Phylogeny of the Neotropical moth tribe Josiini (Notodontidae: Dioptinae): a hidden case of Miillerian mimicry

The Neotropical moth tribe Josiini (Notodontidae: Dioptinae) contains over 100 described species in 11 genera. All are diurnal, with brightly-coloured, presumably aposematic wing patterns. Larval hostplants are exclusively in the genus Passiflora (Passifloraceae) except for two new records, reported here, from Tumera (Turneraceae). A comparative morphological study of 26 representative josiine species yielded 86 characters from adults, larvae and pupae, all of which are figured and discussed. Phylogenetic analysis of these data produced a single most-parsimonious cladogram. According to the phylogenetic results: (1) monophyly of the Josiini is strongly supported; (2) the currently accepted generic classification is in disarray; (3) morphological character variation is extensive, and adult traits reflect phylogeny more effectively than do those of immature stages; (4) wing pattern types have undergone convergent evolution. A rare phenotype, longitudinal wing stripes, appears in two widely divergent clades, suggesting the evolution of Miillerian mimicry within the Josiini.     

Phylogeny of Grumichellini Morse, 1981 (Trichoptera: Leptoceridae) with the description of a new genus from southeastern Peru

Osflintia manu, new genus, new species, of long-horned caddisfly (Leptoceridae: Triplectidinae: Grumichellini) is described and illustrated from southeastern Peru. The phylogeny of Grumichellini Morse (Leptoceridae: Triplectidinae) is revisited and hypotheses of homology of some morphological characters are reconsidered. The monophyly of the tribe is corroborated and the phylogenetic relationships of its included genera are inferred to be (Triplexa (Gracilipsodes ((Grumichella, Amazonatolica) (Atanatolica, Osflintia, n. gen.)))) from adult and larval characters. Diagnostic characters of the new genus include the following: reduced tibial spur formula (2, 2, 2), loss of forewing crossvein sc-r1, hind wing discoidal cell closed, hind wing fork IV present, pair of long setae on tergum IX of the male genitalia, and pair of processes on the apex of segment X.     

Evolution and phylogeny of the scarab subtribe Anisopliina (Coleoptera: Scarabaeidae: Rutelinae: Anomalini)

Abstract.  The subtribe Anisopliina (Scarabaeidae: Rutelinae: Anomalini) is associated with grasses, and its species are distributed in the Palaearctic, Oriental, Ethiopian, Nearctic and Neotropical biogeographical regions. Phylogenetic analysis of adult morphological characters was conducted to examine the monophyly and classification of the group, as well as to examine characters associated with grass pollinivory and graminivory. We review the biology, phylogeny and classification of the Anisopliina and provide an overview of each genus. The analysis of ninety-one morphological characters using parsimony does not support the monophyly of the subtribe Anisopliina. Instead, the results provide support for a group referred to here as the anisopliine clade, a circum-Mediterranean group, forming an internal clade within the well-supported tribe Anomalini. Sister group relationships are discussed, possibly being associated with a New World anomaline taxon. Character states associated with grass herbivory, including mouthpart and leg characters, are discussed based on the phylogenetic analysis. Within the Anomalini, an evolutionary shift from generalized leaf feeding to grass associations and grass pollen feeding is supported.     

The evolution of social behaviour in Blaberid cockroaches with diverse habitats and social systems: phylogenetic analysis of behavioural sequences

The adequacy and utility of behavioural characters in phylogenetics is widely acknowledged, especially for stereotyped behaviours. However, the most common behaviours are not stereotyped, and these are usually seen as inappropriate or more difficult to analyze in a phylogenetic context. A few methods have been proposed to deal with such data, although they have never been tested on samples larger than six species, which limits their evolutionary interest. In the present study, we perform behavioural observations on 13 cockroach species and derive behavioural phylogenetic characters with the successive event‐pairing method. We combine these characters with morphological and molecular data (approximately 6800 bp) in a phylogenetic study of 41 species. We then reconstruct ancestral states of the behavioural data to study evolution of social behaviour in these insects with regard to their social systems (i.e. solitary, gregarious, and subsocial) and diversity of habitat choice. We report for the first time that nonstereotyped behavioural data are adequate for phylogenetic analyses: they are no more homoplastic than traditional data, and support several phylogenetic relationships that we discuss. From an evolutionary perspective, we show that the solitary species Thanatophyllum akinetum does not display original behavioural interactions, suggesting phylogenetic inertia of interactive behaviours despite a radical change in social structure. Conversely, the subsocial species Parasphaeria boleiriana shows original behavioural interactions, which could result from its peculiar social system or habitat. We conclude that phylogenetic approaches in studies of behaviour are useful for deciphering evolution of behaviour and discriminating between its different modalities, even for nonstereotyped characters. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 58–77.     

Character- and tree-based delimitation of species in the 'Cichlasoma' facetum group (Teleostei, Cichlidae) with the description of a new genus

The 'Cichlasoma' facetum group is part of the taxonomically complex group of Neotropical cichlid fishes of the tribe Heroini. Many species groups and unplaced species of heroines are still left without a generic name following the revision of the genus Cichlasoma . We describe here the 'Cichlasoma' facetum group as a new genus, Australoheros , and provide evidence for its monophyly based on phylogenetic analyses of morphological and mtDNA characters. Australoheros is morphologically characterized by the lowest values in meristic characters among heroines and by three apomorphic characters in coloration pattern. In addition to the three described species of Australoheros , our results of species delimitation based on a combination of tree- and character-based approaches identify seven putatively new species of Australoheros . Several coding schemes of morphological characters are used to recover the intrageneric relationships within the genus, resulting in very similar topologies. Discovery of additional species within the genus is expected once material from the whole distribution area is studied.     

The geometry of the marmot (rodentia: sciuridae) mandible: phylogeny and patterns of morphological evolution

Marmots have a prominent role in the study of mammalian social evolution, but only recently has their systematics received the attention it deserves if sociobiological studies are to be placed in a phylogenetic context. Sciurid morphology can be used as model to test the congruence between morphological change and phylogeny because sciurid skeletal characters are considered to be inclined to convergence. However, no morphological study involving all marmot species has ever been undertaken. Geometric morphometric techniques were applied in a comparative study of the marmot mandible. The adults of all 14 living marmot species were compared, and mean mandible shape were used to investigate morphological evolution in the genus Marmota. Three major trends were observed. First, the phylogenetic signal in the variation of landmark geometry, which describes mandible morphology, seems to account for the shape differences at intermediate taxonomic levels. The subgenera Marmota and Petromarmota, recently proposed on the basis of mitochondrial cytochrome b sequence, receive support from mandible morphology. When other sciurid genera were included in the analysis, the monophyly of the genus Marmota and that of the tribe Marmotini (i.e., marmots, prairie dogs, and ground squirrels) was strengthened by the morphological data. Second, the marmotine mandible may have evolved as a mosaic of characters and does not show convergence determined by size similarities. Third, allopatric speciation in peripheral isolates may have acted as a powerful force for modeling shape. This hypothesis is strongly supported by the peculiar mandible of M. vancouverensis and, to a lesser degree, by that of M. olympus, both thought to have originated as isolated populations in Pleistocene ice-free refugia.     

Systematic position of Lythriini revised: transferred from Larentiinae to Sterrhinae (Lepidoptera, Geometridae)

The tribe Lythriini is a small group of diurnally active geometrid moths consisting of a single Palaearctic genus Lythria with five species. The systematic placement of Lythriini has remained controversial: though traditionally it has been placed into the subfamily Larentiinae, a number of morphological characters link this tribe with the subfamily Sterrhinae. A molecular phylogenetic study was conducted to verify the systematic position of Lythriini, using sequences of both mitochondrial and nuclear genes: elongation factor 1α ( EF-1α ), wingless ( wgl ), 28S rRNA expansion segment D2 ( 28S D2 ), cytochrome oxidase subunit 1 ( COI ) and NADH dehydrogenase subunit 1 ( ND1 ) (a total of 3784 bp). Phylogenetic analysis reliably demonstrated that Lythriini belong to the subfamily Sterrhinae. Therefore, we propose to remove tribe Lythriini from Larentiinae and unite it with Sterrhinae. Moreover, our analysis supports the monophyly of both Sterrhinae and Larentiinae. However, although both morphological data and interspecific genetic distances insinuated that Lythria cruentaria and L. sanguinaria are sister species, the latter formed a clade of sister taxa together with L. purpuraria .     

Phylogenetic relationships,subdivision, and biogeography of the cyprinid tribe Labeonini (sensu ) (Teleostei: Cypriniformes), with comments on the implications of lips and associated structures in the labeonin classification

The Labeonini (sensu Rainboth, 1991) is a tribe of the subfamily Cyprininae, the largest subfamily of Cypriniformes. With around 400 species in 34 genera, this tribe is widely distributed in the freshwaters of tropical Africa and Asia. Most species are adapted to fast-flowing streams and rivers, and exhibit unique morphological modifications associated with their lips and other structures around the mouth. The monophyly of this tribe has been tested and generally accepted in previous morphological and molecular studies. The major objectives of this study were to reconstruct the phylogenetic relationships within the tribe Labeonini, test its monophyly and explore the taxonomic subdivisions, intrarelationships and biogeography of the group. The value of the morphological characters associated with the lips and other associated structures in the taxonomic classification of labeonins was also discussed. Nucleotide sequences (3867 bp) of four unlinked nuclear loci were obtained from 51 species in 18 Labeonini genera from throughout the range of the tribe. Maximum parsimony, partitioned maximum likelihood and partitioned Bayesian analyses were used for phylogenetic inference from combined and separate gene data sets. Based on our results, the monophyly of Labeonini was well supported. Two major clades could be recovered within the tribe. Three subclades could further be recognized from the first clade. These clades/subclades are not consistent with groupings of any of previous workers using either morphological or molecular characters for phylogenetic inference. Only five currently recognized genera in this analysis are monophyletic. The similarity between some lips and associated structures (e.g. suctorial discs) of labeonins may due to convergence or parallelism instead of common ancestry. Labeonins of Southeast Asia, India and China are closely related to each other; the multiple clades of African taxa do not form a single monophyletic group, indicating multiple, independent dispersal events of labeonins into Africa from Asia.     

Insights into the phylogenetic relationships within Cixiidae (Hemiptera: Fulgoromorpha): cladistic analysis of a morphological dataset

Abstract.  According to the most recent classifications proposed, the planthopper family Cixiidae comprises three subfamilies, namely Borystheninae, Bothriocerinae and Cixiinae, the latter with 16 tribes. Here we examine morphological characters to present the first phylogenetic reconstructions within Cixiidae derived from a cladistic analysis. We scored 85 characters of the head, thorax, and male and female genitalia for 50 taxa representative of all cixiid subfamilies and tribes and for six outgroup taxa. Analyses were based on maximum parsimony – using both equally weighted and successive weighting procedures – and Bayesian inferences. The monophyly of most currently accepted tribes and subfamilies was investigated through Templeton statistical tests of alternative phylogenetic hypotheses. The cladistic analyses recover the monophyly of Cixiidae, the subfamily Bothriocerinae, and the tribes Pentastirini, Mnemosynini, and Eucarpiini. Successive weighting and Bayesian inference recover the monophyly of the tribe Gelastocephalini, but only Bayesian inference supports the monophyly of Semoniini. The relationships recovered support the groups [Stenophlepsini (Borystheninae + Bothriocerinae)] arising from the tribe Oecleini, and [Andini + Brixiidini + Brixiini (polyphyletic) + Bennini]. Templeton tests reject the alternative hypothesis of a monophyletic condition for the tribe Pintaliini as presently defined.     

Phylogenetic relationships in Dichroplus Stål (Orthoptera: Acrididae: Melanoplinae) inferred from molecular and morphological data: testing karyotype diversification

The neotropical genus Dichroplus and related genera are characterized by a relatively uniform external morphology and a remarkably divergent male genitalia and hence its taxonomy is controversial. It also shows an extreme karyotypic diversification. In this study we used molecular and morphological characters to test the monophyly of the genus and to evaluate chromosome evolution. Twenty‐seven species from Dichroplus and related genera were included in the analysis. Morphological characters refer to the general morphology, male genitalia and female structures. Molecular studies were performed, sequencing part of two mitochondrial genes, cytochrome oxidase I and II. Independent and combined phylogenetic analyses of the data were performed under maximum parsimony. The karyotypic characters (rearrangements) were either mapped onto the combined topology or combined with the other data sets. While the molecular analysis confirms some results attained with morphology, some others do not. All point towards the paraphyly of the genus. Our results show the relevance of morphological data in phylogenetic studies because morphology and molecules supply complementary evidence. The mapping of chromosome characters on the combined tree shows that the most extreme karyotype, in D. silveiraguidoi, is a derived condition, probably reached through several centric fusions, and that X‐autosome centric fusions were recurrently fixed during the evolution of the group. © The Willi Hennig Society 2005.     

Antlers honestly advertise sperm production and quality

Evolutionary theory proposes that exaggerated male traits have evolved via sexual selection, either through female mate choice or male-male competition. While female preferences for ornamented males have been amply demonstrated in other taxa, among mammals sexual characters are commonly regarded as weapons whose main function is to enhance male competitiveness in agonistic encounters. One particularly controversial hypothesis to explain the function of male sexual characters proposes that they advertise male fertility. We test this hypothesis in red deer (Cervus elaphus), a species where sexual characters (antlers) reach an extreme degree of elaboration. We find that a global measure of relative antler size and complexity is associated with relative testes size and sperm velocity. Our results exclude the possibility that condition dependence, age or time of culling, drive these associations. Red deer antlers could signal male fertility to females, the ability to avoid sperm depletion throughout the reproductive season and/or the competitive ability of ejaculates. By contrast, male antlers could also signal to other males not only their competitive ability at the behavioural level (fighting ability) but also at the physiological level (sperm competition).     

The relevance of the mesosomal internal structures to the phylogeny of Augochlorini bees (Hymenoptera: Halictinae)

The use of internal skeletal structures is valuable to phylogenetic reconstruction within Hymenoptera; however, these structures were not further investigated for Augochlorini. Previous phylogenetic studies of Augochlorini based only on external morphology were poorly resolved. The objectives of this work are to explore the comparative morphology of internal structures of the mesosoma and to evaluate their potential as information sources for the phylogenetic relationships within the tribe. The internal structures of the Pseudaugochlora graminea (Fabricius, 1804) mesosoma are described in detail and compared with 16 other genera. We propose 24 new character statements for the Augochlorini phylogeny based on internal structures of mesosoma. Prosternum, propleuron and meso/metafurca provided a great number of informative characters. On the other hand, the mesophragma, metanotum and propodeum were less variable. The monophyly of Augochlorini and of all genus groups was corroborated in a parsimony analysis of internal and external morphological characters. Characters derived from internal structures provided the first‐known morphological synapomorphies for the clades: Megaloptidia group + others, Neocorynura group + others and Augochlora group + Megalopta group. These characters helped to elucidate the evolution of the group when analysed together with external morphological characters.     

Phylogeny of the genus Palmanura (Collembola: Neanuridae)

In order to assess the phylogenetic structure of the springtail genus Palmanura, as well as to test the monophyly of the tribe Sensillanurini (Neanuridae: Neanurinae), a data matrix of morphological (chaetotactic and other) characters of members of this group was assembled and analysed in the light of Wagner parsimony. The data matrix included all the known members of the Neotropical genus Palmanura, plus representatives of Sensillanura and Americanura. Although not all the clades obtained were highly supported by bootstrap resampling, some structures were relatively constant under different approaches. Alternative analyses (unordered and ordered character states, rescaled weighting procedure) were applied. While alternative solutions were obtained, a number of structures were shared by the results irrespective of the method used. On this basis, the results suggest that some further reassessment is required to confirm formally the monophyly of the tribe Sensillanurini. The genera Palmanura and Americanura are mutually poly/paraphyletic; we thus suggest that Palmanura should be considered as a synonym of Americanura, although some character reassessment and more varied outgroup species may be necessary before a formal generic redefinition can be proposed. Finally, a comparison of the performance of the characters under Wagner parsimony analysis indicated that differences in the characters’ retention indexes are due not to the topological (tagmal) position of the traits involved, but to character coding: the characters describing quantitative features (generally numbers of setae) generally performed worse than other types of characters under parsimony. An updated list of the known members of the Sensillanurini (Collembola: Neanuridae: Neanurinae) is presented.
© The Willi Hennig Society 2009.     

Integrative taxonomy approach for analysing evolutionary history of the tribe Euseiini Chant & McMurtry (Acari: Phytoseiidae)

Phylogenetic relationships within the mite Family Phytoseiidae are little known. The presently accepted classification is based on the opinion of specialists, but not on cladistics analysis. The present paper focuses on the tribe Euseiini, containing 271 species, three subtribes and 10 genera. It aims to determine phylogenetic relationships between these taxa and test their monophyly. Molecular analysis combining six markers has been carried out for taxa we succeeded in collecting. Morphological, biogeographic and ecological data have been analysed to determine how these factors can explain the evolutionary relationships emphasized on the phylogenetic tree. Those analyses have been carried out for the taxa available for the molecular study, but also for all species of the tribe. The tribe Euseiini and the two subtribes considered are monophyletic (at least considering the available taxa), supporting the present hypothesis on Phytoseiidae classification. However, the genus Iphiseius seems to not be valid and its unique species is included in the genus Euseius. Clades that were observed within the genus Euseius do not match with recent work on species groups within this genus. It seems that some morphological features such as an insemination apparatus shape and seta length on the dorsal shield constitute some elements explaining the clusters within the genus Euseius. Biogeographic and ecological data analysis led us to hypothesize a west Gondwanian origin of the tribe Euseiini (Africa and Neotropical areas) on Rosids plants (especially of the Orders Malpiphiales and Fabales: subclass Fabidae). Further analyses are still required to (i) take into account more taxa (especially rare ones and species from the Ethiopian part), (ii) to consider more accurate morphological features through more powerful microscopic apparatus, and (iii) to associate a phylogenetic and evolutionary scenario to life traits (pollen feeders).     

Comparative detailed morphology of the Heteroderinae Filip'ev & Schuurmans Stekhoven, 1941, sensu Luc et al. (1988): phylogenetic systematics and revised classification

Taxonomic schemes for the Heteroderinae Filip'ev & Schuurmans Stekhoven, 1941, sensu Luc et al., (1988) have been unstable due to the large number of genera and the paucity of known reliable characters. Reliable characters are essential when using phylogenetic inference in developing a natural classification. Morphological and developmental studies using light, scanning and transmission electron microscopy have revealed the new characters of host response, en face patterns, phasmid structure and female cuticular layers. These techniques also gave us insight into the homoplasy and polarity of many characters, revealed previously undetected character states and clarified misinterpreted character states. A matrix with the 19 most reliable characters is proposed for 20 operational taxonomic units (OTUs) and we employ this matrix for comparing computer generated phylogenetic analyses of the PHYLIP and PAUP packages. PAUP was deemed the more reliable parsimony algorithm for phylogenetic analysis of the Heteroderinae (Fink, 1986; Platnick, 1987). Monophyly of Atalodera + Sherodera + Thecavermiculatus (tribe Ataloderini), and Cactodera + Heterodera + Afenestrata, as well as Punctodera + Globodera + Dolichodera is supported by both programs. Most importantly, analyses strongly support monophyly of all cyst-forming genera (tribe Heteroderini) contrary to previous hypotheses of repeated evolution of the cyst (Wouts, 1985). In addition, monophyly of the Heteroderini with the Ataloderini is demonstrated. PAUP indicates monophyly of Sarisodera + Rhizonema + Bellodera + Hylonema and Ekphymatodera (tribe Sarisoderini new rank). Monophyly of the Sarisoderini was at first only weakly supported, but, subsequently, the reduced width of the submedial lips of second stage juveniles and males was recognized as a synapomorphy which strengthened subsequent PAUP trees and monophyly of the tribe. The present study rejects as paraphyletic or polyphyletic several previously proposed combinations, including Thecavermiculatus sequoiae (versus Rhizonema sequoiae), Sarisodera africana (versus Afenestrata africana), Dolichodera andinus (versus Thecavermiculatus andinus). The question whether T. andinus is a distinct genus, was not resolved due to insufficient data. PAUP supports our previous observations that Cactodera betulae is intermediate in a transformation series between other Cactodera and Heterodera: it also indicates these species as bring monophyletic with Heterodera + Afenestrata, but not with other Cactodera. Although these phylogenetic analyses strongly support some relationships, they indicate unresolved alternative hypotheses for others. Meloidodera (tribe Meloidoderini) and Cryphodera (tribe Cryphoderini) must be investigated for consideration of a possible synapomorphy not included in the present data matrix. Future studies are proposed to more clearly define the monophyly of the Heteroderini, as well as the Sarisoderini. Tests are also proposed to clarify questions of the monophyly of Verutus (tribe Verutini new rank) with the Heteroderinae versus other Tylenchida.     

Phylogenetic relationships of North American Antirrhinum (Veronicaceae)

Species of the genus Antirrhinum (Veronicaceae) provide excellent opportunities for research on plant evolution given their extensive morphological and ecological diversity. These opportunities are enhanced by genetic and developmental data from the model organism Antirrhinum majus. The genus Antirrhinum includes 15 New World species in section Saerorhinum and 21 Old World species in sections Antirrhinum and Orontium. Phylogenetic analyses of sequences of the internal transcribed spacer region (ITS) of nuclear ribosomal DNA were conducted for 19 Antirrhinum species, including all species from the New World, and 13 related genera in the tribe Antirrhineae. These analyses confirm the monophyly of Antirrhinum given the inclusion of the small genus Mohavea and exclusion of A. cyathiferum. The New World species, all of which are tetraploid, form a clade that is weakly supported as sister to the Old World sect. Orontium. The Old World species in sect. Antirrhinum form a well-supported clade that is sister to the remainder of the genus. In addition, both molecular and morphological data are used in the most comprehensive effort to date focused on recovering the phylogenetic relationships among the extremely diverse species in section Saerorhinum.     

Phylogenetic analysis of Cyprichromini (Perciformes: Cichlidae) endemic to Lake Tanganyika and validation of the genus <Emphasis Type="Italic">Paracyprichromis</Emphasis>

The phylogenetic relationships among two Paracyprichromis and five Cyprichromis species, included in the Tanganyikan cichlid tribe Cyprichromini, were investigated using morphological features. The previously proposed diagnostic characters of Paracyprichromis are not synapomorphies, because the nonelongated swim bladder is plesiomorphic, the numbers of dorsal and anal fin rays and scales on longitudinal line and around the caudal peduncle overlap with those of Cyprichromis, and these counts and number of vertebrae are all included within the ranges of other Tanganyikan cichlids. The monophyly of Paracyprichromis is supported by a unique condition of infraorbitals to this genus. Additionally, the monophyly of Cyprichromis was reconfirmed by one of the previously proposed diagnostic characters, the presence of an elongated swim bladder.     

Antler stiffness in moose (Alces alces): Correlated evolution of bone function and material properties?

The material properties of bone can vary considerably among skeletal elements from different parts of the body that serve different functions. However, functional demands placed on a specific type of skeletal element also can vary at a variety of scales, such as between different parts of the element, among individuals of a species, and across species. Variation in bone material properties might be correlated with differing functional demands at any of these scales. In this study we performed three-point bending tests on bone specimens extracted from antlers of moose (Alces alces) to test for three types of variation in bone material stiffness (Young's modulus): within the antler structure, between populations of moose, and between moose and other deer species. Because superficial portions of the antler are exposed to greater bending stress and strain than deeper portions, and because the antler beam (the basal shaft that attaches to the skull) is subjected to greater bending moments than more distal parts of the antler, we predicted that superficial bone and bone from the beam would be stiffer than bone from other parts of the antler. Instead, we identified no significant differences in these comparisons. There were also no significant differences in antler stiffness between moose from Michigan and the Yukon, even though the rapid growth required of antlers from northern latitudes like the Yukon has the potential to compromise bone material properties. However, moose have significantly stiffer antlers (11.6 +/- 0.45 GPa, mean +/- SE) than any other deer in the odocoileine lineage. Moreover, phylogenetic reconstructions of the evolution of antler stiffness in deer indicate a strong potential that high antler stiffness is a derived feature of moose. The unusual palmate shape of moose antlers likely subjects their antler beams to higher bending moments than found in other odocoileines, a factor that may have contributed to the evolutionary divergence of moose antler stiffness from that of other members of this clade. Although similarities in the mineral composition of bone across species likely limit the overall range of phylogenetic variation in bone material properties, our results demonstrate that evolutionary diversity in bone material properties can show correspondence with phylogenetic differences in mechanical or ecological demands on skeletal elements.