Evidence for a pre-PETM dispersal of the earliest European crocodyloids

Crocodyloid remains from the late Paleocene of Mont de Berru (France) hosted in the collections of the Muséum National d’Histoire Naturelle (Paris, France) and in the Institut royal des Sciences naturelles de Belgique (Brussels, Belgium) are described for the first time. This material, although fragmentary, can be clearly referred on a morphological basis to Asiatosuchus depressifrons (Blainville, 1855), a species previously reported from several Eocene Belgian localities thanks to abundant material including a nearly complete skeleton. The Paleocene material shares with A. depressifrons the number of alveoli involved in the dentary symphysis, the exclusion of the splenials from the symphysis, and the presence of a distinct depression on the jugal. The fossil remains from Berru represent the oldest European crocodyloid. Along with the alligatoroid Diplocynodon remensis Martin, Smith, de Lapparent de Broin, Escuillié and Delfino, 2014, previously reported from the same locality, the crocodyloid A. depressifrons indicates that these genera reached Europe before the Paleocene–Eocene Thermal Maximum. Although more complete remains from outside Europe are needed to refine phylogenetic hypotheses, according to the currently established fossil record the forerunners of diplocynodontids likely dispersed from North America, whereas those related to Asiatosuchus likely dispersed from Asia.     

An assemblage of early Oligocene lizards (Squamata) from the locality of Boutersem (Belgium), with comments on the Eocene–Oligocene transition

A diverse, new lizard assemblage from the early Oligocene of Belgium is described. The Boutersem railway local fauna is the most species‐rich lizard assemblage yet reported from the European early Oligocene. Four lizard taxa are present: Lacertidae, Anguidae, Scincoidea and Platynota. One new species is described, Folisaurus boutersemensis sp. nov . This fauna provides new insight into the profound turnover that took place during the Eocene/Oligocene boundary in Europe. The new fauna confirms a marked decrease in diversity across the Eocene/Oligocene boundary. Two groups encountered in the European late Eocene became extinct (Iguanidae*, Glyptosaurinae). Estimates of species‐level extinctions range up to 80%. These estimates include members of virtually all the families present in the late Eocene. The relative importance of climate change and biotic interactions in controlling this pattern is discussed, and negative interactions between lizards and new carnivorous mammals are favoured. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 148–170.     

Revival of the genus Tropicoperdix Blyth 1859 (Phasianidae,Aves) using multilocus sequence data

Although Tropicoperdix has been considered to be either a full genus or a species complex within the Phasianid genus Arborophila (hill partridges), there is long‐standing uncertainty regarding the degree of difference that warrants generic separation, including reported anatomical cranial differences. In addition, the intra‐generic taxonomy remains under dispute. Most studies hypothesize that Tropicoperdix comprises three species, while others postulate from one to four species. However, no molecular study has been performed to clarify the systematic and taxonomic uncertainties surrounding Tropicoperdix. In the present study, we performed a series of molecular phylogenetic analyses of Tropicoperdix and Arborophila taxa based on two mitochondrial genes and five nuclear introns. All the results are consistent with the finding that Tropicoperdix and Arborophila are phylogenetically distinct and distant genera, although the precise phylogenetic position of Tropicoperdix remains undetermined. Retrospective examination of external characteristics also supports the generic separation, as well as providing evidence of remarkable multiple character convergence. We propose that Tropicoperdix comprises at least two full species based on mitochondrial data obtained from museum specimens by using a next‐generation sequencing method. © 2015 The Linnean Society of London     

Crocodylian assemblage from the middle Eocene Ikovo locality (Lugansk Province,Ukraine), with a discussion of the fossil record and geographic origins of crocodyliform fauna in the Paleogene of Europe

The known fossil record of crocodyliforms in Europe during the Paleogene is significantly biased, in that the fauna of Western Europe is far better sampled and understood compared to that of Eastern Europe. We describe in detail all known crocodyliform remains from the middle Eocene (Lutetian) Ikovo locality in Ukraine. We conclude that at least two taxa were present: a moderate to large-sized Tomistominae indet. similar to the basalmost known tomistomines, and the small-sized basal alligatoroid cf. Diplocynodon sp. Despite its scarcity, this is the first basal alligatoroid material reported from Eastern Europe (as part of post-Soviet countries) and the easternmost record of diplocynodontines in Europe so far. An allegedly freshwater cf. Diplocynodon sp. contributes a rare faunal element to the vertebrate assemblage of the Ikovo locality, otherwise dominated by resident or facultative marine taxa. The fossil record and historical paleobiogeography of crocodyliforms from the Paleocene and Eocene of Europe are reviewed. As it has been already known, the middle Eocene fauna of crocodyliforms proves to be taxonomically diverse and complex. Its constituent lineages geographically originated in Asia or North America (Diplocynodontinae, Asiatosuchus-like crocodyloids, Planocraniidae), North America (derived alligatorines), Africa (Tomistominae), and Gondwana (ziphodont mesoeucrocodylians Iberosuchus and Bergisuchus), with possible subsequent speciation in Europe. We propose a novel hypothesis of Asian origins of European diplocynodontines, which will be explicitly tested in future studies. The revealed similarities between crocodylians and turtles from the Ikovo locality and those from Western Europe support the presence of a single Pan-European biogeographical zone during the middle Eocene, distinct from that of Asia.     

Earliest South American paucituberculatans and their significance in the understanding of ‘pseudodiprotodont’ marsupial radiations

We describe the oldest Paucituberculata marsupials, from the La Barda and Las Flores localities (Argentina; Late Palaeocene, and Early–Middle Eocene), as well as from the Itaboraí Basin (Brazil; Late Palaeocene). The new taxa are represented by very scarce, although well‐preserved, dental remains. A parsimony analysis was performed in order to evaluate the phylogenetic affinities of these taxa. Representatives of both Riolestes capricornicus gen. et sp. nov. and Bardalestes hunco gen. et sp. nov. appear to be basal paucituberculatans, and their molar features give clues on the early evolution of the representatives of this order. Within the Paucituberculata we recognize two major clades: Caenolestoidea and Palaeothentoidea. We conclude that ‘pseudodiprotodont’ marsupials of the traditional literature (i.e. Polydolopimorphia + Paucituberculata) do not form a natural group. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 867–884.     

Resurrection in Carapa (Meliaceae): a reassessment of morphological variation and species boundaries using multivariate methods in a phylogenetic context

The taxonomy of the amphi‐Atlantic tree genus Carapa (Meliaceae) has long been controversial. Of the three species currently recognized in the genus, two are known to present substantial morphological variation that has been used in the past to distinguish several taxa, most of which are currently placed in synonymy. Here, a combination of field observations, univariate analyses of leaf, floral and seed characters and principal coordinate analyses of floral characters in the context of a molecular phylogenetic analysis was used to investigate the patterns of variation and delimit morphological species anew in the genus. These results support the recognition of 27 species in Carapa, of which 16 are previously described and 11 are new. In general, phylogenetically related species occurred in the same geographical area, but were morphologically distinct. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 165 , 186–221.     

Checklist of Commelinaceae of Equatorial Guinea (Annobón,Bioko and Río Muni)

A checklist of Commelinaceae of Equatorial Guinea, comprising 46 taxa in 12 genera, is presented. The best represented genus is Palisota, with 11 species. Bibliographical references for Commelinaceae from Equatorial Guinea have been gathered and checked. Eleven species of Commelinaceae are recorded for the first time in the country. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 106–122.     

Provannid and provannid‐like gastropods from the Late Cretaceous cold seeps of Hokkaido (Japan) and the fossil record of the Provannidae (Gastropoda: Abyssochrysoidea)

A fauna of provannid and provannid‐like shells is described from Upper Cretaceous seep carbonates in Hokkaido, Japan. We describe two new provannid species, Provanna tappuensis sp. nov. and Desbruyeresia kanajirisawensis sp. nov. , with preserved protoconchs of unquestionable provannid type with decollate apex. This material confirms the occurrence of Provannidae as early as the Middle Cenomanian. We also describe Hokkaidoconcha gen. nov. and a new family Hokkaidoconchidae fam. nov. , with two named species, H. hikidai sp. nov. and H. tanabei sp. nov . Hokkaidoconchidae are possibly related to the Provannidae, judging from a similar, but not decollate larval shell, although the juvenile teleoconch whorls differ in being of a general cerithimorph appearance and the details of the aperture are unknown. Furthermore, we review the published fossil record of Provannidae and Abyssochrysidae, and we consider that in those older than the Eocene, there is no evidence preserved that unequivocally supports a position there. The Jurassic Acanthostrophia acanthica from Italy seems to be the oldest known record of Abyssochrysidae, and the most reliable occurrence of the family, older than from the Miocene. Other fossil, pre‐Miocene species that have been classified in the Abyssochryssidae are provisionally referred to Hokkaidoconchidae. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154 , 421–436.     

Molecular appraisal of Bunium and some related arid and subarid geophilic Apiaceae–Apioideae taxa of the Ancient Mediterranean

Bunium is unusual in Apiaceae in having a variable cotyledon number and broad infrageneric dysploidy. To test the monophyly of the genus, phylogenetic relationships among 39 Bunium species were investigated with DNA sequence data from nuclear (nrITS) and plastid (psbA‐trnH intergenic spacer) regions. Several other taxa with a similar ecology and geography were also included in the analyses. Our results suggest that Bunium is not monophyletic. Bunium spp. in the eastern part of the study area are more closely related to the Central Asian genera Elaeosticta, Galagania, Hyalolaena, Mogoltavia and Oedibasis than to those in the western part. Our study revealed that molecular, morphological (cotyledon number and width of fruit commissure) and karyological data reveal similar patterns in the taxa studied. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 160 , 149–170.     

Variation in nectar–sugar profile of Anchusa and allied genera (Boraginaceae)

The nectar–sugar profile (fructose, glucose and sucrose) of 14 species of Anchusa and five members of the allied genera Anchusella, Cynoglottis, Hormuzakia and Lycopsis (Boraginaceae: tribe Boragineae) was determined. Most of the species examined (c. 74%) produce sucrose‐dominant nectar, whereas the remaining taxa produce sucrose‐rich nectars. Little variation in nectar–sugar composition was found in some species, even when sampling was repeated in different years and/or localities. Average sucrose concentration was 57.75% (coefficient of variation 19.1%). The only floral morphological character that was correlated with the nectar–sugar profile is the length of the corolla tube, as taxa with relatively long floral tubes produce nectar with lower glucose concentrations. The flowering period is also related to sugar composition, as nectar of late‐flowering species contains lower sucrose concentrations. However, small differences in sugar profiles do not reflect phylogenetic relationships based on molecular studies. It would appear that dry habitats and time of flowering are the main determinants of nectar–sugar composition in the genus Anchusa sensu lato. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162 , 616–627.     

New generic proposal for the European Neogene large testudinids (Cryptodira) and the first phylogenetic hypothesis for the medium and large representatives of the European Cenozoic record

Fossil testudinids are known in Europe since the Eocene, with several taxa of medium size (from more than 0.3 m to less than 0.7 m) recognized in the Palaeogene record, most of them being poorly known. The size of several European Neogene taxa was larger (between 1 and 2 m). These large testudinids were relatively abundant and diverse, ranging from the early Miocene to the Pleistocene. However, there is a nomenclatural gap at the generic level for the Neogene forms, as their generally used assignment to the more primitive Eocene Cheirogaster cannot be sustained. This is because relatively little material has been assigned to the described species, and also because of the absence of a detailed study comparing all of the European taxa. Here, the European Cenozoic taxa are incorporated for the first time in a data matrix, so that a hypothesis on their phylogenetic relationships is justified. This study identified the large testudinids from the Neogene of Europe as belonging to a monophyletic clade, assigned to the new genus T itanochelon . The hitherto poorly understood ‘Testudo’ bolivari, proposed nearly a century ago but lacking diagnosis, is analysed in detail. It is recognized as the best‐represented large testudinid from the European record, and is identified as the type species of T itanochelon gen. nov. Its comparison with the other Neogene species allowed a detailed study of the new genus and an analysis of its phylogenetic relationships with the other European taxa. © 2014 The Linnean Society of London     

Molecular phylogenetics of Uvaria (Annonaceae): relationships with Balonga,Dasoclema and Australian species of Melodorum

An extended molecular phylogenetic analysis of Uvaria (Annonaceae) is presented, using maximum parsimony, maximum likelihood and Bayesian methods, based on sequences of four plastid DNA regions (matK, psbA‐trnH spacer, rbcL and trnL‐F). The additional taxa include the monotypic West African genus Balonga, the monotypic South‐East Asian genus Dasoclema and seven Australian representatives of the genus Melodorum. The results indicate that all of these taxa are nested within a well‐supported clade otherwise consisting of Uvaria species, indicating that their taxonomic treatment needs to be reassessed. The distinguishing morphological characteristics of the taxa are re‐evaluated and interpreted as specialized adaptations of the basic Uvaria structure. The genus Uvaria is accordingly extended following the transfer of these species, necessitating six new nomenclatural combinations and two replacement names. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163 , 33–43.     

A new Transantarctic relationship: morphological evidence for a Rheidae–Dromaiidae–Casuariidae clade (Aves,Palaeognathae, Ratitae)

Although ratites have been studied in considerable detail, avian systematists have been unable to reach a consensus regarding their relationships. Morphological studies indicate a basal split separating Apterygidae from all other extant ratites, and a sister‐group relationship between Rheidae and Struthionidae. Molecular studies have provided evidence for the paraphyly of the Struthionidae and Rheidae, with respect to a clade of Australasian extant ratites. The position of the extinct Dinornithidae and Aepyornithidae also remains hotly debated. A novel pattern of diversification of ratites is presented herein. The phylogenetic analysis is based on 17 taxa and 129 morphological characters, including 77 new characters. The resultant tree yields a sister‐group relationship between New Zealand ratites (Apterygidae plus Dinornithidae) and all other ratites. Within this clade, the Aepyornithidae and Struthionidae are successive sister taxa to a new, strongly supported clade comprising the Rheidae, Dromaiidae, and Casuariidae. The link between South American and Australian biotas proposed here is congruent with numerous studies that have evidenced closely related taxa on opposite sides of the Southern Pacific. These repeated patterns of area relationships agree with current knowledge on Gondwana break‐up, which indicates that Australia and South America remained in contact across Antarctica until the earliest Tertiary. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156 , 641–663.     

Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores,including one new genus and three new species

The Hippasterinae is a subfamily within the Goniasteridae, consisting of five genera and 26 species, which occur in cold‐water settings ranging from subtidal to abyssal depths. All known genera were included in a cladistic analysis resulting in two most parsimonious trees, supporting the Hippasterinae as monophyletic. Our review supports Sthenaster emmae gen. et sp. nov. as a new genus and species from the tropical Atlantic and two new Evoplosoma species, Evoplosoma claguei sp. nov. and Evoplosoma voratus sp. nov. from seamounts in the North Pacific. Hippasteria caribaea is reassigned to the genus Gilbertaster, which previously contained a single Pacific species. Our analysis supports Evoplosoma as a derived deep water lineage relative to its continental‐shelf, shallow water sister taxa. The genus Hippasteria contains approximately 15 widely distributed, but similar‐looking species, which occur in the northern and southern hemispheres. Except for Gilbertaster, at least one species in each genus has been observed or is inferred to prey on deep‐sea corals, suggesting that this lineage is important to the conservation of deep‐sea coral habitats. The Hippasterinae shares several morphological similarities with Circeaster and Calliaster, suggesting that they may be related. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 266–301.     

The origin of a mega‐diverse genus: dating Begonia (Begoniaceae) using alternative datasets,calibrations and relaxed clock methods

Begonia is a mega‐diverse genus comprising c. 1500 species of herbs, shrubs and epiphytes with a near pantropical distribution. Previous date estimates for the most recent common ancestor of Begonia have placed the evolution of this genus into a broad temporal context, but the issue of an absolute date estimate remains open. In this study, we attempt to estimate absolute DNA divergence dates for Begonia and, in doing so, address some of many the factors that can affect such estimates. The largest source of variance in our estimates was because of uncertainty with the calibration constraints and phylogenetic distance between these constraints and Begonia. Another large source of variance was due to the alternative methods of analysis investigated. Less variance was as a result of the alternative DNA datasets and combinations of calibration constraints assessed. Our date estimates suggest that the most recent common ancestor of Begonia could have diversified from the end of the Cretaceous to the beginning of the Neogene, probably during a period of global cooling from the mid Eocene to early Oligocene. These estimates imply that the near pantropical distribution of extant Begonia was generated by intercontinental dispersal after the ancient inferred break up of the supercontinent, Gondwana. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 363–380.     

New information for Ilex phylogenetics based on the plastid psbA‐trnH intergenic spacer (Aquifoliaceae)

The aim of the present work was to clarify the origin and phylogenetic position of the species belonging to the genus Ilex (Aquifoliaceae), especially the South American species. Phylogenetic relationships of the genus Ilex were investigated using the plastid psbA‐trnH intergenic spacer and parsimony and Bayesian analyses. The psbA‐trnH intergenic spacer was shown to evolve slowly within Ilex, but a major gap present in this region was useful in the phylogenetic study of the genus. To obtain more potentially parsimonious characters, atpB‐rbcL intergenic spacer data were combined with those for psbA‐trnH. Many gaps present in the psbA‐trnH region were useful in the phylogenetic study of the genus Ilex. The topology of the trees showed that, in general, the clades are strongly related to geographical areas, a fact especially evident in certain different Asian lineages. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 182–193.     

Diverse labellar secretions in African Bulbophyllum (Orchidaceae: Bulbophyllinae) sections Ptiloglossum,Oreonastes and Megaclinium

Floral food‐rewards of Bulbophyllum range from nectar to protein‐rich mucilage and lipid‐rich labellar secretions. For the first time, the structure of the labellum and the secretory process are investigated for four African Bulbophyllum species. The most specialized type of labellar organization occurred in B. schinzianum, the deep, narrow, median longitudinal groove consisting of palisade‐like secretory cells flanked by trichomes containing lipid droplets, and the copious secretion containing sugar. This groove was absent or poorly defined, shallow and wide, in the remaining taxa, the scant secretion containing lipid. All taxa possessed a striate cuticle lacking cracks and pores, and micro‐channels were present, cuticular blisters occurring only in B. schinzianum. The labellum contained storage parenchyma (B. lupulinum) or mesophyll‐like parenchyma (B. schinzianum), but in section Megaclinium (B. falcatum and B. maximum), these were replaced by aerenchyma. In B. schinzianum, the form of the labellar groove, sweet fragrance and sugary secretion suggest pollination by Hymenoptera, the food‐reward and fragrance indicating that pseudocopulation is unlikely. Conversely, the form of the labellum of taxa having smaller flowers, and the lipid‐rich secretion, suggests pollination by small flies. The labellar aerenchyma may facilitate this process or even aid wind‐assisted pollination. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 266–287.     

Floral biology and mechanisms of spontaneous self‐pollination in five neotropical species of Gentianaceae

Cross‐ and self‐fertilization in angiosperms are regulated by several factors, and a knowledge of the mechanism and time of spontaneous self‐pollination offers opportunities for a better understanding of the evolution of mating systems and floral traits. The floral biology of five species of Gentianaceae found in high‐altitude neotropical grassland is presented, with emphasis on the mechanisms that promote spontaneous self‐pollination. A presumed floral Batesian mimicry system is suggested between the rare and rewardless Zygostigma australe and Calydorea campestris, a species of Iridaceae with pollen‐flowers, pollinated by syrphids and bees. The floral morphology of the other four gentian species points to three different pollination syndromes: melittophily, phalaenophily and ornithophily. However, with the exception of the nocturnal Helia oblongifolia, flowers are nectarless and appear to exhibit non‐model deceptive mechanisms, providing similar floral cues to some sympatric rewarding species with the same syndrome. The similar mechanism of spontaneous self‐pollination in Calolisianthus pedunculatus, Calolisianthus pendulus and H. oblongifolia (Helieae) is based on the stigmatic movements towards the anthers. Selfing is promoted by movements of the style/stigma and of the corolla in Deianira nervosa and Z. australe (Chironieae), respectively. The movements of stamens, style and stigma during anthesis seem to be the most common method of spontaneous self‐pollination in angiosperms. It is suggested that the evolution of delayed spontaneous self‐pollination would be more expected in those taxa with dichogamous flowers associated with herkogamy. Such a characteristic is frequent in long‐lived flowers of certain groups of Asteridae, which comprise most documented cases of autonomous selfing. Thus, the presence of dichogamy associated with herkogamy (which supposedly evolved as a result of selection to promote both separation of male and female functions and the efficient transfer of cross pollen) may be the first step in the adaptive evolution of delayed selfing to provide reproductive assurance. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 160 , 357–368.