Tooth Enamel Microstructure of Living and Extinct Hyracoids Reveals Unique Enamel Types Among Mammals

Among medium- to large-sized terrestrial ‘ungulates,’ there is often a relationship between increasing body size, correlated changes in diet, and increased complexity of the enamel microstructures [notably the development of Hunter-Schreger bands (HSB)]. An exhaustive survey of the enamel microstructures of living and extinct Hyracoidea demonstrates, however, that the Schmelzmuster within this order of mammals is generally one-layered and formed by radial enamel despite a large range of body sizes and dietary adaptations; HSB are remarkably absent. Radial enamel is characteristic of early diverging hyracoids, as well as more derived members of the extinct families Geniohyidae and Pliohyracidae, and the extant Procaviidae. Only some large ‘Saghatheriidae,’ and all members of the family Titanohyracidae, developed a more complex enamel microstructure (i.e., with prisms decussating), a unique condition among Mammalia that we name ‘bundled enamel’ (BE). This structure is reminiscent to some degree of both the ‘Pyrotherium enamel’ and the ‘3D enamel’ of proboscideans. Hyracoids with BE represented a major component of the diversity of mid- to large-sized herbivores during the Paleogene in Africa. Like HSB, which are developed by most other ‘ungulates,’ the BE is regarded as a device for resisting propagation of cracks during mastication. Hyracoids never developed however the ‘modified radial enamel’ that is characteristic of most large and hypsodont perissodactyls and artiodactyls that entered Africa during the Miocene.     

The oldest cynodont: new clues on the origin and early diversification of the Cynodontia

Early mammaliaforms and their extinct relatives, nonmammaliaform cynodonts, have long been the focus of intense research in attempting to unravel how and when major changes toward mammalness occurred. The earliest well-known representatives of cynodonts are latest Late Permian in age. Here, we describe Charassognathus gracilis gen. et sp. nov. , from the early Late Permian of South Africa, representing the oldest cynodont yet found. This specimen displays a notch on the dentary in the same location as the base of the masseteric fossa in the basal cynodonts Procynosuchus and Dvinia , and represents the first indication in theriodonts of an invasion of occlusal musculature onto the dentary. A phylogenetic analysis of seven therocephalians and ten non-mammaliaform cynodonts and equally weighted characters resulted in nine most parsimonious trees, the strict consensus of which shows a basal polytomy in cynodonts, including Charassognathus , Dvinia , Procynosuchus and a clade including the remaining cynodonts. The basal polytomy in the majority rule consensus tree is reduced, as Procynosuchus and Dvinia form a clade. One most parsimonious tree, from an analysis using implied weights, positions Charassognathus as the most basal cynodont. This result implies that the Cynodontia initially diversified in Permian Gondwana, in what is now southern Africa.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 149 , 477–492.     

Was the Early Eocene proboscidean Numidotherium koholense semi-aquatic or terrestrial? Evidence from stable isotopes and bone histology

The Early Eocene deposits of El Kohol, Algeria, have yielded numerous remains of Numidotherium koholense, one of the most primitive and oldest known proboscideans in Africa. The Upper Eocene proboscideans of the Fayum locality (Egypt), Barytherium sp. and Moeritherium sp., were recently interpreted as aquatic or semi-aquatic, according to the stable isotopic compositions (δ¹³C and δ¹⁸O) of their tooth enamel. These data led us to reinvestigate the adaptations of N. koholense. Stable isotopic analysis and observations of histological sections of its long bones reveal that it was essentially terrestrial. According to its position within the phylogenetic tree of Eocene proboscideans, the adaptation to semi-aquatic life appears to have evolved independently in different lineages of Middle and Upper Eocene proboscideans during their adaptive radiation in Africa. Moreover, these new results reopen the debate about the hypothesis that Eocene to Recent proboscideans are derived from semi-aquatic ancestors.     

Two new basal crocodylomorph archosaurs from the Lower Jurassic and the monophyly of the Sphenosuchia

We report on and name two new taxa of basal crocodylomorph archosaurs from the Lower Jurassic, Litargosuchus leptorhynchus gen. et sp. nov. , from the upper Elliot Formation (Stormberg Group) of South Africa, and Kayentasuchus walkeri gen. et sp. nov. , from the Kayenta Formation (Glen Canyon Group) of Arizona, USA. Examination of this material led to a reconsideration of basal crocodylomorph interrelationships. A phylogenetic analysis found no support for the monophyly of Sphenosuchia.  © 2002 The Linnean Society of London. Zoological Journal of the Linnean Society , 2002, 136 , 77–95.     

Long-bone geometry in columnar-limbed animals: allometry of the proboscidean appendicular skeleton

Limb-bone allometry was investigated for 19 species of proboscideans, spanning almost the entire phylogenetic spectrum. More archaic proboscideans ('gompthotheres') have substantially thicker long-bone diaphyses relative to length than elephantids, as has been suggested previously, but contrary to previous suggestions it could not be confirmed that Mammuthus had more massive long-bone diaphyses on average than extant Elephas and Loxodonta . When correcting for phylogeny, the circumference of the limb bones to their length in proboscideans as a group generally scale with negative allometry, becoming stouter with increased length, as would be expected from limb mechanics. Few slopes were, however, statistically significantly negatively allometric. Correcting for phylogeny produced better correlations than traditional regression analyses, in contrast to most other studies where the reverse is the case. Intraspecific analyses of extant Elephas and Loxodonta , in addition to Mammuthus primigenius , Mammut americanum, and Gomphotherium productum , also resulted in negatively allometric regression slopes, frequently conforming to the theory of elastic similarity, as could be expected from the columnar posture of proboscideans. At present the reasons for the more massive limbs of gomphotheres s.l. are not fully understood. © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 149 , 423–436.     

A peculiar climbing Megalonychidae from the Pleistocene of Peru and its implication for sloth history

The Xenarthra, particularly the Tardigrada, are with the Notoungulata and Marsupialia among the most diversified South American mammals. Lujanian South American Land Mammal Age localities from the coastal Piedra Escrita site and Andean Casa del Diablo Cave, Peru, have yielded three specimens of the Megalonychidae Diabolotherium nordenskioldi gen. nov. This singular fossil sloth exhibits a peculiar mosaic of cranial and postcranial characters. Some are considered convergent with those of other sloths (e.g. 5/4 quadrangular teeth, characteristic of Megatheriidae), whereas others clearly indicate climbing capabilities distinct from the suspensory mode of extant sloths. The arboreal mode of life of D. nordenskioldi is suggested by considerable mobility of the elbow, hip, and ankle joints, a posteriorly convex ulna with an olecranon shorter than in fossorial taxa, a radial notch that faces more anteriorly than in other fossil sloths and forms an obtuse angle with the coronoid process (which increases the range of pronation–supination), a proximodistally compressed scaphoid, and a wide range of digital flexion. D. nordenskioldi underscores the great adaptability of Tardigrada: an arboreally adapted form is now added to the already known terrestrial, subarboreal, and aquatic (marine and freshwater) fossil sloths. A preliminary phylogenetic analysis of the Tardigrada confirmed the monophyly of Megatherioidea, Nothrotheriidae, Megatheriidae, and Megalonychidae, in which Diabolotherium is strongly nested.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 149 , 179–235.     

Monophyly and phylogenetic placement of the spider genus Labulla Simon, 1884 (Araneae, Linyphiidae) and description of the new genus Pecado

The genus Labulla Simon is circumscribed in phylogenetic terms to include the species Labulla thoracica (Wider), L. flahaulti Simon and L. machadoi sp. nov. The genital anatomy of the genus is described in detail and the taxonomy of the genus is reviewed. The monophyly of Labulla is supported by numerous morphological apomorphies of the male palp and female epigynum. Based on a cladistic analysis, a new genus, Pecado gen. nov. , is erected to place Labulla impudica Denis, from Northern Africa. Lepthyphantes insularis Saito and ' Labulla ' nepula Tikader, both formerly included in Labulla , are not congeneric with the type species of Labulla .  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 143 , 359–404.     

The genus Pholcus (Araneae, Pholcidae) in the Canary Islands

This paper offers a detailed taxonomic revision of all Canarian Pholcus species described before 2003, all of which are newly described and newly illustrated. Pholcus guadarfia sp. nov. is described, while a neotype for Pholcus malpaisensis Wunderlich, 1992 is also provided. In addition, we propose Pholcus gomerae Wunderlich, 1980 as a senior synonym for Pholcus gomeroides Wunderlich, 1987. More importantly, cladistic analysis based on the morphological characters of the Macaronesian Pholcus species was conducted for the first time. Parsimony analyses of 73 morphological characters revealed the close relationships between those species from the Canary Islands, Madeira and the Macaronesian enclave in Africa (between Agadir and Nouadhibou).  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 151 , 59–114.     

Molecular phylogeny of the genus Tibicina (Hemiptera, Cicadidae): rapid radiation and acoustic behaviour

To estimate the potential contribution of ethological and ecological parameters to the mechanisms of species formation and species isolation in the Palearctic cicada genus Tibicina , we constructed a molecular phylogenetic hypothesis of extant Tibicina species. Seven mitochondrial genes and a fragment of a nuclear gene were sequenced (3046 bp). Mitochondrial genes included 547 informative sites but the nuclear gene was too conserved to be included in the analysis. The tree was characterized by a basal polytomy indicating that Tibicina species arose rapidly. Such rapid radiation might explain the low divergence in the acoustic communication observed between species. Parameters describing habitat selection and acoustic communication were mapped onto the tree. A shift in habitat selection accompanied by acoustic changes might have contributed to one speciation event. The stochastic distribution of the same acoustic characters on the other branches of the tree implies, however, that the subtle acoustic differences between species could be the result of previous speciation events and independent evolutionary histories, rather than having contributed themselves in the speciation and isolation processes.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 611–626.     

Phylogeny of the Zoroasteridae (Zorocallina; Forcipulatida): evolutionary events in deep-sea Asteroidea displaying Palaeozoic features

The Zoroasteridae comprise a small but widespread family of asteroids distributed throughout the deep sea. Although poorly understood, they are often collected in the hundreds, suggesting that they occupy important ecological roles. A phylogenetic analysis including 24 terminal taxa and 70 morphological characters was performed, resulting in a single most-parsimonious tree. The tree separated zoroasterids with open, reticulate skeletons (e.g. Myxoderma ) as more basal than those with more heavily armored, imbricate skeletons (e.g. Zoroaster ), which were more derived. In addition to agreement with established genera, a new genus is supported by the phylogeny as the sister taxon to Myxoderma . The cladistic analysis was performed in conjunction with a revisionary survey of zoroasterid species, resulting in taxonomic changes to species in nearly every genus. Bathymetric and physiographic shifts were observed between the reticulate and imbricate zoroasterid clades. Zoroasterids possess a single marginal plate series, which occurs in basal sister-group neoasteroids (crown-group asteroids). Phylogenetic results suggest that the morphololgical resemblance between zoroasterids and Palaeozoic taxa, such as Calliasterella , is convergent but a paraphyletic Zoroasteride cannot be rejected and remains consistent with basal crown-group affinities. Although the phylogenetic position of the Eocene Zoroaster aff. fulgens was not strongly supported, its presence within a derived cluster of Zoroaster spp. suggests a relatively recent (i.e. Cenozoic) diversification into the deep sea. Taxonomic revisions, and geographical and bathymetric range extensions are also included.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 150 , 177–210.     

Body size in proboscideans, with notes on elephant metabolism

Mass estimates for a number of fossil proboscideans were computed using regression analyses on appendicular bones to body mass, for seven specimens of modern elephants, for which body masses had been recorded prior to death. The marked differences in physical proportions between extant Loxodonta and Elephas , implying substantial differences in body mass at any given shoulder height, were not present in their long bone parameters. Length and least circumferences proved to be the best parameters for prediction of body mass. Some extinct proboscideans, notably certain Mammuthus and Deinotherium , were much larger than extant elephants. Both the basal and the field metabolic rates of extant elephants are lower than predicted for a hypothetical mammal, in accordance with their body size and subsistence on low-quality foods. The feeding quantities often ascribed to extant wild elephants are exaggerated, and would in fact have sufficed to nourish much larger species.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 140 , 523–549.     

A phylogenetic analysis of the genus Eunice (Eunicidae, polychaete, Annelida)

Species of Eunice are distributed worldwide, inhabiting soft and hard marine bottoms. Some of these species play significant roles in coral reef communities and others are commercially important. Eunice is the largest and most poorly defined genus in Eunicidae. It has traditionally been subdivided in taxonomically informal groups based on the colour and dentition of subacicular hooks, and branchial distribution. The monophyly of Eunice and of its informal subgroups is tested here using cladistic analyses of 24 ingroup species based on morphological data. In the phylogenetic hypothesis resulting from the present analyses Eunice and its subgroups are paraphyletic; the genus may be divided in at least two monophyletic groups, Eunice s.s. and Leodice , but several species do not fall inside these two groups. Most of the traditional characters used in the taxonomy of Eunice are homoplasies; however, characters used for the first time in this study, such as certain jaw characters and characters derived from a close examination of chaetal variation along the body, are promising sources of phylogenetic signal.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 150 , 413–434.     

Systematics of the genera of Bodotriidae (Crustacea: Cumacea)

The cumacean family Bodotriidae includes 382 species in 31 genera grouped in three subfamilies: Bodotriinae, Mancocumatinae and Vaunthompsoniinae. Generic diagnoses are based on few characters that often have overlapping states among genera, complicating the understanding of the relationships within the group. The goals of this study are to illuminate the phylogenetic relationships among the genera of the Bodotriidae using morphological characters and to review the systematics of the family. For this purpose, all species within each genus were studied from the literature to code all the variability of genera for 109 variable morphological characters. Phylogenetic analyses show that there is independent reduction of the pleopods in two clades from a plesiomorphic state of five pairs, while the number of exopods of peraeopods has been reduced gradually in more derived groups of bodotriids. The subfamily Bodotriinae is the most derived and the Vaunthompsoniinae the most basal, and is paraphyletic with the Mancocumatinae embedded within it. No discriminatory characters were found between the subfamilies Mancocumatinae and Vaunthompsoniinae and they are not clearly separated in the phylogeny. Mancocumatinae is synonymized with Vaunthompsoniinae and all the genera of the former Mancocumatinae should be included within Vaunthompsoniinae. Analyses of character evolution justify a few other taxonomic changes. All genera were redescribed based on all the coded characters and a complete list of all valid species for each genus is included. Finally, dichotomous keys for identification to genus level are provided.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 151 , 1–58.     

Enamel Structure of Cuvieronius hyodon (Proboscidea, Gomphotheriidae) with a Discussion on Enamel Evolution in Elephantoids

Dental material of the South American elephantoid Cuvieronius hyodon from the Late Pleistocene of the Tarija Basin, Bolivia was sampled for a comprehensive analysis of the microstructure of the enamel. To examine variability at the dentition level, enamel samples of the upper incisor, second deciduous premolar, and molars were sectioned. The incisor and cheek teeth enamel is compared to that of other proboscideans in order to reveal phylogenetically and functional informative features useful to reconstruct the evolution of elephantoid enamel. Studies of the adaptations and evolution of proboscidean enamel have focused so far on molars. Nevertheless, given the possibility of an independent evolution of the enamel at different tooth positions, the variation of the enamel throughout the dentition needs to be taken into consideration when using enamel microstructural characters to infer proboscidean diversity and phylogeny. The results obtained from this study demonstrate the generality, among elephantoids, of the basic microstructural features of Cuvieronius hyodon enamel, allowing the characterization of the Elephantoid Enamel (EE). The differentiation between incisor and molar enamel seen in elephantoids is shown to represent a primitive elephantiform trait, as it also occurs in Phiomia. The three-layered enamel of the cheek teeth appears as the sole synapomorphy of the Elephantoidea, though the character might be homoplastic within the Proboscidea. Characters of the prisms cross-section might be used, on the other hand, to define less inclusive clades within the Elephantoidea.     

Infraspecific variation and phylogeography of the high-polyploid Iberian endemic Anthoxanthum amarum Brot. (Poaceae; Pooideae) assessed by random amplified polymorphic DNA markers (RAPDs) and morphology

High morphological and ecological diversity has been observed in the north-west Iberian Peninsula endemic Anthoxanthum amarum Brot., for which two different morphotypes (northern and southern) have been described on the basis of qualitative traits and geographical origin. In the present study, a combined molecular and morphological method was applied to ten populations of this species with the following aims: (1) to assess whether the variation observed was taxonomically meaningful; (2) to assess the influence of the environment on the variation in the morphological characters; and (3) to track the potential phylogeographical information deduced from our random amplified polymorphic DNA marker (RAPD) data in order to draw inferences about the past history of this species in the north-west Iberian Peninsula. To achieve these aims, 26 macromorphological characters were recorded in 279 specimens, and 77 RAPD phenotypes were identified in the 79 plants studied. The association analyses performed using the morphological and molecular data showed that no clear separation existed between the morphs, and a strong correlation between qualitative characters and the environment was detected. Moreover, both the multivariate analyses and the assignment test based on RAPD data revealed that the genetic variation was hierarchically structured, and three genetically distinct groups could be identified. Two of these clusters might correspond to different expansion routes proposed in the literature for different plant species in the north-west Iberian Peninsula.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 179–192.     

An extraordinary new rheophyte in the genus Leptactina (Rubiaceae, Pavetteae) from Rio Muni (Equatorial Guinea)

A distinctive new species of Rubiaceae from Rio Muni, Equatorial Guinea, is described and illustrated. Leptactina rheophytica is the only rheophyte known in the genus. Its diagnostic characters are elucidated, its taxonomic affinities are discussed, and notes on its conservation status are provided.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 109–113.     

The performance of morphological characters in broad-scale phylogenetic analyses

In view of their propositional content (i.e. they can be right or wrong), character statements (i.e. statements that predicate characters of organisms) are treated as low-level hypotheses. The thesis of the present study is that such character statements, as do more complex scientific theories, come with variable scope. The scope of a hypothesis, or theory, is the domain of discourse over which the hypothesis, or theory, ranges. A character statement is initially introduced within the context of a certain domain of discourse that is defined by the scale of the initial phylogenetic analysis. The doctrine of 'total evidence' requires the inclusion of previously introduced characters in subsequent studies. As a consequence, the initial scope of character statements is widened to the extent that the scale of subsequent analyses is broadened. Scope expansion for character statements may result in incomplete characters, in the subdivision of characters, or in ambiguity of reference (indeterminacy of the extension of anatomical terms). Character statements with a wide scope are desirable because they refer to characters with the potential to resolve deep nodes in phylogenetic analyses. Care must be taken to preserve referential unambiguity of anatomical terms if the originally restricted scope of a character statement is expanded to match a broad-scale phylogenetic analysis.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 297–308.