Phylogeny and classification of Ochlerotatus and allied taxa (Diptera: Culicidae: Aedini) based on morphological data from all life stages

The phylogenetic relationships and generic assignments of ‘Ochlerotatus’ and related taxa of uncertain taxonomic position in the classification of Aedini previously proposed by the authors in 2004 and 2006 are explored using 297 characters from eggs, fourth‐instar larvae, pupae, adults and immature habitat coded for 158 exemplar species. The ingroup comprises 54 species and the outgroup includes four non‐aedine species and 100 aedine species, 21 of which were previously classified as incertae sedis. Data are analysed in a total‐evidence approach using implied weighting. The analysis produced 158 most parsimonious cladograms. The strict consensus tree (SCT) corroborates the monophyly of the 30 generic‐level taxa recognized previously that are included in the analysis. Overall, the results show remarkable congruence with those obtained previously despite differences in the taxa and morphological characters analysed in this and the two previous studies. All species of Ochlerotatus s.s., subgenus ‘Ochlerotatus’sensu auctorum, Geoskusea, Levua, Pseudoskusea and Rhinoskusea included in the analysis fall within a single clade that is treated as genus Ochlerotatus; thus, the last four taxa are restored to their previous subgeneric rank within this genus. Nine additional subgenera, of which four are new, are proposed for monophyletic clades of Ochlerotatus species based on the strength of character support and application of the principle of equivalent rank. Acartomyia stat. nov. , Culicelsa stat. nov. , Gilesia stat. nov. , Protoculex stat. nov. and Chrysoconops stat. nov. are resurrected from synonymy with Ochlerotatus; and Empihals subgen. nov. (type species: Culex vigilax Skuse), Pholeomyia subgen. nov. (type species: Aedes calcariae Marks), Buvirilia subgen. nov. (type species: Aedes edgari Stone & Rosen) and Sallumia subgen. nov. (type species: Aedes hortator Dyar & Knab) are described as new. The sister group of Ochlerotatus includes a number of species that were previously regarded as incertae sedis in ‘Oc. (Finlaya)’ and ‘Oc. (Protomacleaya)’. Based on previous observations, refined relationships and new character support, three additional genera are recognized for species previously included in ‘Finlaya’, i.e. Danielsia stat. nov . (type species: Danielsia albotaeniata Leicester), Luius gen. nov. (type species: Aedes fengi Edwards) and Hopkinsius gen. nov. (type species: Aedes ingrami Edwards). Additionally, Alloeomyia subgen. nov. (type species: Culex pseudotaeniatus Giles) and Yamada subgen. nov. (type species: Aedes seoulensis Yamada) are introduced as subgenera of Collessius and Hopkinsius, respectively. As is usual with generic‐level groups of Aedini, the newly recognized genera and subgenera are polythetic taxa that are diagnosed by unique combinations of characters. The analysis corroborates the previous observation that ‘Oc. (Protomacleaya)’ is a polyphyletic assemblage of species. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 153 , 29–114.     

Description of a new diatom genus Dorofeyukea gen. nov. with remarks on phylogeny of the family Stauroneidaceae

Type material of Navicula kotschyi was studied, and this species was transferred to Dorofeyukea gen. nov. as D. kotschyi comb. nov. Dorofeyukea was described on the basis of DNA sequence and morphological data. Additional species assigned to this genus that were previously included in Navicula include: D. ancisa comb. nov., D. grimmei comb. nov., D. ivatoensis comb. nov., D. orangiana comb. nov., D. rostellata comb. nov. & stat. nov., D. savannahiana comb. nov., D. tenuipunctata comb. nov., and D. texana comb. nov. All Dorofeyukea species share the same morphological features, including having a narrow stauroid fascia surrounded by 1–3 irregularly shortened striae, uniseriate, and weakly radiate striae, circular, or rectangular puncta in the striae that are covered internally by dome‐shaped hymenes, presence of a pseudoseptum at each apex and absence of septa. Partial DNA sequences of SSU and rbcL loci show Dorofeuykae belongs to the clade of stauroneioid diatoms together with Stauroneis, Prestauroneis, Craticula, Karayevia, Madinithidium, Fistulifera, Parlibellus, and, possibly, Schizostauron. A new species from the monoraphid genus Madinithidium, M. vietnamica sp. nov., was described based on valve and chloroplast morphology as well as DNA sequence data.     

Phylogeny and classification of tribe Aedini (Diptera: Culicidae)

The phylogeny and classification of tribe Aedini are delineated based on a cladistic analysis of 336 characters from eggs, fourth‐instar larvae, pupae, adult females and males, and immature stage habitat coded for 270 exemplar species, including an outgroup of four species from different non‐aedine genera. Analyses of the data set with all multistate characters treated as unordered under implied weights, implemented by TNT version 1.1, with values of the concavity constant K ranging from 7 to 12 each produced a single most parsimonious cladogram (MPC). The MPCs obtained with K values of 7–9 were identical, and that for K = 10 differed only in small changes in the relationships within one subclade. Because values of K < 7 and > 10 produced large changes in the relationships among the taxa, the stability of relationships exemplified by the MPC obtained from the K = 9 analysis is used to interpret the phylogeny and classification of Aedini. Clade support was assessed using parsimony jackknife and symmetric resampling. Overall, the results reinforce the patterns of relationships obtained previously despite differences in the taxa and characters included in the analyses. With two exceptions, all of the groups represented by two or more species were once again recovered as monophyletic taxa. Thus, the monophyly of the following genera and subgenera is corroborated: Aedes, Albuginosus, Armigeres (and its two subgenera), Ayurakitia, Bothaella, Bruceharrisonius, Christophersiomyia, Collessius (and its two subgenera), Dahliana, Danielsia, Dobrotworskyius, Downsiomyia, Edwardsaedes, Finlaya, Georgecraigius (and its two subgenera), Eretmapodites, Geoskusea, Gilesius, Haemagogus (and its two subgenera), Heizmannia (and subgenus Heizmannia), Hopkinsius (and its two subgenera), Howardina, Hulecoeteomyia, Jarnellius, Kenknightia, Lorrainea, Macleaya, Mucidus (and its two subgenera), Neomelaniconion, Ochlerotatus (subgenera Chrysoconops, Culicelsa, Gilesia, Pholeomyia, Protoculex, Rusticoidus and Pseudoskusea), Opifex, Paraedes, Patmarksia, Phagomyia, Pseudarmigeres, Rhinoskusea, Psorophora (and its three subgenera), Rampamyia, Scutomyia, Stegomyia, Tanakaius, Udaya, Vansomerenis, Verrallina (and subgenera Harbachius and Neomacleaya), Zavortinkius and Zeugnomyia. In addition, the monophyly of Tewarius, newly added to the data set, is confirmed. Heizmannia (Mattinglyia) and Verrallina (Verrallina) were found to be paraphyletic with respect to Heizmannia (Heizmannia) and Verrallina (Neomacleaya), respectively. The analyses were repeated with the 14 characters derived from length measurements treated as ordered. Although somewhat different patterns of relationships among the genera and subgenera were found, all were recovered as monophyletic taxa with the sole exception of Dendroskusea stat. nov. Fifteen additional genera, three of which are new, and 12 additional subgenera, 11 of which are new, are proposed for monophyletic clades, and a few lineages represented by a single species, based on tree topology, the principle of equivalent rank, branch support and the number and nature of the characters that support the branches. Acartomyia stat. nov. , Aedimorphus stat. nov. , Cancraedes stat. nov. , Cornetius stat. nov. , Geoskusea stat. nov. , Levua stat. nov. , Lewnielsenius stat. nov. , Rhinoskusea stat. nov. and Sallumia stat. nov., which were previously recognized as subgenera of various genera, are elevated to generic status. Catageiomyia stat. nov. and Polyleptiomyia stat. nov. are resurrected from synonymy with Aedimorphus, and Catatassomyia stat. nov. and Dendroskusea stat. nov. are resurrected from synonymy with Diceromyia. Bifidistylus gen. nov. (type species: Aedes lamborni Edwards) and Elpeytonius gen. nov. (type species: Ochlerotatus apicoannulatus Edwards) are described as new for species previously included in Aedes (Aedimorphus), and Petermattinglyius gen. nov. (type species: Aedes iyengari Edwards) and Pe. (Aglaonotus) subgen. nov. (type species: Aedes whartoni Mattingly) are described as new for species previously included in Aedes (Diceromyia). Four additional subgenera are recognized for species of Ochlerotatus, including Oc. (Culicada) stat. nov. (type species: Culex canadensis Theobald), Oc. (Juppius) subgen. nov. (type species: Grabhamia caballa Theobald), Oc. (Lepidokeneon) subgen. nov. (type species: Aedes spilotus Marks) and Oc. (Woodius) subgen. nov. (type species: Aedes intrudens Dyar), and seven are proposed for species of Stegomyia: St. (Actinothrix) subgen. nov. (type species: Stegomyia edwardsi Barraud), St. (Bohartius) subgen. nov. (type species: Aedes pandani Stone), St. (Heteraspidion) subgen. nov. (type species: Stegomyia annandalei Theobald), St. (Huangmyia) subgen. nov. (type species: Stegomyia mediopunctata Theobald), St. (Mukwaya) subgen. nov. (type species: Stegomyia simpsoni Theobald), St. (Xyele) subgen. nov. (type species: Stegomyia desmotes Giles) and St. (Zoromorphus) subgen. nov. (type species: Aedes futunae Belkin). Due to the unavailability of specimens for study, many species of Stegomyia are without subgeneric placement. As is usual with generic‐level groups of Aedini, the newly recognized genera and subgenera are polythetic taxa that are diagnosed by unique combinations of characters. The analysis corroborates the previous observation that ‘Oc. (Protomacleaya)’ is a polyphyletic assemblage of species.     

Why are there so many mimicry rings? Correlations between habitat,behaviour and mimicry in Heliconius butterflies

In the new world tropics there is an extravagant array of sympatric butterfly mimicry rings. This is puzzling under strictly coevolutionary (Müllerian) mimicry: all unpalatable species should converge as ‘co-mimics' to the same pattern. If mimicry has usually evolved in unpalatable species by one-sided (Batesian) evolution, however, it is easy to see that mimicry rings centred on different models could remain distinct. If mimicry rings were also segregated by habitat, a diversity of mimicry rings could be stabilized. In this paper we report correlations between behaviour and mimicry of nine unpalatable Heliconius species. It is already known that co-mimics fly in similar habitats, and non-mimics fly in different habitats, although there is much overlap. Contrary to a previous report, we find little difference in flight heights of heliconiine mimicry rings; all species fly from ground level to the canopy. However, co-mimics roost at night in similar habitats and at similar heights above the ground, but in different habitats and at different heights from species in other mimicry rings. Heliconius (especially the erato taxonomic group) are renowned for roosting gregariously; and co-mimics roost gregariously with each other more often than with non-mimics. Gregarious roosting is therefore common between species, as well as within species. There are thus strong links between mimicry and behavioural ecology in Heliconius. The paradoxical correlation between nocturnal roosting and visual mimicry is presumably explained by bird predation at dusk when roosts are forming, or at dawn before they have disbanded. Direct evidence of predation is lacking, but there are high rates of disturbance by birds at these times. These results, together with knowledge of the phylogeny of Heliconius, suggest that species from the melpomene-group of Heliconius have radiated to occupy mimetic niches protected by model species in the Ithomiinae and the erato-group of Heliconius. A variety of sympatric mimicry rings is apparently maintained because key models fail to converge, while more rapidly-evolving unpalatable mimics evolve towards the colour patterns of the models. The maintenance of mimetic diversity would be aided by the habitat and behavioural differences between mimicry rings revealed here, provided that different predators are found in different habitats. This explanation for the maintenance of multiple mimicry rings is more plausible for Heliconius mimicry than alternatives based on visual mating constraints, thermal ecology, or camouflage.     

Aphis (Hemiptera: Aphididae) species groups found in the Midwestern United States and their contribution to the phylogenetic knowledge of the genus

A phylogeny of the genus Aphis Linnaeus, 1 758 was built primarily from specimens collected in the Midwest of the United States. A data matrix was constructed with 68 species and 41 morphological characters with respective character states of alate and apterous viviparous females. Dendrogram topologies of analyses performed using UPGMA (Unweighted Pair Group Method with Arithmetic Mean), Maximum Parsimony and Bayesian analysis of Cytochrome Oxidase I, Elongation Factor 1‐α and primary endosymbiont Buchnera aphidicola 16S sequences were not congruent. Bayesian analysis strongly supported most terminal nodes of the phylogenetic trees. The phylogeny was strongly supported by EF1‐α, and analysis of COI and EF1‐α molecular data combined with morphological characters. It was not supported by single analysis of COI or Buchnera aphidicola 16S. Results from the Bayesian phylogeny show 4 main species groups: asclepiadis, fabae, gossypii, and middletonii. Results place Aphis and species of the genera Protaphis Börner, 1952, Toxoptera Koch, 1856 and Xerobion Nevsky, 1928 in a monophyletic clade. Morphological characters support this monophyly as well. The phylogeny shows that the monophyletic clade of the North American middletonii species group belong to the genus Protaphis: P. debilicornis (Gillette & Palmer, 1929 ), comb. nov., P. echinaceae (Lagos and Voegtlin, 2009 ), comb. nov., and P. middletonii (Thomas, 1879 ). The genus Toxoptera should be considered a subgenus of Aphis (stat. nov.). The analysis also indicates that the current genus Iowana Frison, 1954 should be considered a subgenus of Aphis (stat. nov.).     

The first fossil spider cricket (Orthoptera: Gryllidae: Phalangopsinae): 20 million years of troglobiomorphosis or exaptation in the dark?

A new spider cricket (Orthoptera: Gryllidae: Phalangopsinae) is described from an adult female preserved in Early Miocene (Burdigalian) amber from the Dominican Republic. Araneagryllus dylani gen. et sp. nov. represents the first fossil record of Phalangopsinae, and is assigned to the tribe Luzarini, subtribe Amphiacustina stat. nov. A cladistic analysis of Amphiacustina places Araneagryllus gen. nov. within a clade comprising Arachnopsita, Leptopedetes, Longuripes, Mayagryllus, Nemoricantor, and Prolonguripes. This clade is the sister group to a clade comprising Amphiacusta, Cantrallia, and Noctivox. The results of this analysis suggest that: (1) the common ancestor of all Amphiacustina was epigean, and was likely to have been cavicolous and/or straminicolous; (2) strict troglobitism evolved twice within Amphiacustina, once in the lineage leading to Noctivox and again in the clade comprising Mayagryllus, Arachnopsita, Longuripes, and Prolonguripes; and (3) Prolonguripes is secondarily epigean, having reverted to life above ground. The occurrence of Araneagryllus gen. nov. in amber indicates that it was not troglobitic, but was instead more likely to have been straminicolous, living on the ground and foraging amongst leaf litter. Araneagryllus gen. nov. possesses a number of characters that are usually considered to be adaptive to a troglobitic life history, suggesting that many so‐called troglobiomorphies are not adaptations to life in caves, but are instead likely to have been exaptive. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 56–65.     

Molecular phylogenetics and evolution of Orchidinae and selected Habenariinae (Orchidaceae)

Internal transcribed spacer (ITS nuclear rDNA) data have been obtained from 190 terrestrial orchid species, encompassing all genera and the great majority of the widely recognized species of Orchidinae, a heterogeneous selection of species of Habenariinae, and single species of Satyriinae and Disinae (the latter serving as outgroup). The resulting parsimony‐based phylogeny reveals 12 well‐resolved clades within the Orchidinae, based on Anacamptis s.l., Serapias, Ophrys, Steveniella–Himantoglossum s.l. (including ‘Comperia’ and ‘Barlia’, most species being 2n = 36), Neotinea s.l., Traunsteinera–Chamorchis, Orchis s.s., Pseudorchis–Amerorchis–Galearis–Neolindleya–Platanthera s.l. (most 2n = 42), Dactylorhiza s.l., Gymnadenia s.l. (most 2n = 40, 80), Ponerorchis s.l.–Hemipilia s.l.–Amitostigma–Neottianthe, and Brachycorythis (most 2n = 42). Relationships are less clearly resolved among these 12 clades, as are those within Habenariinae; the subtribe appears either weakly supported as monophyletic or as paraphyletic under maximum parsimony, and the species‐rich genus Habenaria is clearly highly polyphyletic. The triphyly of Orchis as previously delimited is confirmed, and the improved sampling allows further generic transfers to Anacamptis s.l. and Neotinea s.l. In addition, justifications are given for: (1) establishing Steveniella as the basally divergent member of an appreciably expanded Himantoglossum that incorporates the former genera ‘Barlia’ and ‘Comperia’, (2) reuniting ‘Piperia’ with a broadly defined Platanthera as section Piperia, necessitating ten new combinations, (3) broadening Ponerorchis to include Chusua, and Hemipilia to include single ‘orphan’ species of Ponerorchis and Habenaria, and (4) recognizing ‘Gymnadenia’camtschatica as the monotypic Neolindleya camtschatica within the Pseudorchis～Platanthera clade. Few further generic transfers are likely in Orchidinae s.s., but they are anticipated among habenariid genera, on acquisition of additional morphological and molecular evidence; one probable outcome is expansion of Herminium. Species‐level relationships are also satisfactorily resolved within most of the major clades of Orchidinae, with the notable exceptions of Serapias, the derived sections of Ophrys, Himantoglossum s.s., some sections within Dactylorhiza, the former genus ‘Nigritella’ (now tentatively placed within Gymnadenia s.l.), Hemipilia s.l., and possibly Ponerorchis s.s. Relationships among the 12 major clades broadly accord with bona fide records of intergeneric hybridization. Current evidence supports the recently recognized 2n = 36 clade; it also indicates a 2n = 40 clade that is further diagnosed by digitate root‐tubers, and is derived relative to the recently recognized clade of exclusively Asian genera (Ponerorchis s.l.–Hemipilia s.l.–Amitostigma–Neottianthe). This in turn appears derived relative to the Afro‐Asiatic Brachycorythis group; together, these two clades identify the plesiomorphic chromosome number as 2n = 42. If the African genus Stenogolottis is correctly placed as basally divergent within a monophyletic Habenariinae, the tribe Orchideae and subtribes Orchidinae and Habenariinae could all have originated in Africa, though in contrast the Asiatic focus of the basally divergent members of most major clades of Orchidinae suggests an Asiatic radiation of the subtribe. Morphological characters informally ‘mapped’ across the molecular phylogeny and showing appreciable levels of homoplasy include floral and vegetative pigmentation, flower shape, leaf posture, gynostemium features, and various pollinator attractants. Qualitative comparison of, and reciprocal illumination between, degrees of sequence and morphological divergence suggests a nested set of radiations of progressively decreasing phenotypic magnitude. Brief scenarios, both adaptive and non‐adaptive, are outlined for specific evolutionary transitions. Recommendations are made for further species sampling, concentrating on Asian Orchidinae (together with the Afro‐Asiatic Brachycorythis group) and both Asian and Southern Hemisphere Habenariinae, and adding plastid sequence data. Taxonomic changes listed are: Anacamptis robusta (T.Stephenson) R.M.Bateman, comb. nov. , A. fragrans (Pollini) R.M.Bateman, comb. nov. , A. picta (Loiseleur) R.M.Bateman, comb. nov. , Neotinea commutata (Todari) R.M.Bateman, comb. nov. , N. conica (Willdenow) R.M.Bateman, comb. nov. , Platanthera elegans Lindley ssp. maritima (Rydberg) R.M.Bateman, comb. nov. , P. elegans Lindley ssp. decurtata (R.Morgan & Glicenstein) R.M.Bateman, comb. nov. , P. elongata (Rydberg) R.M.Bateman, comb. nov. , P. michaelii (Greene) R.M.Bateman, comb. nov. , P. leptopetala (Rydberg) R.M.Bateman, comb. nov. , P. transversa (Suksdorf) R.M.Bateman, comb. nov. , P. cooperi (S.Watson) R.M.Bateman, comb. nov. , P. colemanii (R.Morgan & Glicenstein) R.M.Bateman, comb. nov. , P. candida (R.Morgan & Ackerman) R.M.Bateman, comb. nov. and P. yadonii (R.Morgan & Ackerman) R.M.Bateman, comb. nov. © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 142 , 1–40.     

Molecular systematics and phylogeny of the ‘Marbled Whites' (Lepidoptera: Nymphalidae,Satyrinae, Melanargia Meigen)

Abstract. We investigated genetic divergence and phylogenetic relationships amongst all known species of Palaearctic butterflies of the genus Melanargia using sequence information from three genes [mitochondrial cox1 barcode region (658 bp), ribosomal 16S rRNA (c. 518 bp), and nuclear wg (404 bp)]. Results show a lack of DNA divergence among several poorly characterized taxa, as well as deep divergences within and between others. We corroborated the molecular information with morphological and genitalic characters as well as with geographic data. We revise the taxonomy of Melanargia, and propose a new systematic scheme for the group. We revive some previous synonymies (M. lucasi meadwaldoi stat. rev. , M. ines fathme stat. rev. , M. ines jahandiezi stat. rev. , M. meridionalis tapaishanensis stat. rev. ), revise the status of some subspecies into species (M. transcaspica stat. nov. , M. lucida stat. nov. , M. wiskotti stat. nov. ) and of several species into subspecies of other taxa (M. evartianae sadjadii stat. nov. , M. larissa hylata stat. nov. , M. larissa grumi stat. nov. , M. larissa syriaca stat. nov. , M. larissa titea stat. nov. , M. lugens montana stat. nov. , M. epimede ganymedes stat. nov. ), revise the status of subspecies and transfer them to other species (M. larissa lorestanensis stat. nov. , M. larissa iranica stat. nov. , M. larissa karabagi stat. rev. , M. larissa kocaki stat. nov. , M. transcaspica eberti stat. nov. ), and propose new synonymies (M. larissa titea = M. titea standfussi syn. nov. = M. titea titania syn. nov. , M. leda leda = M. leda yunnana syn. nov. , M. lugens lugens = M. lugens ahyoui syn. nov. , M. lugens hengshanensis = M. lugens hoenei syn. nov. , M. halimede halimede = M. halimede gratiani syn. nov. , M. asiatica asiatica = M. asiatica dejeani syn. nov. , = M. asiatica elisa syn. nov. , = M. asiatica sigberti syn. nov. ).     

Phylogenetics and taxonomy of the New World leafy spurges,Euphorbia section Tithymalus (Euphorbiaceae)

The 480 species of leafy spurges, Euphorbia subgenus Esula, represent the main temperate radiation in the large genus Euphorbia. This group is distributed primarily in temperate Eurasia, but with smaller, disjunct centres of diversity in the mountains of the Old World tropics, in temperate southern Africa and in the New World. The majority of New World diversity (32 species) occurs in a single section, section Tithymalus. We analysed sequences of the nrITS and plastid ndhF, trnH‐psbA, trnS‐trnG and trnD‐trnT regions to reconstruct the phylogeny of section Tithymalus and to examine the origins and diversification of the species native to the New World. Our results indicate that the New World species of section Tithymalus form a clade that is sister to the widespread, weedy E. peplus. The New World species fall into two primary groups: a ‘northern annual clade’ from eastern North America and a diverse clade of both annual and perennial species that is divided into three subgroups. Within the second group, there is a small ‘southern annual clade’ from Texas and northern Mexico, a perennial ‘Brachycera clade’ from the western United States and northern Mexico, and a perennial ‘Esuliformis clade’ from montane areas of Mexico, Guatemala, Honduras and the Caribbean island of Hispaniola. Ancestral state reconstructions indicate that the annual habit probably evolved in the ancestor of E. peplus and the New World clade, with a subsequent reversal to the perennial habit. In conjunction with this phylogenetic framework, the New World species of section Tithymalus are comprehensively reviewed. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 191–228.     

Genetic differentiation without mimicry shift in a pair of hybridizing Heliconius species (Lepidoptera: Nymphalidae)

Butterflies in the genus Heliconius have undergone rapid adaptive radiation for warning patterns and mimicry, and are excellent models to study the mechanisms underlying diversification. In Heliconius, mimicry rings typically involve distantly related species, whereas closely related species often join different mimicry rings. Genetic and behavioural studies have n how reproductive isolation in many pairs of Heliconius taxa is largely mediated by natural and sexual selection on wing colour patterns. However, recent studies have uncovered new cases in which pairs of closely related species are near‐perfect mimics of each other. Here, we provide morphometric and genetic evidence for the coexistence of two closely related, hybridizing co‐mimetic species on the eastern slopes of the Andes, H. melpomene amaryllis and H. timareta ssp. nov. , which is described here as H. timareta thelxinoe . A joint analysis of multilocus genotyping and geometric morphometrics of wing shape shows a high level of differentiation between the two species, with only limited gene flow and mixing. Some degree of genetic mixing can be detected, but putative hybrids were rare, only one of 175 specimens being a clear hybrid. In contrast, we found phenotypic differentiation between populations of H. timareta thelxinoe , possibly indicative of strong selection for local mimicry in different communities. In this pair of species, the absence of breakdown of genetic isolation despite near‐identical wing patterns implies that factors other than wing patterns keep the two taxa apart, such as chemical or behavioural signals, or ecological adaptation along a strong altitudinal gradient. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 830–847.     

Systematics of the Miocene–Recent bryozoan genus Pentapora (Cheilostomata)

The four existing species of the ascophoran bryozoan Pentapora Fisher, 1807 are revised, and two new fossil species are introduced: Pentapora lacryma sp. nov. from the Pliocene Coralline Crag Formation of Suffolk, and Pentapora clipeus sp. nov. from the Pliocene of Emilia, Italy. The Arctic species Pentapora boreale Kuklinski & Hayward possesses a lyrula, does not belong in Pentapora, and is a junior synonym of Raymondcia rigida (Lorenz). The morphology of the autozooids is relatively uniform within the genus, and the main distinguishing characters are those of the ovicells and, particularly, the giant avicularia that are developed sporadically in all species apart from Pentapora foliacea, popularly known as ‘Ross coral’. A phylogenetic analysis based on skeletal characters returned a single shortest tree in which the three species of Pentapora from the North Atlantic (P. foliacea, Pentapora pertusa, and P. lacryma sp. nov. ) form a clade crownward of the three basal species from the Mediterranean (Pentapora ottomulleriana, Pentapora fascialis, and P. clipeus sp. nov. ). © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 17–39.     

An unusual edentulous pterosaur from the Early Cretaceous Romualdo Formation of Brazil

Numerous taxa make up the Early Cretaceous fauna of Brazil, including Ornithocheiroidea, Tapejaridae, Thalassodromidae, Chaoyangopteridae and a purported member of Azhdarchidae. Dsungaripteridae has only been tentatively assumed to be present in the form of ‘Santanadactylus’ spixi. New study of NMSG SAO 251093 (a specimen referred to Thalassodromeus sethi) suggests it is a previously unknown species of dsungaripterid, Banguela oberlii, tax. nov., differing from Thalassodromeus and other pterosaurs from the Early Cretaceous of Brazil by a unique combination of characters, including an upturned jaw tip, a short dorsal mandibular symphyseal shelf (dmss), and an autapomorphic thin crest placed halfway along the fused mandibular symphysis without a keel along the ventral margin of the jaw. B. oberlii, tax. nov., is referred to Dsungaripteridae based on a dmss no longer than the ventral shelf, U-shaped caudal margin of the ventral shelf and lateral margins of the mandibular symphysis concave in dorsal view. B. oberlii, tax. nov., is the youngest known dsungaripterid, and expands known morphological diversity in the clade as well as the Early Cretaceous pterosaur fauna of South America.     

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.     

Egg shape mimicry in parasitic cuckoos

Parasitic cuckoos lay their eggs in nests of host species. Rejection of cuckoo eggs by hosts has led to the evolution of egg mimicry by cuckoos, whereby their eggs mimic the colour and pattern of their host eggs to avoid egg recognition and rejection. There is also evidence of mimicry in egg size in some cuckoo–host systems, but currently it is unknown whether cuckoos can also mimic the egg shape of their hosts. In this study, we test whether there is evidence of mimicry in egg form (shape and size) in three species of Australian cuckoos: the fan‐tailed cuckoo Cacomantis flabelliformis, which exploits dome nesting hosts, the brush cuckoo Cacomantis variolosus, which exploits both dome and cup nesting hosts, and the pallid cuckoo Cuculus pallidus, which exploits cup nesting hosts. We found evidence of size mimicry and, for the first time, evidence of egg shape mimicry in two Australian cuckoo species (pallid cuckoo and brush cuckoo). Moreover, cuckoo–host egg similarity was higher for hosts with open nests than for hosts with closed nests. This finding fits well with theory, as it has been suggested that hosts with closed nests have more difficulty recognizing parasitic eggs than open nests, have lower rejection rates and thus exert lower selection for mimicry in cuckoos. This is the first evidence of mimicry in egg shape in a cuckoo–host system, suggesting that mimicry at different levels (size, shape, colour pattern) is evolving in concert. We also confirm the existence of egg size mimicry in cuckoo–host systems.     

Speciation in Caucasian lizards: climatic dissimilarity of the habitats is more important than isolation time

Given sufficient time and limited gene flow, evolutionary lineages tend to transform into separate species. Mechanisms preventing assimilation during repeated gene‐flow events include divergent adaptations and the development of pre‐ or postzygotic isolation. We analysed the morphological and genetic boundaries of three species of the rock lizard clade Darevskia ‘rudis’ (Darevskia rudis, Darevskia valentini, and Darevskia portschinskii) in relation to the environment, and tried to reconstruct evolutionary pathways underlying the observed separation among the species. We studied the geographic distribution of the scalation traits, microsatellite genotypes, and mitochondrial haplotypes. Our analyses showed consistent morphological and genetic patterns at the centre of the ranges for each species, but asymmetric distribution of alleles and scalation characters within the current contact zones among the species. The genetic and morphological diversification of the clade has been shaped during glacial isolation in an area of Southern Caucasus, away from the Black Sea Coast. The ancestral lineage of D. portschinskii separated from the common D. rudis–D. valentini lineage in the middle Pleistocene, and the two latter lineages separated in relatively recent geological time. Neither of the lineages attained complete lineage sorting; moreover, isolation and migration modelling have helped to detect recombinant gene flow from D. rudis to D. portschinskii (but not to D. valentini). This is most likely linked with climatically more similar suitable habitats between D. rudis and D. portschinskii than between D. valentini and the other two species. In itself, the isolation period was insufficient for the development of intrinsic isolation mechanisms in the system studied. Thus, differential landscape‐dependent selection within the contact zones is likely to have triggered the rapid development of isolation mechanisms. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 876–892.     

Molecular phylogenetic relationships reveal taxonomic and biogeographic clades in Dianella (flax lilies; Asphodelaceae,Hemerocallidoideae)

A phylogeny of Dianella is presented based on Bayesian and maximum parsimony analyses of a combined molecular data set using three chloroplast markers (trnQ^UUG–5'rps16, 3'rps16–5'trnK^(UUU) and rpl14–rps8–infA–rpl36) and two nuclear markers (ITS and ETS). Accessions included most Dianella species, including all species from Australia, the centre of diversity for the genus, and related outgroup genera Eccremis, Stypandra, Thelionema and Herpolirion. The phylogeny showed Stypandra sister to Herpolirion + Thelionema, and confirmed the monophyly of Dianella. Within Dianella, a number of clades were resolved that revealed biogeographic relationships. Accessions from south-western Australia (extending into South Australia) formed the earliest diverging clade, followed by D. serrulata from New Guinea, sister to all other clades of Dianella from Australia and other regions. Tropical North Queensland species, including the D. pavopennacea complex, were related to a clade of accessions from New Caledonia and the Hawaiian Islands in the Pacific, and a clade that included samples of D. carolinensis (Caroline Islands) and the widespread D. ensifolia from South-East Asia and across the Indian Ocean to Mauritius and Madagascar. However, D. ensifolia is not monophyletic, with accessions from Japan and Taiwan related to a clade of Queensland samples that are part of the D. revoluta complex. Three New Zealand species (diploid, 2n?=?16) were found to be related to Norfolk Island D. intermedia (type locality; octoploid, 2n?=?64). In contrast ‘D. intermedia’ from Lord Howe Island was resolved as sister to the eastern Australian D. caerulea complex. The phylogenetic results indicate the need for taxonomic revision, particularly revision of the species ‘complexes’ D. longifolia and D. caerulea in Australia, and recognition of more than one species within D. ensifolia and within D. sandwicensis on the Hawaiian Islands.     

GENE SEQUENCE DIVERSITY AND THE PHYLOGENETIC POSITION OF ALGAE ASSIGNED TO THE GENERA PHAEOPHILA AND OCHLOCHAETE (ULVOPHYCEAE,CHLOROPHYTA)1

The phylogenetic position of microfilamentous marine green algae assigned to the species Phaeophila dendroides, Entocladia tenuis (Phaeophila tenuis, and Ochlochaete hystrix was examined through phylogenetic analyses of nuclear‐encoded small subunit rDNA and chloroplast‐encoded tufA gene sequences. These analyses placed the P. dendroides strains within the Ulvophyceae, at the base of a clade that contains representatives of the families Ulvaceae, Ulvellaceae, and the species Bolbocoleon piliferum, supporting an earlier hypothesis that P. dendroides constitutes a distinct lineage. Substantial divergence in both nuclear and plastid DNA sequences exists among strains of P. dendroides from different geographic localities, but these isolated strains are morphologically indistinguishable. The lineage may have an accelerated rate of gene sequence evolution relative to other microfilamentous marine green algae. Entocladia tenuis and O. hystrix are placed neither in the P. dendroides clade nor in the Ulvellaceae as previous taxonomic schemes predicted but instead form a new clade or clades at the base of the Ulvaceae. Ruthnielsenia gen. nov. is proposed to accommodate Kylin's species, which cannot be placed in Entocladia (=Acrochaete), Phaeophila, or Ochlochaete. Ruthnielsenia tenuis (Kylin) comb. nov., previously known only from Atlantic coasts, is reported for the first time from the Pacific coast of North America (San Juan Island, WA, USA). Isolates of R. tenuis from the Atlantic and Pacific coasts of North America have identical small subunit rDNA and tufA gene sequences.     

Studies in the genus Hypericum L. (Clusiaceae). 1 Section 9. Hypericum sensu lato (part 3): Subsection 1. Hypericum series 2. Senanensia,subsection 2. Erecta and section 9b. Graveolentia

Abstract

Part 4(3) of this monographic series of papers on the genus Hypericum is prefaced by an introduction to the genus and a summary of the aims and methods of the project. This is followed by treatments of the remaining parts of sect. 9. Hypericum sensu stricto and the last segregate section from the original sect. Hypericum, sect. 9b. Graveolentia. Both hitherto untreated parts of the reduced sect. Hypericum are mainly Japanese, but some species extend in distribution as far as Kamchatka, eastern Siberia, central China, and Sabah (Mt. Kinabalu). Sect. Graveolentia is North and Central American. Sect. Hypericum subsect. Hypericum series Senanensia contains seven species from northern Japan and adjacent areas, including H. pibairense (Miyabe & Y. Kimura) N. Robson, stat. nov., H. nakaii subsp. miyabei (Y. Kimura) N. Robson, comb. et stat. nov., H. nakaii subsp. tatewakii (S. Watanabe) N. Robson, comb. et stat. nov. and H. senanense subsp. mutiloides (R. Keller) N. Robson, comb. et stat. nov. Sect. Hypericum subsect. Erecta contains 23 species and one hybrid from Japan, Korea, central China, Taiwan, Luzon, Sabah and Sumatera, including H. kawaranum N. Robson, stat. et nom. nov., H. watanabei N. Robson, stat. et nom. nov., H. kimurae N. Robson, stat. et nom. nov., H. pseudoerectum stat. et nom. nov., H. kitamense (Y. Kimura) N. Robson, stat. nov., H. kurodakeanum N. Robson, stat. et nom. nov., H. furusei N. Robson, sp. nov., H. nuporoense N. Robson, sp. nov. and H. ovalifolium subsp. hisauchii (Y. Kimura) N. Robson, stat. nov. Sect. Graveolentia contains nine species and one hybrid from southeastern Canada, the eastern half of the United States, Mexico and western Guatemala, including H. oaxacanum subsp. veracrucense N. Robson, subsp. nov. and H. macvaughii N. Robson, sp. nov.