New land tortoises (Testudines: Testudinidae) from the Miocene of Africa

The described fossil testudinids from the Miocene of Africa are reviewed. Geochelone stromeri sp. nov . is described from Lower Pliocene (Langebaanweg) and Miocene (Namib) specimens. Kinixys erosa , an extant species, is reported from Songhor Hill. Chersina sp. is reported from Arrisdrift. Impregnochelys pachytectis gen. et sp. nov . is described from Rusinga Island, Kenya, and is unique in having the anterior shell opening orientated ventrally and in having struts on the internal surface of the neurals, but shares with Kinixys a unique epiplastron shape, a high number of axillary scutes and unique orientation of the head of the femur.     

The dazed and confused identity of Agassiz's land tortoise, Gopherus agassizii (Testudines, Testudinidae) with the description of a new species, and its consequences for conservation

We investigate a cornucopia of problems associated with the identity of the desert tortoise, Gopherus agassizii (Cooper). The date of publication is found to be 1861, rather than 1863. Only one of the three original cotypes exists, and it is designated as the lectotype of the species. Another cotype is found to have been destroyed in the 1906 San Francisco earthquake and subsequent fire. The third is lost. The lectotype is genetically confirmed to be from California, and not Arizona, USA as sometimes reported. Maternally, the holotype of Gopherus lepidocephalus (Ottley & Velázques Solis. 1989) from the Cape Region of Baja California Sur, Mexico is also from the Mojavian population of the desert tortoise, and not from Tiburon Island, Sonora, Mexico as previously proposed. A suite of characters serve to diagnose tortoises west and north of the Colorado River, the Mojavian population, from those east and south of the river in Arizona, USA, and Sonora and Sinaloa, Mexico, the Sonoran population. Species recognition is warranted and because Gopherus lepidocephalus is from the Mojavian population, no names are available for the Sonoran species. Thus, a new species, Gopherus morafkaisp. n., is named and this action reduces the distribution of Gopherus agassizii to only 30% of its former range. This reduction has important implications for the conservation and protection of Gopherus agassizii, which may deserve a higher level of protection.     

Hesperotestudo (Testudines: Testudinidae) from the Pleistocene of Bermuda, with comments on the phylogenetic position of the genus

A recently discovered fossil land tortoise (Testudines: Testudinidae) is described from the Pleistocene of Bermuda. Its morphology is sufficiently well preserved to allow assignment to the extinct North American genus Hesperotestudo. However, several features of this tortoise are unique and it is named Hesperotestudo bermudae sp. nov. A review of the phylogenetic relationships of the better known genera of the Testudinidae suggests that the affinities of Hesperotestudo lie with other North American tortoises (Gopherus) and not with Geochelone or other testudinines; thus, Hesperotestudo is reassigned to the Xerobatinae. This is at least the fifth documentation of a testudinid dispersing over open ocean to an oceanic island (the first for Hesperotestudo) and it corroborates the hypothesis that members of this family are well suited to over-water dispersal.     

The first endocast of the extinct dodo (Raphus cucullatus) and an anatomical comparison amongst close relatives (Aves,Columbiformes)

The dodo (Raphus cucullatus) became extinct only 100 years after humans first arrived on the Indian Ocean island of Mauritius. Even though it has become an example of oddity, obsolescence, stupidity, and extinction, most aspects of its biology are still unknown. We used high‐resolution X‐ray computed tomography (CT) scanning to examine the endocranial morphology of the dodo and compare this virtual endocast to eight close relatives. Enlarged olfactory bulbs are a shared characteristic of the Raphinae and posteriorly angled semicircular canals are particular to the dodo compared with the other eight species sampled here. A regression of log endocranial volume against log body size shows that the dodo has an endocranial volume on par with other pigeons. Aspects of the dodo's biology are discussed in relation to these endocranial features.     

The Tipulidae (Diptera) of northern Morocco with a focus on the Rif region,including the description of a new species of Tipula (Lunatipula) and an updated checklist for Morocco

An overview of the Tipulidae known to occur in northern Morocco with an emphasis on the Rif mountains is given, incorporating new distribution data based on recently collected material in the area. Dolichopeza (Dolichopeza) hispanica, Tipula (Lunatipula) subpustulata, and Tipula (Yamatotipula) afriberia afriberia are recorded for the first time for the Rif. Tipula (L.) stimulosa Mannheims, 1973 and T. (Vestiplex) vaillanti vaillanti Theowald, 1977 are reported for the first time for the Rif and Morocco. Tipula (Lunatipula) pseudocinerascens Strobl, 1906 and Tipula (Savtshenkia) confusa van der Wulp, 1883 are recorded for the first time for the Rif, Morocco and North Africa. A new species of the subgenus Lunatipula, T. (L.) pjotri n. sp., is described and illustrated. Nephrotoma exastigma, previously reported for the Rif, seems to be absent in Morocco. Reports of Tipula (Emodotipula) obscuriventris Strobl, 1900 for Morocco actually refer to T. (E.) leo. This brings the number of Tipulidae for Morocco to 39 species and for the Rif to 28. An updated checklist of the Tipulidae of Morocco is provided.     

Functional and ecological relevance of intraspecific variation in body size and shape in the lizard Podarcis melisellensis (Lacertidae)

Within populations, individual animals may vary considerably in morphology and ecology. The degree to which variation in morphology is related to ecological variation within a population remains largely unexplored. We investigated whether variation in body size and shape among sexes and age classes of the lizard Podarcis melisellensis translates in differential whole-animal performance (sprint speed, bite force), escape and prey attack behaviour in the field, microhabitat use and diet. Male and female adult lizards differed significantly in body size and head and limb proportions. These morphological differences were reflected in differences in bite strength, but not in sprint speed. Accordingly, field measurements of escape behaviour and prey attack speed did not differ between the sexes, but males ate larger, harder and faster prey than females. In addition to differences in body size, juveniles diverged from adults in relative limb and head dimensions. These shape differences may explain the relatively high sprint and bite capacities of juvenile lizards. Ontogenetic variation in morphology and performance is strongly reflected in the behaviour and ecology in the field, with juveniles differing from adults in aspects of their microhabitat use, escape behaviour and diet.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 251–264.     

Mitochondrial phylogeography and subspecies of the wide‐ranging sub‐Saharan leopard tortoise Stigmochelys pardalis (Testudines: Testudinidae) – a case study for the pitfalls of pseudogenes and GenBank sequences

The leopard tortoise (Stigmochelys pardalis) is the most widely distributed sub‐Saharan tortoise species, with a range extending from the Horn of Africa all over eastern Africa to the Republic of South Africa, Namibia and southernmost Angola. Using 1938 bp of mitochondrial DNA (cyt b gene, partial ND4 gene plus adjacent tRNA genes) from a nearly range‐wide sampling, we examined its phylogeographic structure and compared our findings with previously published GenBank sequences. We identified seven major clades that are largely parapatrically distributed. A few records of distinct haplotypes at the same locality or in close proximity could be the result of translocation of tortoises by man. The greatest diversity occurs in the south of the species’ range, with five out of the seven clades. Testing for isolation‐by‐distance suggests that the observed phylogeographic structure is the result of restricted geographical gene flow and not of historical vicariance. This is in sharp contrast to wide‐ranging thermophilic reptiles from the western Palaearctic, whose phylogeographic structure was significantly shaped by Pleistocene range interruptions, but also by earlier dispersal and vicariant events. Most cyt b sequences of S. pardalis from GenBank turned out to be nuclear pseudogenes, or to be of chimerical origin from such pseudogenes and authentic mitochondrial sequences, which argues for caution regarding uncritical usage of GenBank sequences. The recent revalidation of the two subspecies of S. pardalis was based on such a chimerical sequence that was erroneously identified with the subspecies S. p. babcocki. Furthermore, according to our data, the distribution of mitochondrial clades does match neither the traditional subspecies ranges nor the pronounced geographical size variation of leopard tortoises. We conclude that there is no rationale for recognizing subspecies within S. pardalis.     

Isolation and characterization of eight microsatellite loci for the study of gene flow between Testudo marginata and Testudo weissingeri (Testudines: Testudinidae)

The taxonomic status of the Greek Testudo species, marginated tortoise (Testudo marginata Schoepff, 1793) and dwarf marginated tortoise (Testudo weissingeri Bour, 1995), is controversed. In order to study the gene flow among these Greek Testudo species, eight polymorphic microsatellite markers were characterized and screened for 32 individuals. The number of alleles observed per locus ranged from two to 19. Observed heterozygosities varied from 0.235 to 0.830. These microsatellite loci were also successfully tested on seven closely related species of Testudinidae. This set of microsatellites offers an efficient tool to investigate genetic differentiation among testudinid species.     

The osteology of the golden grey mullet Liza aurata (Teleostei: Mugiliformes: Mugilidae) including interactive three-dimensional reconstructions

Grey mullets are remarkably characterized by their overall uniform external morphology. Identifying species as well as positioning the Mugiliformes in a phylogenetic context is rather difficult. Most recently they were placed in the newly erected Ovalentaria, but more detailed relationships to potential sister taxa were not resolved. Studying the internal morphology, especially the osteology, might provide new insights into the evolution of the Mugiliformes as well as help clarify their phylogenetic position within the Ovalentaria. A detailed osteology of the golden grey mullet Liza aurata is presented. The use of cleared and stained specimens allowed for a complete examination of bony and cartilaginous structures, and a 3D reconstruction from a μCT data set provided additional information on the positional relationships of the bones. Following this, the data obtained were compared with different mugilid species, particularly with the flathead grey mullet Mugil cephalus. Several differences between these species could be identified, such as the position of the basisphenoid, the shape of the hyomandibular and the composition of the branchial arches. These characters might help in understanding the evolutionary changes happening within the mugiliforms and will provide the basis to study this taxon in detail, finally allowing the reconstruction of the body plan of grey mullets.     

Digital cranial endocast of Riograndia guaibensis (Late Triassic,Brazil) sheds light on the evolution of the brain in non-mammalian cynodonts

Abstract

A digital cranial endocast of the specimen UFRGS-PV-596-T of Riograndia guaibensis was obtained from μCT scan images. This is a small cynodont, closely related to mammaliaforms, from the Late Triassic of Brazil. Riograndia has large olfactory bulb casts and the cerebral hemispheres region is relatively wider than in other non-mammaliaform cynodonts. Impressions of vessels were observed and a conspicuous mark on the dorsal surface was interpreted as the transverse sinus. The calculated encephalization quotient is greater than the range seen in most other non-mammaliaform cynodonts. The ratios between linear and area measurements of the dorsal surface suggest four evolutionary changes from a basal eucynodont morphology to mammaliaforms, involving an evolutionary increase of the relative size of the olfactory bulbs and the width of the cerebral hemispheres and cerebellum. The data supports the hypothesis of the neurological evolution of the mammalian lineage starting with a trend for an increase of the olfactory bulbs, which is associated with adaptations in the nasal cavity. This trend is suggested to be linked to the selective pressures for small-sized faunivorous, and probably nocturnal, animals, and represents an initial improvement of the sensory receptor system, subsequently leading to further development of the ‘superior’ structures for sensorial processing and integration.     

The Arcoidea (Mollusca: Bivalvia): a review of the current phenetic-based systematics

The diversity and adaptive radiations of modern Arcoidea, here considered to contain the families Arcidae, Noetiidae, Cucullaeidae, and Glycymerididae, are reviewed. Most fall into either epibyssate or endobyssate life habits with only the Glycymerididae living as free burrowers. The phenetic characters of the families within the Arcoida are reviewed and the families are shown to be supported by very few synapomorphic characters. Homoplasy is shown to be widespread and is illustrated in a series of discussions on the ligament, epibyssate–endobyssate radiations, and possible parallelism within genera, and in a review of arcoid anatomical characters. Previously published molecular data are reviewed and these support the inclusion of the Glycymerididae in the Arcoidea. They also indicate, however, that polyphyly is probably widespread at the subfamily level.  © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society , 2006, 148 , 237–251.     

The evolution of Synchaetidae (Rotifera: Monogononta) with a focus on Synchaeta: An integrative approach combining molecular and morphological data

This study represents the first phylogenetic reconstruction of Synchaetidae (Rotifera, Monogononta) and combines one morphological and two molecular data sets to derive the most comprehensive database for this taxon to date. Overall, 19 species were examined both morphologically (light- and scanning electron microscopy of habitus and trophi) and genetically (COI and 18S rRNA genes), thereby delivering new morphological information as well as the first molecular data for several species. Our results strongly support the monophyly of each of the three genera within Synchaetidae, with Synchaeta being the sister taxon of Polyarthra and Ploesoma. Resolution within each genus, however, was more poorly supported, possibly because of high levels of missing data for the more poorly known species that were also not re-discovered and revised in the present study. Nevertheless, an evolutionary reconstruction based on the total evidence topology indicates that the common ancestor of Synchaetidae was a rotatorivorous and pelagic freshwater rotifer exhibiting slightly enlarged lamellar trophi that were adapted to an enhanced pumping function. These features together with several, additional key character transformations (e.g., hatching independently from benthic or periphytic habitats) probably account for the success of Synchaetidae in pelagic environments and their being one of the most widely distributed and abundant rotifer taxa.     

Late Emsian Rutoceratoidea (Nautiloidea) from the Prague Basin,Czech Republic: morphology,diversity and palaeoecology

Abstract: Nautiloids of the superfamily Rutoceratoidea from the late Emsian (late Early Devonian) of the Prague Basin (Czech Republic) are commented upon. Species recognized include the hercoceratids Hercoceras mirum, H.? transiens, Ptenoceras proximum, P. nudum, P. minusculum and Anomaloceras anomalum, as well as the rutoceratids Adelphoceras bohemicum, Homoadelphoceras devonicans, Pseudorutoceras bolli and Goldringia? devonicans. In addition, four new species are described: Parauloceras regulare sp. nov., Roussanoffoceras chlupaci sp. nov., Otomaroceras sp. nov. and Goldringia sp. nov. Morphology and distribution patterns of Pragian and late Emsian rutoceratoid faunas from the Prague Basin are compared. They show that an increased diversity was accompanied by a higher level of specialization of rutoceratoids, which manifested itself in low abundance, increased facies dependence and greater variation in shell size during the Early Devonian. The evolution of sculpture and a contracted aperture, both regarded as protective adaptive features, was also examined, but no adaptive trend towards more pronounced sculpture and constriction of the aperture was found to have occurred in the Early Devonian. A more distinctive sculpture was, however, observed in shallow‐water assemblages of P. proximum in comparison with deeper‐water faunules, and two additional cephalopod species were examined in order to obtain comparative data. The presence of distinct sculpture patterns in coeval shallow‐ and deeper‐water assemblages suggests limited migration between them and consequently reflects some degree of territoriality in Devonian nautiloids. New data on early shell development in P. proximum are presented. During the Chote? Event, rutoceratoid generic diversity dropped dramatically, one family became extinct and the Early Devonian diversification of the group came to an end. The recovery of nautiloid faunas was slower than that of other cephalopods and associated, unrelated invertebrates. The absence of change in abundance patterns between Pragian and late Emsian rutoceratoid faunas, i.e. prior to and subsequent to ammonoid radiation, suggests that the appearance and radiation of the latter group in the early Emsian did not affect the structure of nautiloid assemblages, i.e. these two clades did not occupy the same niches.     

First record of Isocapnia (Plecoptera: Capniidae) from China with description of a new species

Summary

The winter stonefly genus Isocapnia Banks, 1938??? is mainly distributed in western North America and eastern Asia, but has never been reported from China. In this paper, we report, describe and illustrate a new species of this genus, I. anguis n. sp. from Sichuan Province of China. This is the first record for the genus Isocapnia from China, extending the distribution of Isocapnia into the southwestern China. An identification key to all the Isocapnia species from Asia is provided.???     

Pollen morphology of Campanula (Campanulaceae) and allied genera in Iran with special focus on its systematic implication

Pollen grains of 47 taxa of Campanulaceae including 35 taxa of Campanula that represent its five subgenera and nine sections are investigated. Moreover, five species and three subspecies representing three sections in Asyneuma and one species of each genera Legousia, Michauxia, Zeugandra and Theodorovia were studied by light and scanning electron microscopy. The basic shape of the pollen grains was spheroidal. The apertures vary from tri- to hexa-porate. The sculpturing pattern of exine was rugulate-echinate, rugulate-microechinate or in few species rugulate-microreticulate and microechinate. The most valuable characters for subgeneric classification were the length and density of echini. The length of echini were significantly long (> 2 μm) in C. sclerotricha, Legousia falcata and Michauxia laevigata. Pollen grains show low variation in different species of subgen. Rapunculus, but were variable among different species of some groups, such as sect. Rupestres, probably indicating their non-monophyly despite homogeneity with respect to other morphological characters. Pollen morphology does not support recognition of Asyneuma, Legousia, Michauxia, Symphyandra, Theodorovia, and Zeugandra as separated from Campanula, since none of them exhibit any unique feature.