A new species of Orycteropodidae (Mammalia, Tubulidentata) in the Mio-Pliocene of northern Chad

A new species of fossil Tubulidentata has been found by the Mission Paléoanthropologique Franco-Tchadienne in Northern Chad. It is the first fossil Orycteropodidae (aardvark) from the Mio-Pliocene of Central Africa. The new taxon, Orycteropus abundulafus sp. nov. , is considered in the framework of the available Orycteropodidae fossil record. The Chadian specimen is characterized by the highest dental robustness index among all Tubulidentata, the presence of crests on the pterygoid, the triangular-shaped olecranon fossa and the reduction of the deltoid crest. All of these characters are linked to a less fossorial animal that had a tougher diet. This new African species is closer to the Eurasian O. gaudryi than to any other Tubulidentata. Together they form a clade distinct from that which includes O. afer . This is the first evidence of a relationship for aardvarks between Africa and Eurasia. An initial step is made towards revision of the phylogeny of the order.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 143 , 109–131.     

The tooth of perfection: functional and spatial constraints on mammalian tooth shape

This paper addresses the question of how close mammalian teeth are to ideal functional forms. An 'ideal' form is a morphology predicted to be the best functional shape according to information of the relationships between shape and function. Deviations from an ideal form are likely to indicate the presence of developmental or genetic constraints on form. Model tools were constructed to conform to functional principles from engineering and dental studies. The final model shapes are very similar to several mammalian tooth forms (carnassial teeth and tribosphenic-like cusps), suggesting that these tooth forms very closely approach ideal functional forms. Further evidence that these tooth forms are close to ideal comes from the conservation over 140 million years, the independent derivation and/or the occurrence over a size range of several orders of magnitude of these basic tooth forms. One of the main functional shapes derived here is the 'protoconoid', a fundamental design for double-bladed tools that fits a large number of functional parameters. This shape occurs in tooth forms such as tribosphenic, dilambdodont and zalambdodont. This study extends our understanding of constraints on tooth shape in terms of geometry (how space influences tooth shape) and function (how teeth divide food).  © 2003 The Linnean Society of London . Biological Journal of the Linnean Society , 2003, 78 , 173–191.     

Dietary adaptations in the teeth of murine rodents (Muridae): a test of biomechanical predictions

Functional dental theory predicts that tooth shape responds evolutionarily to the mechanical properties of food. Most studies of mammalian teeth have focused on qualitative measures of dental anatomy and have not formally tested how the functional components of teeth adapt in response to diet. Here we generated a series of predictions for tooth morphology based on biomechanical models of food processing. We used murine rodents (Old World rats and mice) to test these predictions for the relationship between diet and morphology and to identify a suite of functional dental characteristics that best predict diets. One hundred and five dental characteristics were extracted from images of the upper and lower tooth rows and incisors for 98 species. After accounting for phylogenetic relationships, we showed that species evolving plant‐dominated diets evolved deeper incisors, longer third molars, longer molar crests, blunter posteriorly angled cusps, and more expanded laterally oriented occlusal cusps than species adapting to animal‐dominated diets. Measures of incisor depth, crest length, cusp angle and sharpness, occlusal cusp orientation, and the lengths of third molars proved the best predictors of dietary adaptation. Accounting for evolutionary history in a phylogenetic discriminant function analysis notably improved the classification accuracy. Molar morphology is strongly correlated with diet and we suggest that these dental traits can be used to infer diet with good accuracy for both extinct and extant murine species.     

Ontogenetic transition from unicuspid to multicuspid oral dentition in a teleost fish: Astyanax mexicanus, the Mexican tetra (Ostariophysi: Characidae)

Teleost fishes display a remarkable diversity of adult dentitions; this diversity is all the more remarkable in light of the uniformity of first-generation dentitions. Few studies have quantitatively documented the transition between generalized first-generation dentitions and specialized adult dentitions in teleosts. We investigated this transition in the Mexican tetra, Astyanax mexicanus (Characidae), by measuring aspects of the dentition in an ontogenetic series of individuals from embryos to 160 days old, in addition to adults of unknown age. The first-generation dentition and its immediate successors consist of small, unicuspid teeth that develop extraosseously. Multicuspid teeth first appear during the second tooth replacement event, and are derived from single tooth germs, rather than from the fusion of multiple conical tooth germs. We document that the transition from unicuspid to multicuspid teeth corresponds to a change in the location of developing tooth germs (from extraosseous to intraosseous) and in patterns of tooth replacement (from haphazard to simultaneous within a jaw quadrant). In addition, while the size of the largest teeth scales with positive allometry to fish size, the transition to multicuspid teeth is accompanied by an exceptionally large increase in tooth size.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 145 , 523–538.     

Sex-specific strategies of dentine depletion in red deer

Worn teeth in herbivore ungulates may be related to lower efficiency in mastication and hence lower performance. However, selection should favour maximal performance in terms of body mass and reproductive capacity during reproductive lifespan, when permanent teeth are already partially worn. We hypothesize that wear rate may respond to a strategy of use of tooth materials (notably dentine), which balances instantaneous wear rate and performance against tooth preservation for future performance and reproduction. In the present study, we investigated 4151 carcasses of Iberian red deer Cervus elaphus hispanicus and show that more worn molars were not related to lower performance throughout age. By comparing between sexes, tooth wear rates were smaller in females than in males, but the relationship between tooth wear and body performance also differed between the sexes: females did not show a significant relationship between tooth wear and performance but males with more worn teeth were in general heavier and had larger antlers until senile age, when more depleted teeth were related to smaller antlers. These results reveal, for the first time, sex-specific lifetime strategies of dentine expenditure: maintenance of performance ability throughout a longer reproductive lifespan in females, compared with maximizing current performance by depleting dentine reserves within a shorter lifespan in males.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 487–497.     

Ontogeny, anatomy and attachment of the dentition in mosasaurs (Mosasauridae: Squamata)

Aspects of mosasaur dental ontogeny are well preserved in many fossils of these giant marine squamates. Replacement teeth on the tooth-bearing elements (TBEs) first appear as small enamel crowns positioned posterolingual to the attached tooth (posterolabial for the pterygoid). Several developing crowns, of progressively larger size, are aligned in rows relating to a specific tooth position. The crowns rest in a dental groove that varies in width and depth depending on the TBE. The crown closest to the attached tooth is always the largest and is found in a small resorption pit. As resorption proceeds, the pit expands in volume (cementum and alveolar bone), and the crown increases in size and settles into the pit. Once mature crown size is achieved, the dentine root and cementum portion of the root develop rapidly, the attached tooth is lost and the replacement tooth erupts out of the alveolus. Mosasaurid teeth develop along a 'zig-zag'-shaped movement path: horizontally along the dental groove, down into the alveolus, and up and out of the alveolus prior to attachment to the alveolar wall. At no point in mosasaurid tooth development are the crowns observed in a horizontal position. The mosasaurid dental lamina appears to have been a continuous strip of dental epithelium as it is in other squamates. Mosasaurid tooth attachment is thecodont (histologically and geometrically) not subpleurodont. Most aspects of mosasaurid tooth attachment and ontogeny are autapomorphic for the group.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 149 , 687–700.     

Factors affecting habitat selection in a specialist fossorial skink

Habitat specialists maximize their fitness by using a subset of the habitats that are potentially available to them and fare poorly if they move elsewhere. The factors that constrain habitat use are diverse and often difficult to identify, but are important to distinguish if we are to understand the trade-offs that drive species to become specialists. In the present study, we investigated habitat use in a fossorial skink, Lerista labialis , and explore the factors that confine it to the crests of sand dunes in the Simpson Desert, central Australia. Models positing that L. labialis selects dune crests because of their sparse cover of vegetation, more favourable temperatures, and greater abundance of preferred prey, received no support. Instead, a model positing that dune crests provide soft and less compacted sand that facilitates movement by L. labialis , was strongly supported. Sand on the crests was consistently softer that that on the sides and swales of the dunes; the skinks preferred soft rather than hard sand for movement in captivity, and were captured more often on experimentally softened sand than on compacted sand in the field. There was no evidence that L. labialis responds to attributes of the substrate other than softness because captive animals used loose sand from the dune crests, sides, and swales equally. We suggest that the dune crest environment allows L. labialis to reduce the energetic costs of locomotion, provides priority of access to the subterranean galleries of its termite prey, and also a secure refuge from surface-active predators and extreme surface temperatures.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 531–544.     

Six new Mexican species of Bernardia (Euphorbiaceae)

Six new species of Bernardia from Mexico are described and illustrated. Species are distinguished by a combination of characters such as habit, leaf shape and size, indumentum, venation pattern, gland position, length of staminate inflorescence, number of bracts, number of flowers per bract, size of bracts and tepals, number of stamens, style type, fruit dehiscence, size, pubescence, trichome type, endocarp texture, seed shape, size and ornamentation, raphe length and micropyle position. Bernardia macrocarpa belongs to section Alevia ; B. chinantlensis to section Polyboea , which is recorded for the first time in Mexico; B. chiangii , B. mcvaughii , B. rzedowskii and B. valdesii belong to section Tyria , which is the most diverse section in Mexico. With the recognition of these six new species, Mexico becomes the area with the highest diversity for Bernardia .  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 149 , 241–256.     

A new traversodontid cynodont from the Santa Maria Formation (Ladinian-Carnian) of southern Brazil, with a phylogenetic analysis of Gondwanan traversodontids

The Traversodontidae is a Triassic family of nonmammalian cynodonts, recorded worldwide, characterized by its bucco-lingually expanded postcanines. A recently discovered fauna in the locality of Santa Cruz do Sul, corresponding to the Santa Maria Formation of the Brazilian Middle/Upper Triassic (Ladinian/Carnian), is made up exclusively of nonmammalian cynodonts with an abundance of traversodontids. A new taxon, Santacruzodon hopsoni gen. et sp. nov . , the most commonly recorded traversodontid cynodont in the fauna, is described here. It is diagnosed by an autapomorphy, a very large posterolabial cusp in the upper postcanine, representing more than half of the labial crest, and a suite of traits present in other traversodontids. Among these features are the descendent flange of the jugal developed as a ball-shaped projection, also known in the Madagascan traversodontid Dadadon isaloi , and the presence of symmetrical incisors with numerous mesial and distal marginal cuspules, feature shared with the North American taxon Arctotraversodon plemmyridon . A phylogenetic analysis of traversodontids from Gondwana, based on a data matrix of 28 characters (mostly dental) and 15 terminals (13 traversodontids, including the new species, plus Diademodon and Trirachodon ) was performed. The new traversodontid appears as the sister taxon of Dadadon . A monophyletic Carnian group composed of Exaeretodon , Menadon and Scalenodontoides , with the Carnian Gomphodontosuchus , and the Ladinian Massetognathus as successive outgroups is also supported. In line with a recent phylogenetic hypothesis, a nonmonophyletic relationship for Scalenodon 'species' from the Anisian Manda beds also results from our analysis. Based on the cladogram obtained, morphological trends of change in the dentition of traversodontids throughout the Triassic are suggested.  © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society, 2003, 139 , 529−545.     

Unravelling the Functional Biomechanics of Dental Features and Tooth Wear

Most of the morphological features recognized in hominin teeth, particularly the topography of the occlusal surface, are generally interpreted as an evolutionary functional adaptation for mechanical food processing. In this respect, we can also expect that the general architecture of a tooth reflects a response to withstand the high stresses produced during masticatory loadings. Here we use an engineering approach, finite element analysis (FEA), with an advanced loading concept derived from individual occlusal wear information to evaluate whether some dental traits usually found in hominin and extant great ape molars, such as the trigonid crest, the entoconid-hypoconulid crest and the protostylid have important biomechanical implications. For this purpose, FEA was applied to 3D digital models of three Gorilla gorilla lower second molars (M₂) differing in wear stages. Our results show that in unworn and slightly worn M₂s tensile stresses concentrate in the grooves of the occlusal surface. In such condition, the trigonid and the entoconid-hypoconulid crests act to reinforce the crown locally against stresses produced along the mesiodistal groove. Similarly, the protostylid is shaped like a buttress to suffer the high tensile stresses concentrated in the deep buccal groove. These dental traits are less functional in the worn M₂, because tensile stresses decrease physiologically in the crown with progressing wear due to the enlargement of antagonistic contact areas and changes in loading direction from oblique to nearly parallel direction to the dental axis. This suggests that the wear process might have a crucial influence in the evolution and structural adaptation of molars enabling to endure bite stresses and reduce tooth failure throughout the lifetime of an individual.     

Modelling intraspecific resting egg shape variation in a freshwater fairy shrimp Tanymastix stagnalis (L., 1758) (Crustacea, Branchiopoda)

The shape of the resting eggs of a large branchiopod crustacean, the Anostraca Tanymastix stagnalis , is represented very accurately by analytical expressions. The occurrence of atypical shape of some T. stagnalis eggs may be viewed as a simple change of the analytical expression describing the usual egg shape. Their unusual shape may be explained by a higher embryo volume within an envelope of a given size. Biological implications are briefly discussed and hypothesized in an evolutionary point of view.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 55–60.     

Postcranial anatomy of Haldanodon exspectatus (Mammalia, Docodonta) from the Late Jurassic (Kimmeridgian) of Portugal and its bearing for mammalian evolution

The postcranium of the Late Jurassic docodont Haldanodon exspectatus Kühne & Krusat, 1972 is represented by a partial skeleton and isolated bones of other individuals from the Late Jurassic (Kimmeridgian) of the Guimarota coal mine in Portugal. Haldanodon exhibits adaptations for a fossorial lifestyle such as stout and short limb bones and humeri with greatly expanded distal joints and strong deltopectoral crests. Short first and second phalanges and moderately curved and laterally compressed terminal phalanges with lateral grooves suggest that Haldanodon was a scratch-digger. The dorso-ventrally elongated, triangular scapula has a convex transverse profile with strongly laterally reflected anterior and posterior scapula margins, enclosing a deep trough-like 'infraspinous fossa'. A supraspinous fossa is not developed. The saddle-shaped glenoid facet is mainly formed by the coracoid and orientated antero-ventrally indicating a sprawling gait. No epiphyses were detected and the wide size range of humerus and femur possibly indicate a lifelong growth. Haldanodon is more derived than Morganucodon by complete reduction of the procoracoid, absence of the procoracoid foramen, and a peg-like coracoid. It shares with monotremes a postscapular fossa that is absent in Morganucodon . A PAUP analysis based on 280 cranio-dental and postcranial characters corroborated the position of Haldanodon above morganucodontids and below Hadrocodium .  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 145 , 219–248.     

Detecting Inter-Cusp and Inter-Tooth Wear Patterns in Rhinocerotids

Extant rhinos are the largest extant herbivores exhibiting dietary specialisations for both browse and grass. However, the adaptive value of the wear-induced tooth morphology in rhinos has not been widely studied, and data on individual cusp and tooth positions have rarely been published. We evaluated upper cheek dentition of browsing Diceros bicornis and Rhinoceros sondaicus, mixed-feeding R. unicornis and grazing Ceratotherium simum using an extended mesowear method adapted for rhinos. We included single cusp scoring (EM(R)-S) to investigate inter-cusp and inter-tooth wear patterns. In accordance with previous reports, general mesowear patterns in D. bicornis and R. sondaicus were attrition-dominated and C. simum abrasion-dominated, reflecting their respective diets. Mesowear patterns for R. unicornis were more attrition-dominated than anticipated by the grass-dominated diet, which may indicate a low intake of environmental abrasives. EM(R)-S increased differentiation power compared to classical mesowear, with significant inter-cusp and inter-tooth differences detected. In D. bicornis, the anterior cusp was consistently more abrasion-dominated than the posterior. Wear differences in cusp position may relate to morphological adaptations to dietary regimes. Heterogeneous occlusal surfaces may facilitate the comminution of heterogeneous browse, whereas uniform, broad grinding surfaces may enhance the comminution of physically more homogeneous grass. A negative tooth wear gradient was found in D. bicornis, R. sondaicus and R. unicornis, with wear patterns becoming less abrasion-dominated from premolars to molars. No such gradients were evident in C. simum which displayed a uniform wear pattern. In browsers, premolars may be exposed to higher relative grit loads, which may result in the development of wear gradients. The second premolar may also have a role in food cropping. In grazers, high absolute amounts of ingested abrasives may override other signals, leading to a uniform wear pattern and dental function along the tooth row, which could relate to the observed evolution towards homodonty.     

Ontogenesis of internal secretory cells in leaves of Lantana camara (Verbenaceae)

Internal secretory cells may be morphologically indistinguishable from their neighbours except for the presence of secreted material, or they may differ to such an extent that could be classified as secretory idioblasts. Several authors have reported the presence of glandular trichomes secreting essential oil in Verbenaceae, including Lantana . However, none have reported internal secretory cells. Anatomical and histochemical methods applied to Lantana camara leaves revealed the occurrence of internal secretory cells whose ontogenesis and chemical nature are described in this paper. According to leaf developmental analysis, L. camara secretory cells originated from the ground meristem, started to differentiate in the third node leaves, and were actively secreting in the fourth node leaves. The content of the secretory cells was of a lipidic nature, and a terpenoid essence of their secretion was also identified. Based upon differences in size and shape from neighbouring cells and on detection of nonvolatile terpenes, they were confirmed as true internal secretory idioblasts.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 427–431.     

Occlusal relief changes with molar wear in Pan troglodytes troglodytes and Gorilla gorilla gorilla

Most research on primate tooth form-function relationships has focused on unworn teeth. This study presents a morphological comparison of variably worn lower second molars (M(2)s) of lowland gorillas (Gorilla gorilla gorilla; n=47) and common chimpanzees (Pan troglodytes troglodytes; n=54) using dental topographic analysis. High-resolution replicas of occlusal surfaces were prepared and scanned in 3D by laser scanning. The resulting elevation data were used to create a geographic information system (GIS) for each tooth. Occlusal relief, defined as the ratio of 3D surface area to 2D planometric area of the occlusal table, was calculated and compared between wear stages, taxa, and sexes. The results failed to show a difference in occlusal relief between males and females of a given taxon, but did evince differences between wear stages and between taxa. A lack of significant interaction between wear stage and taxon factors suggests that differences in occlusal relief between chimpanzees and gorillas are maintained throughout the wear sequence. These results add to a growing body of information on how molar teeth change with wear, and how differences between primate species are maintained at comparable points throughout the wear sequence. Such studies provide new insights into form-function relationships, which will allow us to infer certain aspects of diet in fossils with worn teeth.     

Comparative morphology of seeds of four closely related species of Aristolochia subgenus Siphisia (Aristolochiaceae, Piperales)

The primary purpose of this study was to determine whether morphological divergence has occurred in seeds of species in a subclade of Aristolochia subgenus Siphisia . Thus, we compared seed mass, surface area, and wings (presence or absence), embryo shape and state, and embryo length to seed length (E:S) ratio in A. californica Torr., endemic to California, A. macrophylla Lamk. and A. tomentosa Sims of eastern USA, and A. manshuriensis Komarov of eastern Asia. We also compared capsule length and number of seeds per capsule in the three US species. All four species have linear, underdeveloped embryos, but for the other morphological features statistically significant differences occurred among the taxa. However, none of the seed morphological features differed between the eastern North American species A. macrophylla and its eastern Asian sister species A. manshuriensis.   © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 433–436.     

Explaining patterns in modern ruminant diversity: contingency or constraint?

Bovid species outnumber those of cervids by over two and half to one. The present study attempts to explain this difference using two principle concepts: contingency and constraint. The concept of contingency considers the role of biogeography and the distributions and timings of major dispersals seen in the two families. Although this historical context is considered vital for understanding the problem, cervids are also characterized by their global absence from open and arid grassland habitats. This is explained in terms of phylogenetic constraint. One proposal considers the deciduous antlers of cervids as constraining the group to higher quality forage that is not generally associated with open habitats. Alternatively and more generally applicable to all ruminants is the ‘cusp fusion hypothesis’, presented here for the first time. The hypothesis seeks to explain how the primitive ruminant molar state, as seen in modern cervids and giraffids, prevents the attainment of very high molar crowns and, with it, the ability to cope with high rates of dietary tooth wear. Conversely, the derived condition of precocial cusp fusion seen in the Bovidae and Antilocapridae has enabled higher tooth crowns, their expansion into open habitats, and an increased diversity potential. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 657–672.     

Description of a new Eocene osteoglossid fish and additional information on †Singida jacksonoides Greenwood and Patterson, 1967 (Osteoglossomorpha), with an assessment of their phylogenetic relationships

A new genus and species of osteoglossomorph fish is described from the Eocene Mahenge site in north central Tanzania. The new species is sympatric with the previously described osteoglossomorph † Singida jacksonoides , but is easily distinguished from it by the presence in the new species of teeth on the jaws and entopterygoid. The new species shares characters with the Heterotidinae, such as the two arms of the preopercle being of similar length and the posterior end of the maxilla lying on the dentary, and other characters with the Osteoglossinae (including † Phareodus ), for example the enlarged pectoral fin ray and the shape of the opercular bone. New specimens of † S. jacksonoides also have been recovered from the site, providing more information on the osteology of this fish. The relationships of these two Mahenge osteoglossomorphs are assessed, with both fishes placed within the Osteoglossidae, but of uncertain position in the family. Addition of the new species to previous cladistic data sets, and the revised information for † S. jacksonoides , causes the subfamily Osteoglossinae to be nonmonophyletic.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144 , 213–228.     

A morphometric study on morphological plasticity of shell form in crevice-dwelling Pterioida (Bivalvia)

Relative warp analysis and eigenshape analysis were used to explore the patterns of variation of erratic shell shapes in and among six species of crevice-dwelling pterioid bivalves. The results of morphometric analyses revealed that a great deal of the variance of shell shape within each species can be reduced into principal components which display patterns of variation common to all species examined. In the species with striking variability, a few sets of principal components account for most of the variance of shell shape. On the other hand, in the species lacking considerable variability, several components contribute to form a given variety. Comparison of relative warp and eigenshape scores with centroid sizes indicates that the direction of ontogenetic shape change has been modified through evolution to produce adaptations to habitats such as crevices or the undersides of rocks.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79, 285–297.