Gavin J. Svenson

A mitochondrial genome phylogeny of termites (Blattodea: Termitoidae): Robust support for interfamilial relationships and molecular synapomorphies define major clades

Despite their ecological significance as decomposers and their evolutionary significance as the most speciose eusocial insect group outside the Hymenoptera, termite (Blattodea: Termitoidae or Isoptera) evolutionary relationships have yet to be well resolved. Previous morphological and molecular analyses strongly conflict at the family level and are marked by poor support for backbone nodes. A mitochondrial (mt) genome phylogeny of termites was produced to test relationships between the recognised termite families, improve nodal support and test the phylogenetic utility of rare genomic changes found in the termite mt genome. Complete mt genomes were sequenced for 7 of the 9 extant termite families with additional representatives of each of the two most speciose families Rhinotermitidae (3 of 7 subfamilies) and Termitidae (3 of 8 subfamilies). The mt genome of the well supported sister-group of termites, the subsocial cockroach Cryptocercus, was also sequenced. A highly supported tree of termite relationships was produced by all analytical methods and data treatment approaches, however the relationship of the termites+Cryptocercus clade to other cockroach lineages was highly affected by the strong nucleotide compositional bias found in termites relative to other dictyopterans. The phylogeny supports previously proposed suprafamilial termite lineages, the Euisoptera and Neoisoptera, a later derived Kalotermitidae as sister group of the Neoisoptera and a monophyletic clade of dampwood (Stolotermitidae, Archotermopsidae) and harvester termites (Hodotermitidae). In contrast to previous termite phylogenetic studies, nodal supports were very high for family-level relationships within termites. Two rare genomic changes in the mt genome control region were found to be molecular synapomorphies for major clades. An elongated stem-loop structure defined the clade Polyphagidae + (Cryptocercus+termites), and a further series of compensatory base changes in this stem-loop is synapomorphic for the Neoisoptera. The complicated repeat structures first identified in Reticulitermes, composed of short (A-type) and long (B-type repeats) defines the clade Heterotermitinae+Termitidae, while the secondary loss of A-type repeats is synapomorphic for the non-macrotermitine Termitidae.

Phylogeny of Dictyoptera: Dating the Origin of Cockroaches, Praying Mantises and Termites with Molecular Data and Controlled Fossil Evidence

Understanding the origin and diversification of organisms requires a good phylogenetic estimate of their age and diversification rates. This estimate can be difficult to obtain when samples are limited and fossil records are disputed, as in Dictyoptera. To choose among competing hypotheses of origin for dictyopteran suborders, we root a phylogenetic analysis (~800 taxa, 10 kbp) within a large selection of outgroups and calibrate datings with fossils attributed to lineages with clear synapomorphies. We find the following topology: (mantises, (other cockroaches, (Cryptocercidae, termites)). Our datings suggest that crown-Dictyoptera—and stem-mantises—would date back to the Late Carboniferous (~ 300 Mya), a result compatible with the oldest putative fossil of stem-dictyoptera. Crown-mantises, however, would be much more recent (~ 200 Mya; Triassic/Jurassic boundary). This pattern (i.e., old origin and more recent diversification) suggests a scenario of replacement in carnivory among polyneopterous insects. The most recent common ancestor of (cockroaches + termites) would date back to the Permian (~275 Mya), which contradicts the hypothesis of a Devonian origin of cockroaches. Stem-termites would date back to the Triassic/Jurassic boundary, which refutes a Triassic origin. We suggest directions in extant and extinct species sampling to sharpen this chronological framework and dictyopteran evolutionary studies.

Family-level relationships of the spittlebugs and froghoppers (Hemiptera: Cicadomorpha: Cercopoidea)

The spittlebug superfamily Cercopoidea (Hemiptera: Cicadomorpha) comprises approximately 3000 phytophagous species (including some economically important pests of grass crops) classified among the families Cercopidae, Aphrophoridae, Epipygidae, Clastopteridae and Machaerotidae. However, the monophyly of these taxa has never been tested and the evolutionary relationships among these major lineages are unknown. Presented here are the results of the first ever phylogenetic investigation of the higher‐level relationships within Cercopoidea, based on DNA nucleotide sequence data from six loci (18S rDNA, 28S rDNA, histone 3, wingless, cytochrome oxidase I and cytochrome oxidase II) generated from exemplars of 109 spittlebug species representing all five described families, seven of eight subfamilies and 61 genera (eight additional exemplars, representing a selection of other Auchenorrhyncha taxa, were included as outgroups). The resulting topologies are used to evaluate the monophyly of each cercopoid family, and further to calculate divergence date estimates to examine the chronological origins and historical diversification of Cercopoidea. The results of this investigation suggest that: (i) four of the five described families are monophyletic; Epipygidae was recovered consistently as originating within Aphrophoridae; (ii) the exclusively Old World Machaerotidae is the most anciently diversified family of extant spittlebugs; (iii) New World Cercopidae (i.e. Ischnorhininae) constitute a derived monophyletic lineage; (iv) the genus Microsargane Fowler, classified currently within Aphrophoridae, actually belongs within Cercopidae; and (v) the origins of the major spittlebug lineages probably coincided with the breakup of Pangaea and, subsequently, Gondwana, as well as major floristic diversification such as the rise of angiosperms.

Of flowers and twigs: phylogenetic revision of the plant-mimicking praying mantises (Mantodea: Empusidae and Hymenopodidae) with a new suprageneric classification

We present a suprageneric revision of the plant‐mimicking mantis families Empusidae and Hymenopodidae on the basis of a total‐evidence phylogenetic analysis with complete generic representation. We coded 124 characters from external morphology and used these together with a DNA dataset comprising ten gene fragments for an aligned matrix with 7514 nucleic acid sites to estimate phylogenetic relationships. We recovered largely congruent topologies across molecular, morphological and total evidence analyses. Empusidae and its assigned subfamilies were recovered as monophyletic. Hymenopodidae was recovered as paraphyletic with respect to Sibyllidae and Phyllothelyinae (Mantidae) whereas a small assemblage of hymenopodid taxa (Galinthias Stål, Congoharpax La Greca, Pseudoharpax Saussure and Harpagomantis Kirby) were recovered outside the family. The Caribbean genus Epaphrodita Audinet‐Serville was also recovered far from the rest of the hymenopodid subfamily Epaphroditinae, in which it is traditionally classified. The nonmonophyly of Acromantinae was recovered with some species nested within Oxypilinae and others found within Hymenopodinae. We present a new classification scheme that includes an elevated family and subfamily (Galinthiadidae and Phyllocraniinae), two reinstated tribes (Anaxarchini and Otomantini), a family demoted to subfamily rank (Sibyllinae), a newly assigned subfamily to Hymenopodidae transferred from Mantidae (Phyllothelyinae), and new generic assignments to existing higher‐level groups. We used our morphological characters to produce extensive re‐descriptions of all suprageneric groups as well as character diagnoses. We provide a dichotomous key to genera supported by example illustrations of important morphological features as well as dorsal habitus images of representative specimens.

The phylogeny and classification of Embioptera (Insecta)

A phylogenetic analysis of the order Embioptera is presented with a revised classification based on results of the analysis. Eighty‐two species of Embioptera are included from all families except Paedembiidae Ross and Embonychidae Navás. Monophyly of each of the eight remaining currently recognized families is tested except Andesembiidae Ross, for which only a single species was included. Nine outgroup taxa are included from Blattaria, Grylloblattaria, Mantodea, Mantophasmatodea, Orthoptera, Phasmida and Plecoptera. Ninety‐six morphological characters were analysed along with DNA sequence data from the five genes 16S rRNA, 18S rRNA, 28S rRNA, cytochrome c oxidase I and histone III. Data were analysed in combined analyses of all data using parsimony and Bayesian optimality criteria, and combined molecular data were analysed using maximum likelihood. Several major conclusions about Embioptera relationships and classification are based on interpretation of these analyses. Of eight families for which monophyly was tested, four were found to be monophyletic under each optimality criterion: Clothodidae Davis, Anisembiidae Davis, Oligotomidae Enderlein and Teratembiidae Krauss. Australembiidae Ross was not recovered as monophyletic in the likelihood analysis in which one Australembia Ross species was recovered in a position distant from other australembiids. This analysis included only molecular data and the topology was not strongly supported. Given this, and because parsimony and the Bayesian analyses recovered a strongly supported clade including all Australembiidae, we regard this family also as monophyletic. Three other families – Notoligotomidae Davis, Archembiidae Ross and Embiidae Burmeister, as historically delimited – were not found to be monophyletic under any optimality criterion. Notoligotomidae is restricted here to include only the genus Notoligotoma Davis with a new family, Ptilocerembiidae Miller and Edgerly, new family, erected to include the genus Ptilocerembia Friederichs. Archembiidae is restricted here to include only the genera Archembia Ross and Calamoclostes Enderlein. The family group name Scelembiidae Ross is resurrected from synonymy with Archembiidae (new status) to include all other genera recently placed in Archembiidae. Embiidae is not demonstrably monophyletic with species currently placed in the family resolved in three separate clades under each optimality criterion. Because taxon sampling is not extensive within this family in this analysis, no changes are made to Embiidae classification. Relationships between families delimited herein are not strongly supported under any optimality criterion with a few exceptions. Either Clothodidae Davis (parsimony) or Australembiidae Ross (Bayesian) is the sister to the remaining Embioptera taxa. The Bayesian analysis includes Australembiidae as the sister to all other Embioptera except Clothididae, suggesting that each of these taxa is a relatively plesiomorphic representatative of the order. Oligotomidae and Teratembiidae are sister groups, and Archembiidae (sensu novum), Ptilocerembiidae, Andesembiidae and Anisembiidae form a monophyletic group under each optimality criterion. Each family is discussed in reference to this analysis, diagnostic combinations and taxon compositions are provided, and a key to families of Embioptera is included.

Revision of the Neotropical bark mantis genus Liturgusa Saussure, 1869 (Insecta, Mantodea, Liturgusini)

Abstract The praying mantis genus Liturgusa Saussure, 1869 occurs only in Central and South America and represents the most diverse genus of Neotropical Liturgusini (Ehrmann 2002). The genus includes bark dwelling species, which live entirely on the trunks and branches of trees and run extremely fast. All species included within the genus Liturgusa are comprehensively revised with a distribution stretching from central Mexico, the island of Dominica to the southeastern regions of Brazil and southern Bolivia. All known species are redescribed to meet the standards of the new treatment of the genus (11 species). Three new genera are described including Fuga gen. n., Velox gen. n., and Corticomantis gen. n. for species previously included in Liturgusa as well as Hagiomantis. Liturgusa mesopoda Westwood, 1889 is moved to within the previously described genus Hagiomantis Audinet Serville, 1838. A total of 19 species are newly described within Liturgusa, Fuga, and Velox including L. algorei sp. n., L. bororum sp. n., L. cameroni sp. n., L. cura sp. n., L. dominica sp. n., L. fossetti sp. n., L. kirtlandi sp. n., L. krattorum sp. n., L. manausensis sp. n., L. maroni sp. n., L. milleri sp. n., L. neblina sp. n., L. purus sp. n., L. stiewei sp. n., L. tessae sp. n., L. trinidadensis sp. n., L. zoae sp. n., F. grimaldii sp. n., and V. wielandi sp. n. Four species names are synonymized: Liturgusa peruviana Giglio-Tos, 1914, syn. n. = Liturgusa nubeculosa Gerstaecker, 1889 and Hagiomantis parva Piza, 1966, syn. n., Liturgusa sinvalnetoi Piza, 1982, syn. n., and Liturgusa parva Giglio-Tos, 1914, syn. n. = Mantis annulipes Audinet Serville, 1838. Lectotypes are designated for the following two species: Liturgusa maya Saussure & Zehntner, 1894 and Fuga annulipes (Audinet Serville, 1838). A male neotype is designated for Liturgusa guyanensis La Greca, 1939. Males for eight species are described for the first time including Liturgusa cayennensis Saussure, 1869, Liturgusa lichenalis Gerstaecker, 1889, Liturgusa guyanensis La Greca, 1939, Liturgusa maya Saussure & Zehntner, 1894, Liturgusa nubeculosa Gerstaecker, 1889, Fuga annulipes (Audinet Serville, 1838), Corticomantis atricoxata (Beier, 1931), and Hagiomantis mesopoda (Westwood, 1889). The female of Fuga fluminensis (Piza, 1965) is described for the first time. Complete bibliographic histories are provided for previously described species. The spelling confusion surrounding Liturgusa/Liturgousa is resolved. Full habitus images for males and females are provided for nearly all species. Habitus and label images of type specimens are provided when possible. Diagnostic illustrations of the head and pronotum for males and females are provided for all species when possible. Illustrations of male genital structures are provided for all species for which males are known. Measurement data, including ranges and averages, are provided for males and females of all species. Combined male and female genus and species level dichotomous keys are provided with a Spanish translation. A complete table of all examined specimens lists label data, museum codes, repositories, and other specimen specific information. A KML file with all georeferenced locality records is downloadable from mantodearesearch.com for viewing in Google Earth. Natural history information is provided for species observed by the author.

Manual of praying mantis morphology, nomenclature, and practices (Insecta, Mantodea)

Abstract This study provides a comprehensive review of historical morphological nomenclature used for praying mantis (Mantodea) morphology, which includes citations, original use, and assignment of homology. All referenced structures across historical works correspond to a proposed standard term for use in all subsequent works pertaining to praying mantis morphology and systematics. The new standards are presented with a verbal description in a glossary as well as indicated on illustrations and images. In the vast majority of cases, originally used terms were adopted as the new standard. In addition, historical morphological topographical homology conjectures are considered with discussion on modern interpretations. A new standardized formulation to present foreleg femoral and tibial spines is proposed for clarity based on previous works. In addition, descriptions for methods of collection, curation, genital complex dissection, and labeling are provided to aid in the proper preservation and storage of specimens for longevity and ease of study. Due to the lack of consistent linear morphometric measurement practices in the literature, we have proposed a series of measurements for taxonomic and morphological research. These measurements are presented with figures to provide visual aids with homologous landmarks to ensure compatibility and comparability across the Order. Finally, our proposed method of pinning mantises is presented with a photographical example as well as a video tutorial available at http://mantodearesearch.com.

A Cretaceous-aged Palaeotropical dispersal established an endemic lineage of Caribbean praying mantises

Recent phylogenetic advances have uncovered remarkable biogeographic histories that have challenged traditional concepts of dispersal, vicariance and diversification in the Greater Antilles. Much of this focus has centred on vertebrate lineages despite the high diversity and endemism of terrestrial arthropods, which account for 2.5 times the generic endemism of all Antillean plants and non-marine vertebrates combined. In this study, we focus on three Antillean endemic praying mantis genera, Callimantis, Epaphrodita and Gonatista, to determine their phylogenetic placement and geographical origins. Each genus is enigmatic in their relation to other praying mantises due to their morphological affinities with both Neotropical and Old World groups. We recovered the three genera as a monophyletic lineage among Old World groups, which was supported by molecular and morphological evidence. With a divergence at approximately 107 Ma, the lineage originated during the break-up of Gondwana. Ancestral range reconstruction indicates the lineage dispersed from an African + Indomalayan range to the Greater Antilles, with a subsequent extinction in the Old World. The profound ecomorphic convergence with non-Caribbean groups obscured recognition of natural relationships within the same geographical distribution. To the best of our knowledge, the lineage is one of the oldest endemic animal groups in the Greater Antilles and their morphological diversity and restricted distribution mark them as a critical taxon to conserve.

The Neotropical ‘polymorphic earless praying mantises’ – Part I: molecular phylogeny and revised higher‐level systematics (Insecta: Mantodea, Acanthopoidea)

We perform phylogenetic analyses of the ‘polymorphic earless praying mantises’, a heterogeneous assemblage comprising c. 55% of mantodean diversity in the Neotropics. Bayesian and maximum‐likelihood were implemented on a DNA dataset of 9949 aligned nucleic acid characters comprising ten mitochondrial and nuclear genes. Our analyses largely resolved congruent relationships with high levels of support for higher‐level taxonomic groups, but revealed extensive inconsistencies between the resolved topology and morphology‐based classification systems. The polymorphic earless praying mantises, now granted superfamily status as the Acanthopoidea stat. n., comprises 8 families, 15 subfamilies and 18 tribes. Our newly revised organization required the following taxonomic changes: (i) Thespidae sensu n., including subfamilies Pseudopogonogastrinae subfam. n., Pseudomiopteryginae sensu n., Bantiinae subfam. n., Miobantiinae sensu n. and Thespinae sensu n. (tribes Musoniellini trib. n. and Thespini sensu n.); (ii) Angelidae stat. n. et sensu n.; (iii) Coptopterygidae stat. n.; (iv) Liturgusidae sensu n.; (v) Photinaidae stat. n., including Macromantinae stat. n., Cardiopterinae stat. n., Photiomantinae subfam. n. and Photinainae sensu n. (tribes Microphotinini trib. n., Orthoderellini stat. n. and Photinaini sensu n.); (vi) Stenophyllidae stat. n.; (vii) Acontistidae stat. n.; and (viii) Acanthopidae sensu n. Our new system also resulted in the reassignment of various genera to new and existing higher‐level taxa, the exclusion of old world genera otherwise traditionally classified among the Thespidae, Liturgusidae and Angelidae, the confirmation of Stenophylla Westwood as member of this clade, and the revalidation of Paradiabantia Piza stat. r. We provide diagnoses for all suprageneric taxa using external morphological characters and male genitalia. A key to higher‐level groups is provided. We incorporate egg case structural variation as a novel approach for taxon delineation.

Visual stimuli that elicit visual tracking, approaching and striking behavior from an unusual praying mantis, Euchomenella macrops (Insecta: Mantodea).

In comparison to other similarly sized mantis species examined in previous studies, Euchomenella macrops has a significantly smaller head, shorter foreleg tibia, but longer prothorax which have been interpreted as specializations for the capture of smaller, slower prey. We tested this conjecture by assessing the rates at which computer generated stimuli elicit visual tracking, approaching, and striking behaviors by adult females. When presented with black disks moving erratically against a white background, strike rate rose progressively as disks enlarged up to 44 deg (visual angle) if the disks moved rapidly (e.g., 143 deg/s); at slower speeds (113, 127 deg/s), smaller disks (<27 deg) were preferred. When black moved linearly from the visual periphery to visual field center (at 73 or 143 deg/s) and then stopped, E. macrops struck consistently at disks as small as 5 deg after movement ceased. E. macrops also struck at higher rates in response to 23 deg erratically moving (subjective) red (versus subjective blue or green) disks that were luminance matched to a grey background although they tracked all colors at equally high rates. Unlike some other species, E. macrops did not strike at higher rates in response to elongated rectangular stimuli moving parallel (versus perpendicular) to their long axis, although the former elicited higher rates of approaching. An analysis of tracking behavior revealed that virtually all tracking movements were a result of head (versus) prothorax rotation.