the Anatolian region became more arid, which may have favoured the annual life cycle that characterizes all species in this clade. To achieve a classification of mutually monophyletic genera in Delphinieae may require transferring the species of Aconitella and Consolida into Delphinium.

Reproductive Biology, Family Conflict, and Size of Offspring in Marine Invertebrates

All organisms face two fundamental trade-offs in the allocation of energetic resources: one between many small versus a few large offspring, and the second between present and future reproduction. Nowhere are these trade-offs more apparent than in the vast range of variation in the sizes of eggs and offspring exhibited among species of marine invertebrates. It has become increasingly clear that, in many taxa of marine organisms, there is also substantial intraspecific variation in the size of eggs and hatchings. This variation has largely been attributed to adaptive maternal effects. In theory, however, the inevitable conflicts of interest that arise in families of sexually reproducing organisms over the optimal distribution of parental resources among siblings could also account for much of this variation in egg and offspring size. Here, we explore the potential impacts of family conflict on offspring traits by comparing the life histories of two exemplar species of marine organisms, the polychaete Boccardia proboscidea and the gastropod Solenosteira macrospira, emphasizing how differences in modes of fertilization and parental care might influence the phenotype and, consequently, the fitness of offspring.

Phylogeny and the evolution of nesting behaviour in the tribe Ageniellini (Insecta: Hymenoptera: Pompilidae)

The Pompilidae (spider hunting wasps) show a marked diversity in their hunting and reproductive ecology, but difficulties in phylogenetic taxonomy have hindered the elucidation of their evolutionary processes. We present here a review of the hunting and reproductive ecology of the pepsine tribe Ageniellini (including the first prey record for Macromerella), and phylogenetic analyses of this group to reconstruct the evolution of nest-constructing behaviour. The maximum parsimony and Bayesian inference analyses are based on 45 adult morphological characters, coded for 41 exemplar species, including five out-group species, representing almost all the genera and subgenera in this tribe. Results supported the following eight clades uniting more than one genus: (1) Cyphononyx + Cryptocheilus + Ageniellini; (2) Ageniellini; (3) Auplopodina, containing Dimorphagenia, Auplopus, Machaerothrix, Phanagenia, Dichragenia, Paragenia, Macromerella, and Macromeris; (4) Phanagenia + Dichragenia + Auplopus artemis + Paragenia + Macromerella + Macromeris; (5) Auplopus artemis + Paragenia + Macromerella + Macromeris; (6) Paragenia + Macromerella + Macromeris; (7) Macromerella + Macromeris; and (8) Ageniella (Lissagenia) + Phanochilus. The monophyly of the genera Auplopus and Ageniella s.l. was not confirmed. The evolution of nesting behaviour was inferred, as burrowing in the soil without using water was ancestral in Ageniellini, and constructing nests by plastering mud obtained by softening soil with water was derived only once at the ancestor of the Auplopodina. The origin of cleptoparasitism was unclear. The origin of communal nesting is also discussed in relation to the evolution of nest-constructing behaviour. The following new combinations are proposed: Ageniella (Alasagenia) sartoriana (Cresson) comb. nov., Cyemagenia certator (Nurse) comb. nov., and Auplopus artemis (Bingham) comb. nov. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160, 88–117.

Towards a phylogeny of the flesh flies (Diptera: Sarcophagidae): morphology and phylogenetic implications of the acrophallus in the subfamily Sarcophaginae

The morphology of the acrophallus, the distal portion of the male phallus carrying the phallotreme, was studied in 72 exemplar species representing 56 genera and subgenera of the flesh fly subfamily Sarcophaginae. For 42 of those species, scanning electron microscopy was used to clarify the phallic morphology. Terms used to describe the male genitalia were updated based on new interpretations of homology. Male genitalic characters, combined with other morphological characters of adult males and females and of larvae, were used to construct a phylogeny. The monophyly of the subfamily was supported, and some generic-level sister-group relationships proposed in the literature, but without previous cladistic analyses, were also supported. The genus Blaesoxipha Loew, as currently recognized, was not monophyletic in our analysis. The genus Helicobia Coquillett is synonymized with Sarcophaga Meigen syn. nov. and treated as a subgenus of the latter. The Sarcophaga subgenera Neobellieria Blanchard and Mehria Enderlein were not monophyletic. Many of the clades in the analysis were supported primarily or exclusively by male genitalic character states, highlighting the importance of the male genitalia as a source of morphological characters for sarcophagine phylogeny.

A phylogeny of <I>Cribrariaceae</I> among <I>Myxomycetes</I> derived from morphological characters

A cladistic study of the Cribrariaceae was performed to examine its phylogenetic position among the Myxomycetes based on a data matrix comprising 54 morphological characters and 55 exemplar myxomycete species. Parsimony ratchet with implied weighting was employed as tree search strategy. Results show the Cribrariaceae as a monophyletic group that includes Cribraria and Dictydium but not Lindbladia and suggest Trichiales as the sister group. Our analyses neither support Dictydium as a genus separated from Cribraria nor the Liceales as monophyletic. This is the first attempt to evaluate the phylogenetic relationships of this group using morphological characters from representative species of all Myxomycetes within a cladistic framework.

Phylogeny and classification of tribe Aedini (Diptera: Culicidae)

The phylogeny and classification of tribe Aedini are delineated based on a cladistic analysis of 336 characters from eggs, fourth-instar larvae, pupae, adult females and males, and immature stage habitat coded for 270 exemplar species, including an outgroup of four species from different non-aedine genera. Analyses of the data set with all multistate characters treated as unordered under implied weights, implemented by TNT version 1.1, with values of the concavity constant K ranging from 7 to 12 each produced a single most parsimonious cladogram (MPC). The MPCs obtained with K values of 7–9 were identical, and that for K = 10 differed only in small changes in the relationships within one subclade. Because values of K 10 produced large changes in the relationships among the taxa, the stability of relationships exemplified by the MPC obtained from the K = 9 analysis is used to interpret the phylogeny and classification of Aedini. Clade support was assessed using parsimony jackknife and symmetric resampling. Overall, the results reinforce the patterns of relationships obtained previously despite differences in the taxa and characters included in the analyses. With two exceptions, all of the groups represented by two or more species were once again recovered as monophyletic taxa. Thus, the monophyly of the following genera and subgenera is corroborated: Aedes, Albuginosus, Armigeres (and its two subgenera), Ayurakitia, Bothaella, Bruceharrisonius, Christophersiomyia, Collessius (and its two subgenera), Dahliana, Danielsia, Dobrotworskyius, Downsiomyia, Edwardsaedes, Finlaya, Georgecraigius (and its two subgenera), Eretmapodites, Geoskusea, Gilesius, Haemagogus (and its two subgenera), Heizmannia (and subgenus Heizmannia), Hopkinsius (and its two subgenera), Howardina, Hulecoeteomyia, Jarnellius, Kenknightia, Lorrainea, Macleaya, Mucidus (and its two subgenera), Neomelaniconion, Ochlerotatus (subgenera Chrysoconops, Culicelsa, Gilesia, Pholeomyia, Protoculex, Rusticoidus and Pseudoskusea), Opifex, Paraedes, Patmarksia, Phagomyia, Pseudarmigeres, Rhinoskusea, Psorophora (and its three subgenera), Rampamyia, Scutomyia, Stegomyia, Tanakaius, Udaya, Vansomerenis, Verrallina (and subgenera Harbachius and Neomacleaya), Zavortinkius and Zeugnomyia. In addition, the monophyly of Tewarius, newly added to the data set, is confirmed. Heizmannia (Mattinglyia) and Verrallina (Verrallina) were found to be paraphyletic with respect to Heizmannia (Heizmannia) and Verrallina (Neomacleaya), respectively. The analyses were repeated with the 14 characters derived from length measurements treated as ordered. Although somewhat different patterns of relationships among the genera and subgenera were found, all were recovered as monophyletic taxa with the sole exception of Dendroskusea stat. nov. Fifteen additional genera, three of which are new, and 12 additional subgenera, 11 of which are new, are proposed for monophyletic clades, and a few lineages represented by a single species, based on tree topology, the principle of equivalent rank, branch support and the number and nature of the characters that support the branches. Acartomyia stat. nov., Aedimorphus stat. nov., Cancraedes stat. nov., Cornetius stat. nov., Geoskusea stat. nov., Levua stat. nov., Lewnielsenius stat. nov., Rhinoskusea stat. nov. and Sallumia stat. nov., which were previously recognized as subgenera of various genera, are elevated to generic status. Catageiomyia stat. nov. and Polyleptiomyia stat. nov. are resurrected from synonymy with Aedimorphus, and Catatassomyia stat. nov. and Dendroskusea stat. nov. are resurrected from synonymy with Diceromyia. Bifidistylus gen. nov. (type species: Aedes lamborni Edwards) and Elpeytonius gen. nov. (type species: Ochlerotatus apicoannulatus Edwards) are described as new for species previously included in Aedes (Aedimorphus), and Petermattinglyius gen. nov. (type species: Aedes iyengari Edwards) and Pe. (Aglaonotus) subgen. nov. (type species: Aedes whartoni Mattingly) are described as new for species previously included in Aedes (Diceromyia). Four additional subgenera are recognized for species of Ochlerotatus, including Oc. (Culicada) stat. nov. (type species: Culex canadensis Theobald), Oc. (Juppius) subgen. nov. (type species: Grabhamia caballa Theobald), Oc. (Lepidokeneon) subgen. nov. (type species: Aedes spilotus Marks) and Oc. (Woodius) subgen. nov. (type species: Aedes intrudens Dyar), and seven are proposed for species of Stegomyia: St. (Actinothrix) subgen. nov. (type species: Stegomyia edwardsi Barraud), St. (Bohartius) subgen. nov. (type species: Aedes pandani Stone), St. (Heteraspidion) subgen. nov. (type species: Stegomyia annandalei Theobald), St. (Huangmyia) subgen. nov. (type species: Stegomyia mediopunctata Theobald), St. (Mukwaya) subgen. nov. (type species: Stegomyia simpsoni Theobald), St. (Xyele) subgen. nov. (type species: Stegomyia desmotes Giles) and St. (Zoromorphus) subgen. nov. (type species: Aedes futunae Belkin). Due to the unavailability of specimens for study, many species of Stegomyia are without subgeneric placement. As is usual with generic-level groups of Aedini, the newly recognized genera and subgenera are polythetic taxa that are diagnosed by unique combinations of characters. The analysis corroborates the previous observation that ‘Oc. (Protomacleaya)’ is a polyphyletic assemblage of species.

Parental and Mating Effort: Is There Necessarily a Trade‐Off?

AbstractOne of the common assumptions in the study of the evolution of parental care is that trade‐offs exist between parental investment and other fitness‐related traits. In general, this body of work follows the traditional definition that parental investment (in the current offspring) decreases that individual’s ability to invest in future reproduction (Trivers 1972). However, examination of the empirical evidence shows that assuming a trade‐off between parental and mating effort is not always appropriate. This overemphasis on a trade‐off between mating and parental effort has arisen in part because of an oversimplification of female reproductive strategies, a failure to consider interactions between the sexes, and a tendency to consider behaviours as unifunctional, thereby ignoring the more complex relationship between mating and parental effort in many species. Here, we first examine the empirical evidence for trade‐offs between mating and parental effort in males and females to ask when trade‐offs occur and what pattern they take. By highlighting a number of exemplar species, we then explore how the presence or absence of trade‐offs relates to mate choice and sexual selection in both sexes. Finally, we highlight the importance of considering individual variation, which has been particularly overlooked in examinations of female investment, and how preferences in one sex may influence the existence and our interpretation of apparent trade‐offs in the other sex.

Surface ultrastructure and evolution of tarsal attachment structures in Plecoptera (Arthropoda: Hexapoda)

The plantar surfaces of the stonefly tarsomeres and pretarsus are examined chiefly using scanning electron microscopy (SEM) of 39 Plecoptera exemplar species representing each family within the two suborders Arctoperlaria and Antarctoperlaria. Features examined include the shape and size of the tarsomeres as well as external features of the plantar surfaces of the tarsomeres, unguitractor plate, claws, arolium and orbicula. Characteristics presumed to be part of the stonefly ground pattern were determined by mapping these features on a cladogram of stonefly phylogeny (Zwick 2000, Annual Review of Entomology 45:709–746). This analysis indicates that the following characters are part of the stonefly ground pattern: (1) cylindrical shape of the tarsomeres, (2) an elongate tarsomere 1, (3) a short tarsomere 2, (4) presence of hairy euplantulae on the plantar surfaces of tarsomeres 1 and 2, (5) absence of specialised attachment surfaces on the plantar area of tarsomere 3, (6) absence of setae on the arolium plantar surface, and (7) presence of setae on the orbicula surface. Hairs covering the euplantulae of tarsomeres 1 and 2 and the median longitudinal unscelerotised region of tarsomeres 1–3 are non-setal cuticular projections and appear to be acanthae. Hairy euplantulae are unusual and are only present in one other insect order, the Mantophasmatodea. Specialised tenent setae were not found. The presence of hairy euplantulae as part of the stonefly ground pattern contradicts the recent view (Beutel and Gorb 2006, Arthropod Systematics & Phylogeny 64:3–25) that stoneflies lack pad-like euplantulae and that the ancestral condition for tarsomeres 1 and 2 may be a narrow median longitudinal unsclerotised band such as that found in representatives of the Austroperlidae. The hypothesis that Plecoptera are the sister clade to the remaining Neoptera suggests that euplantulae might be a synapomorphy of both clades. On the other hand, placement of Plecoptera within a monophyletic Polyneoptera clade within the Neoptera suggests that euplantulae could be an autapomorphy of this subordinate clade.

Genetic engineering of fungal biocontrol agents to achieve greater efficacy against insect pests

Molecular biology methods have elucidated pathogenic processes in several fungal biocontrol agents including two of the most commonly applied entomopathogenic fungi, Metarhizium anisopliae and Beauveria bassiana. In this review, we describe how a combination of molecular techniques has: (1) identified and characterized genes involved in infection; (2) manipulated the genes of the pathogen to improve biocontrol performance; and (3) allowed expression of a neurotoxin from the scorpion Androctonus australis. The complete sequencing of four exemplar species of entomopathogenic fungi including B. bassiana and M. anisopliae will be completed in 2010. Coverage of these genomes will help determine the identity, origin, and evolution of traits needed for diverse lifestyles and host switching. Such knowledge combined with the precision and malleability of molecular techniques will allow design of multiple pathogens with different strategies to be used for different ecosystems and avoid the possibility of the host developing resistance.

Out-of-Africa again: A phylogenetic hypothesis of the genus Charaxes (Lepidoptera: Nymphalidae) based on five gene regions

Despite the long popularity of Charaxes among collectors and researchers, their evolutionary history is largely unknown. The current and accepted species groupings and relationships within the genus are based exclusively on adult morphology and life histories. Here, we examine the monophyly and evolutionary affinities of the species-groups within the genus Charaxes and explore how they relate to members of their closest genera ( Euxanthe, Polyura and Palla) using 4167 bp of sequence data from five (1 mitochondrial and 4 nuclear) gene regions. Within the proposed phylogenetic framework, we estimate ages of divergence within the genus and also reconstruct their historical biogeography. We included representatives of all known species-groups in Africa and Asia, all known species of Euxanthe and Palla and two exemplar species of Polyura. We found the genus Charaxes to be a paraphyletic group with regard to the genera Polyura and Euxanthe, contrary to the earlier assumption of monophyly. We found that 13 out of 16 morphologically defined species-groups with more than one species were strongly supported monophyletic clades. Charaxes nichetes is the sister group to all the other Charaxes. Polyura grouped with the Zoolina and Pleione species-groups as a well-supported clade, and Euxanthe grouped with the Lycurgus species-group. Our results indicated that the common ancestor of Charaxes diverged from the common ancestor of Palla in the mid Eocene (45 million years ago) in (Central) Africa and began diversifying to its extant members 15 million years later. Most of the major diversifications within the genus occurred between the late Oligocene and Miocene when the global climates were putatively undergoing drastic fluctuations. A considerable number of extant species diverged from sister species during the Pliocene. A dispersal–vicariance analysis suggests that many dispersal rather than vicariance events resulted in the distribution of the extant species. The genus Polyura and the Indo-Australian Charaxes are most likely the results of three independent colonizations of Asia by African Charaxes in the Miocene. We synonymize the genera Polyura (syn. nov.) and Euxanthe (syn. nov.) with Charaxes, with the currently circumscribed Charaxes subdivided into five subgenera to reflect its phylogeny.

Generic Revision of the Dioptinae (Lepidoptera: Noctuoidea: Notodontidae) Part 1: Dioptini

Abstract The moth subfamily Dioptinae is almost entirely neotropical. One species—Phryganidia californica—occurs on the west coast of the US, while the remaining taxa are found from Mexico south to northern Argentina and Uruguay. None is known from the Old World. Most dioptines are diurnal as adults, and many exhibit aposematic coloration. A few taxa are nocturnal. Their larval hosts include the families Passifloraceae, Violaceae and Poaceae, plants famous for their use by important butterfly groups. In this paper, a revised generic classification for the Dioptinae is presented. Nearly 17,000 specimens, assembled from 38 private and institutional collections worldwide, form the basis for the first comprehensive analysis of adult morphology in the subfamily. A subset of 115 exemplar species, chosen to represent structural diversity across the Dioptinae, is subjected to detailed morphological study and cladistic analysis. The resulting matrix includes 305 characters delineated by 938 character states. Cladi...

The Neotropical genus Stibadocerina Alexander and its phylogenetic relationship to other Stibadocerinae genera: further evidence of an ancestral trans‐Pacific biota (Diptera: Cylindrotomidae)

Abstract Stibadocerina Alexander, a monotypic genus, includes the only known Neotropical species of the family Cylindrotomidae, S. chilensis Alexander, 1929, from South Central Chile (ca. 36°50′S–42°17′S). In this paper, Stibadocerina chilensis is redescribed and illustrated in detail. A study of wing‐vein homology in the subfamily Stibadocerinae is provided, to identify the components of the reduced radial sector in Stibadocerina and related taxa. The proposed hypotheses of wing‐vein homology are tested, and the systematic position of Stibadocerina is assessed through a cladistic analysis of 13 characters of the male imago, scored for exemplar species of the four genera included in the Stibadocerinae. A single most parsimonious tree supports the monophyly of the Stibadocerinae and the following relationships among its included genera: Stibadocerodes [Stibadocera (Stibadocerella +Stibadocerina)]. The subfamily includes one example of a vicariant distribution with a sister‐group relationship between South Central Chilean and East Asian taxa, and supports a biogeographical interpretation of an ancestral trans‐Pacific biota.