New data on the braincase of the aetosaurian archosaur (Reptilia: Diapsida) Stagonolepis robertsoni Agassiz

New data on the braincase of the aetosaurian archosaur Stagonolepis robertsoni Agassiz are presented, based on new preparation, synthetic casting, and interpretation of fossil material from the Triassic Elgin Sandstones, Scotland. The metotic fissure is not divided by bone. The perilymphatic foramen is completely bound by bone, and faces away from the otic capsule in a posterolateral direction. A prominent subvertical ridge on the anterolateral edge of the exoccipital and upper part of the basioccipital cannot be directly associated with the subcapsular process of the chondrocranium of extant crocodilians. This ridge projects laterally beyond the ventral ramus of the opisthotic, and lies anterior to the external foramina for the hypoglossal nerve. The overall structure of the braincases (especially the otic region) of S. robertsoni and other aetosaurians, where known, is more similar (in terms of derived archosaurian characters) to those of crocodylomorphs than are the braincases of other major suchian groups. This provides evidence for the currently unorthodox hypothesis that, among major suchian clades, Aetosauria and Crocodylomorpha are each others' closest relatives. Support for this hypothesis is found in features of the palatine and prefrontal that have not been considered in recent studies of suchian phylogeny. This alternative phylogenetic hypothesis demands further investigation but, combined with the new morphological data that it explains, it provides a framework for the understanding of the evolution of the derived and distinctive braincase structure of extant crocodilians.  © 2002 The Linnean Society of London, Zoological Journal of the Linnean Society, 2002, 136 , 7−23.     

Ontogeny of the partial secondary wall of the otoccipital region of the endocranium in prehatching Alligator mississippiensis (Archosauria, Crocodylia)

The ontogeny of the posterior otic and anterior occipital portions of the neural endocranium of prehatching Alligator mississippiensis was investigated by reconstruction from sectioned material. In Stage 6 of this species, in which the endochondral ossification of the otoccipital region of the neural endocranium is only in its very early stage, two bony outgrowths-laminae-are present at the external wall of the posterior portion of the neural endocranium. The anterior lamina arises from the external surface of the basal plate at the level of the posterior margin of the subcapsular process; the posterior lamina arises from the external surface of that portion of the pila occipitalis that forms the posteroventral wall of the metotic fissure. During ontogeny, both laminae lying in the anteroposterior sequence ossify in membrane, fuse together, grow laterodorsally, and fuse with the lateral wall of the lateral semicircular canal and the crista parotica. This lamina forms a new, secondary wall enclosing the posterior section of the otic capsule and contains the large external jugular foramen (or foramen vagi) in its basal portion. The laminae, designated lamina juxtaotica anterior and posterior (lamina juxtaotica when fused together), have not been recorded previously in crocodylians and are absent in all other Recent reptiles. From the functional point of view, the juxtaotic lamina 1) forms the margins of the external jugular foramen, and 2) forms the floor of the posterior section of the Eustachian tube. In birds, the structure called the metotic cartilage, which arises in ontogeny as an independent element, has a similar position as the juxtaotic lamina. However, the two structures differ in their developmental origins and their relation to the Eustachian tube and the ramus hyomandibularis of the facialis nerve. Moreover, there is no external jugular foramen in birds.     

The otic region of Doleserpeton (Temnospondyli) and its implications for the evolutionary origin of frogs

There is increasing evidence that the Palaeozoic temnospondyl amphibians had a frog‐like tympanic hearing system. For this reason, the otic region of Doleserpeton is described and compared with modern anurans. The otic capsules are expanded laterally and ventrally relative to other temnospondyls. The opisthotic has a bulbous ventral region resembling the ventrolateral ledge in modern frogs. Two lateral processes are located on the paroccipital process. Comparison with the condition in modern anurans with a tympanic hearing system shows that this may have been the attachment site for the tympanic annulus. Parts of the osseous labyrinth are also described. The inner ear shows numerous features resembling the condition found in frogs. These include strong evidence for the presence of a lissamphibian‐type perilymphatic duct most closely resembling that of anurans. This is the first time such a perilymphatic system has been described in any Palaezoic form. The posterior part of the braincase shows a jugular foramen closely associated with the perilymphatic foramen, as in anurans. Although the distribution of these traits among other temnospondyl groups remains little known, the sum of the evidence points to affinities between anurans and temnospondyls, and adds to the evidence for a close relationship between anurans and the Permian amphibamid Doleserpeton. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154 , 738–751.     

Holocephalan Embryo Provides New Information on the Evolution of the Glossopharyngeal Nerve,Metotic Fissure and Parachordal Plate in Gnathostomes

The phylogenetic relationships between the different groups of Paleozoic gnathostomes are still debated, mainly because of incomplete datasets on Paleozoic jawed vertebrate fossils and ontogeny of some modern taxa. This issue is illustrated by the condition of the glossopharyngeal nerve relative to the parachordal plate, the otic capsules and the metotic fissure in gnathostomes. Two main conditions are observed in elasmobranchs (shark and rays) and osteichthyans (bony fishes and tetrapods). The condition in the other chondrichthyan taxon, the holocephalans, is still poorly known, and without any information on this taxon, it remains difficult to polarize the condition in gnathostomes. Based on the anatomical study of an embryo of the holocephalan Callorhinchus milii by means of propagation X-Ray Synchrotron phase contrast microtomography using both holotomography and single distance phase retrieval process, we show that, contrary to what was previously inferred for holocephalans (i.e. an osteichthyan-like condition), the arrangement of the glossopharyngeal nerve relative to the surrounding structure in holocephalans is more similar to that of elasmobranchs. Furthermore, the holocephalan condition represents a combination of plesiomorphic characters for gnathostomes (e.g., the glossopharyngeal nerve leaves the braincase via the metotic fissure) and homoplastic characters. By contrast, the crown osteichthyans are probably derived in having the glossopharyngeal nerve that enters the saccular chamber and in having the glossopharyngeal foramen separated from the metotic fissure.     

Phylogenetic implications of the morphology of the braincase of caecilian amphibians (Gymnophiona)

Caecilian morphology is strongly modified in association with their fossorial mode of life. Currently phylogenetic analyses of characters drawn from the morphology of caecilians lack resolution, as well as complementarity, with results of phylogenetic analyses that employ molecular data. Stemming from the hypothesis derived from the mammal literature that the braincase has the greatest potential (in comparison to other cranial units) to yield phylogenetic information, the braincase and intimately associated stapes of 27 species (23 genera) of extant caecilians were examined using images assembled via microcomputed tomography. Thirty‐four new morphological characters pertaining to the braincase and stapes were identified and tested for congruence with previously recognized morphological characters. The results reveal that when added to previous character matrices, characters of the braincase and stapes resolve generic‐level relationships in a way that is largely congruent with the results of molecular analyses. Analysis of a combined data set of molecular and morphological data provides a framework for conducting ancestral character state reconstructions, which resulted in the identification of 95 new synapomorphies for various clades and taxa, 27 of which appear to be unique for the taxa that possess them. Together these data demonstrate the utility of the application of characters of the braincase and stapes for resolving phylogenetic relationships for a group whose morphology is largely confounded by functional modifications. In addition this study provides evidence of the utility of the braincase in resolving problematic morphology‐based phylogeny outside of Amniota, in an amphibian group. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166 , 160–201.     

Phylogeny and biogeography of the Vitrinidae (Gastropoda: Stylommatophora)

The phylogeny of the Vitrinidae is reconstructed in a cladistic analysis based on characters of the genitalia, the copulation behaviour and the radula. The genera with an atrial stimulator turned out to be the earliest branches of the Vitrinidae, whereas the genera with a glandula amatoria form a monophyletic, taxonomically apomorphic group. The differences between the proposed phylogeny and previous hypotheses are discussed. The ancestral areas of the Vitrinidae and its sister group, the limacoid slugs Boettgerillidae–Limacidae–Agriolimacidae, are estimated using weighted ancestral area analysis. The Vitrinidae and the limacoid slugs might have originated by a vicariance event between Central Europe and the Near East. © 2002 The Linnean Society of London. Zoological Journal of the Linnean Society , 2002, 134 , 347–358.     

The effects of experimental unilateral anotia on skull development in the chick embryo. III. Chondrocranial development in anotic embryos of 7-20 days of incubation.

The study of the development of of the chondrocranium in chick embryos with unilateral (right-sided) anotia revealed the following main characteristics. 1. The median axes of the chordal and the prechordal part of the cranial base are not in a straight line but show a deviation toward the right side. The angle between the two axes has its vertex in the region of the foramen hypophyseos. 2. The metotic cartilage and the foramina of the IXth and Xth cranial nerves are normal in position. 3. The tectum synoticum develops later and to a lesser extent than normal. 4. Between the basal plate, the metotic cartilage, the occipital arch and the supracapsular cartilage a foramen is formed which, later in development, is closed by outgrowths of the metotic cartilage and the basal plate. 5. The "optic area" shows a practically normal appearance which indicates that the cartilaginous ventral wall of the lagenal capsule is of basal plate origin. 6. The pro-otic process develops practically normal and, hence, is independent of the ear capsule. 7. The quadrate cartilage and the right lower jaw are displaced ventro-posteriorward. The earliest development of the perichondral bones shows some particularities which are closely correlated with the development of the various cartilaginous structures.     

Disparity and geometry of the skull in Archosauria (Reptilia: Diapsida)

A metric comparison of 155 fossil and extant species in lateral view based on the proportions of three homologous units (braincase, orbit and rostrum) reveals the existence of an archosaurian skull geometry. An empirical morphospace depicting skull proportions shows that the most variable unit is the rostrum. Three skull types based on rostral proportion are proposed: meso-, longi- and brevirostral. These types depend, on one hand, on a direct numerical relationship between the braincase and the orbit, with a mean ratio of 1:1; never surpassing a 2:1 or 1:2 ratio limit. On the other hand, skull types show a significant negative correlation between braincase and rostrum proportions. Close relationships have been obtained between orbit and the rostrum, although with lower significance and a geometric meaning specific to each group. Skull types depend mainly on the proportional relationship between the rostrum and the braincase. Mesorostral types account for more natural occurrences within morphospace, implying a plesiomorphic condition in Archosauria. Skulls with highest longirostral values (flying forms) display a more restrictive braincase–orbit ratio relationship. Brevirostrals are limited to the smallest skull lengths, up to approximately 180 mm. 85% of brevirostral modern birds have altricial post-hatchling development. General allometric pattern is very similar for all sampled archosaurs, although giant taxa (i.e. non-avian theropods) display a different type of skull proportional growth, closer to isometry. Results reveal the existence of a constructional skull geometry, highlighting the importance of the deviance of the structural design from adaptive explanations on craniofacial morphology in macroevolution.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80, 67–88.     

Phylogeny of a paradigm lineage: the Drosophila melanogaster species group (Diptera: Drosophilidae)

Although Drosophila melanogaster is a paradigm eukaryote for biology, relationships of this species and the other 174 species in the melanogaster species group are poorly explored and ambiguous. Gene regions of Cytochrome oxidase II (mt:CoII ), Alcohol dehydrogenase ( Adh ) and hunchback ( hb ) were sequenced and analysed phylogenetically to test prior hypotheses of relationships for the group based on chromosomes, morphology, and 28S rRNA gene sequences. A simultaneous cladistic analysis of the three newly sequenced gene regions produced a single well-resolved phylogeny for 49 exemplar species representing eight subgroups. Monophyly of each of the ananassae , melanogaster , montium , and takahashii subgroups is supported; the suzukii subgroup is polyphyletic. This phylogeny is consistent with variation in significant morphological structures, such as the male sex comb on the fore tarsus. The broad range of morphological variation among these species is interpreted and the applicability to evolution and developmental investigations is discussed. This phylogeny facilitates comparative investigations, such as gene family evolution, transposable element transmission, and evolution of morphological structures. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 76 , 21–37.     

Body mass and habitat correlates of song structure in a primitive group of birds

We assessed relationships between acoustic frequency, body mass, and habitat in tinamous. This monophyletic group of primitive birds comprises c. 47 ground dwelling species whose habitats range from dense humid forest to open grasslands. The relationship between frequency and body mass was found to be negative, while the songs of open-habitat species exhibited higher frequencies and a wider bandwidth than the closed-habitat ones. Residual variation in song frequency, after controlling for the effect of body mass and phylogeny, tends to differ among habitats. However, a statistical test of this pattern was not possible because of the existence of only five pairs of sister species differing in habitat. In spite of this, positive contrasts of bandwidth were associated with positive contrasts of habitat, confirming that songs of open-habitat species have a wider bandwidth than those of their more closed habitat relatives.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 423–430.     

Systematics of the tribe Podalyrieae (Fabaceae) based on DNA, morphological and chemical data

Phylogenetic relationships among nine genera and 28 species of the southern African tribe Podalyrieae were estimated from sequences of the internal transcribed spacer (ITS) of nuclear ribosomal DNA as well as morphological and chemical data. Morphological and ITS sequence data produced cladograms with similar topologies, both supporting the monophyly of Podalyrieae (excluding Hypocalyptus ). The combined data sets indicate that subtribe Xiphothecinae are monophyletic, but embedded within Podalyriinae. The high degree of congruence between previous taxonomic hypotheses and those based on DNA data provides further evidence for the utility of ITS sequences in studying phylogeny.  © 2002 The Linnean Society of London , Botanical Journal of the Linnean Society , 2002, 139 , 159–170.     

The braincase and middle ear region of Dendrerpeton acadianum (Tetrapoda: Temnospondyli)

Dendrerpeton acadianum from the Westphalian A (Upper Carboniferous) of Joggins, Nova Scotia, is a phylogenetically and chronologically early temnospondyl. Its external cranial anatomy has been used previously to suggest the presence of a tympanic membrane, and thus of an ear adapted to the perception of airborne sound. However, supporting evidence provided by stapedial and braincase morphology has so far been lacking. The braincase and middle ear region have remained almost wholly unknown. CT scanning and 3-D computer reconstruction of BMNH R.436 have been used to shed light on these important areas. Both stapes prove to be present in the specimen; the right stapes is distorted, but the left stapes lies inside the cranial cavity and is perfectly preserved. The latter resembles the stapes of the relatively few other temnospondyls in which the bone has been described and is most similar to that of Doleserpeton . The morphology and orientation of the stapes provide strong evidence for the presence of an ear adapted to the perception of airborne sound, with similarities to the extant anuran condition. The reconstructed braincase shows a high degree of similarity to that of other adequately known temnospondyls. This gives supporting evidence that D. acadianum is correctly placed in the temnospondyl phylogeny and thus demonstrates one of the earliest hearing systems adapted to the perception of airborne sound that can be homologized with the extant anuran condition.  © 2005 The Trustees of the Natural History Museum, Zoological Journal of the Linnean Society , 2005, 143 , 577−597.     

Evolution and determinants of host specificity in the genus Lamellodiscus (Monogenea)

The evolution and determinants of host specificity in Lamellodiscus species (Monogenea, Diplectanidae) were investigated. The 20 known Mediterranean species were studied, all parasites of fishes from the family Sparidae (Teleostei). An index of specificity, which takes into account the phylogenetic relationships of their fish host species, was defined. The link between specificity and its potential determinants was investigated in a phylogenetic context using the method of independent contrasts. Host specificity in Lamellodiscus species appeared to be highly constrained by phylogeny, but also linked to host size. Mapping specificity onto the parasite phylogenetic tree suggests that specialist species do not represent an evolutionary dead end, and that specialization is not a derived condition. It is hypothesized that the ability to be generalist or specialist in Lamellodiscus is controlled by intrinsic, phylogenetically-related characteristics, and that specialist species tend to use large hosts, which may be more predictable.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77, 431−443.     

Patagial complex evolution in hummingbirds and swifts (Apodiformes): a molecular phylogenetic perspective

Studies of the role of flight in vertebrate evolution often have focused on the propatagial muscle complex because this structure forms the wing's leading edge. However, historical narratives about the evolution of flight anatomy are compromised because traditional higher-level taxonomies typically are based in part on the propatagium itself. To avoid this circularity, I used a consensus molecular phylogeny to examine propatagial evolution in the highly aerial sister groups, hummingbirds and swifts (Apodiformes). Mapping of anatomy on molecular-based phylogeny indicates that structural variation in M .  tensor propatagialis pars brevis (TPB) is congruent with the major subclades of both hummingbirds and swifts. However, the humeral tendon and broad attachment of the fleshy belly of TPB with M .  extensor metacarpi radialis (EMR) most likely underwent parallel change in hummingbirds and swifts, while the distal tendon present only in hummingbirds changed from a thin sheet to a strong tendon. The combination of divergent (within hummingbirds or swifts) and parallel (between hummingbirds and swifts) evolutionary patterns implies that the taxonomic value of the propatagial complex in apodiformes depends on anatomical component and level of divergence. The congruence of anatomy with molecular phylogeny provides independent criteria for designating relatively ancestral versus derived clades of apodiformes. Based on these polarities, living hummingbirds and swifts express additional parallel trends from arboreal to more aerial foraging styles, and from depauperate to species-rich clades. Within apodiformes, the link of flight anatomy with taxonomic and ecologic diversity suggests that elaboration of locomotor modes was important for apodiform diversification, echoing a similar pattern for birds relative to their reptilian ancestors.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 211–219.     

Phylogenetic relationships of the Sparidae (Teleostei: Percoidei) and implications for convergent trophic evolution

A phylogenetic analysis of the majority of sparid genera and representatives of the sparoid families Centracanthidae, Lethrinidae and Nemipteridae is presented using 87 predominately osteological characters. The Sparidae constitute a monophyletic grouping, with the inclusion of the centracanthid Spicara smaris , which nests deep within the ingroup. The phylogeny was then used to investigate agreement with the most recent molecular study, taxonomic stability of subfamilial classification and the evolution of feeding strategies. Results show that the incongruence between morphological and molecular data appears largely to be an artifact of errors in rooting. However, there appears to be real and substantial conflict between the molecular tree and the morphological data, which is not attributable to the different positions of the least congruent taxa. The data support the molecular hypothesis that none of the subfamilial classification, based on dentition and trophic specialization, is monophyletic, and should be rejected pending further taxonomic revision. The phylogeny supports multiple independent origins of trophic types and it is suggested that the evolutionary plasticity of the oral teeth of sparids has been fundamental to the adaptive radiation of this family compared to their closest allies. ©2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 76 , 269–301.     

Locomotion in terrestrial mammals: the influence of body mass, limb length and bone proportions on speed

Traditionally a few limb proportions or total limb lengths have been regarded as indicative of peak running velocity. This is due to physical principles (inferred in- and outvelocities around the joints, stride lengths) and also the observation that fast-moving animals tend to share a number of purported key features which are either absent or not developed to near the same extent in slower moving forms. Previous studies have shown hind limb length and metatarsus/femur ratio to be correlated significantly, albeit modestly with running speed. These studies have nearly all been bivariate analyses. Based on the physical principles, there is reason to suppose that more variables than just m/f ratio could be important as adaptations for fast locomotion, and also that bivariate analyses are too simple. In this study a sample of 76 running mammals was used, with running speeds taken from literature. A number of osteological parameters were discovered to covary significantly with peak running speed, albeit only modestly. Using the information from phylogeny reduced all correlations, often significantly so. Multivariate analyses resulted in markedly higher correlation coefficients. Animals probably do not optimize their anatomy for the purpose of running very fast, which occurs only on rare occasions, but for reducing costs of locomotion.  © 2002 The Linnean Society of London, Zoological Journal of the Linnean Society , 2002, 136 , 685–714.     

The braincase of a specimen of <Emphasis Type="Italic">Proterochampsa</Emphasis> Reig (Archosauriformes: Proterochampsidae) from the Late Triassic of Argentina

The proterochampsids are a Triassic group of superficially crocodile-like forms belonging to the Archosauriformes. In the present contribution, we present new information regarding the braincase of the proterochampsid Proterochampsa Reig 1959, from the Ischigualasto Formation (Carnian) of Argentina, and discuss its phylogenetic considerations. Some unique neurocranial features of Proterochampsa are described, including: the prominence and thickness of the V-shaped ridge that surrounds the basisphenoidal fossa; the medially concave lateral arms of the same ridge; and the semilunar depression on the parabasisphenoid ventrolaterally exposed. Other features are only shared with likely unrelated archosauriforms, including: the great lateral development of the basipterygoid processes and caudal development of its distal end; an eight-shaped metotic foramen; laterally directed basipterygoid processes; and rostral boundary of the basisphenoidal recess V-shaped. Proterochampsa differs in many other aspects from the archosauriform Chanaresuchus, including: a proportionally shorter basioccipital basal tubera; cultriform process ovoid in cross-section; longitudinal sulcus dorsal to the basipterygoid process; deep basisphenoidal recess; and the absence of a prominent intertuberal plate. In many braincase features, Proterochampsa is more similar to archosaurs than to Euparkeria, erythrosuchids and Proterosuchus. They include a reduced semilunar depression. A ventral border of the basioccipital forming a wide convexity and a dorsoventrally thin paroccipital process likely represents a feature shared with Chanaresuchus, but not with Doswellia and other basal archosauriforms.