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     

A study of Sarcocornia A.J. Scott (Chenopodiaceae) from Western Mediterranean Europe

The latest publications on Sarcocornia taxonomy and phylogeny recognize six taxa in this genus on the Iberian Peninsula: S. perennis, S. fruticosa, S. alpini, S. alpini subsp. carinata, S. hispanica, and S. pruinosa. The present study represents a comprehensive revision of the different taxa in the Sarcocornia genus present in Western Mediterranean Europe by means of morphological, micromorphological and phylogenetic internal transcribed spacer (ITS) analysis. Morphological and micromorphological data were studied from Sarcocornia samples from 113 populations in coastal salt marshes and inland salt pans in Portugal, France, Spain and Italy. Sixteen new ITS sequences were obtained from Mediterranean Sarcocornia species and analysed together with previous reported data. Published karyological, ecological and biogeographical data from Western Mediterranean Europe were also reviewed. The results indicate the presence of a new species, S. lagascae, found growing in coastal Mediterranean areas of the Iberian Peninsula. The species S. fruticosa was found to be absent from the Iberian territories.     

Biogeographical research on European species of the lichen genus Physconia

Abstract Aim The (world) distribution of the European species of Physconia Poelt (Lichenes: Physciaceae) [except Ph. distorta (With.) J. R. Laundon and Ph. americana Essl.] was investigated. The study was complemented by data on the ecological behaviour of the species. Location The location of the study is the whole world with a local emphasis on Europe. Methods The geographical distribution of the lichen species under consideration was investigated from collection data of herbaria and field studies in a databank. Additional data were taken from existing literature and critically considered. Distribution maps were created with ArcView GIS. The distribution patterns are expressed as three‐dimensional areal formulas, regarding zonal distribution, altitudinal range and oceanity of the species distributions, using the method of Meusel et al. (1965 ). Results The distribution patterns of the ten Physconia taxa considered are represented in detailed maps and expressed as three‐dimensional areal formulas. The ecological behaviour was discussed. Physconia muscigena and Ph. perisidiosa are species of bipolar distribution type, the other species are holarctic types. Physconia detersa and Ph. enteroxantha occur in both Eurasia and North America, the remaining species (Ph.grisea ssp. grisea, Ph. grisea ssp. lilacina, Ph. petraea, Ph. servitii, Ph. subpulverulenta and Ph.venusta) are pure Old world taxa with a much more limited distribution.The majority of the European Physconia taxa are distributed meridionally to submeridionally and occur preferentially in suboceanic areas. Main conclusions Some of the species under consideration have a wide distribution, nevertheless they show clear affinities to ecological conditions that influence their distribution patterns. A number of species show not only zonal and regional restrictions, but also affinities to the western climatic type. The Mediterranean is the centre of diversity of the genus Physconia in Europe.     

Comparative morphology of pond,stream and phytotelm‐dwelling tadpoles of the South American Redbelly Toads (Anura: Bufonidae: Melanophryniscus)

We present a comprehensive review of larval morphology in the Neotropical toad genus Melanophryniscus. The taxa studied included 23 species with representatives of recognized phenetic groups and different larval ecomorphological guilds: pond, stream, and phytotelm‐dwelling tadpoles. Their external morphology variation is congruent with current phenetic arrangement based on adult features, but also reflects the habitat where larvae develop. Lotic tadpoles (i.e. M. tumifrons group and M. krauczuki) in general exhibit a more depressed body, a longer tail with lower fins, and larger oral discs than lentic forms (i.e. M. stelzneri group, M. moreirae, M. sanmartini, and M. langonei). Despite their peculiar, confined microhabitat, phytotelm larvae do not diverge markedly from non‐arboreal species. The distinctive features of all species are the presence of a pineal end organ and the placement of the intestinal reversal point at the left of the abdomen in typical larval stages. The buccal cavity and musculoskeletal anatomy are quite conserved between species, yet some characteristics differ from those of other bufonids. The presence of one pair of subhyoid muscles is apparently an exclusive trait of Melanophryniscus among Bufonidae. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 112 , 417–441.     

Dispersal,fragmentation, and isolation shape the phylogeography of the European lineages of Polyommatus (Agrodiaetus) ripartii (Lepidoptera: Lycaenidae)

Polyommatus ripartii is a biogeographically and taxonomically poorly understood species of butterfly with a scattered distribution in Europe. Recently, it has been shown that this species includes several European endemic and localized taxa (galloi, exuberans, agenjoi) that were previously considered species and even protected, a result that poses further questions about the processes that led to its current distribution. We analysed mitochondrial DNA and the morphology of P. ripartii specimens to study the phylogeography of European populations. Three genetically differentiated but apparently synmorphic lineages occur in Europe that could be considered evolutionarily significant units for conservation. Their strongly fragmented and counterintuitive distribution seems to be the result of multiple range expansions and contractions along Pleistocene climatic oscillations. Remarkably, based on the 79 specimens studied, these genetic lineages do not seem to extensively coexist in the distributional mosaic, a phenomenon most evident in the Iberian Peninsula. One of the important gaps in the European distribution of P. ripartii is reduced by the discovery of new Croatian populations, which also facilitate a better understanding of the biogeography of the species. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 817‐829.     

Molecular phylogeny and taxonomy of the genus Chaetophora (Chlorophyceae,Chlorophyta), including descriptions of Chaetophoropsis aershanensis gen. et sp. nov.

The broadly defined genus Chaetophora consisted of species with minute, uniseriate branching filaments enveloped in soft or firm mucilage forming macroscopic growths that are spherical, hemispherical, and tubercular or arbuscular, growing epiphytically on freshwater aquatic plants and other submerged surfaces in standing or fast‐flowing water. Recent molecular analyses clearly showed that this genus was polyphyletic. In this study, eight strains of Chaetophora and three strains of Stigeoclonium were identified and successfully cultured. In combination with the morphological data, a concatenated data set of four markers (18S + 5.8S + ITS2+ partial 28S rDNA) was also used to determine their taxonomic relationships and phylogenetic positions. The molecular analysis resolved the broadly defined Chaetophora to at least two genera. Species with a globose thallus of genus Chaetophora formed a separate monophyletic clade, which clearly separated from, a type of lobe‐form Chaetophora species. Therefore, we propose to erect a new genus, Chaetophoropsis, which includes all globose species of the Chaetophora. Chaetophoropsis aershanensis was determined to be a new species, based on its special characteristic of profuse long rhizoids. Stigeoclonium polyrhizum, as the closest relative to Chaetophoropsis, revealed its distant relationships to other species of Stigeoclonium. A globose thallus with a thick, soft mucilage matrix, and special rhizoidal branches lent further support to the placement of S. polyrhizum in the genus Chaetophoropsis and had the closest relationship to C. aershanensis. Taxonomic diversity was proven by distinctive morphological differences and by phylogenetic divergence in the broadly defined Chaetophora identified herein.     

Phylogeographic analyses reveal distinct lineages of the liverworts Metzgeria furcata (L.) Dumort. and Metzgeria conjugata Lindb. (Metzgeriaceae) in Europe and North America

Seed plant genera often exhibit intercontinental disjunctions where different species are found on different continents. Many morphologically circumscribed bryophyte species exhibit similar disjunctions. We used nucleotide sequences from the plastid and nuclear genomes to test hypotheses of phylogeography within representatives of the genus Metzgeria: Metzgeria furcata, Metzgeria conjugata, and Metzgeria myriopoda. The first two species have sexual and asexual populations, exhibit disjunctions between North America and Europe, and have been split into separate species, numerous subspecies or varieties. The third species occurs in eastern North America but is not reported from Europe. Phylogenetic analyses resolved three distinct lineages within the morphologically defined species, M. furcata: one in North America, and two in Europe. Similarly, three morphologically cryptic clades of M. conjugata were resolved by the molecular data: northern North America, Europe, and south‐eastern North America. For both species, molecular divergence among taxa occurred in the absence of morphological change. In the case of M. myriopoda, all plants from eastern North America were both morphologically uniform and genetically homogeneous (although not identical). The present study provides significant insight into a plant group with complex taxonomy, and indicates that these liverwort taxa with wide distributions, extreme sex ratios, and continental disjunctions harbor cryptic lineages. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98 , 745–756.     

Phylogenetics and morphological evolution of coral‐dwelling barnacles (Balanomorpha: Pyrgomatidae)

Pyrgomatid barnacles are a family of balanomorphs uniquely adapted to symbiosis on corals. The evolution of the coral‐dwelling barnacles is explored using a multi‐gene phylogeny (COI, 16S, 12S, 18S, and H3) and phenotypic trait‐mapping. We found that the hydrocoral associate Wanella should be excluded, while some archaeobalanids in the genus Armatobalanus should be included in the Pyrgomatidae. Three well supported clades were recovered: clade I is the largest group and is exclusively Indo‐West Pacific, clade II contains two plesiomorphic Indo‐West Pacific genera, while clade III is comprised of East and West Atlantic taxa. Some genera did not form reciprocally monophyletic groups, while the genus Trevathana was found to be paraphyletic and to include members of three other apomorphic genera/tribes. The highly unusual coral‐parasitic hoekiines appear to be of recent origin and rapidly evolving from Trevathana sensu lato. Pyrgomatids include six‐, four‐, and one‐plated forms, and exhibit convergent evolutionary tendencies towards skeletal reduction and fusion, loss of cirral armature, and increased host specificity. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 162–179.     

Phylogenetics and evolution of phyllotaxy in the Solomon's seal genus Polygonatum (Asparagaceae: Polygonateae)

Polygonatum is the largest and most complex genus in tribe Polygonateae, comprising approximately 57 species widely distributed in the warm temperate, subtropical and boreal zones of the Northern Hemisphere. However, phylogenetic relationships in the genus remain poorly understood. The objectives of this study were to reconstruct the phylogenetic relationships of the genus using four plastid markers, and to examine the evolution of leaf arrangement in Polygonatum in the phylogenetic context of its closely related taxa. Thirty Polygonatum species were sampled to infer phylogenetic relationships using maximum‐likelihood and Bayesian analyses. The evolution of leaf arrangements was reconstructed using Bayesian, parsimony and likelihood methods. The phylogenetic analyses supported the current generic delimitation of Polygonatum, with Heteropolygonatum recognized as a distinct genus. Three major lineages in Polygonatum were well supported, largely correlated with geographical distribution and the most recent classification at the sectional level. However, our results did not support the currently recognized series, especially the two large series Verticillata and Alternifolia. Bayesian analyses support the alternate‐leaf arrangement as the ancestral state for Polygonatum, but parsimony and maximum‐likelihood analyses suggest an equivocal state for crown Polygonatum. Leaf arrangement was found to be evolutionarily labile. A new nomenclatural combination was made: P olygonatum section S ibirica (L.I.Abramova) Y.Meng, comb. nov. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176 , 435–451.     

Unravelling the stylar polymorphism in Melochia (Malvaceae): reciprocity and ancillary characters

Heterostyly is a genetic polymorphism in which plant populations possess two (distyly) or three (tristyly) morphs with flowers differing reciprocally in stigma and anther height. Sex organ deployment has been described as being highly variable among and within species of several distylous taxa belonging to different taxonomic groups. However, the number of studies considering within‐species disparities is still limited. For a better overview of the existing amount of variation that can occur within and between heterostylous species, we sampled 46 populations of six Melochia spp., a style‐polymorphic genus in Cuba. We characterized the floral morphology in all populations and described a set of ancillary characters per species. All of these Melochia spp. are distylous, except for the monomorphic M. nodiflora. The S‐morph produces fewer, larger pollen than L‐morphs, and has verrucose ornamentation. The L‐morph produces reticulate pollen and has larger stigmatic papillae than the S‐morph. The monomorphic M. nodiflora shows ancillary characteristics that are similar to the L‐morph individuals in the related species. As expected, there are differences in ancillary characters among species and also dissimilarities in reciprocity among and within species of Melochia. Our results highlight the importance of considering intraspecific variation in the morphometric characterization of heterostylous taxa. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176 , 147–158.     

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.     

Morphological and molecular evidence for phylogeny and classification of South American pitvipers,genera Bothrops,Bothriopsis, and Bothrocophias (Serpentes: Viperidae)

Species in the genus Bothrops s. l. are extraordinarily variable in ecology and geography, compared with other genera in the subfamily Crotalinae. In contrast to the trend of splitting large and variable groups into smaller, more ecologically and phenotypically cohesive genera, the genus Bothrops has remained speciose. In addition, previous phylogenetic analyses have found Bothrops to be paraphyletic with respect to the genus Bothriopsis. Taxonomic arguments exist for synonymizing Bothriopsis with Bothrops, and for splitting Bothrops into smaller genera, but the greatest hindrance to taxonomic revision has been incomplete phylogenetic information. We present a phylogeny of Bothrops, Bothriopsis, and Bothrocophias based on 85 characters of morphology and 2343 bp of four mitochondrial gene regions, and with significantly greater taxonomic coverage than previous studies. The combined data provide improved support over independent datasets, and support the existence of discrete species groups within Bothrops. The monophyly and distinctness of these groups warrant recognition at the generic level, and we propose a new taxonomic arrangement to reflect these findings. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156 , 617–640.     

How common are dot‐like distributions? Taxonomical oversplitting in western European Agrodiaetus (Lepidoptera: Lycaenidae) revealed by chromosomal and molecular markers

Approximately 50 taxa of butterflies in Western Europe have been described as new species or elevated to the level of species during the last 40 years. Many, especially those belonging to the genus Agrodiaetus, have unusually localized, ‘dot‐like’ distributional ranges. In the present study, we use a combination of chromosomal and molecular markers to re‐evaluate the species status of Agrodiaetus distributed west of the 17th meridian. The results obtained do not support the current designations of Agrodiaetus galloi, Agrodiaetus exuberans, and Agrodiaetus agenjoi as endemic species with highly restricted distribution ranges, but indicate that these taxa are more likely to be local populations of a widely distributed species, Agrodiaetus ripartii. Agrodiaetus violetae is shown to be a polytypic species consisting of at least two subspecies, including Agrodiaetus violetae subbaeticus comb. nov. and Agrodiaetus violetae violetae. Agrodiaetus violetae is genetically (but not chromosomally) distinct from Agrodiaetus fabressei and has a wider distribution in southern Spain than previously believed. Agrodiaetus humedasae from northern Italy is supported as a highly localized species that is distinct from its nearest relatives. We propose a revision of the species lists for Agrodiaetus taking these new data into account. The results reported in the present study are relevant to animal conservation efforts in Europe because of their implications for IUCN Red List priorities. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 130–154.     

Consistent and variable leaf anatomical characters in Carex (Cyperaceae)

The anatomy and morphology of leaves in Carex have the potential to be taxonomically useful. However, studies on the variability of leaf characteristics in the genus are sparse. Researchers therefore risk using leaf anatomical characters without the knowledge of whether they are consistent in a species. We examined 22 qualitative and seven quantitative leaf anatomy characters from transverse leaf sections to test their consistency across 11 Carex spp. The characters were clearly described and primarily microscopic. Some characters were found to exhibit high levels of intraspecific variation, whereas other characters exhibited high levels of consistency in a species, including the shape of the leaf section, the density of papillae and the size of epidermal cells. Caution must be applied when choosing leaf anatomy to delimit taxa because of the intraspecific variability found in some characters, but sufficient numbers of invariant characters exist to provide useful taxonomic separation. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172 , 371–384.     

The utility of leaf flavonoids as taxonomic markers for some Malaysian species of the tribe Shoreae (Dipterocarpaceae)

A flavonoid survey was carried out on 45 taxa from the genera Shorea, Hopea, Parashorea, Neobalanocarpus, and Dryobalanops of the tribe Shoreae in the Dipterocarpaceae. The study showed significant chemotaxonomic differences in leaf flavonoid aglycone patterns and the presence of tannins in these taxa. The flavonoid patterns are useful in the delimitation of some taxa. For example, the genus Parashorea is distinguished by the universal presence of kaempferol 3‐methyl ether, and the monotypic genus Neobalanocarpus is unique in not producing ellagic and gallo tannins. The presence of chalcones and flavone C‐glycosides supports the separation of the genus Hopea into two sections, section Dryobalanoides and section Hopea in Ashton's classification, which is based on the type of venation. The flavonoid distributions in this study show that they can be very useful for differentiating between the Balau group in the genus Shorea and some scaly barked Hopea species, particularly H. helferi (lintah bukit), H. nutans (giam), and H. ferrea (malut). © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157 , 755–762.     

Descriptions and phylogenetic relationships of a new genus and two new species of Oligo‐Miocene cormorants (Aves: Phalacrocoracidae) from Australia

Tertiary cormorant fossils (Aves: Phalacrocoracidae) from Late Oligocene deposits in Australia are described. They derive from the Late Oligocene – Early Miocene (26–24 Mya) Etadunna and Namba Formations in the Lake Eyre and Lake Frome Basins, South Australia, respectively. A new genus, Nambashag gen. nov. , with two new species ( Nambashag billerooensis sp. nov. , 30 specimens; Nambashag microglaucus sp. nov. , 14 specimens), has been established. Phylogenetic analyses based on 113 morphological and two integumentary characters indicated that Nambashag is the sister taxon to the Early Miocene Nectornis miocaenus of Europe and all extant phalacrocoracids. As Nambashag, Nectornis, and extant phalacrocoracids constitute a strongly supported clade sister to Anhinga species, the fossil taxa have been referred to Phalacrocoracidae. Sulids and Fregata were successive sister taxa to the Phalacrocoracoidea, i.e. phalacrocoracids + Anhinga. As phalacrocoracids lived in both Europe and Australia during the Late Oligocene and no older phalacrocoracid taxa are known, the biogeographical origin of cormorants remains unanswered. The phylogenetic relationships of extant taxa were not wholly resolved, but contrary to previous morphological analyses, considerable concordance was found with relationships recovered by recent molecular analyses. Microcarbo is sister to all other extant phalacrocoracids, and all Leucocarbo species form a well‐supported clade. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 277–314.     

A molecular phylogenetic appraisal of the systematics of the Aglaopheniidae (Cnidaria: Hydrozoa,Leptothecata) from the north‐east Atlantic and west Mediterranean

The hydrozoan family Aglaopheniidae (Cnidaria) is widespread worldwide and contains some of the most easily recognizable hydroids because of their large colony size and characteristic microscopic structure. The systematics of the group has, however, been controversial and dedicated molecular analyses are lacking. We therefore analysed existing and new 16S rRNA sequences of Aglaopheniidae, in a total of 98 16S sequences corresponding to 25 putative species (25 nominal and three undescribed) from seven genera. The monophyly of the subfamilies Gymnangiinae and Aglaopheniinae, and tribes Aglaopheniini and Cladocarpini were not verified with 16S sequence data. The genera Gymnangium and Aglaophenia can only be considered valid if both Gymnangium gracicaule and Aglaophenia latecarinata are removed from their respective genera. The phenotypically similar Cladocarpus and Streptocaulus are probably monophyletic and clearly distinct genetically. The genus Lytocarpia may be polyphyletic. The nominal species Aglaophenia pluma, Aglaophenia tubiformis, and Aglaophenia octodonta are probably conspecific, as are also the species Aglaophenia acacia and Aglaophenia elongata. The 16S data revealed the existence of two potentially unnamed species of Aglaophenia respectively from the Azores and Madeira. The phylogeographical structure of the taxa with the greatest representation of haplotypes from the north‐east Atlantic and Mediterranean, revealed the influence of Mediterranean waters in Madeira and the Azores, and gene flow between deep waters of the Mediterranean and Atlantic. The last glaciations in Europe may have caused genetic bottlenecks but also high intraspecific haplotype diversity. Finally, Macrorhynchia philippina was detected in samples from Madeira and possibly represents an invasive species. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 717–727.     

Female polyandry and size‐assortative mating in isolated local populations of the Japanese common toad Bufo japonicus

In anurans, female polyandry under male harassment is distributed across taxa because of external aquatic fertilization. According to the sexual selection theory, male–male competition for access to females is affected by the operational sex ratio (OSR) and population density. The Japanese common toad, Bufo japonicus, is widespread in mainland Japan, and like the European common toad, B. bufo, it engages in explosive breeding. In this study, we observed the breeding behaviour of B. japonicus in isolated local populations for over four years in two breeding ponds with different population sizes and densities: large‐low (L‐pond) and small‐high (S‐pond). We analysed the relative polyandry ratio in egg clutches laid by females and estimated the size‐assortative mating pattern to be an indicator of male–male competition in the two ponds. Both ponds tended to exhibit a size‐assortative mating pattern; however, the frequency of polyandry was different in the two ponds (L‐pond = 20% and S‐pond = 90%). Our results showed that polyandry could occur without multiple amplexus with a high population density, i.e. eggs were often fertilized by free‐swimming sperm in the small shallow pond. We propose that high female polyandry ratios without continuous male harassment are generated because of a male‐biased OSR and a high population density in the small pond. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 236–242.