Morphology and development of additional bony elements in the genus Brachycephalus (Anura: Brachycephalidae)

We describe the extra bony elements, plates, and osteoderms present in species of the genus Brachycephalus. Samples of eight species of Brachycephalus, including seven populations of Brachycephalus ephippium, were examined. The large additional elements associated with the skull (parotic plate) and vertebrae (vertebral and paravertebral plates) all comprise intramembranous bone, similar to that of the frontoparietal or nasal bones of the skull of most of frogs. Additionally, in the dermis of one unnamed species, we discovered and described true osteoderms. We discuss the morphological nature and diversity of theses elements and their importance as evidence of phylogenetic relationship within Brachycephalus. In summary, three distinct conditions of extra bony elements occur in the genus Brachycephalus: (1) bony plates may be present or absent in species of the genus; (2) a few, small bony plates may be developed and these may be represented by (a) paravertebral plates small and restricted to the distal ends of the transverse processes of the presacral IV, (b) parotic plates small and not covering the tops of the squamosals, and (c) ornamented spinal plates on all vertebrae; and (3) well‐developed bony plates may be present as (a) paravertebral plates forming a ‘bone‐shield’ on the dorsal surface of the trunk, ornamented, and visible through the integument, (b) parotic plates covering the tops of the squamosals, and (c) spinal plates associated with all vertebrae, and ornamented on vertebrate I–VI. Although the phenomenon of miniaturization may be associated with the appearance of new elements in at least some of the species in the genus, the traditional rule may not be universally applicable. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 752–767.     

Hyperossification in miniaturized toadlets of the genus Brachycephalus (Amphibia: Anura: Brachycephalidae): Microscopic structure and macroscopic patterns of variation

Species of the genus Brachycephalus, have a snout‐vent length of less than 18 mm and are believed to have evolved through miniaturization. Brachycephalus ephippium, is particularly interesting; because its entire skull is hyperossified, and the presacral vertebrae and transverse processes are covered by a dorsal shield. We demonstrate in this paper that, at the macroscopic level, a completely hyperossified skull and dorsal shield occur only in B. ephippium, but not in B. ferruginus, B. izechsohni, B. pernix, B. pombali, B. brunneus, B. didactylus, and B. hermogenesi. An intermediate condition, in which the skull is hyperossified but a dorsal shield is absent, occurs in B. vertebralis, B. nodoterga, B. pitanga, and B. alipioi. The microscopic structure of hyperossification was examined in skulls of B. ephippium and B. pitanga, revealing a complex organization involving the presence of Sharpey fibers, which in humans are characteristic of periodontal connections. J. Morphol., 2009. © 2009 Wiley‐Liss, Inc.     

A new metriorhynchid crocodylomorph from the Lower Kimmeridge Clay Formation (Late Jurassic) of England,with implications for the evolution of dermatocranium ornamentation in Geosaurini

A new metriorhynchid crocodylomorph from the Lower Kimmeridge Clay Formation (Kimmeridgian, Upper Jurassic) of England is described. This specimen, a three‐dimensionally preserved skull and left mandibular ramus, is referred to a new species: T orvoneustes coryphaeus sp. nov. Within the genus Torvoneustes, T . coryphaeus sp. nov. is unique as it has a long anteromedial process of the frontal, ornamented dermatocranium, and the supraorbital notch forms a strongly acute angle. Our phylogenetic analysis confirms the placement of this specimen in Torvoneustes. The dentition of T . coryphaeus sp. nov. , like that of the type species, has a blunt apex, crown basal–mid regions with numerous tightly packed apicobasally aligned ridges, and apical region with an anastomosed pattern of ridges that interact with the carinae. Within Thalattosuchia these dental characteristics are only found in Torvoneustes and the teleosaurid Machimosaurus. The heavily ornamented dermatocranium of T . coryphaeus sp. nov. is in contrast to the unornamented (nasals and frontal)–lightly ornamented (maxillae and premaxillae) pattern seen in Torvoneustes carpenteri. Curiously, this pattern of reduction and loss of dermatocranium ornamentation is also observed in Metriorhynchus, Dakosaurus, and the subclade Rhacheosaurini. We hypothesize that the ‘smooth’ dermatocranium of Late Jurassic metriorhynchids evolved independently in each subclade (parallel evolution), and would have reduced drag, thereby making locomotion through water more energy efficient. © 2013 The Linnean Society of London     

EXISTENCE OF TWO STOMATAL SHAPES IN THE GENUS DENDROBIUM (ORCHIDACEAE) AND ITS SYSTEMATIC SIGNIFICANCE

On the basis of the shape and size of the outer stomatal ledge, 153 species of the genus Dendrobium (Orchidaceae) were divided into three groups, i.e., species group I, species group II, and section Grastidium. Student's t-test and principal components analysis using seven stomatal characters suggest that the first two groups are quite distinct from each other. The values for section Grastidium, however, are scattered and indicative of heterogeneity. Our data suggest a high probability that species groups I and II are monophyletic sister groups. However, section Grastidium, in which stomatal diversification has occurred, may be para- or polyphyletic relative to the genus Dendrobium.     

Saprolegnia semihypogyna sp. nov., a saprolegniaceous oomycete isolated from soil in Japan

Saprolegnia semihypogyna, a saprolegniaceous oomycete (Saprolegniales, Oomycetes), is described based on two strains isolated from soil samples collected in Japan. It is characterized by having a new combination of sexual characters in the genus Saprolegnia, namely, producing mostly monosporous oogonia with subeccentric oospores and semihypogynous or androgynous antheridial branches. The species is morphologically similar to Scoliolegnia subeccentrica, although the oogonial wall of the former is smooth and that of the latter is densely ornamented. Morphological comparison of the two species indicates that the generic status of the genus Scoliolegnia is questionable. Received: February 22, 2001 / Accepted: September 5, 2001     

Permian ammonoids of the Kolyma-Omolon Region: Kyrian association

This paper describes new taxa of the earliest Permian ammonoids of the Kolyma-Omolon Region: the genus Kolymoglaphyrites gen. nov. with the type species K. lazarevi sp. nov. and the species Uraloceras margaritae sp. nov., recognized as the Kyrian ammonoid association. Judging from the analysis of the main characters, the genus Kolymoglaphyrites is included in the mainly Carboniferous family Glaphyritidae, which expands the interval of the vertical distribution of this family to include the Sakmarian. Uraloceras margaritae is considered as the initial genus in the lineage of weakly ornamented species of Uraloceras. Apparently, this species evolved from Paragastrioceras sterlitamakense at the Asselian-Sakmarian boundary to become later ancestral for U. simense. Based on the phylogenetic position of U. margaritae the Kyrian ammonoid association is dated as Early Sakmarian.     

Phylogeny of the tribe Abrotrichini (Cricetidae,Sigmodontinae): integrating morphological and molecular evidence into a new classification

The tribe Abrotrichini (five genera and 14 living species) is a small clade within the speciose subfamily Sigmodontinae (Rodentia, Cricetidae), representing one of the extant successful radiations of mammals at southern high latitudes of the Neotropics. Its distribution is mostly Andean, reaching its greatest diversity in southern Argentina and Chile. We evaluate the phylogenetic relationships within this tribe through parsimony and Bayesian approaches based on 99 morphological characters (including 19 integumental characters, 38 skull characters, 31 dental characters, three postcranial skeletal characters, seven from the male accessory glands and phallus and one from the digestive system) and six molecular markers (one mitochondrial and five nuclear). We include representatives of all, except one, of the currently recognized species of living Abrotrichini plus one fossil form. Based on total evidence, we recovered a primary division between the genus Abrothrix and a group including the long‐clawed Abrotrichini, Chelemys, Geoxus, Notiomys and Pearsonomys. Both clades are recognized and named here as subtribes. The large degree of morphological variation observed within Abrothrix suggests that species in the genus fall into four groups, which we recognize as subgenera. In addition, the two known species of Chelemys do not form a monophyletic group, and Geoxus was recovered as paraphyletic with respect to Pearsonomys. To reconcile classification and phylogenetics, we describe a new genus for Chelemys macronyx and include Pearsonomys as a junior synonym of Geoxus. Our results highlight the importance of both morphology and molecules in resolving the phylogenetic relationships within this tribe. Based on biogeographical analyses, we hypothesize that Abrotrichini originated in south‐western South America by vicariance and then diversified mostly by successive dispersal events.     

Molecular phylogeny of the diverse parasitoid wasp genus Ophion Fabricius (Hymenoptera: Ichneumonidae: Ophioninae)

Ophion Fabricius is a diverse genus of nocturnal ichneumonid wasps (Insecta: Hymenoptera) that is particularly species‐rich in temperate areas, yet has received little taxonomic attention in the Holarctic region, where most species occur. While there have been some attempts to divide Ophion into monophyletic species groups, the vast majority of species have been lumped into a single, paraphyletic group, the O. luteus species group, which is defined only by the lack of characters specific to the other groups. The challenging morphology of this large catch‐all group has limited attempts to subdivide it, and no phylogenetic hypothesis has been proposed for the genus as a whole. In this study, we use DNA sequence data [28S ribosomal RNA (28S), cytochrome oxidase 1 (COI) and internal transcribed spacer 2 (ITS2)] to present the first molecular phylogeny of Ophion. We also describe the secondary structure of ITS2 for the first time in Ichneumonidae, and explore its implications for phylogeny estimation. We define 13 species groups, nine of which were previously considered part of the O. luteus species group s.l. The included species groups are the O. minutus, O. areolaris, O. scutellaris, O. flavidus, O. parvulus, O. slossonae, O. nigrovarius, O. pteridis, O. luteus s.s., and O. obscuratus species groups, along with three groups lacking described species (Species group 1, New Zealand group, Madagascar group). This study provides a framework for future studies of this diverse and morphologically challenging genus.     

Evolutionary history of the burnet moth genus Zygaena Fabricius, 1775 (Lepidoptera: Zygaenidae) inferred from nuclear and mitochondrial sequence data: phylogeny, host–plant association, wing pattern evolution and historical biogeography

Burnet moths of the genus Zygaena are a striking group of primarily diurnal Lepidoptera displaying an exceptional phenotypic plasticity. Previous attempts to elucidate the phylogenetic history of the group had been confounded by a perplexing pattern of characters or insufficient taxon sampling. In the present study, we infer a phylogeny of the genus Zygaena by analysing 5.4 kb of their nuclear and mitochondrial DNA. Eighty‐four of the 98 currently recognized species in this genus are considered, including representatives of all described species groups. RNA coding sequences are aligned with reference to zygaenoid moth specific secondary structure models of corresponding molecules. We conduct phylogenetic analyses within a Bayesian framework applying partition specific substitution parameters; covariation of paired sites in RNA gene sequences is accommodated by using doublet substitution models. The molecular data reveal that a considerable number of currently recognized species groups in Zygaena are not monophyletic. The traditional subgeneric classification proves to be artificial as well; Agrumenia and Zygaena (sensu stricto) are polyphyletic. Only the subgenus Mesembrynus can be confirmed as a monophyletic species cluster. Optimization of larval host–plant associations and forewing patterns on sampled trees of the Bayesian analyses suggest convergent evolution of similar wing pattern types in distantly related species clusters and a shift from cyanogenic to acyanogenic host‐plants. The phylogenetic results challenge the classic assumption that early species diversification in Zygaena took place in the Irano–Turkestanian region. Rather, the molecular data point to the western Mediterranean area as the geographical origin of the group and imply a subsequent colonization of the Middle East and Central Asia. We discuss the apparently convergent evolution of similar wing patterns in context with the chemical defence system of burnet moths and suggest a species group concept for the genus Zygaena that accounts for the recent findings. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 92 , 501–520.     

DNA barcodes for the pipefish genus Corythoichthys (Actinopterygii: Syngnathiformes) from the Indian Ocean provide insights into cryptic diversity

The syngnathiform genus Corythoichthys comprises a group of taxonomically complex, tail-brooding (Syngnathinae) pipefishes widely distributed in the Indo-Pacific region. Due to the presence of overlapping interspecific morphological characters, reliable taxonomic information on Corythoichthys is still lacking. Using 52 CO1 sequences, including seven newly generated, a phylogenetic analysis was carried out to understand the genetic diversity, distribution and ‘species groups’ within the genus Corythoichthys. Species delimitation using Automatic Barcode Gap Discovery (ABGD) analysis confirmed the presence of 13 species which include ‘species-complexes’ previously considered as a single taxon. Our results revealed the presence of three species groups, ‘C. amplexus’, ‘C. conspicillatus’ and ‘C. haematopterus’ and four unidentified/undescribed species in the wider Indo-Pacific realm. Interestingly, 60 sequences and a mitogenome identified as Corythoichthys in GenBank are misidentified at the genus level. Based on our findings, we suggest that the taxonomy and systematics of Corythoichthys need to be re-examined and validated using integrative methods, and care should be taken while selecting specimens for genetic studies.     

Production of plumage ornaments among males and females of two closely related tropical passerine bird species

The evolution of elaborate secondary sexual traits (i.e., ornaments) is well‐studied in males but less so in females. Similarity in the appearance of ornaments between males and females supports the view that female ornaments arise as a neutral byproduct of selection on male traits due to genetic correlation between sexes, but recent research suggests an adaptive function of female ornaments in at least some contexts. Information on the degree to which production of ornaments differs between the sexes can shed light on these alternative perspectives. We therefore characterized the structural underpinnings of melanin‐based plumage production in males and females of two closely related passerine bird species (genus Malurus). Importantly, both ornamented and unornamented phenotypes in each sex are present between these two species, providing an opportunity to test the null expectation of equivalent modes of production in male and female ornamented phenotypes. In Malurus alboscapulatus, ornamented females are qualitatively similar to males, but we describe a distinctive ornamented female phenotype that differs from that of males in lacking a blue sheen and in lower feather barbule density. In M. melanocephalus, unornamented males and females are also similar in appearance, and we describe a similarity between unornamented phenotypes of males and females in both color and underlying feather barbule structure and pigment composition. Unornamented male M. melanocephalus can flexibly transition to the ornamented phenotype in weeks, and we found extreme differences in color and feather structure between these two alternative male phenotypes. These results contradict the idea that female ornaments have evolved in this system following a simple switch to male‐like plumage by demonstrating greater complexity in the production of the ornamented phenotype in males than in females.     

Phylogeny and biogeography of the dung beetle genus Phanaeus (Coleoptera: Scarabaeidae)

Abstract The dung beetle genus Phanaeus as currently recognized by Edmonds (1994) consists of 51 species placed in 13 species groups and two subgenera. Here, I examine the phylogeny and biogeography of this genus by analysing the mitochondrial cytochrome oxidase subunit I (530 bp), nuclear large subunit ribosomal RNA (28S, D2 region), and 67 morphological characters for 28 species of Phanaeus. Both maximum parsimony and Bayesian analyses from the combined data yielded well‐resolved trees, although low bootstrap and posterior probability support were found for basal nodes. The phylogenetic hypotheses presented here suggest that the subgenera Phanaeus s.str. and Notiophanaeus should each be elevated to the status of full genus. With the exception of the eucraniine outgroups, the paleano species group of the genus Phanaeus is recovered as sister to all other taxa, including the outgroups Oxysternon, Sulcophanaeus and Coprophanaeus. High bootstrap values and posterior probabilities supported the species groups endymion, tridens and vindex. Biogeographical analyses suggest an ancestral distribution for Phanaeus in the Andes in South America, although numerous dispersal events evidently have produced a complicated biogeographical history.     

A total evidence analysis of the phylogeny of hatchet-faced treefrogs (Anura: Hylidae: Sphaenorhynchus)

The Neotropical hylid genus Sphaenorhynchus includes 15 species of small, greenish treefrogs widespread in the Amazon and Orinoco basins, and in the Atlantic Forest of Brazil. Although some studies have addressed the phylogenetic relationships of the genus with other hylids using a few exemplar species, its internal relationships remain poorly understood. In order to test its monophyly and the relationships among its species, we performed a total evidence phylogenetic analysis of sequences of three mitochondrial and three nuclear genes, and 193 phenotypic characters from all species of Sphaenorhynchus. Our results support the monophyly of Sphaenorhynchus with molecular and phenotypic evidence, with S. pauloalvini as the earliest diverging taxon, followed by S. carneus, as the sister taxon of all remaining species of the genus. We recognize three species groups in Sphaenorhynchus (the S. lacteus, S. planicola and S. platycephalus groups), to facilitate its taxonomic study; only three species (S. carneus, S. pauloalvini and S. prasinus) remain unassigned to any group. Sequence data were not available for only two species (S. bromelicola and S. palustris) for which we scored phenotypic data; wildcard behaviour was detected only in S. bromelicola nested inside the S. platycephalus group. On the basis of the resulting phylogenetic hypothesis, we discuss the evolution of oviposition site and a number of phenotypic characters that could be associated with heterochronic events in the evolutionary history of this group.     

Description of <Emphasis Type="Italic">Gosztonyia antarctica</Emphasis>, a new genus and species of Zoarcidae (Teleostei: Perciformes) from the Antarctic Ocean

A new genus and species of zoarcid fish, Gosztonyia antarctica, is described on the basis of four specimens collected from the Bellingshausen Sea, Antarctic Ocean, at a depth of 615 m. Gosztonyia is placed in the subfamily Lycodinae and can be distinguished from all other zoarcid genera by the following combination of characters: seven branchiostegal rays, interdigitating ceratohyal–epihyal articulation, palatal arch reduced, posterior hyomandibular ramus longer than anterior, cranium narrowed, supratemporal commisure and occipital pores absent. A new species, Gosztonyia antarctica, is described and the relationships of the new genus are discussed.     

SPECIATION AND FEATHER ORNAMENTATION IN BIRDS

The hypothesis that sexual selection promotes speciation has rarely been tested. We identified 70 evolutionarily independent events of feather ornaments in birds. For each focal species we noted the number of ornamented and nonornamented species belonging to its genus and its number of subspecies, as well as its mating system and the extent of its geographic range. For purposes of comparison, we randomly chose a second, nonornamented species for which we obtained information on the number of subspecies, and in cases in which the nonornamented species was in the same genus, we chose a third, nonornamented species in a related genus and obtained the same information. We then noted the number of species in each genus and the difference in numbers of species, or species richness, between paired genera. For the genera of the focal ornamented species, we regressed number of ornamented species on number of nonornamented species and found a positive relationship. As number of species per genus rose, number of ornamented species per genus rose more rapidly, indicating that more speciose genera have a higher proportion of ornamented species than less speciose genera. We then took the deviations from this regression, the residual number of species, and regressed them on the differences in species richness between the paired genera. This relationship was positive indicating that ornamented genera with more than the expected number of ornamented species were more speciose with respect to their paired genera than were genera with fewer than the expected number of ornamented species. Finally, we compared the deviations from this regression, the residual number of ornamented species, with species' mating system and found a greater residual number of ornamented species among species whose mating system is associated with greater skew in male mating success and thus more intense sexual selection. Ornamented species had more subspecies than nonornamented species, even when controlling for geographic range, suggesting an association between subspeciation and ornaments.     

石山苣苔属(苦苣苔科)花形态演化及分类学意义

石山苣苔属(苦苣苔科)约30种,主要分布于我国南部的石灰岩地区.目前该属已知物种数虽少但花形态极其多样,是该科中分类最为困难的类群之一.基于分子证据,其它8个属中花形态迥异的一些物种被并入石山苣苔属.然而,该属花形态的演化趋势缺乏系统性的研究,传统分类对属的界定与分子系统学研究结果相矛盾的原因,以及是否有形态特征支持新界定的石山苣苔属还不清楚.该研究中,总共编码了19种石山苣苔属植物和9种报春苣苔属植物的35个形态特征,其中包括26个花部形态特征,在分子系统树上追踪了它们的演化路径.结果表明:无论属内还是属间,多数花部形态特征,尤其以往属的分类界定特征,在演化过程中变化频繁且发生了高度同塑性演化,这是导致传统形态分类不自然的关键因素.此外,在观察研究的所有特征中,花丝和柱头的差异可能在石山苣苔属植物共同祖先中经历了演变,或可用于区分石山苣苔属与其姐妹报春苣苔属的大多数种类.因此,在苦苣苔科植物的分类学研究中应当慎用这些花部性状作为分类依据,而且应对形态特征进行广泛地观察研究,在密集的取样和分辨率更高、更可靠的系统树上追踪它们的演化规律.更为重要的是,需要进一步研究导致复杂形态性状演化的内在分子调控机理和外在的自然选择动力,最终更加深入地理解石山苣苔属等典型喀斯特植物的演化过程和机理.     

Four new species of Typhlocharis (baetica group) (Coleoptera: Carabidae: Anillini) from southwestern Iberian Peninsula with notes on their biogeographical and morphological implications

Four new species of Typhlocharis (Carabidae: Anillini) are described from the southwest of the Iberian Peninsula, assigned to the baetica species group. T. prima sp.n. , T. secunda sp.n. and T. tertia sp.n. (from La Palma del Condado, Huelva, Spain) are syntopic, and T. quarta sp.n. (from Barrancos, Beja, Portugal) is the first species of the group with umbilicate series 4 + 4. They represent, respectively, the southernmost and westernmost distribution data for the group. The baetica group remains well defined and includes ten species. The diagnosis is simplified, with the confirmation that many morphological features characteristic of this group are widespread within the genus. The defining character is the presence of denticles in apical margin of elytra, not associated to the seventh stria or to the elytral suture. The new species provide morphological data that suggest the baetica and silvanoides groups are closely related. Distribution of both groups in the south of the Iberian Peninsula and the presence of morphological characters that are considered plesiomorphic are coherent with the hypothesis of betic‐riffain origin of the genus. They also support the hypothesis of easy alteration of the umbilicate pattern between closely related species and the independent origin of apical denticles in the group compared with those present in other species of the genus. The presence of three well‐differentiated size ranges in syntopic populations suggest ecological diversification to avoid interspecific competition.     

Revision for the swordtail cricket genus Homoeoxipha (Grylloidea: Trigonidiidae: Trigonidiinae) from China

Species of Homoeoxipha are very small and always live on leaves of shrubs and grasses. This genus contains ten species worldwide and is distributed across five zoological regions, from Africa to New Guinea. Homoeoxipha lycoides is the type species of this genus and possesses the characteristic coloration for the genus. These coloration features resulted in some synonyms and disputes over species, because most Homoeoxipha species are ornamented with a similar coloration pattern. We compared and documented the differences between H. lycoides and its relatives in China. A new important character, the epiphallic transversal suture, is used to distinguish them. Based on this work, all of the Chinese species, H. lycoides, H. obliterata and H. nigripes, and one new species, H. eurylobus, with similar coloration to H. lycoides, are described and illustrated.