Size and shape interspecific divergence patterns partly reflect phylogeny in an Onthophagus species‐complex (Coleoptera: Scarabaeidae)

Polyphenism has been suggested as an accelerator for morphological evolution and speciation. In the dung beetles of the genus Onthophagus, horn expression is polyphenic: large males develop horns whereas smaller males express greatly reduced or no horns. Horn static allometries seem to diverge rapidly amongst extant taxa, a process which might trigger changes in the male genital morphology, thus possibly promoting speciation as a by‐product. It can therefore be hypothesized that interspecific distances in allometries and, possibly, in other morphological traits mirror phylogenetic distances. In this study we first assessed the phylogenetic relationships amongst three closely related taxa belonging to the so‐called ‘Onthophagus fracticornis‐similis‐opacicollis’ species‐complex by sequencing the mitochondrial gene cytochrome oxidase subunit 1 (cox1). Biomolecular results indicated three independent lineages, the closest relationships being found between Onthophagus similis and Onthophagus opacicollis. Then we assessed the extent to which divergence pattern of horn static allometries and size and shape divergence patterns of one genital (paramere) and two nongenital (head and epipharynx) structures mirrored the phylogenetic relationships. Interspecific divergence patterns of horn static allometries, paramere, and head shape were found to be congruent with the evolutionary relationships inferred from biomolecular data. Nevertheless, paramere size and epipharynx shape showed patterns not consistent with the phylogeny. Furthermore, the relative size of nongenital structures showed little interspecific divergence compared to their shapes. Our results suggest that size and shape interspecific divergence mirror phylogeny only in part; they also indicate that distinct morphological traits may differ in their tendency to evolve in concert, and that size and shape of the same trait can evolve independently across species. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162 , 482–498.     

Combination of geometric morphometric and genetic approaches applied to a debated taxonomical issue: the status of Onthophagus massai (Coleoptera, Scarabaeidae) as an endemic species vicarious to Onthophagus fracticornis in Sicily

The present study deals with the phenomenon of insular speciation and discusses, as a case study, the debated taxonomical issue of the status of Onthophagus massai (Coleoptera, Sarabaeidae) as an endemic species vicarious to Onthophagus fracticornis in Sicily. The authors investigated the differentiation patterns between an insular population belonging to the supposed species O. massai (collected in its locus typicus, Piano Battaglia) and three Italian O. fracticornis populations (collected along a N–S latitudinal gradient). These patterns are described and analysed using multiple approaches: the qualitative inspection of the microsculpture of elytral surfaces, considered a diagnostic character for O. massai identification; the comparison of horn static allometries, known to be a good indicator of divergence processes between closely related species or isolated populations of the same species; the comparison of the patterns of shape and size difference of the head, epipharynx and genitalia attained with a combination of traditional and geometric (landmark and semilandmark) morphometric methods; and, finally, the estimation of the genetic relationships between Sicilian and continental populations obtained by analysing cytochrome oxidase subunit 1 mitochondrial gene sequences. The integration of the results of these approaches indicates that there is not sufficient evidence to vindicate the species status for O. massai, which should more likely be considered a small-sized version of O. fracticornis (a possible case of insular dwarfism). However, the complex pattern of shape, size and genetic variation observed between the populations analysed hinted at the possibility that a diversification process is ongoing, but not only between insular and continental populations; each population showed a tendency to evolve as an evolutionarily independent unit.     

Intraspecific and interspecific variation of female genitalia in two species of watersnake

The morphological differences in female genitalia within and between species are little studied and poorly understood, yet understanding patterns of variation in female genitalia can provide insights into mechanisms of genital evolution. The present study aimed to explore the patterns of intraspecific and interspecific variation in female genitalia in two sister taxa of watersnake (Nerodia sipedon and Nerodia fasciata) that have similar genital shape. We used a geometric morphometric (GM) approach to study variation in shape of the vagina between and within two sister species. We examined genital shape in female watersnakes ranging from small, sexually immature females to large reproductive females that had recently given birth. We found that shape variation of genitalia is strongly correlated with body size, where larger but not smaller females have a bifurcation in the vagina. However, we also found significant shape variation in the structure of the vagina between the two species, where N. fasciata has narrower genitalia with more prominent bifurcation, whereas N. sipedon has wider genitalia with less marked bifurcation. Using GM allowed us to detect significant differences in genital shape that were not apparent upon visual examination alone, suggesting that shape variation in female genitalia may be greater than previously assumed. Additional study of morphological differences in male reproductive organs for these species would help to determine whether there has been genital co‐evolution, and potentially mechanical reproductive isolation, in these two closely‐related and occasionally sympatric species. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111, 183–191.     

The falcifolia syndrome of Oenothera: IV. Loss of falcifolia-determining factors

Summary Inheritance of the falcifolia syndrome is explained by the hypothesis that this developmental disturbance is caused by the interaction of a non-chromosomal determinant (sigma) of Oenothera suaveolens with a complementary determinant (lambda) of O. lamarckiana when these two species are crossed. Localization of the determinants in the cell, the mechanism of their expression, and their molecular nature have not been established. Somatic segregation within the hybrids may lead to a separation of the two components, as suggested by the different reactions of sister plants in backcross tests and on selfing. The present paper presents results showing that in rare cases the fal determinants may be eliminated from the hybrid's cytoplasm. Thus, plants are produced that are phenotypically identical with the original parental strains but in crosses react like species that lack such determinants.     

Phylogeny of scolopendromorph centipedes (Chilopoda): morphological analysis featuring characters from the peristomatic area

Alternative schemes for the higher‐level systematics of the centipede order Scolopendromorpha have been established from characters of trunk segmentation, including the segmental position of spiracles, and the presence or absence of eyes. A comparative survey of the preoral chamber by light and scanning electron microscopy contributes 16 new characters of the epipharynx and hypopharynx, sampled from 26 species that represent 20 genera. These new data together with 49 additional morphological characters permit cladistic analysis of the major scolopendromorph groups. The shortest cladograms resolve blind Scolopendromorpha as a basal grade within which a clade now classified as Plutoniuminae + Scolopocryptopidae (supported by unreversed characters from the preoral chamber) is sister to the remaining scolopendromorphs. A unique row of bullet‐shaped sensilla between the labral and clypeal parts of the epipharynx provides a new autapomorphy of the Scolopendromorpha. Either 21 or 23 trunk segments optimize at the base of the Scolopendromorpha but in either case homoplasy is forced on the cladogram. New characters from the epipharynx give additional support for the monophyly of several traditional groupings, including Cryptopinae, Scolopendridae, Otostigmini, and Scolopendrini, and a basal resolution of Asanadini within Scolopendridae. Of the two competing hypotheses for the position of the enigmatic Mediterranean Plutonium zwierleini—being either sister to the cryptopid Theatops or sister to all other Scolopendromorpha—the former hypothesis is strongly supported; spiracles on all trunk segments in Plutonium are homoplastic with the state in Geophilomorpha. Observations on feeding behaviour are needed to illuminate convergence in characters of the epipharynx and mandible in Edentistoma (Otostigminae) and Campylostigmus (Scolopendrini). © The Willi Hennig Society 2008.     

Phylogenetic relationships among Boleosoma darter species (Percidae: Etheostoma)

Darters represent a species rich group of North American freshwater fishes studied in the context of their diverse morphology, behavior, and geographic distribution. We report the first molecular phylogenetic analyses of the Boleosoma darter clade that includes complete species sampling. We estimated the relationship among the species of Boleosoma using DNA sequence data from a mitochondrial (cytochrome b) and a nuclear gene (S7 ribosomal protein intron 1). Our analyses discovered that the two Boleosoma species with large geographic distributions (E. nigrum and E. olmstedi) do not form reciprocally monophyletic groups in either gene trees. Etheostoma susanae and E. perlongum were phylogenetically nested in E. nigrum and E. olmstedi, respectively. While analysis of the nuclear gene resulted in a phylogeny where E. longimanum and E. podostemone were sister species, the mitochondrial gene tree did not support this relationship. Etheostoma vitreum was phylogenetically nested within Boleosoma in the mitochondrial DNA and nuclear gene trees. Our analyses suggest that current concepts of species diversity underestimate phylogenetic diversity in Boleosoma and that Boleosoma species likely provide another example of the growing number of discovered instances of mitochondrial genome transfer between darter species.     

Mandibular Shape, Ontogeny and Dental Development in Bonobos (Pan paniscus) and Chimpanzees (Pan troglodytes)

The postnatal ontogenetic patterns and processes that underlie species differences in African ape adult mandibular morphology are not well understood and there is ongoing debate about whether African ape faces and mandibles develop via divergent or parallel trajectories of shape change. Using three-dimensional (3D) morphometric data, we first tested when in postnatal development differences in mandibular shape are initially evident between sister species Pan troglodytes and P. paniscus. Next, we tested whether each species has a distinct and non-parallel trajectory of mandibular development. Mandibles sampled across a broad developmental range of wildshot bonobos (n = 44) and chimpanzees (n = 59) were radiographed and aged from their dental development. We then collected 3D landmark surface data from all the mandibles. A geometric morphometric analysis of size-corrected 3D data found that bonobos and chimpanzees had parallel and linear ontogenetic trajectories of mandibular shape change. In contrast, mandibular shape was statistically different between P. paniscus and P. troglodytes as early as infancy, suggesting that species shape differences are already established near or before birth. A linear and stable trajectory of shape change suggests that mandibular ontogeny in these apes is unimpacted by non-linear variation in tooth developmental timing.     

北京山区栓皮栎林优势种群点格局分析

采用点格局分析方法对北京山区栓皮栎林主要种群和不同发育阶段种群的空间分布格局及相互关系进行分析。结果表明:(1)群落物种组成简单,栓皮栎和油松是群落的优势种群,优势种群以中径木为主,缺乏可供更新的幼苗,种群呈现出衰退趋势;(2)在研究尺度内,栓皮栎、油松单种的空间分布格局以聚集型分布为主,随着尺度的增大呈现出随机分布和均匀分布的趋势;栓皮栎和油松呈现显著负相关;(3)优势种群不同发育阶段的空间分布格局与研究尺度紧密相关,栓皮栎小树和中树以聚集型分布为主、大树呈随机分布,而油松不同发育阶段均以随机分布为主;(4)栓皮栎、油松种群不同发育阶段之间的空间关联主要发生在小尺度范围内,小树和中树以正关联为主,而中树和大树、小树和大树以负关联为主;随着研究尺度的增大趋于无关联。     

Interspecific shape divergence in Aphodiini dung beetles: the case of <Emphasis Type="Italic">Amidorus obscurus</Emphasis> and <Emphasis Type="Italic">A. immaturus</Emphasis> (Coleoptera: Scarabaeoidea)

The dung beetles Amidorus obscurus and A. immaturus are nearly indistinguishable, being characterized by a marked constancy in external morphological traits and little sexual dimorphism in adults. We studied two syntopic populations from the Italian Alps by means of geometric morphometric analyses. To identify possible undetected shape differences between species, we focused on the head, pronotum and scutellum (three external traits) and the epipharynx. Results indicate that the external traits are rather similar in the two species, whereas the epipharynx is clearly different. Interspecific differences in the aedeagus were also taken into account; these are noteworthy because parameres of A. immaturus differ in shape and are at least three times longer than those of A. obscurus. If it is assumed that the diversification of the two species took place during the quaternary ice age, A. immaturus would have evolved these marked differences rather quickly, in keeping with the hypothesis of rapid genital evolution. In an ontogenetic trajectory framework, we also considered the morphology of larvae. Interspecific divergence in the shape of the epipharynx is already evident at the preimaginal stage, whereas that of the genital disc is not. Accordingly, we hypothesise that the feeding and reproductive traits of these two species diverged morphologically when they become functional. Finally, by considering recent advances in ecological and evolutionary knowledge of dung beetles, the pattern of relative constancy in external morphology exhibited by the tribe Aphodiini, and that of great morphological diversification displayed by Onthophagini, were compared, and hypotheses about the origins of these differences discussed.     

Building Developmental Integration into Functional Systems: Function-Induced Integration of Mandibular Shape

The mammalian mandible is a developmentally modular but functionally integrated system. Whether morphological integration can evolve to match the optimal pattern of functional integration may depend on the developmental origin of integration, specifically, on the role that direct epigenetic interactions play in shaping integration. These interactions are hypothesized to integrate modules and also to be highly conservative, potentially constraining the evolution of integration. Using the fox squirrel (Sciurus niger) mandible as a model system, we test five a priori developmental hypotheses that predict mandibular integration and we also explore for correlations between shapes of mandibular regions not anticipated by any of the developmental models. To determine whether direct epigenetic interactions are highly conserved in rodents, we examine the correlation structure of fluctuating asymmetry, and compare integration patterns between fox squirrels and prairie deer mice (Peromyscus maniculatus bairdii). In fox squirrels, we find a correlation structure unanticipated by all a priori developmental models: adjacent parts along the proximodistal jaw axis are correlated whereas more distant ones are not. The most notable exception is that the shape of the anterior incisor alveolus is correlated with the shape of the ramus (FA component) or coronoid (symmetric component). Those exceptions differ between species; in prairie deer mice, the molar alveolus is connected to more parts, and the incisor alveolus to fewer, than in fox squirrels. The structure of integration suggests that the mandible cannot be decomposed into parts but rather is a single connected unit, a result consistent with its functional integration. That match between functional and developmental integration may arise, at least in part, from function-induced growth, building developmental integration into the functional system and enabling direct epigenetic interactions to evolve when function does.     

Genomic,ecological, and morphological approaches to investigating species limits: A case study in modern taxonomy from Tropical Eastern Pacific surgeonfishes

A wide variety of species are distinguished by slight color variations. However, molecular analyses have repeatedly demonstrated that coloration does not always correspond to distinct evolutionary histories between closely related groups, suggesting that this trait is labile and can be misleading for species identification. In the present study, we analyze the evolutionary history of sister species of Prionurus surgeonfishes in the Tropical Eastern Pacific (TEP), which are distinguished by the presence or absence of dark spots on their body. We examined the species limits in this system using comparative specimen‐based approaches, a mitochondrial gene (COI), more than 800 nuclear loci (Ultraconserved Elements), and abiotic niche comparisons. The results indicate there is a complete overlap of meristic counts and morphometric measurements between the two species. Further, we detected multiple individuals with intermediate spotting patterns suggesting that coloration is not diagnostic. Mitochondrial data recovered a single main haplotype shared between the species and all locations resulting in a complete lack of structure (Φ_ST = 0). Genomic analyses also suggest low levels of genetic differentiation (F_ST = 0.013), and no alternatively fixed SNPs were detected between the two phenotypes. Furthermore, niche comparisons could not reject niche equivalency or similarity between the species. These results suggest that these two phenotypes are conspecific and widely distributed in the TEP. Here, we recognize Prionurus punctatus Gill 1862 as a junior subjective synonym of P. laticlavius (Valenciennes 1846). The underlying causes of phenotypic variation in this species are unknown. However, this system gives insight into general evolutionary dynamics within the TEP.     

Evolutionary relationships among aquatic anamorphs and teleomorphs: Lemonniera, Margaritispora, and Goniopila

The hypothesis that similar conidial morphologies in aquatic hyphomycetes are a result of convergent evolution was tested using molecular sequence data. Cladistic analyses were performed on partial sequences of 28S rDNA of seven species of Lemonniera, one species of Margaritispora and one species of Goniopila. Lemonniera has tetraradiate conidia with long arms, whereas Margaritispora and Goniopila have typically globose (isodiametric) conidia, with short conical protuberances in a stellate or quadrangular arrangement. Lemonniera and Margaritispora have phialidic conidiogenesis and both produce dark, minute sclerotia in culture whereas Goniopila has holoblastic conidiogenesis and does not produce sclerotia in culture. Goniopila produces a microconidial phialidic synanamorph in culture. All three genera have schizolytic conidial secession. Molecular analyses demonstrate that Lemonniera species are placed in two distinct clades: one within Leotiomycetes; the other within Pleosporales, Dothideomycetes. Margaritispora is placed with Lemonniera species within Leotiomycetes. Goniopila and Lemonniera pseudofloscula are placed within Dothideomycetes. No morphological character was entirely congruent with the molecular derived phylogeny. This suggests that for the group of species studied, conidial shape is not a reliable indicator of phylogeny but more likely the result of convergent evolution in response to the aquatic environment.     

地黄属分子系统学分析

地黄属是中国准特有属,属内种间关系仍待澄清。该研究基于多个个体的叶绿体与核基因片段对地黄属进行系统发育重建,探讨属内物种分化与可能历史。结果表明:(1)地黄属为单系群,天目地黄为本属原始类群,并与湖北地黄互为姐妹群或连续姐妹群,裂叶地黄-高地黄、地黄-茄叶地黄分别组成姐妹群;(2)多个个体构建的系统树更能揭示本属物种关系的复杂性;(3)祖先分布区重建表明本属经历过3次扩张与2次隔离过程,物种形成过程与历史气候变化密切相关。最后该研究提出了地黄属物种形成研究亟待解决的几个重要问题。     

Alkaloid metabolism by cytochrome P-450 enzymes in Drosophila melanogaster

The cytochrome P-450 family of enzymes is the primary means of foreign compound detoxification in virtually all organisms. Cytochrome P-450s have been strongly implicated in the metabolism of cactus alkaloids, and consequently, the observed patterns of host plant utilization by cactophilic species of Drosophila in the Sonoran Desert. The current study looked for evidence of alkaloid-metabolizing P-450 enzymes in a non-cactophilic species, D. melanogaster. The results of in vitro metabolism assays indicate the presence of a phenobarbital-inducible P-450 in adult D. melanogaster which is capable of metabolizing alkaloids. P-450 quantification data suggest that the enhanced level of metabolism is not the result of an overall increase in total P-450 content. Results from larval viability and adult longevity studies indicate that D. melanogaster's in vitro activity does not produce an enhanced in vivo tolerance of alkaloids.     

The phylogenetic placement of the Leucogastrales, including Mycolevis siccigleba (Cribbeaceae), in the Albatrellaceae using morphological and molecular data

The morphology of many hypogeous fungi converges on a homogeneous reduced form, suggesting that disparate lineages are subject to a uniform selection pressure. The primary goal of this study was to evaluate the morphology and infer the phylogeny of the Leucogastrales with Mycolevis siccigleba using a Bayesian methodology. A comprehensive morphological assessment was used for an a priori phylogenetic inference to guide the sequencing effort. All structures except spore ornamentation pointed to the Albatrellaceae as the most likely sister taxon. Polyporoletus sublividus, a close relative of Albatrellus, produces ornamented basidiospores with a similar structure to M. siccigleba basidiospores. The ITS from 30 taxa was used for the molecular phylogenetic analysis. P. sublividus was found sister to Mycolevis. Leucophleps spinispora and L. magnata formed a group sister to the Polyporoletus/Mycolevis group, whereas Leucogaster was polyphyletic with respect to the core of the Leucogastrales and sister to A. caeruleoporus. This relationship was expected as previously undescribed chlamydospores produced by members of Albatrellus had a similar morphology to the basidiospores of L. rubescens.     

A phylogenetic overview of the family Pyronemataceae (Ascomycota, Pezizales)

Partial sequences of nuLSU rDNA were obtained to investigate the phylogenetic relationships of Pyronemataceae, the largest and least studied family of Pezizales. The dataset includes sequences for 162 species from 51 genera of Pyronemataceae, and 39 species from an additional 13 families of Pezizales. Parsimony, ML, and Bayesian analyses suggest that Pyronemataceae is not monophyletic as it is currently circumscribed. Ascodesmidaceae is nested within Pyronemataceae, and several pyronemataceous taxa are resolved outside the family. Glaziellaceae forms the sister group to Pyronemataceae in ML analyses, but this relationship, as well as those of Pyronemataceae to the other members of the lineage, are not resolved with support. Fourteen clades of pyronemataceous taxa are well supported and/or present in all recovered trees. Several pyronemataceous genera are suggested to be non-monophyletic, including Anthracobia, Cheilymenia, Geopyxis, Humaria, Lasiobolidium, Neottiella, Octospora, Pulvinula, Stephensia, Tricharina, and Trichophaea. Cleistothecial and truffle or truffle-like ascomata forms appear to have evolved independently multiple times within Pyronemataceae. Results of these analyses do not support previous classifications of Pyronemataceae, and suggest that morphological characters traditionally used to segregate the family into subfamilial groups are not phylogenetically informative above the genus level.     

Molecular phylogeny of the yuhinas (Sylviidae: Yuhina): a paraphyletic group of babblers including Zosterops and Philippine Stachyris

Mitochondrial sequences (2,379 bp) from cytochrome b, ND3, 12s and 16s rRNA were analyzed in order to reconstruct the phylogenetic relationships within the yuhinas (Yuhina), including the chestnut-faced babbler Stachyris whiteheadi which is endemic to the Philippines, the Japanese white-eye Zosterops japonicus, the chestnut-flanked white-eye Z. erythropleurus, and the oriental white-eye Z. palpebrosus. The results showed strong support for the idea that S.whiteheadi and three white-eye species form a clade embedded within the Yuhina clade. The robustness of the Yuhina/Philippine Stachyris/Zosterops clade rejected the validity of the monotypic genus Staphida, which was proposed for the striated yuhina Yuhina castaniceps based on its peculiar morphological traits. Sister group relationships between the striped-throated yuhina Yuhina gularis and the rufous-vented yuhina Yuhina occipitalis and between the whiskered yuhina Yuhina flavicollis and the white-naped yuhina Yuhina bakeri were discovered. The sympatric patterns of the above two sister groups in the Himalayas is most likely due to secondary contact. The molecular phylogeny also suggests that crestlessness was derived just once for the Zosterops/Philippine Stachyris clade. The Zosterops/Philippine Stachyris clade in our study also implies that an ancestor of Zosterops/Philippine Stachyris derived the ability to disperse over long distances, so that it could fly over the sea and arrive at the Philippines.     

择伐对思茅松自然种群结构和空间分布格局的影响

思茅松是云南省重要的材脂兼用树种,而择伐是思茅松天然林管理的有效手段,也是获取木材的重要途径。基于对云南省景谷县思茅松天然林的样地调查数据,采用种群径级结构代替年龄结构、Ripley的L函数点格局等方法,对思茅松天然林在不同强度择伐10a后的种群结构和空间分布格局及其不同发育阶段在不同尺度下的相互关系进行了系统研究和分析。结果表明:择伐10a后3种类型的思茅松自然种群的胸径、树高之间呈显著的正相关且可用幂指数模拟预测模型。未择伐的思茅松种群呈衰退型,轻度择伐(12.9%)后种群呈稳定型,中度择伐(29.3%)后的种群属增长型,适当的中度择伐有利于自然种群的更新;未择伐与中度择伐后的思茅松种群分布格局整体呈随机型,但后者较前者更趋向于均匀分布,轻度择伐后的思茅松种群分布格局随着尺度的增加由随机转向聚集分布。择伐对不同径级思茅松天然林种群空间分布格局有重要影响,未择伐的思茅松自然种群随着发育阶段的发展,多数尺度上由随机分布逐渐转变为聚集分布,至大树则又以随机分布为主;轻度择伐和中度择伐的思茅松自然种群则随着发育阶段的发展而由完全的随机分布逐渐转变为聚集分布,中度择伐的聚集分布则更为明显。思茅松不同发育阶段间在所有尺度上基本都呈显著正相关,表明思茅松种群种内竞争弱,有利于种群的维持。     

Cortinarius badiolaevis, a new conifer-associated, darkening species in the subgenus Telamonia (Basidiomycota, Agaricales)

A new species, Cortinarius badiolaevis Niskanen, Liimat., Mahiques, Ballarà, Kyt?v. in subgenus Telamonia, is described based on morphological and molecular data. Typical for the species is a dark brown cap, pale flesh, indistinct universal veil, small amygdaloid to ellipsoid spores [7.0–8.5 × 4.5–5.0(−5.5) μm], and dark exsiccata. It occurs in mesic to dryish coniferous forests, and grows in calcareous soil presumably with Pinus and Picea. So far, it is only known from Spain and Sweden, and considered rare. Based on the phylogenetic analyses of the ITS regions, it does not belong to any of the known sections, nor does it have any very closely related sister species.