Evolution of the syrinx of Apodiformes, including the vocal-learning Trochilidae (Aves: Strisores)
The vocal organ of birds, the syrinx, represents a key innovation in the evolutionary history of vertebrate communication. Three major avian clades: passerines, parrots, and hummingbirds, independently acquired both specialized syringeal structures and vocal-production learning, between which a functional relationship has been proposed but remains poorly understood. In hummingbirds, the syrinx has never been studied comparatively alongside non-learning relatives in the parent clade Strisores. Here we describe the anatomy of the syrinx in three swift species using enhanced-contrast computed tomography, which reveals structures previously unreported in the clade. We also tested for correlations between syringeal and acoustic traits in a sample of hummingbirds and swifts using phylogenetically informed regressions. The swift syrinx presents lateral labia located on the first pair of bronchial half-rings, which are present in hummingbirds and may be ancestral to Strisores. The further enlarged lateral labia of hummingbirds are found to be significantly correlated to the reduction in length of their trachea and m. tracheolateralis. Acquisition of intrinsic muscles and loss of the sternotrachealis muscle co-occur with these shifts. We recover a significant negative correlation between tracheal elongation and maximum vocalization frequency, suggesting that tracheal shortening in hummingbirds facilitated the acquisition of high-frequency vocalizing.
A cryptic radiation of Caribbean sea slugs revealed by integrative analysis: '' (Sacoglossa: Caliphyllidae) is six distinct species
Integrative studies have revealed cryptic radiations in several Caribbean lineages of heterobranch sea slugs, raising questions about the evolutionary mechanisms that promote speciation within the tropical Western Atlantic. Bergh, 1871 is a genus comprising 12 named species in the family Caliphyllidae that lack the photosynthetic ability of other sacoglossans but are noted for vibrant colours on the large cerata (dorsal leaf-like appendages) that characterize many species. Two species are widely reported from the Caribbean: (Trinchese, 1881) and Engel, 1927. Here, we present an integrative assessment of diversity in Caribbean . Four methods of molecular species delimitation supported seven species in samples from the Caribbean and adjacent subtropical Western Atlantic. Six delimited species formed a monophyletic lineage in phylogenetic analyses but were > 9% divergent at the barcoding locus and could be differentiated using ecological, reproductive and/or morphological traits. We redescribe , a senior synonym for the poorly known Ortea & Templado, 1988, and describe five new species. Evolutionary shifts in algal host use, penial armature and larval life history might have acted synergistically to promote the rapid divergence of endemic species with restricted distributions in this radiation, substantially increasing global diversity of the genus.
Jumping continents and major host lineages: phylogeny and diversity of the enigmatic Cryptotropidae (Platyhelminthes: Digenea)
Members of several genera in the digenean superfamily Microphalloidea, namely , , , and , are characterized by an unusual dorsal position of the genital atrium. In the absence of phylogenetic data, their systematic position has been unstable. In the most recent taxonomic revision of the Microphalloidea, they were housed in three different families. We studied the morphology and obtained DNA sequences of several microphallolideans with a dorsal genital pore, collected from vertebrates in Southeast Asia and South America. We used sequences of the nuclear 28S gene to infer a superfamily-level phylogeny and sequences of the mitochondrial gene for family-level phylogeny and species-level comparisons. Based on the combination of molecular phylogenetic data and morphological features, we restore the Cryptotropidae and synonymize the Renschetrematidae with the Cryptotropidae. We erect new genera and , provide amended diagnoses of the Cryptotropidae and and provide keys to the identification of genera within the family. We resurrect the genus , previously synonymized with , and remove from . We describe four new species from lizards in Vietnam and birds in the Philippines and Peru. The unique morphological features and unusually broad host associations and geographical distributions of cryptotropids are discussed.
Molecular phylogeny supports invalidation of and (Digenea: Diplostomidae) and reveals a from mammals
, and are among genera of diplostomid digeneans known to parasitize mammalian definitive hosts. Despite numerous recent molecular phylogenetic studies of diplostomids, limited DNA sequence data is available from diplostomids parasitic in mammals. Herein, we provide the first 28S rDNA and mtDNA sequences from morphologically identified, adult specimens of and . Newly generated 28S sequences were used to infer the phylogenetic interrelationships of these two genera among other major lineages of diplostomoideans. The phylogeny based on 28S and a review of morphology clearly suggests that should be considered a junior synonym of , while should be considered a junior synonym of . (type species), and were transferred into as , and ; (type species) and were transferred into as and In addition, (formerly included in ) was restored and transferred into based on a morphological study of well-fixed, adult specimens and the comparison of DNA sequences among spp. The diplostomid genus was restored based on review of morphology.
Burmese amber reveals a new stem lineage of whirligig beetle (Coleoptera: Gyrinidae) based on the larval stage
Burmese amber is well known for preserving unique extinct lineages of insects. Here, we describe a new fossil beetle in its larval stage from Burmese amber. Bayesian and parsimony phylogenetic analysis of 50 morphological characters support this fossil as being sister to both the tribes Dineutini and Orectochilini, representing an extinct stem lineage in Gyrininae. It is described here as a new genus and species of whirligig beetle, , a taxon that preserves remarkable intermediate features between the whirligig beetle tribe Gyrinini and the crown Orectochilini and Dineutini. This new taxon preserves key features for studying the evolution of characters within the larval stage of the Gyrinidae and highlights the importance of Burmese amber for preserving both stem and crown lineages present during the mid-Cretaceous, before the end-Cretaceous mass extinction event.
Molecular and morphological systematics of Gray, 1847 (Mollusca: Gastropoda: Heterobranchia: Aplysiomorpha)
Molecular and morphological data from newly collected specimens and a review of the literature and type material indicate that the widespread tropical sea hare is a species complex of five genetically distinct taxa. The name is retained for a widespread species in the Indo-Pacific tropics. is endemic to the eastern Pacific. , and a new species described herein are restricted to the tropical Atlantic, with partially overlapping ranges in the Caribbean region and St. Helena. The temperate Pacific species is also distinct and endemic to temperate southeastern Australia and New Zealand. These species of constitute highly divergent lineages and most contain unique internal anatomical characteristics, particularly in the male reproductive morphology and shell shape, making them relatively easy to identify upon dissection. However, externally all these species are extremely variable in colour pattern and morphology and are virtually indistinguishable. This is particularly problematic for identification in the Atlantic Ocean where three species co-occur in the Caribbean region.
The first definitive Middle Jurassic atoposaurid (Crocodylomorpha, Neosuchia), and a discussion on the genus
Atoposaurids were a clade of semiaquatic crocodyliforms known from the Late Jurassic to the latest Cretaceous. Tentative remains from Europe, Morocco, and Madagascar may extend their range into the Middle Jurassic. Here we report the first unambiguous Middle Jurassic (late Bajocian-Bathonian) atoposaurid: an anterior dentary from the Isle of Skye, Scotland, UK. A comprehensive review of atoposaurid specimens demonstrates that this dentary can be referred to based on several derived characters, and differs from the five previously recognized species within this genus. Despite several diagnostic features, we conservatively refer it to sp., pending the discovery of more complete material. As the oldest known definitively diagnostic atoposaurid, this discovery indicates that the oldest members of this group were small-bodied, had heterodont dentition, and were most likely widespread components of European faunas. Our review of mandibular and dental features in atoposaurids not only allows us to present a revised diagnosis of , but also reveals a great amount of variability within this genus, and indicates that there are currently five valid species that can be differentiated by unique combinations of dental characteristics. This variability can be included in future broad-scale cladistics analyses of atoposaurids and closely related crocodyliforms, which promise to help untangle the complicated taxonomy and evolutionary history of Atoposauridae. © 2015 The Authors. Zoological Journal of the Linnean Society published by John Wiley & Sons Ltd on behalf of The Linnean Society of London.
Phylogeny of the deep-sea cirripede family Scalpellidae (Crustacea, Thoracica) based on shell capitular plate morphology
A cladistic analysis of 23 extant species of the deep-sea pedunculate cirripede family Scalpellidae was undertaken, based on 61 shell plate characters, and taking the Jurassic-Cretaceous scalpellomorph genus as an out-group. The consensus tree shows progressive morphological change from basal to more derived taxa, but a derived group is marked by major morphological innovation, including 27 character state changes that permit subdivision of the family into two sharply demarcated clades - the more basal group is here placed within a redefined Scalpellinae ( , , , , , and ), and a more derived group named that shows numerous progressive trends in morphology, permitting the recognition of three genera ( , , and ). The phylogeny is independently supported by a recently published multiple DNA marker-based molecular phylogeny. The more basal Scalpellinae appeared in the Aptian (Early Cretaceous, 120 Mya), and derived Amigdoscalpellinae were already present by the Campanian (Late Cretaceous, 78 Mya), represented by and from the UK Chalk. Specialized receptacles to accommodate dwarf males in the apical interior of the scutum evolved at least three times during the history of the scalpellids. © 2015 The Linnean Society of London.
Two new cryptic and sympatric species of the king crab parasite Briarosaccus (Cirripedia: Rhizocephala) in the North Pacific
Rhizocephalan barnacles have been reported to parasitize a wide range of king crab species (Lithodidae). So far all these parasites have been assigned to a single species, Boschma, 1930, which is assumed to have a global distribution. Here we investigate specimens from three different king crab hosts from the fjord systems of Southeastern Alaska: Benedict, 1895, (Tilesius, 1815), and (Brandt, 1850). Using molecular markers and by morphological comparison we show that specimens from these three commercial exploited king crabs are in fact morphologically distinct from , and further represent two separate species which we describe. The two new species, n. sp. and n. sp., are cryptic by morphological means and were identified as distinct species by the use of genetic markers (COI and 16S). They occur sympatrically, yet no overlap in king crab hosts occurs, with n. sp. only found on , and n. sp. as parasite of the two hosts.
Remarkable ancient divergences amongst neglected lorisiform primates
Lorisiform primates (Primates: Strepsirrhini: Lorisiformes) represent almost 10% of the living primate species and are widely distributed in sub-Saharan Africa and South/South-East Asia; however, their taxonomy, evolutionary history, and biogeography are still poorly understood. In this study we report the largest molecular phylogeny in terms of the number of represented taxa. We sequenced the complete mitochondrial cytochrome gene for 86 lorisiform specimens, including ∼80% of all the species currently recognized. Our results support the monophyly of the Galagidae, but a common ancestry of the Lorisinae and Perodicticinae (family Lorisidae) was not recovered. These three lineages have early origins, with the Galagidae and the Lorisinae diverging in the Oligocene at about 30 Mya and the Perodicticinae emerging in the early Miocene. Our mitochondrial phylogeny agrees with recent studies based on nuclear data, and supports as the oldest galagid lineage and the polyphyletic status of . Moreover, we have elucidated phylogenetic relationships for several species never included before in a molecular phylogeny. The results obtained in this study suggest that lorisiform diversity remains substantially underestimated and that previously unnoticed cryptic diversity might be present within many lineages, thus urgently requiring a comprehensive taxonomic revision of this primate group.
An integrative study of Thomson (Hymenoptera: Eulophidae) associated with invasive pests in Europe and North America: taxonomic and ecological implications
The species of attacking two invasive pests of tomato and canola in Europe and North America, respectively, (Meyrick) (Lepidoptera: Gelechiidae) and (Marsham) (Coleoptera: Curculionidae), have been revised using an integrative taxonomy approach. Molecular data from the mitochondrial cytochrome oxidase subunit I and the nuclear D2 expansion region of the 28S ribosomal subunit and internal transcribed spacer 2, the discovery of new morphological features, and study of type material resulted in the delineation of three species groups, the , , and groups, the discovery of four new species, and the resurrection of three taxa from synonymy. Lectotypes have been designated for 13 species originally described in by Walker. Although has not been revised, an illustrated key is given to differentiate 23 recognized European species. The key, type images, and treatments of the three species groups will enable more accurate identification of the valid species of in the future. They will also benefit biological control practitioners of pest species. The ecological consequences of the new taxonomic concepts are discussed.
The characters of Palaeozoic jawed vertebrates
Newly discovered fossils from the Silurian and Devonian periods are beginning to challenge embedded perceptions about the origin and early diversification of jawed vertebrates (gnathostomes). Nevertheless, an explicit cladistic framework for the relationships of these fossils relative to the principal crown lineages of the jawed vertebrates (osteichthyans: bony fishes and tetrapods; chondrichthyans: sharks, batoids, and chimaeras) remains elusive. We critically review the systematics and character distributions of early gnathostomes and provide a clearly stated hierarchy of synapomorphies covering the jaw-bearing stem gnathostomes and osteichthyan and chondrichthyan stem groups. We show that character lists, designed to support the monophyly of putative groups, tend to overstate their strength and lack cladistic corroboration. By contrast, synapomorphic hierarchies are more open to refutation and must explicitly confront conflicting evidence. Our proposed synapomorphy scheme is used to evaluate the status of the problematic fossil groups Acanthodii and Placodermi, and suggest profitable avenues for future research. We interpret placoderms as a paraphyletic array of stem-group gnathostomes, and suggest what we regard as two equally plausible placements of acanthodians: exclusively on the chondrichthyan stem, or distributed on both the chondrichthyan and osteichthyan stems.
Overview of the taxonomy of zooxanthellate Scleractinia
Coral taxonomy has entered a historical phase where nomenclatorial uncertainty is rapidly increasing. The fundamental cause is mandatory adherence to historical monographs that lack essential information of all sorts, and also to type specimens, if they exist at all, that are commonly unrecognizable fragments or are uncharacteristic of the species they are believed to represent. Historical problems, including incorrect subsequent type species designations, also create uncertainty for many well-established genera. The advent of studies in the 1970s revealed these issues; now molecular technology is again changing the taxonomic landscape. The competing methodologies involved must be seen in context if they are to avoid becoming an additional basis for continuing nomenclatorial instability. To prevent this happening, the International Commission on Zoological Nomenclature (ICZN) will need to focus on rules that consolidate well-established nomenclature and allow for the designation of new type specimens that are unambiguous, and which include both skeletal material and soft tissue for molecular study. Taxonomic and biogeographic findings have now become linked, with molecular methodologies providing the capacity to re-visit past taxonomic decisions, and to extend both taxonomy and biogeography into the realm of evolutionary theory. It is proposed that most species will ultimately be seen as operational taxonomic units that are human rather than natural constructs, which in consequence will always have fuzzy morphological, genetic, and distribution boundaries. The pathway ahead calls for the integration of morphological and molecular taxonomies, and for website delivery of information that crosses current discipline boundaries.
The phylogenetic significance of colour patterns in marine teleost larvae
Ichthyologists, natural-history artists, and tropical-fish aquarists have described, illustrated, or photographed colour patterns in adult marine fishes for centuries, but colour patterns in marine fish larvae have largely been neglected. Yet the pelagic larval stages of many marine fishes exhibit subtle to striking, ephemeral patterns of chromatophores that warrant investigation into their potential taxonomic and phylogenetic significance. Colour patterns in larvae of over 200 species of marine teleosts, primarily from the western Caribbean, were examined from digital colour photographs, and their potential utility in elucidating evolutionary relationships at various taxonomic levels was assessed. Larvae of relatively few basal marine teleosts exhibit erythrophores, xanthophores, or iridophores (i.e. nonmelanistic chromatophores), but one or more of those types of chromatophores are visible in larvae of many basal marine neoteleosts and nearly all marine percomorphs. Whether or not the presence of nonmelanistic chromatophores in pelagic marine larvae diagnoses any major teleost taxonomic group cannot be determined based on the preliminary survey conducted, but there is a trend toward increased colour from elopomorphs to percomorphs. Within percomorphs, patterns of nonmelanistic chromatophores may help resolve or contribute evidence to existing hypotheses of relationships at multiple levels of classification. Mugilid and some beloniform larvae share a unique ontogenetic transformation of colour pattern that lends support to the hypothesis of a close relationship between them. Larvae of some tetraodontiforms and lophiiforms are strikingly similar in having the trunk enclosed in an inflated sac covered with xanthophores, a character that may help resolve the relationships of these enigmatic taxa. Colour patterns in percomorph larvae also appear to diagnose certain groups at the interfamilial, familial, intergeneric, and generic levels. Slight differences in generic colour patterns, including whether the pattern comprises xanthophores or erythrophores, often distinguish species. The homology, ontogeny, and possible functional significance of colour patterns in larvae are discussed. Considerably more investigation of larval colour patterns in marine teleosts is needed to assess fully their value in phylogenetic reconstruction.
A new genus of miniaturized and pug-nosed gecko from South America (Sphaerodactylidae: Gekkota)
Sphaerodactyl geckos comprise five genera distributed across Central and South America and the Caribbean. We estimated phylogenetic relationships among sphaerodactyl genera using both separate and combined analyses of seven nuclear genes. Relationships among genera were incongruent at different loci and phylogenies were characterized by short, in some cases zero length, internal branches and poor phylogenetic support at most nodes. We recovered a polyphyletic Coleodactylus, with Coleodactylus amazonicus being deeply divergent from the remaining Coleodactylus species sampled. The C. amazonicus lineage possessed unique codon deletions in the genes PTPN12 and RBMX while the remaining Coleodactylus species had unique codon deletions in RAG1. Topology tests could not reject a monophyletic Coleodactylus, but we show that short internal branch lengths decreased the accuracy of topology tests because there were not enough data along short branches to support one phylogenetic hypothesis over another. Morphological data corroborated results of the molecular phylogeny, with Coleodactylus exhibiting substantial morphological heterogeneity. We identified a suite of unique craniofacial features that differentiate C. amazonicus not only from other Coleodactylus species, but also from all other geckos. We describe this novel sphaerodactyl lineage as a new genus, Chatogekko gen. nov. We present a detailed osteology of Chatogekko, characterizing osteological correlates of miniaturization that provide a framework for future studies in sphaerodactyl systematics and biology.
Molecular systematics of the Philippine forest skinks (Squamata: Scincidae: ): testing morphological hypotheses of interspecific relationships
Skinks of the genus are the most diverse clade of squamates in the Philippine Archipelago. Morphological examination of these species has defined six phenotypic groups that are commonly used in characterizations of taxonomic hypotheses. We used a molecular phylogeny based on four mitochondrial and two nuclear genes to assess the group's biogeographical history in the archipelago and examine the phylogenetic validity of the currently recognized Philippine species groups. We re-examined traditional characters used to define species groups and used multivariate statistics to quantitatively evaluate group structure in morphometric space. Clustering analyses of phenotypic similarity indicate that some (but not all) members of previously defined species groups are phenotypically most similar to other members of the same group. However, when species group membership was mapped on our partitioned Bayesian phylogenetic hypothesis, only one species group corresponds to a clade; all other species group arrangements are strongly rejected by our phylogeny. Our results demonstrate that (1) previously recognized species group relationships were misled by phenotypic convergence; (2) is widely paraphyletic; and (3) multiple lineages have independently invaded the Philippines. Based on this new perspective on the phylogenetic relationships of Philippine , we revise the archipelago's diverse assemblage of species at the generic level, and resurrect and/or expand four previously recognized genera, and describe two new genera to accommodate the diversity of Philippine skinks of the group. © 2011 The Linnean Society of London, , 2011, , 1217-1243.
A molecular phylogeny of Equatorial African Lacertidae, with the description of a new genus and species from eastern Democratic Republic of the Congo
Currently, four species of the lacertid lizard genus Adolfus are known from Central and East Africa. We sequenced up to 2,825 bp of two mitochondrial (16S and cyt b) and two nuclear (c-mos and RAG1) genes from 41 samples of Adolfus (representing every species), two species each of Gastropholis and Holaspis, and in separate analyses combined this data with GenBank sequences of all other Eremiadini genera and four Lacertini outgroups. Data from DNA sequences were analyzed with maximum parsimony (PAUP), maximum-likelihood (RAxML) and Bayesian inference (MrBayes) criteria. Results demonstrated that Adolfus is not monophyletic: A. africanus (type species), A. alleni and A. jacksoni are sister taxa, whereas A. vauereselli and a new species from the Itombwe Plateau of Democratic Republic of the Congo are in a separate lineage. Holaspis and Gastropholis were recovered in separate clades. Based on this molecular data, relatively substantial sequence divergence and multiple morphological differences, we describe a new genus of lacertid for the lineage including A. vauereselli and the new Itombwe species. The recognition of this new, endemic genus underscores the conservation importance of the Albertine Rift, especially the Itombwe Plateau, a unique region that is severely threatened by unchecked deforestation, mining and poaching.
Previously undocumented diversity and abundance of cryptic species: a phylogenetic analysis of Indo-Pacific Arminidae Rafinesque, 1814 (Mollusca: Nudibranchia) with descriptions of 20 new species of Dermatobranchus
The phylogenetic relationships amongst the Arminidae were analysed based upon morphological characters of 58 presently described species or nudibranchs, including 35 previously described Arminidae and 20 new species of Dermatobranchus. From the literature review and anatomical examinations, 43 characters were considered for 78 taxa. These characters were polarized using Berthella canariensis as the outgroup taxon and the type species of several other genera identified from recent publications. The resulting phylogeny supports the monophyly of Arminidae, Dermatobranchus, Doridina, and Proctonotidae. The paraphyly of the Arminina is further demonstrated in this study. Two previously described, but poorly known, species of Indo-Pacific Armina are redescribed, Armina magnaBaba, 1955 and Armina paucifoliataBaba, 1955. The anatomy and taxonomic status of nine previously described species of Dermatobranchus were examined in this study. The anatomy of Dermatobranchus pustulosus (van Hasselt, 1824) has been overlooked since Bergh (1888) illustrated the radula of van Hasselt's specimen. It is redescribed and its range is extended to several new localities in the western Pacific. Dermatobranchus pulcherrimus Miller & Willan, 1986 is considered here as a new synonym of Dermatobranchus rubidus (Gould, 1852). The following 20 species of Dermatobranchus are new and are described in the present paper: Dermatobranchus albineus sp. nov., Dermatobranchus arminus sp. nov., Dermatobranchus caesitius sp. nov., Dermatobranchus caeruleomaculatus sp. nov., Dermatobranchus cymatilis sp. nov., Dermatobranchus dendonephthyphagus sp. nov., Dermatobranchus diagonalis sp. nov., Dermatobranchus earlei sp. nov., Dermatobranchus fasciatus sp. nov., Dermatobranchus funiculus sp. nov., Dermatobranchus kalyptos sp. nov., Dermatobranchus kokonas sp. nov., Dermatobranchus leoni sp. nov., Dermatobranchus microphallus sp. nov., Dermatobranchus oculus sp. nov., Dermatobranchus phyllodes sp. nov., Dermatobranchus piperoides sp. nov., Dermatobranchus rodmani sp. nov., Dermatobranchus semilunus sp. nov., and Dermatobranchus tuberculatus sp. nov. Eighteen of these new taxa are found in the Indo-Pacific tropics and two are found in temperate South Africa, D. albineus and D. arminus. Unique combinations of morphological characters distinguish these as new species of Dermatobranchus. Several species that are externally similar have radically divergent internal morphology, are members of different clades of Dermatobranchus, and represent cryptic species. Especially important is the radular morphology, which shows remarkable diversity of form, probably related directly to the diversification of feeding of members of this clade on various octocorals.
The hog-badger is not an edentate: systematics and evolution of the genus (Mammalia: Mustelidae)
Hog-badgers (mustelid carnivorans classified in the genus ) are distributed throughout East and Southeast Asia, including much of China, the eastern Indian Subcontinent, Indochina and the large continental Asian island of Sumatra. is usually regarded as monotypic, comprising the single species . F. Cuvier, 1825, but taxonomic boundaries in the genus have never been revised on the basis of sizeable series from throughout this geographical range. Based on a review of most available specimens in world museums, we recognize three distinctive species within the genus, based on craniometric analyses, qualitative craniodental features, external comparisons, and geographical and ecological considerations. (Blyth, 1853) is a shaggy-coated, medium-sized badger widely distributed in temperate Asia, from Tibet and the Himalayan region to eastern and southern China. F. Cuvier, 1825, is an extremely large, shorter-haired badger, distributed throughout Southeast Asia, from eastern India to Myanmar, Thailand, Vietnam, Cambodia and Laos. The world's largest extant badger, . co-occurs with . in eastern India and probably in southern China, and fossil comparisons indicate that its geographical range may have extended into central China in the middle Pleistocene. The disjunctly distributed species (Hubrecht, 1891), originally described within the order 'Edentata' by a remarkable misunderstanding, is the smallest and darkest member of the genus and is endemic to the Barisan mountain chain of Sumatra. Apart from . , no other occurs on the Sunda Shelf below peninsular Thailand. The natural history of each species of , so far as is known, is briefly reviewed. No claim to original US Government works.
Higher-order phylogeny of modern birds (Theropoda, Aves: Neornithes) based on comparative anatomy. II. Analysis and discussion
In recent years, avian systematics has been characterized by a diminished reliance on morphological cladistics of modern taxa, intensive palaeornithogical research stimulated by new discoveries and an inundation by analyses based on DNA sequences. Unfortunately, in contrast to significant insights into basal origins, the broad picture of neornithine phylogeny remains largely unresolved. Morphological studies have emphasized characters of use in palaeontological contexts. Molecular studies, following disillusionment with the pioneering, but non-cladistic, work of Sibley and Ahlquist, have differed markedly from each other and from morphological works in both methods and findings. Consequently, at the turn of the millennium, points of robust agreement among schools concerning higher-order neornithine phylogeny have been limited to the two basalmost and several mid-level, primary groups. This paper describes a phylogenetic (cladistic) analysis of 150 taxa of Neornithes, including exemplars from all non-passeriform families, and subordinal representatives of Passeriformes. Thirty-five outgroup taxa encompassing Crocodylia, predominately theropod Dinosauria, and selected Mesozoic birds were used to root the trees. Based on study of specimens and the literature, 2954 morphological characters were defined; these characters have been described in a companion work, approximately one-third of which were multistate (i.e. comprised at least three states), and states within more than one-half of these multistate characters were ordered for analysis. Complete heuristic searches using 10 000 random-addition replicates recovered a total solution set of 97 well-resolved, most-parsimonious trees (MPTs). The set of MPTs was confirmed by an expanded heuristic search based on 10 000 random-addition replicates and a full ratchet-augmented exploration to ascertain global optima. A strict consensus tree of MPTs included only six trichotomies, i.e. nodes differing topologically among MPTs. Bootstrapping (based on 10 000 replicates) percentages and ratchet-minimized support (Bremer) indices indicated most nodes to be robust. Several fossil Neornithes (e.g. Dinornithiformes, Aepyornithiformes) were placed within the ingroup a posteriori either through unconstrained, heursitic searches based on the complete matrix augmented by these taxa separately or using backbone-constraints. Analysis confirmed the topology among outgroup Theropoda and achieved robust resolution at virtually all levels of the Neornithes. Findings included monophyly of the palaeognathous birds, comprising the sister taxa Tinamiformes and ratites, respectively, and the Anseriformes and Galliformes as monophyletic sister-groups, together forming the sister-group to other Neornithes exclusive of the Palaeognathae (Neoaves). Noteworthy inferences include: (i) the sister-group to remaining Neoaves comprises a diversity of marine and wading birds; (ii) Podicipedidae are the sister-group of Gaviidae, and not closely related to the Phoenicopteridae, as recently suggested; (iii) the traditional Pelecaniformes, including the shoebill (Balaeniceps rex) as sister-taxon to other members, are monophyletic; (iv) traditional Ciconiiformes are monophyletic; (v) Strigiformes and Falconiformes are sister-groups; (vi) Cathartidae is the sister-group of the remaining Falconiformes; (vii) Ralliformes (Rallidae and Heliornithidae) are the sister-group to the monophyletic Charadriiformes, with the traditionally composed Gruiformes and Turniciformes (Turnicidae and Mesitornithidae) sequentially paraphyletic to the entire foregoing clade; (viii) Opisthocomus hoazin is the sister-taxon to the Cuculiformes (including the Musophagidae); (ix) traditional Caprimulgiformes are monophyletic and the sister-group of the Apodiformes; (x) Trogoniformes are the sister-group of Coliiformes; (xi) Coraciiformes, Piciformes and Passeriformes are mutually monophyletic and closely related; and (xii) the Galbulae are retained within the Piciformes. Unresolved portions of the Neornithes (nodes having more than one most-parsimonious solution) comprised three parts of the tree: (a) several interfamilial nodes within the Charadriiformes; (b) a trichotomy comprising the (i) Psittaciformes, (ii) Columbiformes and (iii) Trogonomorphae (Trogoniformes, Coliiformes) + Passerimorphae (Coraciiformes, Piciformes, Passeriformes); and (c) a trichotomy comprising the Coraciiformes, Piciformes and Passeriformes. The remaining polytomies were among outgroups, although several of the highest-order nodes were only marginally supported; however, the majority of nodes were resolved and met or surpassed conventional standards of support. Quantitative comparisons with alternative hypotheses, examination of highly supportive and diagnostic characters for higher taxa, correspondences with prior studies, complementarity and philosophical differences with palaeontological phylogenetics, promises and challenges of palaeogeography and calibration of evolutionary rates of birds, and classes of promising evidence and future directions of study are reviewed. Homology, as applied to avian examples of apparent homologues, is considered in terms of recent theory, and a revised annotated classification of higher-order taxa of Neornithes and other closely related Theropoda is proposed. (c) 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 149, 1-95.
Examining the relationship between sexual dimorphism in skin anatomy and body size in the white-lipped treefrog, (Anura: Hylidae)
Amphibians transport water, oxygen, carbon dioxide and various ions (e.g. sodium and potassium) across their skin. This cutaneous permeability is thought to affect their ability to respond to environmental change and to play a role in global population declines. Sexual dimorphism of skin anatomy has been accepted in some species, but rejected in others. The species in which such dimorphism has been detected have all been sexually dimorphic in body size, with males that are smaller and have thinner skin. It is unclear whether this difference in skin thickness manifests a functional difference or if it is related to body size alone. Skin thickness (epidermis, spongy dermis, compact dermis and total thickness) was examined in males and females of the white-lipped treefrog (). Although the skin of males is absolutely thinner than that of females, this difference is explained by body size differences between the sexes. Overall, we conclude that skin thickness in male and female correlates with body size dimorphism and suggest that future studies on amphibian skin anatomy include measures of body size, test the ecological significance of sexually dimorphic skin anatomy and better document the prevalence of sexually dimorphic amphibian skin anatomy.