Divergence between passerine populations from the Malvinas – Falkland Islands and their continental counterparts: a comparative phylogeographical study

Bursts of speciation have followed colonization of remote oceanic islands by diverse taxa, a process evidenced by island endemics around the world. The present study explores whether the Malvinas – Falkland Islands (MFI), a relatively understudied archipelago off the South Atlantic coast of Patagonia, harbour endemic genetic lineages of passerine birds. Nine passerine species nest regularly in the MFI (Cinclodes antarcticus, Muscisaxicola maclovianus, Troglodytes cobbi, Cistothorus platensis, Turdus falcklandii, Anthus correndera, Melanodera melanodera, Sturnella loyca, and Carduelis barbata). Mitochondrial DNA (cytochrome c oxidase I sequences) are used to quantify and compare divergence between insular and continental populations, finding genetic patterns to vary across these nine species. Most MFI passerines do not show significant genetic differentiation from continental populations, whereas C. platensis, M. melanodera, and T. falcklandii are modestly diverged. Finally, T. cobbi differes markedly from its closest continental relative Troglodytes aedon, a result that is confirmed using nuclear and vocal data. The study also identifies broadly divergent lineages within continental populations of C. platensis and T. aedon. Taken together, these results suggest that the land bird populations of the MFI were established at different times. Troglodytes cobbi is the oldest MFI land bird, splitting from continental T. aedon during the Great Patagonian Glaciation of the Pleistocene. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, •• , ••–••.     

Fly pollination in Cypripedium: a case study of sympatric C. sichuanense and C. micranthum

Most Cypripedium spp. are known to be pollinated by bees. However, myiophilous traits are found in some species, especially in sections Trigonopedia and Sinopedilum. Here we chose C. micranthum and C. sichuanense, two sympatric species endemic to Sichuan, China, to test whether these orchids are fly pollinated. Artificial pollination showed that both flowers are self‐compatible but need pollen vectors for successful reproduction. Field observation showed that C. micranthum was pollinated by fruit flies and C. sichuanense by dung flies, both novel pollinators of Cypripedium orchids. These sympatric Cypripedium spp. are also cross‐compatible, but hybrids were not found in nature. The pollination syndromes of C. sichuanense and C. micranthum fit into the complex sapromyiophily pattern. It appears that pollinator specificity is responsible for their reproductive isolation. The discovery of fly pollination in C. sichuanense and C. micranthum, which belong to the related sections Trigonopedia and Sinopedilum, suggests a shift from bee to fly pollination in the genus Cypripedium. Unlike most Cypripedium spp., the anthers of C. micranthum release discrete pollinia with narrow stalks instead of the usual amorphous pollen smears. This ‘proto‐pollinarium’ is described, probably for the first time. These pollinia are most likely an adaptation for pollination by microdiptera, so the fly can carry the contents of both chambers in the same anther. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 170 , 50–58.     

Form and function within a phylogenetic framework: locomotory habits of extant predators and some Miocene Sparassodonta (Metatheria)

In this study, we analysed locomotory habits in extant predators and Sparassodonta species through geometric morphometric techniques and discriminant analyses of the distal humerus in anterior view, proximal ulna in lateral view, and tibia in proximal view. We included a wide sample of extant predators, and considered the phylogenetic and allometric structure in the data sets. We also included some Sparassodonta, a group of carnivorous metatherians that inhabited South America during the Cenozoic, and inferred their locomotory habits. Results suggest the presence of a close relationship between shape and locomotory habits, even after removing the shape component explained by phylogeny in the three postcranial elements. Terrestrial habits were inferred for Arctodictis sinclairi, Borhyaena tuberata, ‘Lycopsis’ longirostrus, and Thylacosmilus atrox. Some degree of cursoriality was highlighted in B. tuberata and T. atrox, and climbing abilities in ‘L.’ longirostrus, and to a lesser degree in B. tuberata. Scansorial habits were inferred for Cladosictis patagonica, Sipalocyon gracilis, Prothylacynus patagonicus, and Pseudonotictis pusillus, and in the case of C. patagonica, some digging ability was also tentatively inferred. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 165 , 224–251.     

A new group of Early Devonian plants with valvate sporangia containing sculptured permanent dyads

As part of a study to explore diversity and disparity in Early Devonian terrestrial vegetation, several hundreds of sporangia with in situ spores have been isolated from a Lochkovian locality in Shropshire. These include a small number (seven) of sporangia showing dehiscence into four valves and containing permanent sculptured dyads, belonging to the Cymbohilates horridus complex and C. cymosus, which are recorded in coeval dispersed spore assemblages. A further, previously described, mesofossil comprises an incomplete sporangium containing C. horridus that terminates a naked isotomously branching stem with stomata. The valvate sporangia are placed in a new genus, Partitatheca, containing four species, P. splendida (type), P. horrida, P. densa and P. cymosa, their names reflecting the names of the dispersed spore species and varieties. Complex ultrastructure in the walls of the dyads is similar to that in earlier dyads in the Dyadospora complex where it provides evidence for a hepatic affinity of the earliest embryophytes, but the new taxa present a combination of bryophyte and tracheophyte characters and are considered to represent a new embryophyte lineage. General discussion includes the development of dyads, more particularly their relevance to understanding the diversity in meiotic processes, and the disappearance of dyads from the dispersed spore record prior to the Middle Devonian. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 168 , 229–257.     

Geometric morphometric analysis of shell shape variation in Conus (Gastropoda: Conidae)

Geometric morphometric techniques allow for the direct quantification and analysis of variation in biological shape and have been used in studies in systematic biology. However, these techniques have not been used for species discrimination in the gastropod genus Conus, a major taxon of significant tropical reef predators recognized for their peptide‐based toxins. Here, we used landmark digitization and analysis to show that five species commonly studied for their conotoxins –Conus consors, Conus miles, Conus stercusmuscarum, Conus striatus, and Conus textile – can be effectively distinguished from each other by their shape, as manifested in the results of a principal components analysis (PCA) and the generated thin‐plate splines. Two piscivorous species, C. stercusmuscarum and C. striatus, show clear overlaps in the PCA plot, although each taxon clusters within itself, as does each of the others. The loadings on the first two principal components show that the forms of the shells' aperture and spire are particularly important for discrimination. Phylogenetic analysis using neighbour‐joining methods shows that group separations are comparable with published phylogenetic schemes based on molecular data and feeding mode (i.e. piscivory, vermivory, molluscivory). The results of this study establish the utility of geometric morphometric methods in capturing the interspecific differences in shell form in the genus Conus. This may lead to the utilization of these methods on other gastropod taxa and the creation of species‐recognition programs based on shell shape. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 165 , 296–310.     

Systematics and cladistics of a new Naupactini genus (Coleoptera: Curculionidae: Entiminae) from the Andes of Colombia and Ecuador

A new genus of broad‐nosed weevils belonging to the tribe Naupactini (Coleoptera: Curculionidae) is described, based on six species from the Páramo–Puna subregion (Andean region) of South America. Three species are new and three have been transferred from the genus Asymmathetes (Asymmathetes nigrans, Asymmathetes rugicollis, and Asymmathetes vulcanorum). The new genus is recognized by the black, denuded, and shiny tegument, the well‐developed pre‐epistome, the elytral base curved backwards on middle, the reduction of the hindwings, and the widely separated procoxae, closer to the anterior than to the posterior margin of the prosternum. A cladistic analysis of the six species of the new genus plus five out‐groups (Amitrus, Amphideritus, Asymmathetes, Melanocyphus, and Trichocyphus), using 49 morphological characters, resulted in a single cladogram. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166 , 54–71.     

Revision of the New World genus Quichuana Knab, 1913 (Diptera: Syrphidae), including descriptions of 24 new species

The species of the New World syrphid genus, Quichuana Knab, 1913 (Diptera: Syrphidae: Eristalini), are revised and the genus is re‐diagnosed. Twenty‐four new species are described, bringing the total number considered valid to 48. New species were mostly reared from Bolivia, Brazil, Costa Rica, El Salvador, Mexico, Peru, Trinidad, and Venezuela. The male of Quichuana picadoi is described for the first time. Quichuana championi is proposed as a synonym of Quichuana cincta, Quichuana aurata as a synonym of Quichuana angustiventris, and Quichuana sepiapennis as a synonym of Quichuana calathea. Quichuana inca var. brevicera is raised to specific status. Lectotypes are designated for Helophilus auratus, Quichuana bezzii, Quichuana fasciata, and Quichuana parisii. A species‐level identification key for both males and females is provided. Male genitalia of 18 species are illustrated. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166 , 72–131.     

Phylogenetic relationships in the genus Agathemera (Insecta: Phasmatodea) inferred from the genes 16S,COI and H3

We present a phylogenetic analysis that includes all known species of Agathemera, using as outgroup four species of the genera Heteronemia, Spinonemia, Monticomorpha, and Anisomorpha. Phylogenetic inference was based on three genes, 16S, COI (mitochondrial markers) and H3 (nuclear marker), based on the maximum‐parsimony, maximum‐likelihood and Bayesian inference methods. Our results show that the genus Agathemera is monophyletic; six of its eight species showed phylogenetic support, while the group A. claraziana + A. millepunctata was not resolved. The resulting topology shows two major clades, the first with A. maculafulgens, A. luteola, A. crassa, A. millepunctata, and A. claraziana, and the second including A. grylloidea, A. mesoauriculae, and A. elegans. Species of the first clade have large mesonotal processes and are found mainly east of the Andes, except for A. crassa. Members of the second clade have small or absent mesonotal processes and are distributed mainly to the west of the Andes. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 165 , 63–72.     

The phylogeny of a reduced ‘sand goby’ group based on behavioural and life history characters

Phylogenetic analysis of 27 behavioural and life history traits for five Mediterranean sand goby species (Perciformes, Gobiidae) produced one tree with a consistency index (excluding uninformative characters) of 0.756. This tree agreed with previous molecular analyses in providing strong support for the monophyly of the sand gobies, indicating that Pomatoschistus and Knipowitschia are paraphyletic and helping to resolve the ambiguous position of Economidichthys pygmaeus, placing it as the basal member of the reduced data set. Although the tree was completely resolved, the branches above E. pygmaeus were only moderately supported in the bootstrap analysis. Overall, the behavioural data provide information that may eventually help clarify the speciation bursts within the Mediterranean sand goby clade as much as is possible. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 165 , 916–924.     

Selection for abdominal tergite pigmentation and correlated responses in the trident: a case study in Drosophila melanogaster

In Drosophila melanogaster, abdominal tergite pigmentation and the appearance of a trident‐shaped thoracic pattern exhibit similar biogeographical variation and sensitivity to temperature. These pigmentation traits may be under common selection pressure in natural populations or may be genetically correlated. To investigate the nature of this interaction, replicated populations of D. melanogaster were selected for increased or decreased melanization of the abdominal tergites for 40 generations. Selection for abdominal tergite pigmentation leads to correlated changes in trident formation. Although selection was performed only on female flies, male pigmentation also responded to selection. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 287–294.     

Diversification of lindsaeoid ferns and phylogenetic uncertainty of early polypod relationships

We analysed one nuclear gene (18S) and seven plastid markers [five protein coding (atpA, atpB, rbcL, rpoC1, rps4) and two non‐coding (trnH‐psbA, trnL‐trnF] for 31 members of Polypodiales and four outgroup taxa. We focused our sampling on the lindsaeoids and associated ferns in order to obtain a better understanding of the diversification of the early polypods. However, the exact phylogenetic position of Saccoloma and Cystodium remained uncertain. Based on relaxed molecular clock analyses, it appears that the crown group lindsaeoids diversified in the Caenozoic, more or less simultaneously with the main radiation of other Polypodiales, even though the original divergence between the lindsaeoid and non‐lindsaeoid polypods occurred before the end of the Jurassic. The current pantropical distribution of lindsaeoids can be explained by either long‐distance dispersal across the oceans or vicariance caused by the retreat of previously widely distributed tropical forests from higher to lower latitudes. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 170 , 489–503.     

Early Palaeogene Louisinidae (Macroscelidea,Mammalia), their relationships and north European diversity

Seventeen species from the Palaeocene and Early Eocene of northern Europe, of which 12 are new, are described belonging to the extinct macroscelidean family Louisinidae, raised here from subfamily rank. These species belong to nine genera, of which five are new. The new genera are Walbeckodon , Berrulestes , Gigarton , Thryptodon , and Prolouisina . The new species are Walbeckodon krumbiegeli , Walbeckodon girardi , Paschatherium levei , Berrulestes phelizoni , Berrulestes pellouini , Berrulestes poirieri , Gigarton meyeri , Gigarton sigogneauae , Gigarton louisi , Thryptodon brailloni , Louisina marci , and Teilhardimys brisswalteri . Prolouisina is erected for ‘Louisina’atavella Russell, 1964. Cladistic analysis was undertaken to understand the relationships within the Louisinidae and between them and the North American family Apheliscidae, in which they had earlier been included as a subfamily. Louisinidae are shown to be sister group to a clade consisting of Apheliscidae plus Amphilemuridae and part of a paraphyletic and polyphyletic Adapisoricidae, all of which are tentatively considered to be stem members of the order Macroscelidea. The most primitive macroscelidid, Chambius, from the Early Eocene of northern Africa is nested within Apheliscidae when postcranial characters were included, but in a majority of cases within the Louisinidae when postcranial characters were excluded. Most species from northern Europe became extinct at the end of the Palaeocene, although the genus Paschatherium survived for much of the Early Eocene and Teilhardimys survived into the earliest Eocene. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 856–936.     

Morphological evolution in the graminid clade: comparative floral anatomy of the grass relatives Flagellariaceae and Joinvilleaceae

New comparative data are presented on the reproductive morphology and anatomy of two genera closely related to grasses, Flagellaria and Joinvillea, in which the flowers are superficially similar, especially in stamen morphology. This investigation demonstrates some anatomical differences between the two genera. For example, both genera depart from the ‘typical’ condition of tepal vasculature (three‐traced outer tepals and one‐traced inner tepals): in Flagellaria, each tepal receives a single vascular bundle and, in Joinvillea, each tepal is supplied by three vascular bundles. Joinvillea possesses supernumerary carpel bundles, as also found in the related family Ecdeiocoleaceae, but not in Flagellaria or grasses. In the anther, the tapetum degenerates early in Flagellaria, and is relatively persistent in Joinvillea, in which the pollen grains remain closely associated with the tapetum inside the anther locule, indicating a correlation between peripheral pollen (a feature that is common in grasses) and a persistent tapetum. This study highlights the presence of a pollen‐tube transmitting tissue (PTTT) or solid style in the gynoecium of Flagellaria, as also in many Poaceae, but not in Joinvillea or Ecdeiocoleaceae. We speculate that the presence of a PTTT could represent one of the factors that facilitated the subsequent evolution of the intimately connected gynoecia that characterize grasses. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 170 , 393–404.     

A taxonomic revision and pollen morphology of the genus Dendrokingstonia (Annonaceae)

The genus Dendrokingstonia (Annonaceae) is taxonomically revised and palynologically studied. Three species are recognized, one of which, D. gardneri , is described as new to science. One new combination, D. acuminata , is made. The genus occurs from southern Thailand to Peninsular Malaysia and Sumatra. On the basis of macromorphology and pollen characters, it is considered to be related to Monocarpia. Both genera show a combination of macromorphological characters that is rare in the family, i.e. considerably enlarged stigmas, leaves with percurrent tertiary veins, a highly reduced number of carpels per flower and relatively large monocarps with a thick, hard wall. Scanning and transmission electron microscopy show that the pollen grains of Dendrokingstonia and Monocarpia are monosulcate monads with a columellate infratectum and a more or less bulging intine at the sulcus. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 168 , 76–90.     

Intraspecific versus interspecific variation in Miocene Great Basin mylagaulids: implications for systematics and evolutionary history

The Mylagaulidae are a family of burrowing rodents abundant in Miocene faunas from western North America. Recent taxonomic revisions of mylagaulids from the Great Plains suggest that their systematics may be best understood on a regional basis. Previous studies addressed the taxonomy and evolutionary history of mylagaulids from the Great Basin, but recent discoveries of specimens, new phylogenetic data, and more detailed stratigraphical information necessitate a thorough reanalysis of their relationships and occurrences. We present a revision of the systematics of the mylagaulids from the Great Basin. In addition to rare large mylagaulids of uncertain taxonomic affinity, we recognize four species of mylagaulids distributed throughout Oregon and Nevada from the late Hemingfordian through to the early late Hemphillian: Alphagaulus vetus, Hesperogaulus gazini, Hesperogaulus wilsoni, and a new species from the genus Hesperogaulus. All species are known from large sample sizes of isolated premolars, allowing consideration of ontogenetic variation in determining the key morphological differences that allow recognition of different species. Although the number of enamel lakes varies within a given taxon, the presence of some of these lakes is taxonomically significant. This result emphasizes the importance of understanding ontogeny in describing species of fossil hypsodont mammals. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 427–450.     

Unravelling the evolutionary history of the polyploid complex Ranunculus parnassiifolius (Ranunculaceae)

Ranunculus L. represents the largest genus within Ranunculaceae, comprising more than 600 species with a worldwide distribution. However, there are still many gaps in our knowledge of the infrageneric taxonomy and evolution of Ranunculus. In this regard, intraspecific variation of the polyploid complex Ranunculus parnassiifolius remains under discussion. To reconstruct the biogeographical history of the polyploid complex R. parnassiifolius, 20 populations distributed throughout the Cantabrian Mountains, Pyrenees, and Alps were investigated. Phylogenetic studies were based on nuclear internal transcribed spacers (ITS) and plastid (rpl32‐trnL, rps16‐trnQ) sequence data, analysed using Bayesian approaches as well as the evolution of morphological characters. Additionally, biogeographical patterns were conducted using statistical dispersal–vicariance analysis. The analyses presented here support the recognition of two evolutionary independent units: R. cabrerensis sensu lato (s.l.) and R. parnassiifolius s.l. Furthermore gradual speciation depending on the biogeographical territory is proposed, and optimal reconstructions have probably favoured the ancestor of Ranunculus parnassiifolius as originating in the Iberian Peninsula. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 107 , 477–493.