Phylogenetic relationships and biogeography of the Ipsiura cuckoo wasps (Hymenoptera: Chrysididae)

Phylogenetic studies addressing relationships among chrysidid wasps have been limited. There are no hypotheses proposed for the Neotropical lineages of Chrysidini other than the classic cladogram published in the 1990s by Kimsey and Bohart. Herein we present a cladistic analysis based on 64 morphological characters coded for 54 species of Chrysidini, 32 of them being Ipsiura and 22 representing Caenochrysis, Chrysis, Exochrysis, Gaullea, Neochrysis, and Pleurochrysis. The species of Ipsiura were recovered as monophyletic and as the sister clade of Neochrysis in all most parsimonious trees. We discuss the high plasticity of some morphological characters as evidenced by their high homoplasy in the phylogenetic results, and we clarify the main morphological changes inferred on the phylogenetic tree for this genus. The effects of the inferred homoplasy were evaluated under an implied weighting cladistic analysis, and from a probabilistic perspective with Bayesian inference. Those alternative strategies did not alter the general conclusions about the monophyly of Ipsiura or the generic relationships in Chrysidini (changes were noticed in the species‐level relationships within certain parts of Ipsiura, where low branch support was common across all approaches). Among the species groups proposed by Linsenmaier (1985), only the marginalis group was recovered as monophyletic. We also evaluated the convoluted biogeographic history of the group. The resulting historical reconstructions indicate a complicated scenario of diversification of these wasps in the Neotropics, and a close association with forested biomes is discussed.     

Morphology and metamorphosis of Eupsophus calcaratus tadpoles (Anura: Leptodactylidae)

Eupsophus calcaratus, a leptodactyloid frog from the austral Andean forests of Argentina and Chile, has endotrophic, nidicolous tadpoles. We studied a metamorphic series from Stages 31 to 46 of Gosner's developmental table (1960). Other than the scarce pigmentation, proportionately large eyes, and massive developing hindlimbs, the remaining external characters are similar to those of generalized, exotrophic larvae. At the same time, internal morphology does not reveal any character state attributable to the endotrophic-nidicolous way of life; conversely, structures such as the hyobranchial skeleton and the mandibular cartilages are similar to those of exotrophic-macrophagous tadpoles. The metamorphic process is characterized by the delayed development of diverse structures (e.g., ethmoid region, palatoquadrate, and hyobranchial apparatus), and the retention of some larval characters (e.g., parietal fenestrae, overall absence of ossification) with the absence of development of some "juvenile" characters (e.g., adult otic process, several bones) in metamorphosed individuals. These heterochronic processes and truncation of larval development are related to a shorter larval life (when compared to other species of the austral Andean region) and to the small size at metamorphosis.     

Congruence between comparative morphology and molecular phylogenies: Evolution of the male genital skeletal/muscular system in the subtribe Polyommatina (Lepidoptera,Lycaenidae)

The skeleton and musculature of male genitalia were studied in species of a model butterfly group (subtribe Polyommatina, Lycaenidae). In total, we analyzed 45 species of the tribe Polyommatini most of which were previously used in the molecular phylogenetic study (Talavera et al., 2013). The studied morphological characters were mapped on the molecular trees, which allowed us to reveal trends of morphological changes and to estimate the age of their origin. As a result, chronology of evolution of skeleton and musculature traits was established. It was shown that periods of slow morphological evolution alternated in the subtribe Polyommatina with those of a high rate of origin of new traits. For example, topography of the intravalvar muscles has not changed for 26 MY preserving their initial fan-shaped attachment. The evolution of intravalvar muscles started 10 MYA, proceeded slowly during the first 5 MY, and then accelerated during the last 5 MY resulting in the extensive splitting of the musculature in most monophyletic lineages. Mapping the morphological characters on the phylogeny demonstrated that the rates of skeleton and muscle evolution within the skeleton/musculature apparatus were different. In most cases the intravalvar musculature evolved much faster than the skeleton. The cladistic interpretation of states of morphological traits was found to be consistent with phylogenetic reconstructions based on analysis of multiple molecular markers. Moreover, morphological synapomorphies were found for the lineages Alpherakya + Glabroculus and Aricia + (Alpherakya + Glabroculus), which had low statistical support in molecular phylogenetic analysis. Additionally, in some cases molecular studies helped to reveal trends in the evolution of morphological traits. For example, the unpaired uncus and the compact juxta are not plesiomorphic for Cupidina as previously thought; instead, they were shown to have evolved secondarily within this subtribe.     

CLADISTIC ANALYSES OF THE COCCINELLIDAE (COLEOPTERA)

Abstract  Numerical cladistic method was used for analyzing the phylogeny of ladybirds, based upon adult and larval characters. The analysis of adult characters for 21 tribes, representing all subfamilies proposed by various authors, exhibited 18 equally parsimonious cladograms with high level of homoplasy. Five lineages were roughly revealed in the strict consensus tree: Sticholotidinae, Epilachninae, Coccinellinae (including Singhikalini), Coccidulinae (excluding Noviini) and Scymninae+Chilocorinae. The taxonomic interpretation of the adult analysis was similar to the classification proposed by Chazeauet al . (1990). Comparison of a preliminary analysis based on the larval characters for 14 tribes revealed only one component consistently supported in both analyses (Serangiini+Sukunahikonini). Larval analysis showed that Epi-lachinae is the most primitive and highly derived group of the Coccinellidae. Larval characters might be more satisfactory in revealing the phylogeny.     

A complex hyobranchial apparatus in a Cretaceous dinosaur and the antiquity of avian paraglossalia

The highly specialized feeding apparatus of modern birds is characterized in part by paraglossalia, triangular bones or cartilages in the tongue that constitute part of the rarely fossilized hyobranchial apparatus. Here, we report on a new, juvenile specimen of the ankylosaurid dinosaur Pinacosaurus grangeri Gilmore, 1933 that provides the first evidence of paraglossalia outside of crown group Aves. The specimen is remarkable in preserving a well‐ossified hyobranchial apparatus, including paired paraglossalia, first and second ceratobranchials, epibranchials, and evidence of a median cartilaginous basihyal. Reassessment of Edmontonia, another ankylosaur, also reveals evidence of bony paraglossalia. Ankylosaur paraglossalia closely resemble those of birds, but are relatively larger and bear prominent muscle scars, supporting the hypothesis that ankylosaurs had fleshy, muscular tongues. The other hyobranchial elements, surprisingly, resemble those of terrestrially feeding salamanders. Ankylosaurs had reduced, slowly replacing teeth, as evidenced from dental histology, suggesting that they relied greatly on their tongues and hyobranchia for feeding. Some curved, rod‐like elements of other dinosaur hyobranchia are reinterpreted as second ceratobranchials, rather than first ceratobranchials as commonly construed. Ankylosaurs provide rare fossil evidence of deep homology in vertebrate branchial arches and expose severe biases against the preservation and collection of the hyobranchial apparatus. In light of these biases, we hypothesize that paraglossalia were present in the common ancestor of Dinosauria, indicating that some structures of the highly derived avian feeding apparatus were in place by the Triassic Period. © 2015 The Linnean Society of London     

五种蝌蚪口器及舌鳃骨的结构比较

采用体视显微镜和骨骼双染色法对5种不同栖息环境的无尾两栖动物蝌蚪的口器和舌鳃骨的形态结构特征进行了观察。5种蝌蚪口器由唇齿行、唇乳突和角质颌等组成。舌鳃骨是由关联骨Ⅰ、关联骨Ⅱ、角舌骨、舌鳃骨盘和角鳃骨等骨骼组成。蝌蚪的梅氏软骨若较发达,其摄食方式可能为刮食;蝌蚪的舌鳃骨发达,其摄食方式则可能为滤食。角质颌、唇齿以及角鳃骨上鳃耙的出现显著增强了蝌蚪主动摄食能力和对食物与非食物的主动选择性。     

Phylogeny of the sea spiders (Arthropoda, Pycnogonida) based on direct optimization of six loci and morphology

Higher‐level phylogenetics of Pycnogonida has been discussed for many decades but scarcely studied from a cladistic perspective. Traditional taxonomic classifications are yet to be tested and affinities among families and genera are not well understood. Pycnogonida includes more than 1300 species described, but no systematic revisions at any level are available. Previous attempts to propose a phylogeny of the sea spiders were limited in characters and taxon sampling, therefore not allowing a robust test of relationships among lineages. Herein, we present the first comprehensive phylogenetic analysis of the Pycnogonida based on a total evidence approach and Direct Optimization. Sixty‐three pycnogonid species representing all families including fossil taxa were included. For most of the extant taxa more than 6 kb of nuclear and mitochondrial DNA and 78 morphological characters were scored. The most parsimonious hypotheses obtained in equally weighted total evidence analyses show the two most diverse families Ammotheidae and Callipallenidae to be non‐monophyletic. Austrodecidae + Colossendeidae + Pycnogonidae are in the basal most clade, these are morphologically diverse groups of species mostly found in cold waters. The raising of the family Pallenopsidae is supported, while Eurycyde and Ascorhynchus are definitely separated from Ammotheidae. The four fossil taxa are grouped within living Pycnogonida, instead of being an early derived clade. This phylogeny represents a solid framework to work towards the understanding of pycnogonid systematics, providing a data set and a testable hypothesis that indicate those clades that need severe testing, especially some of the deep nodes of the pycnogonid tree and the relationships of ammotheid and callipallenid forms. The inclusion of more rare taxa and additional sources of evidence are necessary for a phylogenetic classification of the Pycnogonida. © The Willi Hennig Society 2006.     

Cladistic analysis of the Sepsidae (Cyclorrhapha: Diptera) based on a comparative scanning electron microscopic study of larvae

The result of a phylogenetic analysis of the Sepsidae based on larval characters is presented. It is shown that cyclorrhaphan larvae can be as rich a source of characters as Nematocera immatures when investigated using an SEM. The cladistic analysis comprised fifty-two species in sixteen genera of the Sepsidae and five outgroup species and used fifty-seven morphological characters. It found seven parsimonious trees which only differed with respect to the arrangement of some species within the genus Themira. The basal dichotomies of the phylogenetic trees are particularly well supported, indicating the conservative nature of larval characters. Orygma is confirmed as the sister group of all the remaining sepsids, the Sepsinae. There is good larval evidence that Ortalischema is the sister group of all remaining Sepsinae and that the Toxopodinae constitute an early radiation within the Sepsidae. According to larval data, some genera are paraphyletic ( Themira, Palaeosepsis ), but adult characters appear to contradict these findings. Among the traditionally recognized higher taxa within the Sepsidae, Hennig's Themira species-group and Steysbal's Sepsini have to be rejected as polyphyletic. However, Hennig's Sepsis species-group is confirmed as monophyletic and will probably constitute one major element of a future phylogenetic system of the Sepsidae. States of the strongly modified fore-legs of some adult sepsid males are mapped onto the phylogenetic tree, largely confirming Šulc's ideas about the evolution of these features. The origin and evolution of male sternites with brushes and a gland on the tibiae of the males ('osmeterium') are discussed. Whereas adult characters point to a sister-group relationship between the Sepsidae and the Ropalomeridae, larval characters appear to indicate a sister-group relationship between the Coelopidae and the Sepsidae. The evidence for both hypotheses is critically evaluated.     

Gross morphology and topographical relationships of the hyobranchial apparatus and laryngeal cartilages in the ostrich (Struthio camelus)

The ostrich hyobranchial apparatus consists of the centrally positioned paraglossalia and basiurohyale and paired caudo‐lateral elements (horns), each consisting of the ceratobranchiale and epibranchiale. The paraglossalia lie within the tongue parenchyma and consist of paired, flat, caudo‐laterally directed cartilages joined rostrally. The basiurohyale forms a single dorso‐ventrally flattened unit composed of an octagonal‐shaped body from which extend rostral (the rostral process) and caudal (the urohyale) projections. The laryngeal skeleton consists of cricoid, procricoid and paired arytenoid cartilages. The large ring‐shaped cricoid cartilage displays a body and paired wings which articulate with each other and with the procricoid. The blunt, ossified, rostral projection of the cricoid and the scalloped nature of the arytenoid cartilages are unique to the ostrich. The procricoid is a single structure which links the paired arytenoids and wings of the cricoid. The hyobranchial apparatus is firmly attached to the tongue parenchyma and to the larynx and proximal trachea. In contrast to previous reports in this species, the horns of the hyobranchial apparatus are not related to the skull. Ossification of the body of the basihyale, the ceratobranchials and the rostral process and body of the cricoid cartilage of the larynx lends stability to these structures.     

Phylogenetic relationships of toads of the Rhinella granulosa group (Anura: Bufonidae): a molecular perspective with comments on hybridization and introgression

The Rhinella granulosa group consists of 13 species of toads distributed throughout open areas of South America and Panama. In this paper we perform a phylogenetic analysis considering all but one species of the group, employing five nuclear and four mitochondrial genes, for up to 7910 bp per specimen. Separate phylogenetic analyses under direct optimization (DO) of nuclear and mitochondrial sequences recovered the R. granulosa group as monophyletic and revealed topological incongruence that can be explained mainly by multiple events of hybridization and introgression, both mitochondrial and nuclear. The DO combined analysis, after the exclusion of putatively introgressed or heterozygous genomes, resulted in a phylogenetic hypothesis for the R. granulosa group in which most of the species are recovered as monophyletic, but with interspecific relationships poorly supported. The optimization of morphological (adult and larval), chromosomal, and behavioural characters resulted in 12 putative phenotypic synapomorphies for this species group and some other synapomorphies for internal clades. Our results indicate the need for additional population genetic studies on R. dorbignyi and R. fernandezae to corroborate the taxonomic status of both taxa. Finally, we discuss biological and genetic characteristics of Bufonidae, as possible explanations for the common occurrence of hybridization and introgression observed in some lineages of this family.     

A cladistic analysis of the nomadine bees (Hymenoptera: Apoidea)

Abstract. This study compares the results of Rozen's cladistic analysis of the larvae of fifteen genera of cleptoparasitic bees in the subfamily Nomadinae with an independent data set of adult characters for the same genera. Adult characters exhibited considerably higher levels of homoplasy and poorer resolution of cladistic relationships, with multiple equally parsimonious cladograms. However, comparison of a Nelson consensus tree based on adult characters with the cladogram based on larval characters reveals three components consistently supported in both analyses (the tribes Epeolini and Ammobatini, and Neopasites + Neolarra) , one component supported only by adult characters (Isepeolus + Protepeolus) , and one terminal component supported only by larval characters (Nomada + Ammobatini), as well as several more inclusive groupings based on larval characters that are difficult to compare with the adult consensus tree because it shows so much less resolution. When adult and larval characters are combined in a single data matrix, the resulting cladogram closely resembles the cladogram based on larval characters alone, although levels of homoplasy are considerably higher than in the larval analysis.
A preliminary analysis of adult characters for thirty-four genera in the Nomadinae also exhibited high levels of homoplasy and very large numbers of equally parsimonious cladograms. Nevertheless, certain consistent monophyletic groupings, most notably the Epeolini and Ammobatini, were also supported in this analysis. The one currently recognized tribe whose monophyly has received no support from any analysis is the Nomadini.
The relevance of these phylogenetic hypotheses to our understanding of host associations and variable features of egg morphology and oviposition behaviour in nomadine bees is briefly discussed.     

CLADISTIC ANALYSIS OF METAZOAN RELATIONSHIPS: A REAPPRAISAL

Abstract— A recently published cladistic analysis of metazoan relationships based on 77 embryological and morphological characters is shown to be defective with respect to both methodological issues and the interpretation of certain characters. Consequently, many conclusions of this analysis are not supported by the data. Alternative phylogenetic hypotheses are therefore proposed, based on a re-evaluation of an adapted data set.     

A comparative study of the hyobranchial apparatus in Hynobiidae (Amphibia: Urodela)

The morphology of the adult hyobranchial apparatus has played an important role in understanding the systematics and evolution of urodeles, but the hyobranchial apparatus of hynobiid salamanders has received little attention so far. In this study, the hyobranchial apparatus of eight hynobiid salamanders (Hynobius leechii, Onychodactylus zhangyapingi, Ranodon sibiricus, Batrachuperus pinchonii, Salamandrella keyserlingii, Liua shihi, Pachyhynobius shangchengensis and Pseudohynobius flavomaculatus) is described and compared based on the clearing and double-staining method. The basic elements of the hyobranchial apparatus of the eight species are similar, including one basibranchial, cornua, one pair of radial loops, one pair of ceratohyals, one pair of hypobranchials II, one pair of ceratobranchials II, one urohyal (absent in O. zhangyapingi), one pair of the complex of hypobranchial I and ceratobranchial I (separated in certain species). Although the hyobranchial apparatus is similar among hynobiid salamanders and shows a unique morphological pattern, there are also certain species-specific distinctions that may be used for specific or generic diagnosis. The results of an ancestral state reconstruction of five traits showed that the ossified basibranchial, the presence of a separated hypobranchial I and ceratobranchial I, the absence of a urohyal, the ossified hypobranchial I and the partially ossified ceratohyal are derived traits. The state shown by the traits of each species is consistent with the phylogenetic position of each species. Compared with other Urodela, the hyobranchial apparatus of this group shows certain distinctive features that may represent the diagnostic characters of the family Hynobiidae. The partially ossified ceratohyal is correlated with the habitat and represents an ecological adaptation.     

Phylogenetic analysis of the Pinnotheridae (Crustacea, Brachyura) based on larval morphology, with emphasis on the Dissodactylus species complex

Comparative larval morphology was used to elucidate phylogenetic relationships within the Pinnotheridae and the Dissodactylus species complex. Within the family, seven zoeal and six megalopal characters suggested two equally parsimonious phylogenetic hypotheses for pinnotherid larvae, both with Ostracotheres tridacnae representing the sister group for the Dissodactylus complex. Results indicated that the genus Pinnotheres is a polyphyletic taxon, and that the traditional subfamilial arrangement comprises paraphyletic taxa within the subfamilies Pinnotherinae and Pinnothereliinae. Certain evidence has suggested that Fabia and Juxtafubia should be excluded from the Pinnotherinae and placed into the Pinnothereliinae. Larval and adult morphology suggested that Pinnotheres politus should be included within Tumidotheres. The phylogenetic analysis within the Dissodactylus complex involved one zoeal and 16 megalopal characters. Results suggested a single phylogenetic hypothesis based on larval morphology. Combining adult morphology with larval evidence resulted in two equally parsimonious phylogenetic hypotheses, one of which agreed with a previously suggested hypothesis based only on adult characters.     

Comparative myology of the forelimb of Liolaemus sand lizards (Liolaemidae)

The lizard genus Liolaemus includes numerous constituent clusters of putatively related taxa, one of which is the Liolaemus boulengeri group, which in turn includes the sand lizards (of the Liolaemus wiegmannii subgroup). Members of the sand lizard group exhibit three different modes of burying into sand. The general morphology of the forelimb muscles of those Liolaemus species is analysed. Herein, we present a study of the forelimb musculature of all species considered by Halloy et al. (1998). This study has three principal goals. First, we are seeking myological characters that will be useful in formulating phylogenetic hypothesis about the species of Liolaemus. With these characters, we also wish to compile morphological data that represent the morphological space implied in the diverse locomotor behaviours of these animals. Second, we are looking for derived features that reflect functional changes in the use of forelimb. Third, we wish to provide a cladistic analysis that can be used to test phylogenetic hypothesis derived from other sources of data. We present 48 characters in a data set and analyse it cladistically. We obtained a hypothesis of relationships of the Liolaemus species and compared this with previous hypotheses based on other characters. The trees obtained are not congruent with previously proposed phylogenies. We were unable to identify in our trees nodes that are based on structures reflecting functional changes in the use of the forelimb. The morphological similarities in the forelimb musculature of all species analysed seems to conform a very conservative general anatomical pattern with which Liolaemus sand lizards perform most of their locomotor behaviours.     

Discovery of the Quedius antipodum Sharp larva from New Zealand: phylogenetic test of larval morphology for Staphylinini at the intratribal level (Coleoptera: Staphylinidae)

Quedius antipodum Sharp is an endemic species from New Zealand. Here we describe its larva, the first of the species‐rich group of the south temperate ‘Quedius’ spp. This finding throws light on the controversy between the conventional systematics of Quedius Stephens and newer phylogenetic analyses, both of which are based on non‐larval characters only. We compare the larva of Q. antipodum with those of the north temperate Quedius (Quediina), where it was traditionally placed, and with the known larvae of Amblyopinina, a group where Q. antipodum was placed by recent phylogenetic studies. Sister‐group relationships of Q. antipodum within the tribe Staphylinini are explored based on larvae by means of parsimony analysis: 77 morphological characters scored for 20 species from 17 genera. Consistent with the adult morphology and DNA sequences, larvae‐based cladistic analysis confirms that Q. antipodum should not be placed in the north temperate genus Quedius. However, larval analysis alone remains dubious with respect to finding the exact sister relationships of that species.     

Using Phylogenies to Study Convergence: The Case of the Ant-Eating Mammals

Identifying when homoplasy is due to convergence requires confidencein trees and precise analysis of potentially convergent characters.Some features of mammals that eat mostly ants and termites areused as examples of convergence; the most speciose assemblagesof these mammals are in the orders Xenarthra and Pholidota.My studies on cranial muscles in xenarthrans and pholidotansaim to 1) precisely describe the anatomy in ant-eating and non-ant-eatinglineages, 2) assess variation among ant-eating lineages, and3) compare the most derived conditions (in xenarthran anteatersand pholidotan pangolins). These data clarify the nature ofmorphological adaptation in ant-eating mammals, and when combinedwith accumulating phylogenetic studies, allow us to distinguishfeatures that have evolved convergently from those that arevariable but not correlated with diet. Interpreting the extremesimilarity in anteaters and pangolins remains problematic dueto lingering disagreement among phylogenetic hypotheses. Prevailingopinion favors interpretation of these similarities as convergent.     

Development of the nidicolous tadpoles of Eupsophus emiliopugini (Anura: Cycloramphidae) until metamorphosis,with comments on systematic relationships of the species and its endotrophic developmental mode

Vera Candioti, M.F., Nuñez, J.J. and Úbeda, C. 2011. Development of the nidicolous tadpoles of Eupsophus emiliopugini (Anura: Cycloramphidae) until metamorphosis, with comments on systematic relationships of the species and its endotrophic developmental mode. —Acta Zoologica (Stockholm) 92 : 27–45. Species of Eupsophus are unique within Alsodinae in having nidicolous tadpoles. They are characterized by traits typical of generalized exotrophic (e.g., oral disc and spiracular tube) and endotrophic larvae (e.g., scant pigmentation and large hind limbs). The larval morphology and development of E. emiliopugini, including external, buccal, and musculoskeletal features, is described herein. Like the larvae of other alsodines, these larvae have four lingual and four infralabial papillae, quadratoethmoid process, and an m. rectus cervicis with a double insertion. Among the traits exclusive to the genus are: the absence of the pseudopterygoid process and quadrato‐orbital commissure; presence of the m. subarcualis rectus I with two slips; and presence of the m. subarcualis rectus II–IV inserting on Ceratobranchial II. The development and metamorphosis of Eupsophus include some characters that develop later (e.g., degeneration of mouthparts and chondrocranium with minimum calcification), characters that develop earlier (e.g., hind‐limb appearance and jaw and suspensorium ossification), and characters that develop at the same time (e.g., most external features and cranial muscles) than in most exotrophic species. Some distinctive characters (third lower labial ridge absent, general configuration of the hyobranchial skeleton, skeletal development with retention of larval traits) resemble those of other endotrophic species, and the precocious ossification of jaws and suspensorium is shared with several direct‐developing species among recent amphibians.