Geographic distribution and phenetic skull variation in two close species of Graomys (Rodentia, Cricetidae, Sigmodontinae)

The interspecific differentiation of South American rodents of the genus Graomys was assayed at ecological and morphometric levels in two species. At the ecological level, niche modelling was used. At the morphometric level, the hypothesis that the size and shape of the skull vary with the geographic location was tested using geometric morphometrics by assessing the extent and spatial distribution of phenotypic skull variation within and among two species, Graomys griseoflavus and Graomys chacoensis. Our results of niche modelling indicate the spatial differentiation between the two species, with G. chacoensis inhabiting preferably the Chaco ecoregion and G. griseoflavus inhabiting mainly the Monte ecoregion. In multiple linear regressions, approximately 20% of the skull size variation is explained by latitude, altitude, and temperature seasonality. The partial least square analysis reveals strong correlation between shape and environmental variables, mainly with latitude, annual mean temperature, and annual precipitation. Discrimination between G. griseoflavus and G. chacoensis was highly reliable when using geometric morphometric tools. These results permit us to elucidate some evolutionary processes that have occurred in these species.     

Morphometric analysis of six Gerbillus species (Rodentia,Gerbillinae) from Tunisia

Size and shape changes in the skull of the genus Gerbillus were investigated using geometric morphometrics. Six species from Tunisia were studied (G. gerbillus, G. campestris, G. nanus, G. tarabuli, G. simoni and G. latastei). Statistical analyses of shape variability allowed us to discriminate three morphological groups which are congruent with the three groups suggested by previous morphological and molecular studies. However, our results contrast with previous molecular investigations. In fact, according to results obtained by the use of principal component analysis, canonical variate analysis and UPGMA, we found a higher degree of divergence between the subgenus Dipodillus and the other two subgenera Gerbillus and Hendecapleura. This fact suggests that the morphometric differences observed among species within the genus Gerbillus are not mainly related to phylogeny. To reconciliate the molecular and morphological approaches, we propose a hypothesis of differential rates of phenotypic evolution in the genus Gerbillus. In this view, the species belonging to the subgenus Dipodillus evolved apomorphic features of the skull likely related to a higher degree of habitat specialization. By contrast, the more generalist Gerbillus and Hendecapleura subgenera show less differentiated plesiomorphic morphology.     

Discrimination of the sibling species Apodemus flavicollis and A. sylvaticus (Rodentia,Muridae)

Karyotyping and several molecular methods have allowed successful identification of two morphologically similar wide-ranging Western Palearctic species, the yellow-necked field mouse Apodemus flavicollis (Melchior, 1934) and the long-tailed wood mouse A. sylvaticus (Linnaeus, 1758), but reliable species diagnosis on the basis of morphometric characters is particularly problematic. Although they are easily morphologically distinguishable in Central and Northern Europe, this is not the case in southern parts of their distribution areas. Despite that, we have successfully discriminated A. flavicollis and A. sylvaticus from Serbia (Southern Europe) using geometric and traditional morphometric methods on a data set for ventral crania of specimens previously genotyped by the Inter Simple Sequence Repeat-PCR (ISSR-PCR). Discrimination power of applied approaches was more or less similar. The majority of our results were consistent with those obtained for specimens collected across the Czech Republic (Central Europe). Morphological differences observed herein, as well as those already reported between A. flavicollis and A. sylvaticus from the central and northern parts of their distribution areas, could be the outcome of their biology, i.e. ecological discrepancies, different assumed evolutionary scenarios considering biogeography, phylogeny, history and ontogeny.     

Geometric craniometric analysis of sexual dimorphism and ontogenetic variation: A case study based on two geographically disparate species, Aethomys ineptus from southern Africa and Arvicanthis niloticus from Sudan (Rodentia: Muridae)

Non-geographic morphometric variation, particularly at the level of sexual dimorphism and ontogenetic (age-related) variation, has been documented in rodents, and useful for establishing whether to analyse sexes separately or together, and for selecting adult specimens for subsequent data recording and analysis. However, such studies have largely been based on traditional morphometric analyses of linear measurements that mainly focus on overall size, rather than shape-related morphometric variation. Unit-free, landmark/outline-based geometric morphometric analyses are considered to offer a more appropriate tool for assessing shape-related morphometric variation. In this study, we used geometric cranial morphometric analysis to assess the nature and extent of sexual dimorphism and age variation within the Tete veld rat, Aethomys ineptus (Thomas and Wroughton, 1908) from southern Africa and the African Nile rat, Arvicanthis niloticus (Desmarest, 1822) from Sudan. The results obtained were in turn compared with previously published results based on independent geometric and traditional cranial morphometric data from the same sampled populations examined in the present study. While our geometric morphometric results detected statistically significant sexual dimorphism in cranial shape within Ar. niloticus only, previously published results based on traditional morphometric data failed to detect significant sexual dimorphism within this species. However, similar to previously published traditional morphometric data, our geometric morphometric results detected statistically significant age-related variation in cranial shape and size within both Ae. ineptus and Ar. niloticus, with individuals of age classes 5 and 6 being considered to represent adult specimens. Our results highlight the importance of carefully evaluating both size- and shape-related non-geographic morphometric variation prior to the analysis of geographic variation and the delineation of species. Erroneous conclusions of non-geographic variation may have implications in the interpretation of geographic and evolutionary processes that may be responsible for morphological differences at both the inter- and intra-specific levels.     

Intercontinental effect on sexual shape dimorphism and allometric relationships in the beetle pest Diabrotica virgifera virgifera LeConte (Coleoptera: Chrysomelidae)

Hind wing shape variation was examined in 686 adult Diabrotica virgifera virgifera collected from maize plants in Europe and the USA Corn Belt, using geometric morphometric techniques. Sexual dimorphism at an Intercontinental scale was assessed using canonical variates analysis, a multivariate statistical method used to find the shape characters that best distinguish among groups of specimens. Our results showed that each of the populations of D. v. virgifera investigated in this study showed high levels of sex based hind wing shape dimorphism. In particular a stronger and more obvious pattern of hind wing shape variation was found in the USA than in Europe. These results support previous studies on D. v. virgifera wing shape that show that female D. v. virgifera have more elongated wings than males. These differences raise the question of whether sexual dimorphism may be modulated by natural selection.     

A new species of Golunda (Rodentia, Muridae) from the Late Pleistocene of Indian Himalaya

The youngest fossil Golunda (Rodentia, Muridae) is described from the Late Pleistocene fluvio-lacustrine deposits, exposed at Dulam (Bageshwar), Kumaun Lesser Himalaya, India. The age of fossiliferrous horizon is estimated as 31,000 yr BP. A new species, Golunda dulamensis nov. sp. has highly derived characters, e.g., antero-posteriorly stretched molars, upper molars with more length and less width, stephanodonty, cusps in M₃ strongly inclined backward giving the molars a very stretched aspect, and metaconid and entoconid in M₃ forming almost straight lingual row of the cusps. G. dulamensis nov. sp. is most similar to present day G. ellioti but differs from the later slightly by larger size, a thin connection between t₄ and t₅ in M¹, and comparatively larger entoconid and very weakly developed antero-labial cusp in M₃. We suggest that highly specialized molars of G. dulamensis nov. sp. and present day G. ellioti are derivable through G. kelleri. We also propose that Golunda migrated from Asia to Africa, not from Africa to Asia as was thought by earlier workers.     

Geographic phenetic variation of two eastern-Mediterranean non-commensal mouse species, Mus macedonicus and M. cypriacus (Rodentia: Muridae) based on traditional and geometric approaches to morphometrics

We tested the hypothesis that skull shape within the genus Mus may vary with geographic location by assessing the extent and spatial distribution of phenotypic skull variation within and among two wild mouse species, M. macedonicus and M. cypriacus, using traditional and geometric morphometrics including a rather novel application of sliding semilandmarks. Shape was shown to be significantly correlated both with longitude and latitude in M. macedonicus, yet the correlation between morphometric and geographic distances was not significant, and morphometric differences between Asian and European populations were not higher than those within the particular continents. The phylogenetic signal was found to be stronger in dental characters than in cranial ones, however, overall concordance between the pattern of morphometric variation and the presumed history of M. macedonicus was rather weak. In both species, the dorsal and ventral sides of the skull were shown to covary in many aspects though there were also some differences between them, making the functional interpretation of these differences difficult. Discrimination between M. cypriacus and M. macedonicus as well as discrimination between two M. macedonicus subspecies was highly reliable using both traditional and geometric morphometric tools to analyze skull measurements.     

Phylogenetic relationships in European Ceriporiopsis species inferred from nuclear and mitochondrial ribosomal DNA sequences

The aim of this work was to clarify taxonomy and examine evolutionary relationships within European Ceriporiopsis species using a combined analysis of the large subunit (nLSU) nuclear rRNA and small subunit (mtSSU) mitochondrial rRNA gene sequences. Data from the ITS region were applied to enhance the view of the phylogenetic relationships among different species. The studied samples grouped into four complex clades, suggesting that the genus Ceriporiopsis is polyphyletic. The generic type Ceriporiopsis gilvescens formed a separate group together with Ceriporiopsis guidella and Phlebia spp. in the phlebioid clade. In this clade, the closely related species Ceriporiopsis resinascens and Ceriporiopsis pseudogilvescens grouped together with Ceriporiopsis aneirina. C. resinascens and C. pseudogilvescens have identical LSU and SSU sequences but differ in ITS. Ceriporiopsis pannocincta also fell in the phlebioid clade, but showed closer proximity to Gloeoporus dichrous than to C. gilvescens or C. aneirina–C. pseudogilvescens–C. resinascens group. Another clade was composed of a Ceriporiopsis balaenae–Ceriporiopsis consobrina group and was found to be closely related to Antrodiella and Frantisekia, with the overall clade highly reminiscent of the residual polyporoid clade. The monotypic genus Pouzaroporia, erected in the past for Ceriporiopsis subrufa due to its remarkable morphological differences, also fell within the residual polyporoid clade. Ceriporiopsis subvermispora held an isolated position from the other species of the genus. Therefore, the previously proposed name Gelatoporia subvermispora has been adopted for this species. Physisporinus rivulosus appeared unrelated to two other European Physisporinus species. Moreover, Ceriporiopsis (=Skeletocutis) jelicii grouped in a separate clade, distinct from Ceriporiopsis species. Finally, the ITS data demonstrated the proximity of some Ceriporiopsis species (Ceriporiopsis portcrosensis and Ceriporiopsis subsphaerospora) to Skeletocutis amorpha.     

草兔胚后发育过程中头骨大小与形状变化的几何形态学分析(兔形目:兔科)

在本文中,我们使用几何形态学的研究方法对58 个处于不同年龄阶段的草兔头骨进行了分析。分析中总共使用来自头骨背面、腹面和侧面的180 个标点和半标点。研究结果表明,草兔在胚后发育早期即迅速建立起与成年个体近似的形态结构,组成头骨的不同形态单元存在显著的异速生长现象,主要的形变发生在幼年至年龄1 阶段,即出生后的6 个月以内。从大小的变化来看,鼻骨在胚后发育过程中呈现正的异速生长,额骨和眼眶区则与头骨整体大小变化基本等速,而顶骨、听泡和枕骨大孔则呈现显著的负异速生长。我们亦用几何形态学的方法绘制出头骨在生长发育早期和晚期不同部位的形态变化轨迹,这一结果显示较显著的形状变化发生在鼻骨、前颌骨、眶上突和头骨的纵轴方向。头骨整体形态在胚后发育过程中伸长并变窄。这些变化将有利于幼兔较早实现完善的头部系统的建立,尤其有利于提高呼吸系统的通风能力,提高在高强度的捕食压力下保持警觉,在多样的运动过程中保持身体平稳的能力,也可能进一步提高了对固态食物的处理能力。这种异速生长模式可能是善于奔跑的植食性哺乳动物在功能需求上的一种适应性进化特征。     

Ontogeny of skull size and shape changes within a framework of biphasic lifestyle: a case study in six <Emphasis Type="Italic">Triturus</Emphasis> species (Amphibia,Salamandridae)

As with many other amphibians, Triturus species are characterized by a biphasic life cycle with abrupt changes in the cranial skeleton during metamorphosis. The post-metamorphic shape changes of the cranial skeleton were investigated using geometric morphometric techniques in six species: Triturus alpestris, T. vulgaris, T. dobrogicus, T. cristatus, T. carnifex, and T. karelinii. The comparative analysis of ontogenetic trajectories revealed that these species have a conserved developmental rate with divergent ontogenetic trajectories of the ventral skull shape that mainly reflect phylogenetic relatedness. A striking exception in the ontogenetic pattern was possibly found in T. dobrogicus, characterized by a marked increase in the developmental rate compared to the other newt species. The size-related shape changes explained a large proportion of shape change during post-metamorphic growth within each species, with marked positive allometric growth of skull elements related to foraging.     

Species delimitation in the European species of Clavulina (Cantharellales, Basidiomycota) inferred from phylogenetic analyses of ITS region and morphological data

The identification of the conventionally accepted species of Clavulina (Cantharellales, Basidiomycota) in Europe (Clavulina amethystina, Clavulina cinerea, Clavulina cristata, and Clavulina rugosa) is often difficult and many specimens are not straightforwardly assignable to any of those four species, which is why some authors have questioned their identity. In order to assess the status of those species, a morphological examination was combined with the molecular analysis of the ITS region. The same six major clades were obtained in the Bayesian and parsimony phylogenetic analyses, and all six clades were well-supported at least by one of the analyses. Morphological characters, such as the overall branching pattern, the presence and intensity of grey colour, the cristation of the apices, and basidiospore size and shape were to various extents correlated with the phylogenetic signal obtained from the ITS region. The congruence between the molecular analyses and morphology, rather than geographical origin, suggests the existence of several species that can be delimited using a combined phylogenetic and morphological species recognition. The analyses revealed that C. cristata and C. rugosa are well-delimited species. In contrast, more than one taxa could be subsumed under the names C. amethystina and C. cinerea, the taxonomical complexity of which is discussed. The ITS region is proved to be adequate to separate phylogenetic species of Clavulina.     

The sister-group relationships of the largest family of lichenized fungi,Parmeliaceae (Lecanorales, Ascomycota)

Parmeliaceae is the largest family of lichen-forming fungi. In spite of its importance for fungal diversity, its relationships with other families in Lecanorales remain poorly known. To better understand the evolutionary history of the diversification of lineages and species richness in Parmeliaceae it is important to know the phylogenetic relationships of the closest relatives of the family. A recent study based on two molecular loci suggested that either Protoparmelia s. str. or a group consisting of Gypsoplaca and Protoparmelia s. str. were the possible sister-group candidates of Parmeliaceae, but that study could not distinguish between these two alternatives. Here, we used a four-locus phylogeny (nuLSU, ITS, RPB1, MCM7) to reveal relationships of Parmeliaceae with other potential relatives in Lecanorales. Maximum likelihood and Bayesian analyses showed that Protoparmelia is polyphyletic, with Protoparmelia s. str. (including Protoparmelia badia and Protoparmelia picea) being most closely related to Parmeliaceae s. str., while the Protoparmelia atriseda-group formed the sister-group to Miriquidica. Gypsoplaca formed the sister-group to the Parmeliaceae s. str. + Protoparmelia s. str. clade. Monophyly of Protoparmelia as currently circumscribed, and Gypsoplaca as sister-group to Parmeliaceae s. str. were both significantly rejected by alternative hypothesis testing.     

Hybridizations and genetic relationships among Lindernia species (Scrophulariaceae): L. procumbens and two subspecies of L. dubia

Lindernia procumbens and L. dubia are common annual weeds in flooded rice fields of Japan. Two subspecies of L. dubia, subsp. major and subsp. dubia, are usually recognized in Japan but they are both regarded as synonyms of L. dubia elsewhere. In a cluster analysis based on AFLP, most L. dubia subsp. major formed a separate cluster from L. dubia subsp. dubia although 11% of haplotypes classified using AFLP were not coincident with classification using the shape of leaf bases, which is the commonly used identification trait. Artificial F₁ plants between L. procumbens and L. dubia subsp. major, and between L. procumbens and L. dubia subsp. dubia did not produce seed. Forty percent of capsules produced by F₁ plants from these two subspecies were slimmer and 80% pollen were sterile in slimmer capsules. However, seed number of most F₁ capsules was not different from that of self-fertilized plants, suggesting that there was no complete reproductive isolation between the subspecies. Natural hybridization of these subspecies may have occurred but we are not aware of it because F₁ plants are rare and F₂ plants are indistinguishable from these subspecies.     

Ecological and functional correlates of molar shape variation in European populations of Arvicola (Arvicolinae, Rodentia)

Lower first molar shape in Arvicola spp. at European level has been studied by means of Geometric Morphometrics. We took into account bioclimatic variables, size, lifestyle and phylogenetic relationships. We used Partial Least Squares and Phylogenetic Independent Contrasts in order to assess which factor affects the most molar shape morphology.Once the phylogenetic history was taken into account, climate resulted the most influencing factor in explaining molar morphology, followed by size. Molar shape is not related to lifestyle. Molar shape and size, even if different among species, are not phylogenetically structured. On the opposite, lifestyle depends on the phylogeny, and size and lifestyle are significantly related even considering phylogenetic relationships. Fossorial forms are significantly smaller than semi-aquatic ones, suggesting that they are less subjected to predator pressure of semi-aquatic species and that the two lifestyles are characterized by different allometric patterns.     

Phylogenetic relationships among Southeast Asian climbing bamboos (Poaceae: Bambusoideae) and the Bambusa complex

A phylogenetic analysis of Bambusa and allies based on the plastid DNA non-coding regions rps16-trnQ, trnC-rpoB, trnH-psbA and trnD-T, and a partial nuclear GBSSI gene, was carried out. This included representatives from all four Bambusa subgenera (including type species), a group of segregate Southeast Asian genera distinctive by their climbing–scrambling culms (Dinochloa, Holttumochloa, Kinabaluchloa, Maclurochloa, Soejatmia, Sphaerobambos), and two other Bambusinae genera (Dendrocalamus, Gigantochloa). The results do not support the present subgeneric classification of Bambusa. The climbing Southeast Asian genera, all of which include species previously placed in Bambusa, are distinct from the “core Bambusa group” (type species and alliance) and the Bambusa complex generally.     

Botryosphaeriaceae from tuart (Eucalyptus gomphocephala) woodland, including descriptions of four new species

Eucalyptus gomphocephala (tuart) is a tree native to the southwest coast of Western Australia, where, in some areas, there is a significant decline in the health of tuart. Botryosphaeriaceous taxa have been isolated as endophytes and canker pathogens from numerous hosts in many parts of the world and have been implicated in the decline of E. gomphocephala. In the present study, endophytic fungi were isolated from a wide variety of native woody plant species (Acacia cochlearis, A. rostellifera, Allocasuarina fraseriana, Agonis flexuosa, Banksia grandis, E. gomphocephala, E. marginata and Santalum acuminatum), at two locations in native E. gomphocephala woodland; a site in decline at Yalgorup National Park and a healthy site at Woodman Point Regional Park. Of the 226 isolates obtained, 154 were botryosphaeriaceous taxa, 80 % of which were found to be Neofusicoccum australe, isolated from all hosts at both collection sites. Four new species are described, Dothiorella moneti, Dothiorella santali, Neofusicoccum pennatisporum, and a species belonging to a genus only recently included in the Botryosphaeriaceae, Aplosporella yalgorensis. The other species isolated were Botryosphaeria dothidea on the new hosts A. rostellifera, A. cochlearis and E. marginata and Dichomera eucalypti, on the new host E. marginata. None of the new species formed lesions on excised stems of their host species, E. gomphocephala, or a common plantation species, E. globulus. However, Neofusicoccum australe formed lesions on excised stems of E. globulus and E. gomphocephala.     

Phylogenetic relationship and taxonomic position of Xylanthemum tianschanicum (Krasch.) Muradyan (Compositae,Anthemideae) as inferred from nrDNA ITS data

The central Asian endemic Xylanthemum tianschanicum (Krasch.) Muradyan (Compositae, Anthemideae) is the only species of Xylanthemum Tzvelev with radiate capitula. While the species was formerly found to be closely related to members of the subtribe Handeliinae, other representatives of the genus were considered to be similar to members of Tanacetum. In order to assess its phylogenetic relationships and taxonomic classification, a molecular analysis based on ITS sequences of 13 representatives of the subtribe Handeliinae and 17 species of three other subtribes (Anthemidinae, Artemisiinae and Matricariinae) was performed. Due to the close relationship of the X. tianschanicum to the type species of Richteria (Richteria pyrethroides) in a well supported clade (PP = 1, BS = 100%), which is supported by morphological features concerning the indumentum, capitulum morphology, and pappus structure, the combination Richteria tianschanica (Krasch.) Sonboli & Oberpr. is proposed based on the basionym Pyrethrum tianschanicum Krasch.     

Systematics and evolution of the genus Torrubiella (Hypocreales, Ascomycota)

Torrubiella is a genus of arthropod-pathogenic fungi that primarily attacks spiders and scale insects. Based on the morphology of the perithecia, asci, and ascospores, it is classified in Clavicipitaceae s. lat. (Hypocreales), and is considered a close relative of Cordyceps s. 1., which was recently reclassified into three families (Clavicipitaceae s. str., Cordycipitaceae, Ophiocordycipitaceae) and four genera (Cordyceps s. str, Elaphocordyceps, Metacordyceps, and Ophiocordyceps). Torrubiella is distinguished morphologically from Cordyceps s. lat. mainly by the production of superficial perithecia and the absence of a well-developed stipitate stroma. To test and refine evolutionary hypotheses regarding the placement of Torrubiella and its relationship to Cordyceps s. lat., a multi-gene phylogeny was constructed by conducting ML and Bayesian analyses. The monophyly of Torrubiella was rejected by these analyses with species of the genus present in Clavicipitaceae, Cordycipitaceae, and Ophiocordycipitaceae, and often intermixed among species of Cordyceps s. lat. The morphological characters traditionally used to define the genus are, therefore, not phylogenetically informative, with the stipitate stromata being gained and/or lost several times among clavicipitaceous fungi. Two new genera (Conoideocrella, Orbiocrella) are proposed to accommodate two separate lineages of torrubielloid fungi in the Clavicipitaceae s. str. In addition, one species is reclassified in Cordyceps s. str. and three are reclassified in Ophiocordyceps. The phylogenetic importance of anamorphic genera, host affiliation, and stipitate stromata is discussed.     

Fruit anatomy of species of Solanum sect. Acanthophora (Solanaceae)

The fruits of 10 species of Solanum sect. Acanthophora were studied. Cross and/or longitudinal and/or tangential microtome sections, stained mostly with astra blue/basic fuchsin, were made for microscopic examination. Three different kinds of cells were found in the epidermis, immediately below which a hypodermis, consisting in any of four types of structures, was always found. The mesocarp exhibits two histologically differentiated zones, an external one (formed by normal or spongy parenchyma, according to the species), and an internal one, commonly juicy, and with proliferations among the seeds. The diagnostic value of all these structures is assessed. Morpho-anatomical information is used to define fruit types beyond the berry, traditionally described for Solanum, and the probable dispersal syndrome related to them is discussed. Fruit similarities are slightly noticeable in several cases, while differences may be the result of their sexual system – which affects specially the size – and their histology, which is related to the dispersal syndrome. The comparison of our data with previous molecular phylogeny of the section suggests that a significant morphological variation is not associated with significant DNA sequence changes.