Morphological variability in time and space: an example of patterns within buchiid bivalves (Bivalvia,Buchiidae)

Abstract: We studied morphological variation of the bivalve Buchia over its geographical and temporal range. Buchia was widely distributed during the Late Jurassic and Early Cretaceous, and, while previous quantitative studies have shown that species are characterized by large amounts of variation, there have been no prior attempts to measure how morphology varies geographically. We employed traditional morphometric techniques using nine linear/angular measurements on 1855 buchiid shells from eight localities taken from widely separated, but mostly coeval, sections across its range. Principal component and canonical variate analyses indicate that geographical morphospace of buchiids varied significantly, but we did not find evidence of a latitudinal gradient in shell shape. The amount of variation between localities was similar to the amount of variation between species, indicating the importance of geographical effects on morphology. Disparity (morphological diversity within a taxon, calculated by the sum of variances) and diversity (number of species) were calculated for each location and time period (age). Disparity and diversity reached ultimate lows just before the genus’ extinction in the Hauterivian, and is suggestive that extinction was morphologically selective. We did not find significant trends for either metric, but there were discordances throughout its temporal range. Latitudinal trends of disparity and diversity within Buchia are not apparent. This research adds to the growing body of work on geographical variation and is a preliminary step to understanding the nature and variation of buchiid species and of biodiversity in general.     

Integrated species delimitation and conservation implications of an endangered weevil Pachyrhynchus sonani (Coleoptera: Curculionidae) in Green and Orchid Islands of Taiwan

Oceanic islands are productive habitats for generating new species and high endemism, which is primarily due to their geographical isolation, smaller population sizes and local adaptation. However, the short divergence times and subtle morphological or ecological divergence of insular organisms may obscure species identity, so the cryptic endemism on islands may be underestimated. The endangered weevil Pachyrhynchus sonani Kôno (Coleoptera: Curculionidae: Entiminae: Pachyrhynchini) is endemic to Green Island and Orchid Island of the Taiwan‐Luzon Archipelago and displays widespread variation in coloration and host range, thus raising questions regarding its species boundaries and degree of cryptic diversity. We tested the species boundaries of P. sonani using an integrated approach that combined morphological (body size and shape, genital shape, coloration and cuticular scale), genetic (four genes and restriction site‐associated DNA sequencing, RAD‐seq) and ecological (host range and distribution) diversity. The results indicated that all the morphological datasets for male P. sonani, except for the colour spectrum, reveal overlapping but statistically significant differences between islands. In contrast, the morphology of the female P. sonani showed minimum divergence between island populations. The populations of P. sonani on the two islands were significantly different in their host ranges, and the genetic clustering and phylogenies of P. sonani established two valid evolutionary species. Integrated species delimitation combining morphological, molecular and ecological characters supported two distinct species of P. sonani from Green Island and Orchid Island. The Green Island population was described as P. jitanasaius sp.n. Chen & Lin, and it is recommended that its threatened conservation status be recognized. Our findings suggest that the inter‐island speciation of endemic organisms inhabiting both islands may be more common than previously thought, and they highlight the possibility that the cryptic diversity of small oceanic islands may still be largely underestimated.     

藏南喜马拉雅地区晚侏罗世-早白垩世双壳类雏蛤动物群

藏南喜马拉雅地区晚侏罗世地层中雏蛤化石丰富,其它化石仅有少量箭石和菊石,而且属种单一,多为地方种,无法进一步划分和对比,因此,雏蛤属(Buchia)的组合特征具有重要的地质意义。结合菊石Uhligites和Substeuraces,作者在藏南喜马拉雅地区上侏罗统中识别出:1)Buchia concentrica-Buchia spitiensis;2)Buchiamosquensis- Buchia rugosa;3)Buchia blanfordiana- Buchia piochii等3个雏蛤动物组合;在下白垩统中识别出Buchiasubokensis雏蛤动物组合,其中,Buchia mosquensis-Buchia rugosa组合为证明拉弄拉地区存在晚侏罗世晚基末利期至早提塘期地层提供了新的证据,并与斯匹提地区、北美等地区进行了动物群和地层的对比,初步探讨了本区雏蛤动物群在生物地层方面的研究价值。     

Evolutionary patterns in the terebratulid genus Caryona Cooper (Brachiopoda,Jurassic of France)

The study of ontogenetic and morphological changes in different species of the brachiopod genus Caryona COOPER (Terebratulidae) from the Lower Callovian‐Lower Oxfordian of northern France shows a succession (and an interference) of processes which vary according to the characters studied. This is an example of “mosaic evolution”;. The ontogenetic changes result from heterochrony, especially acceleration and hypermorphism. The main phylogenetic tendency is peramorphosis in most characters. This evolutionary pattern agrees with phyletic gradualism.

Moreover, analysis of the internal characters demonstrates the limited value of these in discrimination of the various genera.     

THE OCCURRENCE OF THE UPPER JURASSIC BIVALVE MALAYOMAORICA MALAYOMAORICA (KRUMBECK) ON THE ORVILLE COAST, ANTARCTICA

Malayomaorica malayomaorica is an important Upper Jurassic bivalvein the southern hemisphere: widely distributed in strata ofEarly - Middle Kimmeridgian age, it is a zone fossil of considerablepotential for regional biostratigraphic correlations. Its commonoccurrence in the Latady Formation of the Orville Coast, Antarctica,indicates that at least part of this stratigraphic unit hasa Kimmeridgian age. Although its precise taxonomic status remainsin some doubt, it would appear to be the earliest buchiid-likebivalve so far recorded from the southern hemisphere. Its verywide distribution around the margins of Gondwana is similarto that established for species of the Late Jurassic bivalvegenera Retroceramus, Buchia and Anopaea (Received 13 April 1981;     

Taxonomic revision of the genus Galictis (Carnivora: Mustelidae): species delimitation,morphological diagnosis,and refined mapping of geographical distribution

Although critical for enabling in‐depth evolutionary, ecological, or conservation‐orientated studies, taxonomic knowledge is still scarce for many groups of organisms, including mammals of the order Carnivora. For some of these taxa, even basic aspects such as species limits and geographical distribution are still uncertain. This is the case for the Neotropical mustelid genus Galictis, considered one of the least studied carnivoran genera in the Americas. To address this issue, we performed a comprehensive assessment of morphological and molecular characters to test the number of species within Galictis, and to characterize their distinctiveness and evolutionary history. In addition, we reviewed and consolidated the available information on the taxonomy of this genus, so as to provide a historical framework upon which we could interpret our data. Our analyses demonstrated that two Galictis species can be clearly delimited and diagnosed using metric and nonmetric morphological characters as well as DNA sequences from mitochondrial and nuclear gene segments. On the basis of this clarified species‐level delimitation, we reassessed the geographical range of each Galictis taxon, identifying possible areas of sympatry between them. These results provide a solid taxonomic framework for Galictis, enabling the development of additional studies focusing on this poorly known taxon. © 2013 The Linnean Society of London, Zoological Journal of the Linnean Society, 2013, 167 , 449–472.     

Unravelling evolutionary lineages among South African velvet worms (Onychophora: Peripatopsis) provides evidence for widespread cryptic speciation

The endemic South African velvet worm genus Peripatopsis currently contains eight recognized species described from variable morphological characters and the current taxonomy is unsatisfactory. In an attempt to investigate evolutionary relationships within Peripatopsis, we collected 137 individuals from 34 sample localities for six of the eight species. Sequence data derived from two partial mitochondrial (mt)DNA gene loci (COI and 12S rRNA), as well as partial sequence data from the ribosomal nuclear 18S rDNA locus in combination with gross morphological characters and scanning electron microscopy (SEM), was used to examine evolutionary relationships. Phylogenetic relationships were investigated using minimum evolution (ME) and Bayesian inferences (BI). Additionally, we also undertook a maximum likelihood (ML) analyses on the combined DNA sequence data set. The combined DNA evidence topologies derived from the ME, BI, and ML was highly congruent and was characterized by the presence of multiple lineages within recognized taxa. Peripatopsis clavigera, Peripatopsis moseleyi, and Peripatopsis sedgwicki each comprised two evolutionary lineages; Peripatopsis capensis comprised three; and Peripatopsis balfouri comprised six operational taxonomic units respectively. Genealogical exclusivity at both mtDNA and nuclear DNA among the geographically coherent groups coupled with pronounced sequence divergence suggested a two‐fold increase in the number of species within Peripatopsis. Previously used gross morphological characters (such as the number of leg pairs and colour) were either highly variable within operational taxonomic units, or were invariant, suggesting that alternative morphological characters are necessary for species discrimination. SEM results revealed potentially useful diagnostic characters that can discriminate between at least discriminate some of the newly‐identified lineages. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society 2009, 97 , 200–216.     

Habitat fragmentation and body‐shape variation of African hind Cephalopholis taeniops (Valenciennes) in an archipelago system (Cape Verde,eastern Atlantic Ocean)

Morphological variations of the serranid fish Cephalopholis taeniops were studied in relation to habitat fragmentation in the Cape Verde Archipelago. While a significant allometric effect existed (11·5% of total body‐shape variation), differences in morphology associated with sex and ontogeny were not significant. MANOVA followed by CVA showed that each island presented a particular allometric pattern. Average body shape for all islands was well discriminated with CVA models. Pair‐wise comparisons of the ontogeny of morphological change between islands revealed that northern islands (Santo Antão, São Vicente and Santa Luzia) along with Boavista Island showed a similar direction in shape ontogeny, while all other paired comparisons indicated different ontogenetic patterns. When comparing directions of inter‐population shape changes, individuals from Fogo Island, the southernmost locality, departed far from the orthogonal relation, suggesting that they were undergoing independent body‐shape trajectories. Physical isolation by geographic distance and depth was positively correlated with morphological divergence among populations from different islands. This finding supports the hypothesis that habitat fragmentation in the Cape Verde Archipelago can be interpreted in terms of marine population structure.     

Genetic and morphological studies of Nothobranchius (Cyprinodontiformes) from Malawi with description of Nothobranchius wattersi sp. nov.

Molecular and morphological data were used to explore evolutionary differentiation among populations of Nothobranchius in the Lake Malawi–upper Shire River and the Lakes Chilwa–Chiuta drainage systems in Malawi. The aim of the study was to test the hypothesis that Nothobranchius of the Malawi–Shire system constitute a separate evolutionary group from Nothobranchius kirki. Mitochondrial and nuclear sequence data show a strongly supported phylogenetic split into two monophyletic groups separating the Lake Malawi basin fish from N. kirki. Unlike N. kirki, Lake Malawi–Shire fish do not deviate from neutrality and express an excess of rare haplotypes and mutations in terminal branches, characteristic of recently expanded populations. Further, the two groups significantly differ in morphology. Two body characters (dorsal‐fin base length and pre‐pelvic–pre‐anal distance) are significantly different between the two species in both sexes. Several other characters are significantly different in either male or female comparisons with respect to both standard and head lengths, and robust morphological differentiation is detected by multivariate analysis. The two groups are readily distinguished on the basis of male colouration, especially in scale centres and the caudal fin. On the basis of this differentiation at the molecular and morphological levels, in addition to colouration, the Lake Malawi–Shire fish are hereby formally recognized as constituting a new species, Nothobranchius wattersi. This distinction is in agreement with the geomorphologic and recent climatic history in the region.     

Evolution of skull and body shape in Triturus newts reconstructed from three‐dimensional morphometric data and phylogeny

To explore the relationship between morphological change and species diversification, we reconstructed the evolutionary changes in skull size, skull shape, and body elongation in a monophyletic group of eight species that make up salamander genus Triturus. Their well‐studied phylogenetic relationships and the marked difference in ecological preferences among five species groups makes this genus an excellent model system for the study of morphological evolution. The study involved three‐dimensional imagery of the skull and the number of trunk vertebrae, in material that represents the morphological, spatial, and molecular diversity of the genus. Morphological change largely followed the pattern of descent. The reconstruction of ancestral skull shape indicated that morphological change was mostly confined to two episodes, corresponding to the ancestral lineage that all crested newts have in common and the Triturus dobrogicus lineage. When corrected for common descent, evolution of skull shape was correlated to change in skull size. Also, skull size and shape, as well as body shape, as inferred from the number of trunk vertebrae, were correlated, indicating a marked impact of species' ecological preferences on morphological evolution, accompanied by a series of niche shifts, with the most pronounced one in the T. dobrogicus lineage. The presence of phylogenetic signal and correlated evolutionary changes in skull and body shape suggested complex interplay of niche shifts, natural selection, and constraints by a common developmental system. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 243–255.     

Evolution of the Juan Fernández diving beetle,Rhantus selkirki (Coleoptera,Dytiscidae)

Here we provide evidence that confinement in Robinson Crusoe Island (located about 660 km west of continental Chile) over evolutionary time leads to strong morphological modifications in diving beetle (Dytiscidae) larvae. We analysed a large set of morphological larval characters for all currently recognised genera of Colymbetinae as a framework, to infer phylogenetic relationships within the large genus Rhantus Dejean, 1833 and, in particular, of the charismatic Juan Fernández diving beetle, Rhantus selkirki Jäch, Balke & Michat, 2015, comparing our results with a recent phylogeny of the Colymbetinae based on DNA sequence data. We suggest that adaptation to the island's particular habitats resulted in the reversal of certain characters of R. selkirki back to the plesiomorphic states. This may cause the species to be erroneously interpreted as more ‘primitive’ if only morphological characters are analysed. Confinement in the particular, shallow and barely vegetated aquatic habitats of Robinson Crusoe Island for a long time seems to have led to this divergent morphology, particularly in characters related to swimming ability such as several leg and urogomphal setae. In this way, R. selkirki larvae secondarily resemble those of some earlier diverging dytiscid lineages such as Agabinae and Copelatinae, which typically creep on the bottom of water bodies and do not swim well.     

A NEW BLACKFLY SPECIES OF SIMULIUM (DIPTERA: SIMULIIDAE) FROM GUIZHOU PROVINCE,CHINA*

Abstract The present paper deals with a new species of Simulium (Simulium) from Leigongshan, Guizhou province, S. (S.) qianense sp. nov. The specific name was given in the place of production. On the basis of the morphological characters, the new species (S.) qianense sp. nov evidently falls into S. variegatum species‐group, is most similar to S. (S.) taiwanicum, 1979, but can be distinguished from the latter by the characteristic pattern of scutum and legs color in the both sexes, the shape of male geritalia and several characters in the larva. The type specimens are kept in the Department of Biology of the Guiyang Medical College.     

Patterns,trends, and rates of evolution within the Actinistia

Synopsis The interrelationships of 31 actinistian species (including Latimeria chalumnae) are analyzed based on a cladistic analysis of 75 osteological characters. Inference of evolutionary trends (e.g., modification of body shape and skull morphology) from the phylogenetic patterns demonstrates that the morphology of actinistians is less conservative than has been proposed previously. This empirical cladistic approach supports two distinct tempos of evolution during an evolutionary history of 380 million years. Along a phylogenetic pathway originating with a Devonian stem-species and ending with the living Latimeria chalumnae (including 101 morphological changes and 18 cladogenetic events), the first tempo occurred during the Devonian — Permian periods as a decreasing rate of morphological changes, which was followed by a stabilizing tempo during the Permian — Recent periods. The decreasing tempo is characterized by a sequence of gradual versus quantum temporal changes and low versus faster rates, whereas the stabilizing tempo primarily is gradual and low. In contrast to a common assumption, no significant correlation was found between the rates of morphological evolution and the temporal diversity of species.     

Morphological divergence of lake and stream Phoxinus of Northern Italy and the Danube basin based on geometric morphometric analysis

Minnows of the genus Phoxinus are promising candidates to investigate adaptive divergence, as they inhabit both still and running waters of a variety of altitudes and climatic zones in Europe. We used landmark‐based geometric morphometric methods to quantify the level of morphological variability in Phoxinus populations from streams and lakes of Northern Italy and the Danube basin. We analyzed body shape differences of populations in the dorsal, lateral, and ventral planes, using a large array of landmarks and semilandmarks. As the species identification of Phoxinus on morphological characters is ambiguous, we used two mitochondrial genes to determine the genetic background of the samples and to ensure we are comparing homogenous groups. We have found significant body shape differences between habitats: Minnow populations inhabiting streams had a deeper body and caudal peduncle and more laterally inserted pectoral fins than minnows inhabiting lakes. We have also found significant body shape differences between genetic groups: Italian minnows had deeper bodies, deeper and shorter caudal peduncles, and a shorter and wider gape than both groups from the Danube. Our results show that the morphology of Phoxinus is highly influenced by habitat and that body shape variation between habitats was within the same range as between genetic groups. These morphological differences are possibly linked to different modes of swimming and foraging in the respective habitats and are likely results of phenotypic plasticity. However, differences in shape and interlandmark distances between the groups suggest that some (though few) morphometric characters might be useful for separating Phoxinus species.     

Realized evolvability: quantifying phenotypic evolution in a Drosophila clade

Understanding the evolutionary potential of morphological evolution is still a major problem in evolutionary biology. In this study, we tried to quantify the amount of variation of different traits among species of a Drosophila clade reared under standard conditions. Nineteen different traits have been measured on nine species of the same clade, the Neotropical saltans group of Drosophila. Measured traits can be distributed into five categories: size‐traits (wing and thorax), shape indices (ratios among the size traits), number of sternopleural bristles on the thorax, number of abdominal bristles on successive sternites, and dorsal pigmentation of abdomen. All species are of medium size with a generally dark pigmentation. A remarkable feature is the presence of numerous bristles on the 6th sternite of the males, while this segment is bare in other Drosophila species. A multivariate analysis revealed that it was possible to discriminate all the investigated species by using the complete set of measured traits. For each trait, phenotypic variability was investigated at the within‐ and between‐species levels. As a rule, the interspecific coefficient of variation (CV) was much greater than the within species CV, and it is proposed to call it realized evolvability. All possible correlations were calculated between traits across species, providing many unexpected results. Size traits were highly correlated among them, but not correlated with shape indices. Abdominal traits (bristles and pigmentation) were correlated, but often in opposite directions, with thoracic shape indices. Tergite pigmentation was negatively correlated with bristle number on sternite. For the moment, most of the correlations cannot be explained by developmental processes or parallel selective pressures. Nonetheless, mapping the evolution of the two characters on a molecular phylogeny of the studied species revealed two opposite phylogenetic trends for abdominal pigmentation and setation, respectively. Our data suggest a need for similar studies in other well‐defined Drosophila clades.