Elasmobranchs from the Lower Triassic Sulphur Mountain Formation near Wapiti Lake (BC, Canada)

Hybodontoid and nonhybodontoid sharks are described from the Lower Triassic Vega‐Phroso Siltstone Member of Sulphur Mountain Formation on the basis of newly discovered material. The age of the classic fossil site ‘Wapiti Lake’ in the Canadian Rocky Mountains is discussed on the basis of new field data and one conodont found in association. Preliminary results suggest that these elasmobranch remains are between early Smithian and Spathian in age. Apart from the enigmatic genus Listracanthus and previously reported edestoids, the shark fauna consists of at least one hybodont, at least two questionable hybodontoid genera and an elasmobranch of enigmatic affinities, represented by peculiar denticles only and described as ‘genus A’incertae sedis. The presence of the only previously reported hybodont genus, cf. Palaeobates, is erroneous. The largest specimen represents the most complete Early Mesozoic shark known. The heterodonty of its dentition, fin spine morphology and the short, robust body shape imply it represents a member of a new family of shark, Wapitiodidae fam. nov. , and is described here as Wapitiodus aplopagus gen. et sp. nov. The unique dental morphology shows affinities to Polyacrodus but clearly differs in the complete lack of side cusps. Wapitiodus gen. nov. possesses a primitive fin spine structure. The tooth crowns are entirely blunt in the distal (posterior) tooth files, and are acuminate‐unicuspid in several anterior files. Tooth morphology, the shape of the basal cartilages, the proximal insertion of the fin spines and the pectoral fin structure are interpreted as diagnostic characters for this new genus, and possibly for the Wapitiodidae fam. nov. The majority of observed characters appear to be primitive and are reminiscent of Palaeozoic sharks, however, and these features include dorsal fin spine morphology and gross skull anatomy. A second species, provisionally placed in the same genus, is described as Wapitiodus homalorhizo sp. nov. Wapitiodus homalorhizo sp. nov. can be distinguished from W. aplopagus gen. et sp. nov. by the proportions of the fin spines, tooth morphology and possibly the body shape. Several isolated teeth and other fragmentary material are referred to either Wapitiodus gen. nov. sp. indet. or to ?Polyacrodus sp. (Polyacrodontidae gen. et sp. indet.). A third genus of elasmobranch (incertae sedis) is described as ‘Genus A’ and is recognized by its peculiar scales. © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 149 , 309–337.     

Systematics and phylogeny of Vasseuromys (Mammalia,Rodentia, Gliridae) with a description of a new species from the late Miocene of eastern Europe

Vasseuromys is a species‐rich genus of small‐ to medium‐sized glirids spanning the latest Oligocene to late Miocene of Europe and western Asia. Despite extensive discoveries over the past 50 years, little phylogenetic work has been done on Vasseuromys. This study presents the first phylogenetic analysis of the genus that includes all the described species and a new taxon Vasseuromys tectus sp. nov. from the late Miocene of eastern Europe, providing the first insights into the evolutionary relationships within the clade. Results suggest that the genus is clearly paraphyletic. Two strongly supported genus‐level clades are recognized within ‘Vasseuromys’: a restricted Vasseuromys clade (containing the three species, V. pannonicus, V. rugosus and V. tectus) and the Glirulus clade that includes ‘Vasseuromys’ duplex. The remaining ‘Vasseuromys’ species are found to constitute a set of paraphyletic taxa, with the polyphyletic ‘Ramys’ nested within it. The genus Gliruloides is synonymized with Glirulus. Vasseuromys tectus sp. nov. is the most derived member of the genus in having a greater number of cheek teeth ridges including constantly present anterotrope, centrotrope, second prototrope on M1–2, third metatrope on M2, two to three posterotropids on p4 and strong ectolophids on lower molars. The results of the study confirm a European origin for Vasseuromys while suggesting that the late Miocene species of the genus dispersed from the east in the early Turolian.     

Evolutionary trends on extant cat skull morphology (Carnivora: Felidae): a three‐dimensional geometrical approach

The skulls of 33 extant cat species were characterized through three‐dimensional geometric morphometrics using 20 landmarks. A principal component analysis (PCA) was performed with Procrustes fitted coordinates, and the PC‐scores were phylogenetically corrected by independent contrasts method. Three PCs allowed for the definition of five cat skull patterns. PC1: ‘snouted/massive‐headed cats’ (genus Panthera) opposing the ‘round‐headed small cats’ (genus Oncifelis, Prionailurus rubiginosus, Prionailurus bengalensis, among other small cats); PC2: ‘tapering‐headed cats’ (Neofelis nebulosa, Herpailurus yagouaroundi, Prionailurus planiceps) opposing the ‘stout‐headed cats’ (Acinonyx jubatus, Uncia uncia, Otocolobus manul, Felis margarita, and Felis nigripes); and PC3: ‘low profiled‐headed cats’ (mostly, Pr. planiceps). A sixth pattern, the ‘generalized skull’, observed in the Caracal lineage, genus Lynx, Leopardus pardalis, and Catopuma temminckii, indicates a morphological convergence among midsized‐cats. The morphological trends ‘snouted/massive’ and ‘round’ clearly denote a co‐evolution between size and shape. The other skull patterns evolved unrelatedly to the size (i.e. their allometric variations are not a size function). Nevertheless, each species comprises an amalgam of these patterns, so the influence of the size permeates, in some extent, the skull morphology along all cat lineages. The felid ecomorphological fit to hypercarnivory is obvious; however, different skull shapes in same‐sized species with similar habits, indicate that the variation in the skull morphology may result from phenotypic fluctuations, whose adaptive value (if indeed there is any) is still obscure. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 176–190.     

Sciadopitys cladodes from Eocene Baltic amber

The Baltic amber deposit represents the largest accumulation of any fossil resin worldwide and hundreds of thousands of entrapped arthropods have been recovered so far. The source plants of Baltic amber, however, are still controversial, and the botanical composition of the ‘Baltic amber forest’ remains poorly studied. Here, we provide the first unequivocal Baltic amber inclusions of the umbrella pine Sciadopitys (Sciadopityaceae), a genus that has been suggested as the source of succinite (the main variety of Baltic amber) based on chemical analyses. As previously suggested sciadopitoid inclusions must be reconsidered as being notional, representing angiosperm leaves instead, the new fossils are the first unambiguous macrofossil evidence of Sciadopitys from the ‘Baltic amber forest’, and the first pre‐Oligocene macrofossil record of Sciadopitys from Europe. The fossil Sciadopitys cladodes provide new insights into the conifer diversity of the ‘Baltic amber forest’ and broaden the picture of its palaeoecology, indicating the presence of humid swamp to raised bog habitats. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 180 , 258–268.     

Multiple nuclear and mitochondrial DNA sequences provide new insights into the phylogeny of South African Lacertids (Lacertidae,Eremiadinae)

Eremiadinae, one of three subfamilies of Lacertidae, are distributed throughout Asia and Africa. Previous phylogenetic studies suggested that one of the main groups of Eremiadinae (the Ethiopian clade) consist of two clades with predominately East‐African and South‐African distribution. Yet, especially the latter one, which includes the genera Pedioplanis, Meroles, Ichnotropis, Tropidosaura and Australolacerta, was not well supported in the molecular phylogenetic analysis. In this study, we analysed the phylogenetic relationships among the genera of the ‘South African clade’ to assess whether this group actually forms a highly supported clade and to address questions concerning the monophyly of the genera. We sequenced sections of the widely used mitochondrial genes coding for 16S rRNA, 12S rRNA and cytochrome b (altogether 2045 bp) as well as the nuclear genes c‐mos, RAG‐1, PRLR, KIF24, EXPH5 and RAG‐2 (altogether 4473 bp). The combined data set increased the support values for several nodes considerably. Yet, the relationships among five major lineages within the ‘South African clade’ are not clearly resolved even with this large data set. We interpret this as a ‘hard polytomy’ due to fast radiation within the South African lacertids. The combined tree based on nine marker genes provides strong support for the ‘South African Clade’ and its sister group relationship with the ‘East African Clade’. Our results confirm the genus Tropidosaura as a monophylum, while Ichnotropis is paraphyletic in our trees: Ichnotropis squamulosa appears more closely related to Meroles than to Ichnotropis capensis. Furthermore, the monophyly of Meroles is questionable as well. Based on our results, I. squamulosa should be transferred from Ichnotropis into the genus Meroles. Also, the two species of Australolacerta (A. australis and A. rupicola) are very distantly related and the genus is perhaps paraphyletic, too. Finally we propose a phylogeographical scenario in the context of palaeoclimatic data and compare it with a previously postulated hypothesis.     

Redescription of the phytosaurs Paleorhinus (‘Francosuchus’) angustifrons and Ebrachosuchus neukami from Germany,with implications for Late Triassic biochronology

Phytosaurs are a diverse and morphologically distinctive clade of superficially crocodile‐like archosauriforms that had a near global distribution during the Late Triassic. Because their remains are among the most abundant vertebrate remains recovered in many Upper Triassic terrestrial formations, phytosaurs are used extensively in long‐range biochronological and biostratigraphic correlations. The biochronologically oldest and earliest branching known phytosaurs include an array of nominal species from the early Late Triassic of the United States, Germany, Poland, Morocco, and India that have been synonymized within the genus Paleorhinus, and subsequently used to define a global ‘Paleorhinus biochron’. However, recent phylogenetic work suggested that the North American species previously referred to Paleorhinus are paraphyletic. Here, we reassess the systematics and anatomy of putative specimens of Paleorhinus from southern Germany. Two well‐preserved basal phytosaur skulls from the Blasensandstein (Carnian) of Bavaria form the holotypes of Francosuchus angustifrons and Ebrachosuchus neukami, both of which were synonymized with Paleorhinus by previous workers. We demonstrate that Francosuchus angustifrons shares unique synapomorphies with specimens referred to Paleorhinus bransoni from the Late Triassic of Texas, and thus refer the species to Paleorhinus. By contrast, the longirostrine Ebrachosuchus is highly distinctive in morphology, and our new cladistic analysis of Phytosauria demonstrates that it represents a valid taxon that is more closely related to Phytosauridae than to Paleorhinus. We provide the first autapomorphy‐based support for a monophyletic but restricted Paleorhinus (supported by a nodal row on the jugal, and low paired ridges on the squamosal) and confirm that previous broader conceptions of Paleorhinus are likely to be paraphyletic. © 2013 The Linnean Society of London     

Phylogenetic relationships in the genus Leontopodium (Asteraceae: Gnaphalieae) based on AFLP data

The genus Leontopodium comprises 30–41 species. The centre of diversity is the Sino‐Himalayan region in south‐western China, where about 15 species occur. The two species native to Europe, L. alpinum (known as the common ‘Edelweiss’) and L. nivale, are part of the cultural heritage of the people living there. Despite its importance, very little is known about the systematics of the genus. Because recent molecular studies have shown that species within this genus are closely related and difficult to distinguish with rDNA and cpDNA data, we used AFLPs to obtain a more detailed understanding of the phylogeny of the genus. Our main aims were as follows: (1) to clarify species relationships within the genus; and (2) to reveal information about the biogeography of the genus. We used AFLPs with six primer combinations to investigate 216 individuals in 38 populations of 16 different species. With AFLPs, we were able to recognize 10 different groups, all of which had strong bootstrap support. These results were also congruent with the morphology‐based taxonomy of the genus. Most private and rare fragments were found in the Yunnan region (south‐western China) relative to Europe and Mongolia/central China, suggesting a long‐lasting in situ history of populations in the centre of diversity of the genus. Our results illustrate the utility of AFLPs to resolve phylogenetic relationships between these closely related species. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 165 , 364–377.     

Molecular dating and diversification of the South American lizard genus Liolaemus (subgenus Eulaemus) based on nuclear and mitochondrial DNA sequences

The temperate South American lizard genus Liolaemus is the one of the most widely distributed and species‐rich genera of lizards on earth. The genus is divided into two subgenera, Liolaemus sensu stricto (the ‘Chilean group’) and Eulaemus (the ‘Argentino group’), a division that is supported by recent molecular and morphological data. Owing to a lack of reliable fossil data, previous studies have been forced to use either global molecular clocks, a standardized mutation rate adopted from previous studies, or the use of geological events as calibration points. However, simulations indicate that these types of assumptions may result in less accurate estimates of divergence times when clock‐like models or mutation rates are violated. We used a multilocus data set combined with a newly described fossil to provide the first calibrated phylogeny for the crown groups of the clade Eulaemus, and derive new fossil‐calibrated substitution rates (with error) of both nuclear and mtDNA gene regions for Eulaemus specifically. Divergence date estimates for each of the crown groups and appropriate rate estimates will provide the foundation for understanding rates of speciation, historical biogeography, and phylogeographical history for various clades in one of the most diverse lizard genera in the poorly studied Patagonian region. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 825–835.     

Phylogenetic relationships and historical biogeography of tribes and genera in the subfamily Nymphalinae (Lepidoptera: Nymphalidae)

We infer for the first time the phylogenetic relationships of genera and tribes in the ecologically and evolutionarily well‐studied subfamily Nymphalinae using DNA sequence data from three genes: 1450 bp of cytochrome oxidase subunit I (COI) (in the mitochondrial genome), 1077 bp of elongation factor 1‐alpha (EF1‐α) and 400–403 bp of wingless (both in the nuclear genome). We explore the influence of each gene region on the support given to each node of the most parsimonious tree derived from a combined analysis of all three genes using Partitioned Bremer Support. We also explore the influence of assuming equal weights for all characters in the combined analysis by investigating the stability of clades to different transition/transversion weighting schemes. We find many strongly supported and stable clades in the Nymphalinae. We are also able to identify ‘rogue’ taxa whose positions are weakly supported (the different gene regions are in conflict with each other) and unstable. Our main conclusions are: (1) the tribe Coeini as currently constituted is untenable, and Smyrna, Colobura and Tigridia are part of Nymphalini; (2) ‘Kallimini’ is paraphyletic with regard to Melitaeini and should be split into three tribes: Kallimini s.s., Junoniini and Victorinini; (3) Junoniini, Victorinini, Melitaeini and the newly circumscribed Nymphalini are strongly supported monophyletic groups, and (4) Precis and Junonia are not synonymous or even sister groups. The species Junonia coenia, a model system in developmental biology, clearly belongs in the genus Junonia. A dispersal‐vicariance analysis suggests that dispersal has had a major effect on the distributions of extant species, and three biotic regions are identified as being centres of diversification of three major clades: the Palaearctic for the Nymphalis‐group, the Afrotropics for Junoniini and the Nearctic for Melitaeini. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 86 , 227–251.     

Phylogenetic analysis of Micrathena and Chaetacis spiders (Araneae: Araneidae) reveals multiple origins of extreme sexual size dimorphism and long abdominal spines

The phylogenetic relationships amongst the New World spiny orb‐weaving spiders Micrathena and Chaetacis were assessed through parsimony and Bayesian analyses of morphological characters. A total of 146 characters was scored for ten outgroup taxa and 37 Micrathena and four Chaetacis species. The results indicate that Chaetacis nests within Micrathena and we propose Chaetacis as a junior synonym of Micrathena. Twelve subgeneric species groups of Micrathena are recognized and diagnosed. Species with extremely long spines evolved at least eight times in the genus and we suggest that this may be related to antipredator defences. Micrathena is primitively sexually monomorphic and extreme sexual size dimorphism has arisen at least six times in the genus. Most of these events are because of enlargement of the female in relation to the ancestral size, although in two cases sexual dimorphism was attained through male reduction, adding more data to the ‘giant females’ vs. ‘dwarf males’ controversy. The genus is probably of South American origin and has repeatedly invaded Central and North America. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166 , 14–53.     

Phylogenetic analysis of higher-level relationships of Odonata

Abstract. This is the most comprehensive analysis of higher‐level relationships in Odonata conducted thus far. The analysis was based on a detailed study of the skeletal morphology and wing venation of adults, complemented with a few larval characters, resulting in 122 phylogenetically informative characters. Eighty‐five genera from forty‐five currently recognized families and subfamilies were examined. In most cases, several species were chosen to serve as exemplars for a given genus. The seven fossil outgroup taxa included were exemplar genera from five successively more distant odonatoid orders and suborders: Tarsophlebiidae (the closest sister group of Odonata, previously placed as a family within ‘Anisozygoptera’), Archizygoptera, Protanisoptera, Protodonata and Geroptera. Parsimony analysis of the data, in which characters were treated both under equal weights and implied weighting, produced cladograms that were highly congruent, and in spite of considerable homoplasy in the odonate data, many groupings in the most parsimonious cladograms were well supported in all analyses, as indicated by Bremer support. The analyses supported the monophyly of both Anisoptera and Zygoptera, contrary to the well known hypothesis of zygopteran paraphyly. Within Zygoptera, two large sister clades were indicated, one comprised of the classical (Selysian) Calopterygoidea, except that Amphipterygidae, which have traditionally been placed as a calopterygoid family, nested within the other large zygopteran clade comprised of Fraser's ‘Lestinoidea’ plus ‘Coenagrionoidea’ (both of which were shown to be paraphyletic as currently defined). Philoganga alone appeared as the sister group to the rest of the Zygoptera in unweighted cladograms, whereas Philoganga + Diphlebia comprised the sister group to the remaining Zygoptera in all weighted cladograms. ‘Anisozygoptera’ was confirmed as a paraphyletic assemblage that forms a ‘grade’ towards the true Anisoptera, with Epiophlebia as the most basal taxon. Within Anisoptera, Petaluridae appeared as the sister group to other dragonflies.     

Reassessment of the Lower Cretaceous (Barremian) pliosauroid Leptocleidus superstes Andrews, 1922 and other plesiosaur remains from the nonmarine Wealden succession of southern England

The Lower Cretaceous (upper Berriasian to lowermost Aptian) nonmarine Wealden succession of southern England has been a prolific source of vertebrate fossils for over 180 years. The sequence is most famous for terrestrial reptiles, particularly dinosaurs; however, significant aquatic tetrapod discoveries including rare nonmarine plesiosaurs have also been reported. The record of Wealden plesiosaurs currently incorporates a single valid taxon, Leptocleidus superstes Andrews, 1922a, based on a partial skeleton and skull from the Barremian Upper Weald Clay Formation of Berwick, Sussex. Traditional classifications place this plesiomorphic pliosauroid with basal Jurassic rhomaleosaurids; however, the genus Leptocleidus has since become a ‘waste basket’ for various Cretaceous rhomaleosaurid‐like plesiosaurs from around the globe. In an attempt to clarify this situation, the type specimen of L. superstes was reexamined and redescribed. Previously unrecorded anatomical features were identified including an elongate, gracile paraoccipital process on the exoccipital‐opisthotic, and tooth ornament comprising widely spaced, coarse striations that are restricted to the lingual surface of the crown (mesiodistal ‘carinae’ are absent). Other indeterminate pliosauroid remains (recovered along with coeval elasmosaurids) from the upper Berriasian–Valanginian Hastings Beds Group also exhibit potentially diagnostic traits: an atlas centrum with no anterolateral exposure and with ventral margin formed by the intercentrum; a single‐headed rib articulation on the atlas centrum extending onto the axis centrum; and epipodials that are longer than broad. The placement of L. superstes is controversial in recent phylogenies. To test the competing hypotheses, L. superstes together with all closely related species were rescored into the most comprehensive published phylogenetic data sets of Plesiosauria and Pliosauroidea. Separate maximum parsimony and Bayesian analyses of each matrix unanimously supported a relationship between L. superstes and pliosauroids but could not confirm placement within either Rhomaleosauridae sensu stricto, or a discrete ‘leptocleidoid’ clade. Examination of character states advocating affinities amongst Leptocleidus spp. suggests homoplasy rather than clear homology between what are potentially palaeobiogeographically disparate genus‐level taxa. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 161 , 663–691.     

Genets (Carnivora, Viverridae) in Africa: an evolutionary synthesis based on cytochrome b sequences and morphological characters

The taxonomy of the genets (genus Genetta) has long been discussed, thus hampering endeavours towards evolutionary reconstruction. Sequence data from the complete cytochrome b gene (cyt b) were generated for 50 specimens representing 15 morphological species in order to allow the production of the first exhaustive molecular phylogeny of the genets. Second, a revised morphological matrix comprising 50 characters was combined with the cyt b data to estimate the level of morphological homoplasy. Phylogenetic analyses were conducted using parsimony, maximum likelihood and Bayesian procedures. Our results based on cyt b contradict a part of the traditional taxonomy of genus Genetta, the servaline and small‐spotted genets being paraphyletic, but confirmed the species status recently re‐investigated for three genets belonging to the large‐spotted complex, including the newly described G. bourloni. The combined analysis yielded similar results although morphological characters were clearly homoplasic. Partitioned Bremer supports indicated conflicting signals between the two data sets throughout the tree, and species‐diagnostic characters, useful for delimiting species boundaries, were significantly correlated to habitat. However, morphological data supported the monophyly of clades (G. victoriae, other genets) (G. servalina, G. cristata), large‐spotted genet complex and forest forms. Our results suggest a complex evolutionary history of the genets in Africa, with a Poiana‐like ancestor inhabiting rain forest, and then a diversification involving two independent invasions of open habitats and one reversion to rain forest. Divergence estimates based on cyt b revealed that splitting events within genets partly follow a climatic speciation model during the cyclical periods of the Quaternary, although ‘primitive’ rain forest lineages diverged earlier, during the Late Miocene and Early Pliocene. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 81 , 589–610.     

First Potwarmus from the Miocene of Saudi Arabia and the early phylogeny of murines (Rodentia: Muroidea)

Potwarmus is recorded for the first time in Saudi Arabia. The material comes from the Middle Miocene of the Hofuf Formation at Al‐Jadidah. This species, Potwarmus flynni sp. nov. , is described and compared with all species currently placed in the genera Potwarmus, Dakkamys, Paradakkamys, Vallaris, and some species of ‘Myocricetodon’ (i.e. the so‐called ‘primitive dendromurids’) as well as with two of the most primitive definitive murines (namely Antemus chinjiensis and Progonomys debruijni). Potwarmus flynni sp. nov. differs from the above‐mentioned species in having the anterior part of the m1 strongly reduced. A cladistic analysis provides evidence that this new Arabian taxon is close to Potwarmus primitivus and Potwarmus sp. nov. from Jebel Zelten (Libya), and also that this genus is unlikely to be the sister taxon to Antemus. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156 , 664–679.     

Evolutionary morphology of whip spiders: towards a phylogenetic system (Chelicerata: Arachnida: Amblypygi)*

The aim of this study is to present a cladogram and phylogenetic system and to use this to discuss the phylogeny and biogeography of the Amblypygi. A total of 29 morphological structures were studied, their plesiomorphic and apomorphic characters or character states were identified, and the resulting data matrix was analysed. As a result, the ‘old’Charontidae or Pulvillata emerge as a paraphyletic group; the genus Paracharon is the sister group of all other amblypygids, which are now termed Euamblypygi. The ‘new’Charontidae (sensu Quintero: the genera Stygophrynus and Charon) are the sister group of the Phrynida or Apulvillata; together they form the Neoamblypygi. The relationships of the genera of the Charinidae cannot be resolved with the available data. They may be a paraphyletic group. The genus Catageus is a possible candidate for being the sister group of the Neoamblypygi. The new system allows a discussion of the phylogeny and biogeography of whip spiders. It also points to unresolved taxa and thus indicates the questions future research should address.     

Congeneric phylogeography: hypothesizing species limits and evolutionary processes in Patagonian lizards of the Liolaemus boulengeri group (Squamata: Liolaemini)

In poorly known groups for which data are insufficient to develop biologically plausible model‐based approaches to phylogeographical analyses, a ‘first hypotheses’ protocol is suggested as offering the best way to generate hypotheses for subsequent model‐based tests. Preliminary hypotheses are formulated about species boundaries and population processes in three species complexes of the Liolaemus boulengeri group in the context of mtDNA ‘congeneric phylogeography’. The temperate South American Liolaemus provides a model with ancient and recent allopatric divergence across ecologically and geologically complex landscapes, incipient speciation, secondary contact, and discordance between molecular and morphological patterns of variation. Moderately dense sampling of widely distributed ‘inertial’ species in the Patagonian Steppe has revealed hidden genetic and probably species diversity, and also hinted at demographic and historical processes that may have shaped the histories of these taxa. Five of the seven focal species of the present study were paraphyletic for mtDNA genealogies, suggesting that they represent complexes of species, and nested clade phylogeographical analysis (NCPA) analyses suggest that different historical and demographic processes have shaped the observed patterns. Introgression and incomplete lineage sorting are hypothesized as being the cause of some of the observed paraphyly. Provisional delimitations of species are proposed and NCPA is used to generate hypotheses of population history, all of which are subject to further testing. Multi‐faceted studies, involving phylogenetic assessments of independent molecular markers and morphological variation across codistributed taxa with estimates of niche breadths in a landscape context, will likely yield the most promising returns for cross‐validation of hypotheses of population and speciation histories. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society, 2006, 89 , 241–275.     

Phylogeny of the platygastroid wasps (Hymenoptera) based on sequences from the 18S rRNA, 28S rRNA and cytochrome oxidase I genes: implications for the evolution of the ovipositor system and host relationships

The Platygastroidea are a diverse group of mostly small to tiny wasps, the common biology for which is endoparasitism of insect and spider eggs. No analytically‐based phylogeny exists for the superfamily, and the current suprageneric classification is flawed in part because of its reliance on homoplasious and pleisiomorphic morphological characters. To determine platygastroid relationships as a basis for investigating host and ovipositor evolution, phylogenies of > 70 in‐group species (representing 55 genera) were reconstructed by parsimony and Bayesian methods using three molecular markers; the mitochondrial cytochrome oxidase I and the nuclear genes 28S and 18S rRNA. The results strongly support the monophyly of the superfamily and one of the two families, Platygastridae, but the Scelionidae are most likely polyphyletic. However, within the Scelionidae, there is a well supported ‘main scelionid clade’ that contains the majority of genera assigned to the family. At the subfamilial level, both putative subfamilies of Platygastridae, the Platygastrinae, and Sceliotrachelinae, are likely to be polyphyletic. Within the Scelionidae, both the Teleasinae and Telenominae are monophyletic, but the Scelioninae is clearly not so. The current tribal classification for the Scelionidae is in need of major reassessment because no tribes, with the exception of the Scelionini s.s., were found to be monophyletic. Further illustrating the problems associated with the current classification is the nonmonophyly of a number of genera, namely Opisthacantha Caloteleia, Telenomus, Trimorus, Teleas and Idris. Analysis of ovipositor evolution in the superfamily revealed that the Ceratobaeus‐type ovipositor system is ancestral; however, this trait was lost prior to the evolution of the main scelionid clade, for which the Scelio‐type ovipositor system is ancestral and defines a mostly paraphyletic assemblage. Ancestral state analysis indicates that the Ceratobaeus‐type ovipositor was subsequently re‐evolved in the main scelionid clade, representing a possible contradiction of Dollo’s law. Previously, the tribal placement has been used to predict the host associations of genera for which host data were unavailable. However, the fact that most tribes are not monophyletic throws into doubt any such speculation based on the current classification. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 91 , 653–669.     

Blastogeny in tabulate corals: case studies using X‐ray microtomography

X‐ray microtomography (XMT) is a non‐invasive and non‐destructive method that has often been used to study fossils. It allows serial sections to be made as little as few micrometers apart; such a resolution is unachievable for classical serial sectioning; moreover, in contrast to the latter, the specimen is not destroyed. Microtomography can, however, be applied only in cases where differences in X‐ray absorption between the skeleton and its infilling are great. We show that this method may be also applied to tabulate corals. Case studies of blastogeny are based on Silurian (Aulopora, Favosites) and Devonian (Thamnopora) specimens from Poland. We show that the sequence of events in the blastogeny of Aulopora sp. is different from that of ‘Aulopora serpens minor’ from the Devonian of the Holy Cross Mountains and similar to auloporids from the Devonian of England. Blastogeny in Favosites is very similar to that known from the related genera Squameofavosites and Thamnopora. This suggests that members of the genus Aulopora may be more diversified within the genus (as presently understood) than genera within the Favositidae.     

The comparative osteology and phylogenetic relationships of African and South American lungfishes (Sarcopterygii: Dipnoi)

Lepidosirenidae is a clade of freshwater lungfishes that include the extant South American Lepidosiren paradoxa Fitzinger, 1837 and African species of the genus Protopterus. These genera have been geographically separated since the break‐up of Gondwana in the Early Cretaceous, but they display similar biology and morphology. Species were distinguished by a combination of features such as head‐to‐body ratios, the number of pairs of ribs, and the presence of external gills, but no discrete skeletal characters were identified, and no comparative studies including all extant species have been published. I used computed tomography (CT), X‐ray photography, and specimens from museum collections to describe the skeletal morphology of all species of lepidosirenid in a comparative context. I digitally disarticulated the bones in each specimen to compile a comparative atlas of the cranial and pectoral elements of all extant lungfishes, which has the potential to increase the correct identifications of specimens in museum collections. The morphology of the frontoparietal, parasphenoid, supraorbital, and suboperculum differ between species. I used those characters, along with molecular sequence data from the ribosomal RNA gene 16S, to run combined morphological and molecular phylogenetic analyses. Lepidosirenidae is monophyletic in all analyses, but the interrelationships of the species of Protopterus vary with the different sources of character data. © 2015 The Linnean Society of London     

Phylogeny and evolution of the South African genus Metalasia (Asteraceae–Gnaphalieae) inferred from molecular and morphological data

Metalasia is a genus in tribe Gnaphalieae (Asteraceae), endemic to South Africa and with its main distribution in the Cape Floristic Region. The genus comprises 57 species and, with a number of closely related genera, it constitutes the ‘Metalasia clade’. A species‐level phylogenetic analysis is presented, based on DNA sequences from two nuclear (internal and external transcribed spacer: ITS, ETS) and two plastid (psbA‐trnH, trnL‐trnF) regions together with morphological data. Analyses combining molecular and morphological data attempt not only to resolve species interrelationships, but also to detect patterns in character evolution. Phylogenetic analyses corroborate our earlier study and demonstrate that Metalasia is formed of two equally sized, well‐supported sister groups, one of which is characterized by papillose cypselas. The results differ greatly from earlier hypotheses based on morphology alone, as few morphological characters support the phylogenetic patterns obtained. The two clades of Metalasia do, however, appear to differ in distribution, corresponding to the different rainfall regimes of South Africa. Analyses show a few taxa to be problematic; one example is the widely distributed M. densa which appears to be an intricate species complex. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 174 , 173–198.