A sisters’ story: comparative phylogeography and taxonomy of Hierophis viridiflavus and H. gemonensis (Serpentes,Colubridae)

We used a multidisciplinary approach to infer the taxonomy and historical biogeography of Hierophis viridiflavus and H. gemonensis, performing molecular analyses of mitochondrial (16S, Cyt‐b, ND4) and nuclear markers (PRLR), a landmark‐based morphometric study and a cytogenetic analysis. Our data distinguished three main groups in the studied species, corresponding to H. gemonensis and to two monophyletic clades (E and W) within H. viridiflavus. Clades E and W display a significant genetic (about 4% for Cyt‐b and ND4) and morphological divergence and a different morphology of the W sex chromosome (submetacentric in clade E and telocentric in clade W). Taking into account the existing divergence, these clades appear to represent independent phylogenetic units, deserving elevation to species status. Specific names should be H. viridiflavus (Lacépède, 1789) and H. carbonarius (Bonaparte 1833) for clades W and E, respectively. The phylogeography of the studied species is only partially concordant with a general pattern of ‘southern richness and northern purity’ of genetic diversity, whereas H. gemonensis exhibits high genetic diversity at low latitudes (especially in the Peloponnese), H. carbonarius shows a number of different haplotypes both at low (along the southern Italian Apennines and in Sicily) and high latitudes in Italy. Furthermore, a relaxed clock model hypothesizes the differentiation between H. gemonensis and H. viridiflavus sensu lato at about 7 Mya, in the Messinian. Subsequently, the speciation involving H. viridiflavus sensu stricto and H. carbonarius took place in the Quaternary, probably as a result of Pleistocene climatic oscillations. Furthermore, our results are consistent with the existence of several ‘refugia within refugia’ in Italy and in the Balkans and depict the major cladogenesis as allopatric events, mainly driven by paleoclimatic and geographical factors.     

Thermal strategies and energetics in two sympatric colubrid snakes with contrasted exposure

The thermoregulatory strategy of reptiles should be optimal if ecological costs (predation risk and time devoted to thermoregulation) are minimized while physiological benefits (performance efficiency and energy gain) are maximized. However, depending on the exact shape of the cost and benefit curves, different thermoregulatory optima may exist, even between sympatric species. We studied thermoregulation in two coexisting colubrid snakes, the European whipsnake (Hierophis viridiflavus, Lacépède 1789) and the Aesculapian snake (Zamenis longissimus, Laurenti 1768) that diverge markedly in their exposure, but otherwise share major ecological and morphological traits. The exposed species (H. viridiflavus) selected higher body temperatures (~30°C) than the secretive species (Z. longissimus, ~25°C) both in a laboratory thermal gradient and in the field. Moreover, this difference in body temperature was maintained under thermophilic physiological states such as digestion and molting. Physiological and locomotory performances were optimized at higher temperatures in H. viridiflavus compared to Z. longissimus, as predicted by the thermal coadaptation hypothesis. Metabolic and energetic measurements indicated that energy requirements are at least twice higher in H. viridiflavus than in Z. longissimus. The contrasted sets of coadapted traits between H. viridiflavus and Z. longissimus appear to be adaptive correlates of their exposure strategies.     

The thermoregulatory strategy of two sympatric colubrid snakes affects their demography

Population dynamics of terrestrial vertebrates are affected by climatic fluctuations, notably in ectotherms. An understanding of the interaction between physiology and demographic processes is necessary to predict the impacts of climate change. Reptiles are particularly sensitive to temperature, but only a few studies have explored the relationship between thermoregulatory strategy and demography in these animals. Using 12 years of mark-recapture data on two sympatric colubrid snakes (Hierophis viridiflavus and Zamenis longissimus), we tested whether demographic parameters are influenced by contrasted thermoregulatory strategies. The thermophilic and conspicuous species (H. viridiflavus) grew faster than the thermoconforming and secretive species (Z. longissimus), and this difference was most pronounced in open habitats, suggesting that the metabolic benefits associated with high thermal preferences depend on environmental factors at small spatial scales. Survival varied annually in both species, but was not lower in H. viridiflavus despite a higher degree of exposure. In Z. longissimus, survival was negatively affected by low temperatures during the active season, possibly underlying an exposure trade-off.     

Directional asymmetry and correlation of tail injury with left-side dominance occur in Serpentes (Sauropsida)

Recent evidence on directional asymmetry (DA) in species of the paraphyletic assemblage “Lacertilia” indicates a possible biological significance of this phenomenon, and we tested its occurrence in Serpentes by examining bilateral morphological characters, such as the numbers of supralabial and infralabial scutes and the diameter of the eyes, on both sides of museum specimens of seven species: Coronella austriaca, Elaphe longissima, Hierophis viridiflavus, Natrix natrix, Natrix tessellata, Platyceps saharicus, and Vipera aspis. Significant DA (not antisymmetry) occurred in at least one sex of four species and in two characters: H. viridiflavus—the number of infralabial scutes in males; N. tessellata—the diameter of the eyes in males; P. saharicus—the number of infralabial scutes in males and the diameter of the eyes in females; and V. aspis—the number of infralabial scutes in the pooled sexes. In N. tessellata and P. saharicus the DA varied geographically. Rank correlation between the DA of different characters, positive or negative, was significant in one character pair in each of three species. In the largest sample (N. tessellata males from the Levant), tail injury correlated with morphological left-dominant asymmetry (a greater number on the left) of infralabial scutes. Four of ten morphologically left-dominant males, but only 1 of 37 symmetrical or right-dominant males, had an injured tail. The syndrome of morphological DA with functional handicap may be genotypical or phenotypical. Our results support three earlier conclusions: bilateral characters should be examined on both sides of an animal; asymmetry should be analyzed separately in each sex; and the accident proneness of left-handed Homo sapiens merits re-evaluation in view of the apparently wide-spread accident proneness in left-biased Sauropsida (exclusive of Aves).     

Chondrocranial morphology of Leptodactylus larvae (Leptodactylidae: Leptodactylinae): Its utility in phylogenetic reconstruction

Chondrocranial morphology of leptodactylid frogs is scarcely known and has not been completely described for any species of Leptodactylus. We describe the diversity of chondrocranial morphology in the genus Leptodactylus based on the analysis of 22 species, representing the four species groups: the fuscus Group, ocellatus Group, melanonotus Group, and pentadactylus Group. Furthermore, 26 characters are identified and used in a phylogenetic analysis. The phylogenetic analysis using Physalaemus, Crossodactylus, and Hylodes as outgroups suggests two monophyletic clades within Leptodactylus: the melanonotus-ocellatus clade and the pentadactylus-fuscus clade. However, it does not support the monophyly of the species groups as currently recognized and it suggests a paraphyletic Leptodactylus. Enforcing the monophyly of the ingroup, i.e., Leptodactylus, results in the same major two clades of Leptodactylus. Leptodactylus riveroi, a taxon previously unassigned to any species group, appears most closely related to the melanonotus-ocellatus clade based on chondrocranial characteristics. J. Morphol. 238:287–305, 1998. © 1998 Wiley-Liss, Inc.     

The skull of the Upper Cretaceous snake Dinilysia patagonica Smith‐Woodward, 1901, and its phylogenetic position revisited

The cranial anatomy of Dinilysia patagonica, a terrestrial snake from the Upper Cretaceous of Argentina, is redescribed and illustrated, based on high‐resolution X‐ray computed tomography and better preparations made on previously known specimens, including the holotype. Previously unreported characters reinforce the intriguing mosaic nature of the skull of Dinilysia, with a suite of plesiomorphic and apomorphic characters with respect to extant snakes. Newly recognized plesiomorphies are the absence of the medial vertical flange of the nasal, lateral position of the prefrontal, lizard‐like contact between vomer and palatine, floor of the recessus scalae tympani formed by the basioccipital, posterolateral corners of the basisphenoid strongly ventrolaterally projected, and absence of a medial parietal pillar separating the telencephalon and mesencephalon, amongst others. We also reinterpreted the structures forming the otic region of Dinilysia, confirming the presence of a crista circumfenestralis, which represents an important derived ophidian synapomorphy. Both plesiomorphic and apomorphic traits of Dinilysia are treated in detail and illustrated accordingly. Results of a phylogenetic analysis support a basal position of Dinilysia, as the sister‐taxon to all extant snakes. The fossil taxa Yurlunggur, Haasiophis, Eupodophis, Pachyrhachis, and Wonambi appear as derived snakes nested within the extant clade Alethinophidia, as stem‐taxa to the crown‐clade Macrostomata. The hypothesis of a sister‐group relationship between Dinilysia and Najash rionegrina, as suggested by some authors, is rejected by the results of our analysis. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 194–238.     

The Entotympanic of Pangolins and the Phylogeny of the Pholidota (Mammalia)

Entotympanics are independent elements present in the auditory bullae of various eutherians. An entotympanic has been reported for extant pangolins of the Order Pholidota, but the actual distribution of this element remains uncertain, in part, because it is a small, loosely attached structure that is often lost in macerated skulls. Consequently, it is unknown whether or not the entotympanic characterizes Pholidota primitively or has evolved within the group. This report addresses the morphology and distribution of the entotympanic among living and extinct pholidotans. An entotympanic occurs in the African pangolins Manis gigantea, M. temminckii, and in one specimen of M. tricuspis. In each, it is a small, nodular bone that occupies a distinct fossa primarily on the basioccipital, the presence of which allows us to assess the occurrence of an entotympanic even in specimens in which the bone has fallen out. Both the entotympanic and the basioccipital facet are lacking in the four remaining extant pangolin species and in the late Eocene pangolin Patriomanis. To assess the significance of this entotympanic distribution, a phylogenetic analysis of extant pangolins plus Patriomanis based on 67 cranial characters was performed. Four different outgroup analyses all resulted in the same single most parsimonious tree, in which the three extant Asian pangolins form a monophyletic clade and the four extant African pangolins fall into a paraphyletic assemblage. Optimization of the entotympanic distribution onto this tree results in two patterns, dependent on the outgroup choice. If Patriomanis is the sole outgroup to the extant pangolins, the entotympanic arises within pangolins as a synapomorphy of Manis gigantea and M. temminckii, convergently acquired in some M. tricuspis. If Xenarthra and Palaeanodonta are employed as outgroups, the entotympanic optimization is ambiguous: the pattern is either as above or the entotympanic is present primitively within Pholidota and lost secondarily in Patriomanis and a clade comprising M. tricuspis, M. tetradactyla, and the Asian forms.     

Skull osteology of Vipera walser (Squamata,Viperidae): Description,variability, ontogeny,and diagnostic characters in comparison to other Italian vipers

Vipera walser is the most recently recognized European viper. This rare species is endemic to a small area in the Piedmont Alps of Italy, but its closest relatives are found among the Caucasian viper species. In order to provide a starting point for a phylogenetic and biogeographic investigation based on osteology, and including fossils remains, we analyzed four specimens of V. walser and compared them with specimens of the four other Italian viper species. Based on these specimens, we improved the diagnosis of V. walser and provided a first evaluation of intraspecific variability and ontogenetic variation. The skull of V. walser is subject to significant variation, most likely related to ontogeny in some cases (i.e., development of the parietal crest, development of the basioccipital process, shape of the posterior margin of the parabasisphenoid, shape of the quadrate). Based on the studied material, it is possible to distinguish V. walser from the other Italian vipers by the shape of the occipital crest of the supraoccipital, which is posteriorly directed, whereas it is laterally directed in the other species. The osteological diagnosibility provides further support for the validity of V. walser as a distinct species from Vipera berus.     

Phylogeny of the large extinct South American Canids (Mammalia,Carnivora, Canidae) using a “total evidence” approach

South America currently possesses a high diversity of canids, comprising mainly small to medium‐sized omnivorous species, but in the Pleistocene there were large hypercarnivorous taxa that were assigned to Protocyon spp., Theriodictis spp., Canis gezi, Canis nehringi and Canis dirus. These fossils have never been included in phylogenies based on quantitative cladistics, but hand‐constructed cladograms published in the 1980s included some of them in the South American canine clade and others in the Canis clade. In this work, the phylogenetic position of the large extinct South American canids was studied using a large sample of living and extinct canids, as well as different sources of characters (e.g. DNA and 133 osteological characters). The phylogenetic analysis corroborates the inclusion of Theriodictis and Protocyon in the “South American clade”, where C. gezi is also included. In addition, the position of C. dirus as a highly derived Canis species is confirmed. The simultaneous analysis supports hypercarnivory having arisen at least three times in Caninae and once in the “South American clade”. The combination of the phylogenetic analyses, the fossil record and divergence dates estimated in previous works suggests that at least three or four independent lineages of the “South American clade” invaded South America after the establishment of the Panama bridge around 3 million years ago, plus other events corresponding to the immigration of Urocyon and Canis dirus.
© The Willi Hennig Society 2009.     

Geographic phenetic variation of two eastern-Mediterranean non-commensal mouse species, Mus macedonicus and M. cypriacus (Rodentia: Muridae) based on traditional and geometric approaches to morphometrics

We tested the hypothesis that skull shape within the genus Mus may vary with geographic location by assessing the extent and spatial distribution of phenotypic skull variation within and among two wild mouse species, M. macedonicus and M. cypriacus, using traditional and geometric morphometrics including a rather novel application of sliding semilandmarks. Shape was shown to be significantly correlated both with longitude and latitude in M. macedonicus, yet the correlation between morphometric and geographic distances was not significant, and morphometric differences between Asian and European populations were not higher than those within the particular continents. The phylogenetic signal was found to be stronger in dental characters than in cranial ones, however, overall concordance between the pattern of morphometric variation and the presumed history of M. macedonicus was rather weak. In both species, the dorsal and ventral sides of the skull were shown to covary in many aspects though there were also some differences between them, making the functional interpretation of these differences difficult. Discrimination between M. cypriacus and M. macedonicus as well as discrimination between two M. macedonicus subspecies was highly reliable using both traditional and geometric morphometric tools to analyze skull measurements.     

Craniodental characters and the relationships of Procyonidae (Mammalia: Carnivora)

Recent phylogenies of Procyonidae based on molecular data differ significantly from previous morphology‐based phylogenies in all generic sister taxon relationships. I have compiled the most comprehensive dataset of craniodental morphology that incorporates previous morphological characters, and with the aid of high‐resolution X‐ray computed tomography, new characters. This expanded craniodental analysis is based on 78 characters and yields new phylogenetic results regarding the ingroup relationships of Procyonidae. These results include Bassariscus astutus as the least derived member of Procyonidae and Ailurus fulgens nested well within the clade. Additionally, there are some similarities to previous morphological analyses of Procyonidae. Although the characters used to unite and diagnose Procyonidae vary depending on the phylogenetic analysis and have ambiguous taxonomic distribution amongst both Procyonidae and Musteloidea, there is significant morphological support for clades within Procyonidae. In addition to the strength of the morphological support within the clade, the disparate topographical regions of the skull from which the characters are derived may indicate that these synapomorphies are indeed the result of homology rather than adaptive convergence, as suggested by analyses based on molecular data. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 669–713.     

Ontogeny and phyletic size change in living and fossil lemurs

Lemurs are notable for encompassing the range of body‐size variation for all primates past and present—close to four orders of magnitude. Benefiting from the phylogenetic proximity of subfossil lemurs to smaller‐bodied living forms, we employ allometric data from the skull to probe the ontogenetic bases of size differentiation and morphological diversity across these clades. Building upon prior pairwise comparisons between sister taxa, we performed the first clade‐wide analyses of craniomandibular growth allometries in 359 specimens from 10 lemuroids and 176 specimens from 8 indrioids. Ontogenetic trajectories for extant forms were used as a criterion of subtraction to evaluate morphological variation, and putative adaptations among sister taxa. In other words, do species‐level differences in skull form result from the differential extension of common patterns of relative growth? In lemuroids, a pervasive pattern of ontogenetic scaling is observed for facial dimensions in all genera, with three genera also sharing relative growth trajectories for jaw proportions (Lemur, Eulemur, Varecia). Differences in masticatory growth and form characterizing Hapalemur and fossil Pachylemur likely reflect dietary factors. Pervasive ontogenetic scaling characterizes the facial skull in extant Indri, Avahi, and Propithecus, as well as their larger, extinct sister taxa Mesopropithecus and Babakotia. Significant interspecific differences are observed in the allometry of indrioid masticatory proportions, with variation in the mechanical advantage of the jaw adductors and stress‐resisting elements correlated with diet. As the growth series and adult data are largely coincidental in each clade, interspecific variation in facial form may result from selection for body‐size differentiation among sister taxa. Those cases where trajectories are discordant identify potential dietary adaptations linked to variation in masticatory forces during chewing and biting. Although such dissociations highlight selection to uncouple shared ancestral growth patterns, they occur largely via transpositions and retention of primitive size‐shape covariation patterns or relative growth coefficients. Am. J. Primatol. 72:161–172, 2010. © 2009 Wiley‐Liss, Inc.     

Osteology and relationships of Thaiichthys nov. gen.: a Ginglymodi from the Late Jurassic – Early Cretaceous of Thailand

Abstract: The osteology of Thaiichthys buddhabutrensis, nov. gen., from the Late Jurassic – Early Cretaceous of Thailand is described on the basis of a collection of well‐preserved specimens. The mode of preservation of the material allows describing the external anatomy, as well as some elements of the internal anatomy (braincase, elements of the vertebral column). Most of the cranial and postcranial skeleton shows a rather conservative anatomy for ‘semionotiformes’, but the jaw apparatus displays specializations. Variations observed in the ossification pattern of the skull roof and of the cheek, in the morphology of the median dorsal scales and in fin rays’ count indicate that caution should be applied when these characters are used in diagnoses and in phylogenetic analyses. A phylogenetic analysis including a set of gars, of ‘semionotiformes’, of Macrosemiiformes and of Halecomorphi shows the following features: (1) the monophyly of Holostei; (2) sister‐pair relationships between Tlayuamichin/Semiolepis, Isanichthys/’Lepidotes’ latifrons and Araripelepidotes/Pliodetes; (3) the latter pair, together with Thaiichthys and possibly ‘Lepidotes’ mantelli, are resolved as stem Lepisosteiformes; and (4) the ‘semionotiformes’ (a group gathering species of Semionotus and Lepidotes) do not form a clade.     

Cranial anatomy and palaeobiology of the Miocene marsupial Hondalagus altiplanensis and a phylogeny of argyrolagids

New cranial material of Hondalagus altiplanensis, from the middle Miocene of southern Bolivia, allows a rediagnosis of the genus and an assessment of its palaeobiology and phylogenetic relationships with other argyrolagid marsupials. The new specimens demonstrate several derived (synapomorphic) cranial features shared by HondalagusArgyrolagus: a globular braincase, ventrally directed occipital condyles, a broad zygomatic arch, and a short, deep dentary with a flat and long coronoid notch. Hondalagus had powerful masticatory muscles and its cementum-encased hypselodont cheek teeth suggests it had a very abrasive diet. The deep fossae on the lateral aspect of the skull of argyrolagids, interpreted by Simpson as large, laterally-facing orbits, are actually sharply margined temporal fossae. Hondalagus has a very large carotid foramen medially situated within the suture of the basisphenoid and basioccipital. A phylogenetic analysis of five argyrolagid genera was conducted using 32 characters (16 cranial, 16 dental) and a didelphid and a caenolestid as outgroups. Hondalagus-Argyrolagus-Microtragulus form a monophyletic group with an undescribed gen. et sp. nov. (MACN-Ch-1305) from the lower Miocene (Colhuehuapian) of Argentina as its sister taxon. Proargyrolagus appears as sister group to the other taxa of argyrolagids.     

Recognition of a new generic‐level swallow taxon from central Africa

The forest swallow Petrochelidon fuliginosa is a little‐known species endemic to lowland forests in central Africa; for lack of access to high‐quality genetic material, the species has been omitted from all previous molecular phylogenetic studies of the swallows. The species is currently placed in the genus Petrochelidon, within the ‘mud‐nester’ clade of swallows, yet its plumage, morphology, and nesting behavior do not align well with those of other major swallow lineages. As a consequence, upon securing recent specimens and high‐quality tissue samples, we sequenced DNA from two mitochondrial genes and one nuclear marker to place this species in the swallow phylogenetic tree. Our results placed the forest swallow firmly within the ‘mud nester’ clade, but outside of the clade corresponding to Petrochelidon. This outcome led us to document and describe formally a distinct, generic‐level lineage of swallow endemic to the Lower Guinean forest region of central Africa.     

Phylogenetic relationships in genus <Emphasis Type="Italic">Geopora</Emphasis> (Pyronemataceae,Pezizales)

Species delimitation in the genus Geopora (Pyronemataceae) is complicated because of small number of differentiating characters, values of which tend to overlap among the species. Current classification relies mainly on size and shape of ascospores and fruit-bodies, position of the apothecia in the ground and length of excipular hairs. We measured ascospores in ca. 90 Geopora specimens. Sequences of internal transcribed spacer (ITS) rDNA gene were obtained to investigate phylogenetic relationships in the genus. Maximum parsimony (MP) and Bayesian analyses reveal that the well-supported clades detected often do not correspond to the species concepts based on morphological characters. Nine out of the ten lineages include specimens which were initially identified as belonging to different species. The dimensions of ascospores vary to great extent within the lineages. The size and shape of fruit-bodies, length of excipular hair and hymenium colour are mostly homogenous within each clade; however, these characters coincide to a great extent among the lineages and the latter can be assessed only from fresh fruit-bodies. Nuclear DNA content, and accordingly, ploidy level do not provide evidence for species distinction. Geopora arenicola, G. tenuis and G. sepulta were recognized as monophyletic species. Geopora foliacea and G. cervina could not be explicitly delimited and the G. cervina complex comprising three clades was introduced.     

Australian Laurencia majuscula (Rhodophyta,Rhodomelaceae) and the Brazilian Laurencia dendroidea are conspecific

Morphological and molecular studies have been undertaken on two species of the red algal genus Laurencia J.V.Lamouroux: Laurencia majuscula (Harvey) A.H.S. Lucas and Laurencia dendroidea J.Agardh, both from their type localities. The phylogenetic position of these species was inferred by analysis of the chloroplast‐encoded rbcL gene sequences from 24 taxa. In all phylogenetic analyses, the Australian Laurencia majuscula and the Brazilian L. dendroidea formed a well‐supported monophyletic clade within the Laurencia sensu stricto. This clade was divided into two subclades corresponding to each geographical region; however, the genetic divergence between Australian L. majuscula and Brazilian L. dendroidea was only 0–1.35%. Examination of the type specimens and sequences of freshly collected samples of both Laurencia majuscula and L. dendroidea show the two to be conspecific despite their disjunct type localities.     

North American freshwater ancyrocephalids (Monogenea) with articulating haptoral bars: phylogeny reconstruction

A phylogenetic (cladistic) analysis was conducted for 15 ancyrocephalid (Monogenea) species parasitizing North American freshwater fishes. The analysis was based on 10 (four binary and six multistate) characters. Vaginal characters were not included due to insufficient data. The resultant, most parsimonious cladogram was 38 steps long with a consistency index of 0.921: reduced consistency was attributed to two cases of homoplasy and one reversal. Three unresolved polychotomies were observed. Three clades were recognized: clade 1 — Actinocleidus species; clade 2 — Anchoradiscoides, Anchoradiscus, Clavunculus species and Syncleithrium; and clade 3 — Crinicleidus species. The protocol for and implications of character identification and polarisation and the establishment of transformation series were discussed.