Sperm morphology of salamandrids (Amphibia, Urodela): implications for phylogeny and fertilization biology

Mature spermatozoa belonging to four salamander species, Salamandrina terdigitata, Triturus alpestris, Triturus carnifex and Triturus vulgaris, have been investigated by electron microscopy. The sperm ultrastructure of these species was compared with that of previously examined urodeles (36 species and 20 genera) and with that of anurans and caecilians. Many phylogenetic considerations may be inferred as a consequence of comparative spermatology. Urodela appears to be a monophyletic order characterized by three sperm synapomorphies: the acrosomal barb, nuclear ridge and marginal filament. Cryptobranchoidea are confirmed to form a monophyletic suborder having two synapomorphic characters: absence of mitochondria in the tail, and cylindrical shape of the tail axial rod. Within the family Salamandridae, sperm morphology confirms the phylogenetic distance between Salamandrina and Triturus, as already pointed out on the basis of molecular and morphological characters. The very complex ultrastructure of spermatozoa confirms a previous opinion that internal fertilization is the ancestral condition of the Amphibia.     

The cranial morphology and relationships of the aberrant Carboniferous amphibian Spathicephalus mirus Watson

The skull of Spathicephalus mirus Watson, an amphibian from the Namurian, basal Upper Carboniferous, of Scotland is described. It shares with the Loxommatoidea a closed palate, palatal ornament and antorbital vacuities, and the family Spathicephalidae is recognized as the sister group of the Loxommatidae. A new diagnosis of the Loxommatoidea is presented together with one of the Spathicephalidae. An analysis of the functional morphology of the Spathicephalus skull suggests that it was incapable of rapid jaw closure required for catching fish. Instead it is proposed that Spathicephalus was a sluggish bottom-dwelling filter-feeder of small, soft invertebrates.     

The cranial morphology and taxonomy of the saurerpetontid Isodectes obtusus comb. nov. (Amphibia: Temnospondyli) from the Lower Permian of Texas

Five saurerpetontid temnospondyl amphibian specimens from the Lower Permian Arroyo Formation of Baylor County, Texas are redescribed under the binomenIsodectes obtususcomb. nov.The material includes the types ofIsodectes megalopsCope,Trimerorhachis conangulusCope,Acheloma?caseiBroili andEobrachyops townendaeWatson. The Arroyo Formation saurerpetontid is indistinguishable from that from the Middle Pennsylvanian of Linton, Ohio, previously known asSaurerpeton obtusumand the two taxa are synonymized here. The Saurerpetontidae is a monophyletic family comprising the generaIsodectesandAcroplousand defined by reduced strip-like tabulars, dorsal exposure of the palatine excluding lacrimal from orbit margin, parasphenoid with «winged» basal plate, rectangular dermal plates bordering medial margin of mandible, and ilium with elongate dorsal blade. Some of the resemblances described by Watson between «Eobrachyops townendae» and the later brachyopid temnospondyls appear to be artefacts of crushing of an individual specimen, and the relationship between the Saurerpetontidae and Brachyopidae requires reconsideration.     

Revision of<Emphasis Type="Italic">Cheliderpeton vranyi</Emphasis><Emphasis Type="SmallCaps">Fritsch</Emphasis>, 1877 (Amphibia,Temnospondyli) from the Lower Permian of Bohemia (Czech Republic)

The temnospondylCheliderpeton vranyi Fritsch, 1877 from the Lower Permian Ruprechtice horizon (Rotliegend) of the Intrasudetic Basin (Bohemia, Czech Republic) is redescribed. Many features of the skeleton permit a new understanding of the type species and consequently of the genus. Diagnostic characters are the narrow and round-tipped snout, straight to convex outline of the skull roof, narrow and long otic notch, posteriorly expanded quadratojugal, and the relatively wide and short rhombic interclavicle. The ilium with a short, expanded dorsal branch and the missing contact of nasal/maxilla are features shared with the related Upper PermianIntasuchus from Russia and the Eryopidae.Actinodon germanicus is a junior synonym ofCheliderpeton vranyi.      

The morphology and relationships of the Carboniferous amphibian Eoherpeton watsoni Panchen

The oldest recorded anthracosaur amphibian Eoherpeton watsoni Panchen, from localities in the Visean and Namurian of the Scottish Carboniferous, is described. It lacks a distinct otic notch, and reconstruction of the position of cranial ligaments and muscles suggest that a tympanum was never developed. On the basis of these observations it is concluded that arguments used to refute a relationship between anthracosaurs and amniotes, based on apparent differences in the morphology of their otic regions, are invalid. The order Anthracosauria is considered to be a monophyletic group characterized by the development of a sutural connection between the tabular and parietal bones on a skull table which includes both the intertemporal and supratemporal as well as the tabular. It is divided into two suborders: Seymouriamorpha characterized by a sutural connection between the squamosal and intertemporal and the presence of a lateral otic tube, and Anthracosauroideae characterized by the absence of post-temporal fossae. The Discosauriscidae, Kotlassiidae and Seymouriidae are included in the Seymouriamorpha, and the Eoherpetontidae, Gephyrostegidae and Embolomeri comprise the Anthracosauroideae. The Proterogyrinidae are considered the most primitive family of embolomeres. A new phylogeny of Amphibia is presented. The old groupings of Labyrinthodontia, Lepospondyli and Lissamphibia are rejected and Amphibia is divided, on the basis of differences in morphology of the neurocranium and its relationship with the surrounding bones of the skull, into two major groups, the first including the Ichthyostcgalia, Temnospondyli, Microsauria and the extant Anura, Apoda and Urodela, while the Aistopoda, Nectridea, Palaeostegalia, Loxommatoidea, Anthracosauria and Diadectomorpha, together with the Amniota, form the second. Similarities in the dentition of the palaeostegalian Crassigyrinus and loxommatoids suggest the two taxa are sister-groups, but the immediate relationships of anthracosaurs remain indeterminate and they here form an unresolved trichotomy with diadectomorphs and amniotes.     

A new Lower Permian trematopid (Temnospondyli: Dissorophoidea) from Richards Spur,Oklahoma

A new trematopid amphibian, Acheloma dunni, is reported based on excellently preserved cranial and postcranial elements recovered from the Lower Permian fissure fill deposits of the Dolese Brothers Co. limestone quarry near Richards Spur, Oklahoma. The new taxon is characterized by lateral exposures of the palatine (l.e.p.) and ectopterygoid (l.e.e.), which are clearly visible externally and completely enclosed within the suborbital elements. This large, terrestrial carnivore may represent the top predator of the Richards Spur assemblage. A phylogenetic analysis including 12 taxa and 53 cranial characters yielded a single most parsimonious tree, placing Ach. dunni within the monophyletic Trematopidae as the sister taxon to Acheloma cumminsi. Furthermore, the analysis includes the enigmatic Ecolsonia and Actiobates within Trematopidae, forming a clade with the Upper Pennsylvanian Anconastes and the Lower Permian Tambachia. The study comprehensively analyses all valid and aberrant forms of Trematopidae. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 161 , 789–815.     

Miniaturization and its effects on cranial morphology in plethodontid salamanders, genus Thorius (Amphibia: Plethodontidae). I. Osteological variation

Cranial skeletal morphology, ontogeny and variation are examined in five species of Thorius , a genus of diminutive plethodontid salamanders that are among the smallest, extant, tailed tetrapods. The skull of adull Thorius is characterized by: (1) limited development or absence of several ossified elements and dentition; (2) increased inter-and intraspecific variability; (3) novel morphological configurations of the braincase and jaw suspensorium. Posthatching cranial mineralization in all species of Thorius is truncated precociously with respect to that typical of larger and more generalized plethodontid genera, such as Pseudoeurycea. These features implicate paedomorphosis as a predominant mechanism responsible for the evolution of decreased size in Thorius from larger plethodontid ancestors. Interspecific differences in cranial morphology are evident; species may be characterized by greater or lesser degrees of truncated development. However, there is no consistent relationship between degree of paedomorphosis and mean adult body size in interspecific comparisons. Adult morphology of several individual elements represent potentially useful taxonomic characters for distinguishing species.
Reduction, increased variability, and morphological novelty are common to many lineages of dwarfed taxa. They represent a null hypothesis for examination of the developmental mechanisms and morphological consequences of miniaturization in other groups.     

Evolutionary relationships of the lungless caecilian Atretochoana eiselti (Amphibia: Gymnophiona: Typhlonectidae)

Appreciation of the diversity of caecilian amphibians has recently been enhanced by the discovery of a radically divergent aquatic caecilian of the Neotropical Typhlonectidae. Atretochoana eiselti is the largest lungless tetrapod and the only lungless caecilian, and it possesses a suite of remarkable cranial modifications that set it apart from all other caecilians. Numerical phylogenetic analyses, using 141 morphological characters, were performed in order to resolve the evolutionary relationships of Atretochoana and representatives of all other typhlonectid genera. These analyses yield a single most parsimonious tree, (Chthonerpeton (Nectocaecilia (Typhlonectes natans, Typhlonectes compressicauda) (Potomotyphlus, Atretochoana)))) , that is both well resolved and, as judged by Bremer support and by bootstrapping, is well supported. This tree is used as a basis for interpreting ecological shifts and associated morphological evolution within the Typhlonectidae. The available data suggest that the rate of morphological evolution in the Atretochoana lineage is significandy greater than that in other typhlonectid lineages.     

Gnathostome vertebrae and the classification of the Amphibia

Four pairs of arcualia were primitively present in each segment of gnathostomes. The individual vertebral ossifications of early temnospondyls are most economically interpreted as the endochondral ossifications of these cartilaginous arcualia. Centra have formed independently on at least two occasions within the tetrapods and arcualia play little or no part in the formation of true centra in any living form. The so called pleuro- and intercentra of the temnospondyls can in no way be homologized with the centra of either lissamphibians or amniotes. The Nectridea are considered to be the sister group of the Lissamphibia and the Aistopoda the sister group of these two. The anthracosaurs, seymourians and microsaurs are regarded as amniotes. There is no evidence for resegmentation in the vertebral column.     

A Relict Rhinesuchid (Amphibia: Temnospondyli) From The Lower Triassic Of South Africa

' Lydekkerina ' putterilli Broom from the Lystrosaurus Assemblage Zone (Middle Beaufort Group, South Africa) is shown to be a paedomorphic rhinesuchid ( Broomistega putterilli gen. nov.) rather than a lydekkerinid or juvenile stage of the rhinesuchid Uranocentrodon , as previously presumed. The most conspicuous characters, not related to its paedomorphic condition, include the shape of the parietals and the structure of the parasphenoid body and exoccipitals. The pattern of cranial growth changes in Broomistega has been investigated by comparison of the holotype with two hitherto undescribed immature specimens. For several characters referring to the size and position of the orbits, and re-modelling of the jugal proportions, retarded timing of growth transformations, compared to ontogenetic patterns in Permian rhinesuchids (typified by Muchocephalus ) is demonstrated. Some features of the evolution of the basicranial region of the skull, in the transition from rhinesuchids to more advanced capitosauroids, are re-evaluated. key words : temnospondyls, rhinesuchids, relict, paedomorphosis, growth changes.     

Evolution of reproduction in the Rhacophoridae (Amphibia, Anura)

Rhacophorid treefrogs have different reproductive modes: some go through a tadpole stage and some have direct development, and the adults of some species produce foam nests. Philautus is the only genus characterized by direct development. The production of foam nests has been reported in the genera Polypedates, Rhacophorus, Chiromantis and Chirixalus. Recent molecular studies did not provide a robust hypothesis concerning the origin of these reproductive modes in the Rhacophoridae. In order to better understand the evolution of these reproductive modes, we tried to clarify relationships within this group, using DNA sequencing. Our data set consists in a large number of new sequences (1676 base pairs corresponding to threee genes) for five outgroup ranoids and 48 Rhacophoridae, including 16 undescribed species from Sri Lanka and southern India, and all homologous data available in Genbank. After the inclusion of Philautus from India, our data show that the separation of Philautus into clades does not coincide with their geographic distribution. Our data point to the existence of a clade, including the genera Rhacophorus, Polypedates, Chiromantis and Chirixalus, which confirms the results of Wilkinson et al. (2002) and suggests that the ability to produce foam nests has emerged only once in the Rhacophoridae, as already stated by these authors.     

The plagiosaurid temnospondyl Plagiosuchus pustuliferus (Amphibia: Temnospondyli) from the Middle Triassic of Germany: anatomy and functional morphology of the skull

The cranial anatomy of the plagiosaurid temnospondyl Plagiosuchus pustuliferus, from the Middle Triassic of Germany, is described in detail on the basis of a newly discovered skull and mandibular material. The highly derived skull is characterized by huge orbitotemporal fenestrae, a reduction of the circumorbital bones – the prefrontal, postfrontal and (probably) postorbital are lost – and the expansion of the jugal to occupy most of the lateral skull margin. Ventrally the extremely long subtemporal vacuities correlate with the elongate adductor fossa of the mandible. The dentition is feebly developed on both skull and mandible. Ossified ?ceratobranchials and ‘branchial denticles’ indicate the presence of open gills clefts in life. The remarkably divergent cranial morphology of P. pustuliferus highlights the extraordinary cranial diversity within the Plagiosauridae, probably unsurpassed within the Temnospondyli. Specific structural aspects of the skull – including an extremely short marginal tooth row, feeble dentition and an elongated chamber for adductor musculature – together with evidence for a hyobranchial skeleton, suggests that P. pustuliferus utilized directed suction feeding for prey capture. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 348–373.     

Cartilage in the cloaca: Phallodeal spicules in Caecilians (Amphibia: Gymnophiona)

The presence of spicules (also termed spines, teeth, or denticles) on the intromittent organ, a character unique to male members of some species in the African caecilian family Scolecomorphidae, has long been known (Noble, '31; Taylor, '68; Wake,'72; Nussbaum,'85). However, their organization and structure has not been examined. Series of males of three species of Scolecomorphus were examined in order to determine the gross and histological organization of the spicules. Spicule morphology changes with ontogeny within species and differs grossly among species. The cellular organization of the spicules is remarkably similar among species, however. The spicules are composed of chondroid cartilage. The large discoid cells are arranged in “stacks” to form the projections, which emerge from an extensive base plate of the same cartilage. The spicules have a connective tissue sheath, which is not continuous with the cloacal epithelium. The sheath of the spicules may or may not be mineralized; the cartilage itself does not appear to be calcified. Mineralization apparently is not correlated with age, reproductive status, or season. Cartilage in the cloaca of these few species poses a series of questions about the interactions of developmental, structural, functional, and phylogenetic properties in the evolution of the reproductive biology of caecilians. J. Morphol. 237:177–186, 1998. © 1998 Wiley-Liss, Inc.     

Adaptive radiation in miniature: the minute salamanders of the Mexican highlands (Amphibia: Plethodontidae: Thorius)

The small size and apparent external morphological similarity of the minute salamanders of the genus Thorius have long hindered evolutionary studies of the group. We estimate gene and species trees within the genus using mitochondrial and nuclear DNA from nearly all named and many candidate species and find three main clades. We use this phylogenetic hypothesis to examine patterns of morphological evolution and species coexistence across central and southern Mexico and to test alternative hypotheses of lineage divergence with and without ecomorphological divergence. Sympatric species differ in body size more than expected after accounting for phylogenetic relationship, and morphological traits show no significant phylogenetic signal. Sympatric species tend to differ in a combination of body size, presence or absence of maxillary teeth, and relative limb or tail length, even when they are close relatives. Sister species of Thorius tend to occupy climatically similar environments, which suggests that divergence across climatic gradients does not drive species formation in the genus. Rather than being an example of cryptic species formation, Thorius more closely resembles an adaptive radiation, with ecomorphological divergence that is bounded by organism‐level constraints. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 622–643.     

The head morphology of Clambidae and its implications for the phylogeny of Scirtoidea (Coleoptera: Polyphaga)

External and internal structures of the head of adults of Clambus are described and illustrated in detail. The results are compared with structural features found in the clambid genus Calyptomerus, in representatives of other scirtoid families, and also in species of other coleopteran suborders, notably Myxophaga. The results tentatively support the monophyly of Scirtoidea and a close relationship between Clambidae and Eucinetidae is suggested by one shared derived feature of the mandible, a long and slender apical tooth with a serrate edge. The monophyly of Clambidae is very strongly supported and Acalyptomerus is probably the sistergroup of a clade Calyptomerus + Clambinae. Potential scirtoid autapomorphies are the loss of the dorsal tentorial arms, a bulging gula, a strongly transverse labrum, and a ridge separating the mediostipes from the lacinia. However, all these features are homoplasious. The monophyly of Clambidae is supported by modifications of the head capsule which is strongly flattened and broadened, by a deep clypeofrontal incision enabling vertical antennal movements, and a series of antennal features. Synapomorphies of Clambinae + Calyptomerus (Clambidae excluding Acalyptomerus) are the conglobate body form with the ventral side of the head capsule in contact with the mesocoxae, and compound eyes integrated in the contour of the head. The completely subdivided eye is an autapomorphy of Clambus. An entire series of features is shared by Clambidae (or Scirtoidea) and Myxophaga. Most of them are apomorphies that apparently evolved independently in both groups. However, the presence of well‐developed maxillary and labial glands is arguably a retained groundplan feature of Coleoptera, with parallel loss in Archostemata, Adephaga and various groups of Polyphaga. J. Morphol. 277:615–633, 2016. © 2016 Wiley Periodicals, Inc.     

Spermatozoa of Pseudinae (Amphibia, Anura, Hylidae), with a test of the hypothesis that sperm ultrastructure correlates with reproductive modes in anurans

We describe, for the first time, the sperm ultrastructure of the two genera of Pseudinae. Based on sperm ultrastructure, the five species herein examined can be separated into three groups: one containing Pseudis paradoxa, P. bolbodactyla, and P. tocantins, the second containing P. minuta, and the third containing Lysapsus laevis. The midpiece is similar in all species and auxiliary fibers and the undulating membrane are absent. In Pseudis a subacrosomal cone and a multilaminar structure (P. minuta) or a granular material (P. paradoxa group) are seen above the nucleus. Lysapsus laevis has only remnants of the subacrosomal cone. All species have peripheral fibers associated with the outer doublets of the axoneme. We tested the hypothesis of correlation between the presence of an undulating membrane and fertilization environments in anurans using a concentrated changes test (CCT) based on the Hay et al. (Mol Biol Evol 1995;12:928-937) hypothesis of phylogenetic relationships among anuran families. Only a subset of the resolved topologies derived from the Hay et al. (1995) cladogram, where Ranoidea is the sister-group of Sooglossidae, produced significant probabilities of the CCT. Therefore, support for the correlation between sperm ultrastructure and fertilization environments in anurans is, at best, equivocal.