Early larval ontogeny of the Permo-Carboniferous temnospondyl Sclerocephalus

The early larval development of the temnospondyl Sclerocephalus sp. is analyzed, based on 38 specimens from the Lower Rotliegend (Permo-Carboniferous boundary) of the Saar-Nahe Basin (south-west Germany). The study focuses on the smallest larval specimens, which exemplify changes in both proportions and ossification patterns. In comparison with dissorophoid larvae, the skull ossifies more fully and at a much faster rate; the smallest specimens already have completely formed circumorbital bones that are sutured throughout. Sculpturing undergoes two marked changes, first from uniformly pitted to pits of variable size and regional differentiation, and finally to the origin of ridges. The palate of small larvae differs from that of larger specimens in patterns of dentition, having more teeth including a denticle field on the cultriform process. The mandible of small larvae is described for the first time, being narrower than in adults and having three dentigerous coronoid elements. The smallest specimens have poorly ossified neural arches, lack vertebral centra, and have faintly ossified humeri, femora, and very poorly developed distal elements. The posterior ribs, metapodia, and phalanges appeared after the dermal elements of the pectoral girdle, whereas the scapulocoracoid and ischium are absent throughout the larval period. Early growth and differentiation of the limbs and the ilium illustrates the developmental patterning of the appendages, which proceeded from proximal to distal. Dermal squamation is uniform in small stages, consisting of round or oval osteoderms with pronounced growth rings; in large larvae, they start to differentiate in certain body regions.     

The systematic relationships of the snake genus Anomochilus

Phylogenetic analysis of 38 skeletal characters, 12 muscular characters and 15 visceral characters in 17 major snake clades plus Anomochilus suggests that Anomochilus is the sister taxon of all other living alethinophidian snakes. However, skeletal, muscular and visceral character sets analysed separately or in pairs give four groups of nonconcordant tree topologies. Based on the cladogram derived from the total evidence, two families are erected to prevent the existing family Uropeltidae from becoming paraphyletic: Anomochilidae, for the Malaysian and Indonesian genus Anomochilus , and Cylindrophiidae, for the Sri Lankan, Southeast Asian and Indonesian genus Cylindrophis and the Upper Eocene fossil Eoanilius.     

Embryos of therizinosauroid theropods from the Upper Cretaceous of China: diagnosis and analysis of ossification patterns

Exceptionally complete, in ovo dinosaur embryos from the Upper Cretaceous of China are analysed. Ossification patterns of these embryos suggest that they died during the final third of their development. The therizinosauroid identity of the embryos follows from: (1) an edentulous premaxilla with a sharp downturned edge; (2) dentary with a lateral shelf; (3) teeth with fan-shaped crowns, with a few marginal cusps; (4) humerus with a massive deltopectoral crest extending proximally, with a pointed proximomedial tuberosity; (5) ilium with an expanded and hooked preacetabular process; (6) strongly curved hypertrophied manual unguals tapering to sharp points. These embryos are closest to two Chinese therizinosauroids, Neimongosaurus yangi Zhang et al . 2001 and Erliansaurus bellamanus Xu et al . 2002 . An elongated narial opening, reduced basipterygoid process, low cervical neural spines, a transversely narrow pubic apron, and a pubic foot expanded anteriorly are found in these embryos and are synapomorphies uniting the Therizinosauroidea and the Oviraptorosauria. Fusion of cervical and caudal neural arches and centra, complete ossification of thoracic ribs and ilium, possible co-ossification of tibia and fibula, fused pubes, complete meta- and acropodial elements, together with small portions of unossified epiphyses of long bones suggest an advanced precociality of these embryos.     

Phylogenetic relationships among species of the genus Issatchenkia Kudriavzev

The phylogenetic relatedness of Issatchenkia spp. was estimated from partial rRNA sequences in two regions of the large subunit and one region of the small subunit. I. terricola was the most divergent species of the genus, differing from other members by 18% nucleotide differences in the highly variable 25S-635 region. These data indicate Issatchenkia to be the most divergent ascomycetous yeast genus presently known.     

Phylogenetic relationships within the Archiloa genus complex (Proseriata,Monocelididae)

Of the seven genera which we have recognised within the Archiloa genus complex sensu Karling (1966) the cosmopolitan genus Archilina is the most primitive and is characterised only by plesiomorphic characters, and has to be considered paraphyletic. All other species of the Archiloa genus complex are hypothesized to be derived from Archilina-like ancestors through different evolutionary lineages. One lineage led to the genera Archiloa, Inaloa, Archilopsis and Monocelopsis, taxa found in the Atlantic and the Mediterranean. These genera are monophyletic and their relationships are analyzed. The genera Mesoda (Brazil) and Tajikina (Northern Pacific) can be considered as two other separate lineages. Similarly, within what we now consider as the genus Archilina different lineages can be recognized in different regions.     

Limb ossification patterns of the ichthyosaur Stenopterygius, and a discussion of the proximal tarsal row of ichthyosaurs and other neodiapsid reptiles

Limb ossification patterns for the Lower Jurassic (Toarcian) ichthyosaur, Stenopterygius , are described. It is found that limb ossification follows a continuous proximal to distal sequence from the propodial elements through to the terminal elements of 1st to 4th digit in the manus and the 1st to 3rd digit in the pes. The 5th manal and 4th pedal digit begin ossification later than more preaxial digits and also show evidence of proximal addition of elements near the distal mesopodial row in a manner consistent with delayed ossification of the 5th distal mesopodial in other diapsids. Ossification of manal elements in the Supernumerary 3–4 (S3-4) digit and the 5th digit appear interdependent; if one or the other is highly ossified, ossification of the other is retarded. The 1st pedal digit is considered to be lost in Stenopterygius and the 4th pedal digit is identified as the 5th digit. Delayed ossification of the mesopodium is not observed. The most preaxial proximal tarsal is identified as the centralc; the remaining proximal tarsals are the astragalus and calcaneum, and it is inferred that the astragalus and calcaneum ossified from within a single proximal cartilage.     

The structure and development of the jaw adductor musculature in the turtle Chelydra serpentina

The investigation of the development of the trigeminal jaw adductor musculature in the turtle Chelydra serpentina documents the early aggregation of muscle rudiments around the innervating nerve branches, probably a consequence of inductive interaction. This may explain the early continuity of the intramandibularis with the intermandibularis muscle. Several aspects of muscle development differ in the turtle as compared to lizards. These differences highlight the fact that conjectures of homology, based on a static topographical correspondence of adult structures, cannot capture the dynamics of the developmental process. The intramandibularis muscle of turtles, comparable to that of crocodiles, represents a plesiomorphous structure which is not homologous to the intramandibularis muscle of lacertoid lizards, a derived feature of the Lacertoidea. A derived feature of the chelonian jaw adductor musculature is the posterodorsal expansion of the external adductor along a supraoccipital crest, developing according to a pattern of Haeckelian recapitulation. Muscle development serves to corroborate the concept of a monophyletic Eureptilia, including diapsids and synapsids, as opposed to the (paraphyletic) Anapsida. The impact of the differentiation of the external adductor into a pulley system on cranial kinesis is analysed in biomechanical terms.     

Phylogenetic relationships within phylum Rotifera: orders and genus Notholca

We investigated evolutionary relationships among orders in phylum Rotifera and among species in genus Notholca (Rotifera) by computing parsimonious cladograms. All of the most-parsimonious cladograms generated for the ordinal level confirm the view that class Monogononta, superclass Eurotatoria, and phylum Rotifera are monophyletic. Species within the genus Notholca were separated into six groups (clades), but some species have been defined based on highly variable characters not reliably studied using cladistics. Therefore, phenetic studies are warranted, especially for species possessing caudal processes.     

The phylogeny and ontogeny of humeral torsion

In a series of specimens extending from fossil material through recent vertebrates including man there occurs a gradual phylogenetic increase in the degree of humeral torsion. A further (ontogenetic) torsion is superimposed upon the phylogenetic one in man which increases from birth until the proximal epiphysial cartilage of the humerus disappears and bony fusion occurs. There is a distinct correlation between the calculated strength of humeral rotator muscles inserting above and below the proximal epiphysis; this suggests that they provide the forces involved in the production of humeral torsion. It is shown that ontogenetic or secondary torsion occurs proximally and not along the shaft of the bone. Differences in the degree of humeral torsion in either sex of adult Whites and Negroes are given and discussed.     

Intra- and inter-specific relationships in the six Italian species of the fairy shrimp genus Chirocephalus: combining allozyme and mtDNA data

This study analysed the levels of genetic differentiation within and among the six Italian species of the fairy shrimp genus Chirocephalus by analysing electrophoretic polymorphisms at 22 enzymatic loci and by sequencing a 665‐bp fragment of the mitochondrial gene encoding for subunit I of cytochrome oxidase. The allozyme data revealed different levels of intra‐specific differentiation; mean θ estimates were low in Chirocephalus salinus, higher and comparable in C. diaphanus and C. kerkyrensis, while C. ruffoi was the most genetically structured species. At the inter‐specific level, C. marchesonii was the most differentiated species, both for allozymes and mtDNA. Phylogenetic relationships deduced from allozymes and mtDNA were not always consistent with each other. This highlights the differences in performance of the two classes of molecular markers and the need of different independent strategies of data analysis to search for possible incongruence. Neither allozymes nor mtDNA supported monophyly of the diaphanus‐group, previously recognized on the basis of the morphology of appendages (antennae and penes). In contrast the molecular results were consistent with the great heterogeneity in resting egg morphology among representatives of the diaphanus‐group.     

Intergeneric phylogenetic relationships in catfishes of the Loricariinae (Siluriformes: Loricariidae), with the description of Fonchiiloricaria nanodon: a new genus and species from Peru

Recent investigations in the upper Río Huallaga in Peru revealed the presence of an intriguing species of the Loricariinae. To characterize and place this species within the evolutionary tree of the subfamily, a molecular phylogeny of this group was inferred based on the 12S and 16S mitochondrial genes and the nuclear gene F-reticulon4. The phylogeny indicated that this distinctive species was a member of the subtribe Loricariina. Given its phylogenetic placement, and its unusual morphology, this species is described as a new genus and new species of Loricariinae: Fonchiiloricaria nanodon. This new taxon is diagnosed by usually possessing one to three premaxillary teeth that are greatly reduced; lips with globular papillae on the surface; the distal margin of lower lip bearing short, triangular filaments; the premaxilla greatly reduced; the abdomen completely covered by plates, with the plates between lateral abdominal plates small and rhombic; a caudal fin with 14 rays; the orbital notch absent; five lateral series of plates; dorsal-fin spinelet absent; preanal plate present, large and solid, and of irregular, polygonal shape, the caudal peduncle becoming more compressed posteriorly for the last seven to 10 plates.     

Ontogeny of the Medial Masseter Muscle, Pseudo-Myomorphy, and the Systematic Position of the Gliridae (Rodentia, Mammalia)

Histological serial sections of fetal stages of various rodent taxa have been studied. We have concentrated on the ontogenetic differentiation of the infraorbital region in some hystricomorphs and myomorphs. The glirid taxa Graphiurus, Eliomys, Glis and Muscardinus show a specific mode of development of the medial masseter muscle that is clearly different from the other groups: In early ontogenetic stages, the muscle invariably has its anterior-most origin at the dorsolateral rim of the infraorbital foramen, and only in later fetal stages migrates to the side of the muzzle. In contrast to the glirines, Graphiurus has an aboral origin of the lateral masseter muscle from the beginning, and we consider this as a plesiomorphic state. In all other taxa showing an enlarged masseter medialis, this muscle bundle originates from the anterior portion of the ascending process or even from the premaxillary. We interpret our ontogenetical findings as support for the hypothesis of the monophyly of the Gliroidea and for the concept of `pseudo-myomorphy' of Vianey-Liaud (1985), which was derived from the fossil record. This evidence suggests that the gliroids are probably not members of the Myomorpha, but have acquired myomorphy independently. This conclusion is also in agreement with many molecular studies that suggest closer affinities of glirids with sciurids.     

Coming of age in fringillid birds: heterochrony in the ontogeny of secondary sexual characters

The great variation in the timing of development of adult plumages in North American male passerine birds has in recent years provoked much theoretical interest. It has commonly been assumed that the heterochronic process involved is invariably a retardation of the development of adult plumages. On the basis of a phylogenetic analysis of the nine-primaried oscines (Fringillidae), I suggest that the heterochronic process that has operated most frequently in North American taxa is not a retardation, but an acceleration. I also suggest that sexual dimorphism in plumage, and late plumage maturation in males, may arise as a secondary result of selection for neoteny in females with a correlated response in males, and not necessarily, as traditionally thought, as a result of selection for bright plumages in males.     

Evolutionary relationships of Eastern Palearctic Brown Frogs, genus Rana: paraphyly of the 24-chromosome species group and the significance of chromosome number change

The Eurasian 'brown frogs' are a morphologically conservative assemblage consisting of the European Rana temporaria and a large number of similar species considered to be related. Although the chromosome number of the great majority of Rana species is 26, there are several species of brown frogs (including R. arvalis, R. chensinensis, R. omativentris and R. dybowskii ) which instead have 24 chromosomes. Yet comparative study of isozyme variation at 25 gene loci from these and seven other European, Caucasian, and east Asian species indicates that the 24-chromosome brown frogs are not a monophyletic group. To the exclusion of R. arvalis , the east Asian 24-chromosome species form a distinct clade related to the 26-chromosome species R. temporaria, R. dalmatina, R. japonica , and R. tagai more closely than to R. arvalis. This result is reinforced by the comparative position of the nucleolar organizer region (NOR) in the karyotypes of these frogs, as shown by NOR-banding of metaphase chromosomes. The NOR is on chromosome No. 2 in R. arvalis but on chromosome No. 10 in the other three 24-chromosome species. The systematic positions of the Causasian species A. camerani and R. macrocnemis relative to the other species are unclear except that these frogs are distinct enough to be considered part of an outgroup clade. However, in general, the systematics of the entire brown frog group remains unresolved due to great genetic divergences between species and the evident homoplasy of the brown frog morphotype. The parallel origins of reduced karyotypes among the brown frogs demonstrate that caution should be exercised when judging systematic relationships based upon gross karyology in the absence of more detailed information.     

Phylogenetic relationships of the sea eagles (genus Haliaeetus): reconstructions based on morphology, allozymes and mitochondrial DNA sequences

The relationships of eight extant species of sea eagles (genus Haliaeetus ) and some potentially related Accipitrid genera were investigated. A phylogenetic hypothesis is developed based on morphological characters and allozyme data, which is then contrasted with a reconstruction based on 1071 nucleotides of mitochondrial DNA (cytochrome b and part of one tRNA gene). Phylogenies derived from the three data sets are largely congruent, although several aspects are supported by only one kind of evidence (but not contradicted by the others). Only with regard to the position of south-east Asian H. leucogaster and its closely related sister species H. sanfordi was there conflict between allozyme and mtDNA data. The position of H. pelagicus relative to H. leucoryphus or H. albicilla/leucocephalus could not be resolved with the mtDNA sequence data, suggesting that some speciation events occurred in relatively quick succession during the early Pliocene or late Miocene periods. Available evidence suggests that extant sea eagles originated at tropical latitudes, the northernmost species appearing most derived.     

Phylogenetic relationships of the monotypic Peruvian genus Laccopetalum (Ranunculaceae)

Laccopetalum giganteum (Ranunculaceae) is a rare endemic buttercup from the montane regions of the Peruvian Andes. The systematic position of this monotypic genus within Ranunculaceae was investigated using cpDNA matK sequence data. Our findings indicate that L. giganteum forms a highly supported clade together with Krapfia. Several morphological characters are shared by these genera; large subglobose and fleshy flowers, presence of androgynophore with a separated staminal and carpellate region and many tiny achenes. The Laccopetalum-Krapfia clade is sister to the core Ranunculus group with a high bootstrap support. The number of sepals and similar characteristics of achene morphology support an affinity of Laccopetalum with Ranunculus s.s tr.