Bumblebee visitation and seedset in Melampyrum pratense and Viscaria vulgaris: heterospecific pollen and pollen limitation

Summary Fruiting and seed set in two bumblebee-pollinated herbs, Melampyrum pratense L. (annual, Scrophulariaceae) and Viscaria vulgaris Bernh. (perennial, Caryophyllaceae) were studied on a dry meadow in southwestern Sweden in June 1986 and 1988. Both species produced seeds by self-fertilization. In Melampyrum (homogamous) fruiting and seed set by selfing were much lower than by natural pollination; in Viscaria (protandrous) fruiting by selfing and by natural polination were similar, but seed set per flower was lower by selfing than by natural pollination. Sequential hand pollinations increased seed set in Melampyrum, but not in Viscaria. Thus, the number of pollinations is important for high seed set in Melampyrum, and number of pollen grains deposited one pollination is important for high seed set in Viscaria. Late flowering resulted in the production of fewer seeds in both species, although the visitation rate in pure Viscaria stands was sufficient, because of limited resources. Pollen was the limiting resource in Viscaria, because hand pollination increased natural seed set. In Melampyrum pollen was limiting in 1988 but so were consumable resources, because the seedset decreased with time despite hand pollination. Pure stands of Viscaria had sced set similar to plants in mixed stands (with Melampyrum and Rhinanthus), although plants in mixed stands received fewer visits. Many seeds produced late in the season are the result of self pollination; emasculated Viscaria flowers had a very low seedset late in the season. Pollen loads containing approximately 50% heterospecific grains did not affect seed set in either species. Application of heterospecific (Lupinus) pollen to receptive Viscaria styles 6 h before conspecific pollen did not affect seed set.     

Phylogenetic and biogeographical relationships of the Sander pikeperches (Percidae: Perciformes): patterns across North America and Eurasia

North America and Eurasia share several closely related taxa that diverged either from the breakup of the Laurasian supercontinent or later closures of land bridges. Their modern population structures were shaped in Pleistocene glacial refugia and via later expansion patterns, which are continuing. The pikeperch genus Sander contains five species – two in North America (S. canadensis and S. vitreus) and three in Eurasia (S. lucioperca, S. marinus, and S. volgensis) – whose evolutionary relationships and relative genetic diversities were previously unresolved, despite their fishery importance. This is the first analysis to include the enigmatic and rare sea pikeperch S. marinus, nuclear DNA sequences, and multiple mitochondrial DNA regions. Bayesian and maximum‐likelihood trees from three mitochondrial and three nuclear gene regions support the hypothesis that Sander diverged from its sister group Romanichthys/Zingel ～24.6 Mya. North American and Eurasian Sander then differentiated ～20.8 Mya, with the former diverging ～15.4 Mya, congruent with North American fossils dating to ～16.3–13.6 Mya. Modern Eurasian species date to ～13.8 Mya, with S. volgensis being basal and comprising the sister group to S. lucioperca and S. marinus, which diverged ～9.1 Mya. Genetic diversities of the North American species are higher than those in Eurasia, suggesting fewer Pleistocene glaciation bottlenecks. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 156–179.     

Taxonomy and molecular phylogeny of the Asian Paraleucophenga Hendel (Diptera,Drosophilidae)

This paper reports on nine Asian species of the genus Paraleucophenga, of which four are new to science: Paraleucophenga brevipenis sp. nov. , Paraleucophenga hirtipenis sp. nov. , Paraleucophenga longiseta sp. nov. , and Paraleucophenga tanydactylia sp. nov. We also report on a new synonym, Paraleucophenga shanyinensis Chen & Toda, 1994 syn. nov. A key to all of the species examined, based on morphological data, is provided, together with a ‘molecular’ key to seven Paraleudophenga species based on DNA sequence data of the mitochondrial NADH dehydrogenase subunit 2 (ND2) gene. The phylogenetic relationships among seven Paraleucophenga species are reconstructed based on DNA sequences of the mitochondrial ND2 gene, using two Leucophenga species as outgroups. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 615–629.     

A revised classification of subtribe Helianthinae (Asteraceae: Heliantheae) II. Derived lineages

Sequence data for internal transcribed spacer (ITS) and partial external transcribed spacer (ETS) regions were combined in a phylogenetic analysis with previously obtained plastid DNA restriction site data to provide a comprehensive molecular phylogenetic hypothesis for derived members of subtribe Helianthinae. Analyses of the two molecular datasets provided conflicting evidence on relationships among some groups, supporting the hypothesis that hybridization has played a significant role in the divergence of the subtribe. A revised generic‐level classification is presented that divides the approximately 350 species of the subtribe among 21 genera. The paraphyletic Viguiera is narrowed to embrace only the type species, V. dentata. Four newly described genera, Dendroviguiera, Gonzalezia, Heiseria and Sidneya, are composed of species formerly included in Viguiera. Aldama is expanded to include 118 species extending from southwestern North America and Mexico to South America. This requires 116 new combinations, including 58 that were recently transferred into Rhysolepis, which is a synonym of Aldama, based on molecular phylogenetic results. One species of Viguiera is transferred to Tithonia, and two combinations in Hymenostephium are validated. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 167 , 311–331.     

The genus Callicarpa (Lamiaceae) on Borneo

The Bornean species of Callicarpa (Lamiaceae) are revised. Twenty three species are recognized: of these, four are described as new ( C. argentii , C. coriacea , C. subaequalis and C. teneriflora ) and two are raised to species level from variety ( C. endertii and C. hispida ). All 23 species described are assessed for World Conservation Union (IUCN) conservation status. Distribution maps and an identification key are provided. The historic confusion between the genera Geunsia and Callicarpa is discussed. Geunsia is included as a synonym of Callicarpa based on the examination of morphological characters and molecular data. Geunsia is suggested to be nested within Callicarpa based on phylogenetic analyses of nuclear ribosomal internal transcribed spacer (nrITS), partial matK and trnD‐T sequences of 30 taxa using maximum parsimony. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 416–455.     

Neurocranial osteology and systematic relationships of Varanus (Megalania) prisca Owen, 1859 (Squamata: Varanidae)

The neurocranial osteology of the giant monitor lizard Varanus (Megalania) prisca Owen, 1859 is described in detail for the first time. Optimization of neurocranial characters onto phylogenetic topologies for varanoids, including Lanthanotus, Heloderma, and Varanus species nests V. prisca within an Indo‐Australian clade of Varanus on the basis of characters of the otic capsule. A sister‐taxon relationship between V. prisca and Varanus komodoensis Ouwens, 1912 is proposed based on apomorphies of the crista prootica, fenestra vestibuli, occipital recess, and supraoccipital. These results support a monophyletic clade of giant monitors among Indo‐Australian species, and unambiguously synonymize Megalania with Varanus at both generic and subgeneric levels. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 445–457.     

Phylogeography and phylogenetic relationships of Malagasy tree and ground boas

Three species of boid snakes are recognized in Madagascar, namely the genus Sanzinia (one species and two subspecies) and the genus Acrantophis (two species). In the present study, we studied the patterns of genetic variation of these species across Madagascar using a fragment of the mitochondrial 16S rRNA gene in 77 specimens. To support the phylogenetic relationships of the lineages identified, three further gene fragments (cytochrome b, 12S rRNA and c‐mos) were analyzed in a reduced but representative set of samples. The results obtained corroborate that the genus Sanzinia includes two highly divergent mitochondrial lineages that evolved independently from each other on the east versus the west side of Madagascar. Each of these lineages presents a further subdivision that separates northern from southern groups. The nuclear marker showed no variation among the Malagasy boas, indicating either very low substitution rates in this gene or relatively recent speciation events coupled with high mitochondrial substitution rates. Because the broad geographic sampling detected no admixture among haplotypic lineages within Sanzinia, it is hypothesized that these may represent distinct species. Deviant haplotypes of snakes morphologically similar to Acrantophis dumerili indicate that this taxon may be a complex of two species as well. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 640–652.     

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.     

Systematic and ecological wood anatomy of Neotropical Schefflera (Araliaceae), with an emphasis on the Didymopanax group

Wood anatomy has been investigated from 35 species belonging to the Neotropical clade of the polyphyletic genus Schefflera (Araliaceae), representing three of the five subgroups (Didymopanax, Crepinella and Sciodaphyllum). The species examined are rather uniform in their wood structure, sharing the presence of scalariform and simple perforation plates, septate fibres and scanty paratracheal axial parenchyma. The observed variation in many wood characters showed statistically significant differences relative to latitude, climate and, especially, vegetation types. In particular, the intervessel pits are larger in species from higher latitudes and in seasonally dry habitats than those from lower latitudes and rainforests. Latitudinal and ecological trends in the variation of vessel element lengths, bar numbers on perforation plates, intervessel pit sizes and ray widths may be at least partially explained as effects of adaptation to drier environments in the course of dispersal outside the Amazonian region and diversification in the Atlantic Forest subclade and the Savannic subclade within the Didymopanax group. The occurrence of a granular annulus on the intervessel pit membranes in S. chimantensis and S. sprucei (both of the Crepinella group) is the first record of this feature in Araliaceae. In comparisons of Neotropical Schefflera with the other major clades of Schefflera sensu lato, wood anatomical diversity is consistent with the polyphyly of this genus based on molecular phylogenetic analyses. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173 , 452–475.     

Multiple introgression events and range shifts in Schizocodon (Diapensiaceae) during the Pleistocene

Range shifts during the Pleistocene shaped the unique phylogeographical structures of numerous species. Accompanying species migration, sister taxa may have experienced multiple introgression events. Here, we report the signature of introgression events in multiple areas in Schizocodon, herbs endemic to Japan, using amplified fragment length polymorphism (AFLP) fingerprinting and plastid DNA haplotyping in 48 populations. Although the present distributions of S. soldanelloides and S. ilicifolius are mainly allopatric, the species share plastid DNA haplotypes in each region (north‐eastern, north‐central, south‐central and south‐western Japan); in contrast, the specific groups were highly supported by AFLP analyses. These results support the occurrence of multiple introgression events in Schizocodon. Notably, the disjunct plastid haplotypes found only in S. ilicifolius var. intercedens suggest complete plastid DNA replacement at local areas from S. soldanelloides into S. ilicifolius var. ilicifolius. Furthermore, we found that S. soldanelloides experienced range contraction and expansion during glacial and interglacial cycles based on mismatch distribution analysis and ecological niche modelling. Based on several pieces of evidence, our study supports the idea that historical range shifts associated with Pleistocene climatic oscillations favoured multiple and regional introgression events in Schizocodon. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173 , 46–63.     

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.     

The Sphyraena obtusata group (Perciformes: Sphyraenidae) with a description of a new species from southern Japan

Sphyraena iburiensis sp. nov. is described, and taxonomic reviews are provided for S. obtusata and S. pinguis. These species, characterized by having 2 gill rakers, are defined as the S. obtusata group. Sphyraena iburiensis, known only from the Pacific coast of southern Japan, is characterized by 8.5–9.5 scales above the lateral line, a single row of scales in the groove along the lower margin of the suborbital region from the posterior tip of the maxilla to below the eye (=suborbital groove) not covered with skin, 2 distinct longitudinal stripes on the lateral surface of the body when fresh (upper stripe usually lost in preserved specimens), the lower stripe reaching the caudal-fin base just below the lateral line. Sphyraena obtusata, distributed in the Indo-West Pacific, is characterized by 5–7.5 scales above the lateral line, a single row of scales in the suborbital groove covered with skin, 2 somewhat indistinct longitudinal stripes on the lateral surface of the body when fresh (upper stripe usually lost in preserved specimens), the lower stripe joining the lateral line midway between the end of the second dorsal-fin base and caudal peduncle and extending to the middle of the caudal-fin base. Sphyraena pinguis, distributed in the Indo-West Pacific, is characterized by 7.5–9.5 scales above the lateral line, a single row of scales in the suborbital groove not covered with skin, and a single longitudinal stripe on the lateral surface of the body joining the lateral line slightly before or just below the end of the second dorsal-fin base and extending to the middle of the caudal-fin base. Seven (S. aureoflammea, S. brachygnathos, S. flavicauda, S. grandisquamis, S. langsar, S. lineata, and S. strenua) and 2 (S. chrysotaenia and S. schlegelii) nominal species are regarded as junior synonyms of S. obtusata and S. pinguis, respectively. In addition, lectotypes are designated for S. flavicauda, S. langsar, S. lineata, and S. obtusata. A key to the three species of the S. obtusata group is provided.     

Phylogeny,phylogeography, and systematics of the American pea crab genus Calyptraeotheres Campos, 1990, inferred from molecular markers

We used mitochondrial cytochrome oxidase I (COI) and the large ribosomal subunit (16S) genes to establish evolutionary relationships amongst species of Calyptraeotheres, evaluate their usefulness as DNA‐barcoding genes, and assess molecular diversity at the population level within Calyptraeotheres garthi. Bayesian, maximum likelihood, and maximum parsimony phylogenies confirmed the monophyly of Calyptraeotheres, showing that the ancestor of C. garthi, Calyptraeotheres hernandezi, and Calyptraeotheres granti radiated after the formation of the Panamanian isthmus. This finding contradicts the austral/tropical hypothesis previously proposed based on morphological data. The COI and 16S distance matrices supported separation of species as well as the genera, and corroborated that DNA barcoding is a useful tool and complements the classical taxonomy in Pinnotheridae. Phylogenetic and genetic distance analyses suggested that C. hernandezi is a junior synonym of C. garthi. Finally, C. garthi did not show a population structure across its distribution range, and showed a pattern consistent with a recent population expansion event that began 230–300 Kya. © 2013 The Linnean Society of London     

Harvesting Betulaceae sequences from GenBank to generate a new chronogram for the family

Betulaceae, with 120–150 species in six genera, are a family of Fagales that occurs mainly in the Northern Hemisphere. Previous studies of the evolution of Alnus, Betula, Carpinus, Corylus, Ostrya and Ostryopsis have relied on a relatively small number of sequence data and molecular clocks with fixed‐point calibrations. We exploited GenBank to construct Betulaceae matrices of up to 900 sequence accessions and 9300 nucleotides of nuclear and plastid DNA; we also computed species consensus sequences to build 46‐ and 29‐species matrices that strike a balance between species sampling and nucleotide sampling. Trees were rooted on Ticodendraceae and Casuarinaceae, and divergence times were inferred under relaxed and strict molecular clocks, using alternative fossil constraints. The data support the traditional two subfamilies, Betuloideae (Alnus, Betula) and Coryloideae, and show that Ostryopsis is sister to Ostrya/Carpinus. The fossil record and molecular clocks calibrated with alternating fossils indicate that the stem lineage of Betulaceae dates back to the Upper Cretaceous, the two subfamilies to the Palaeocene and the most recent common ancestors of each of the living genera to the mid‐ to late Miocene. A substitution rate shift in Coryloideae between 25 and 15 Mya preceded the mid‐Miocene climatic optimum and may be linked to temperate niches that became available following the mid‐Miocene. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172 , 465–477.     

Competition for bumblebee visitation between Melampyrum pratense and Viscaria vulgaris with healthy and Ustilago-infected flowers

Summary We studied bumblebee foraging on two sympatrically and simultaneously flowering species, Melampyrum pratense (Scrophulariaceae) and Viscaria vulgaris (Caryophyllaceae) during the flowering season of Viscaria in south-west Sweden. We distinguished between healthy and Ustilago-infected Viscaria plants. Both species shared the main insect visitor, queens of Bombus hortorum, which collected nectar from both species but pollen from Melampyrum only. The pattern of visitation changed over the season: bumblebees preferred Viscaria early on, but changed to Melampyrum later in the season, probably because of the higher sugar content of Melampyrum nectar and the possibility of collecting both nectar and pollen from the same flower. Pollen collecting is probably of increasing importance since the need of pollen for the developing larvae will increase with time. Flowers of Viscaria received fewer visits in plots with other species than in pure Viscaria plots during one year and received more visits early than late in the season during both years. Melampyrum flowers received similar amounts of visits in mixed and pure environments. They also received more visits early than late, although this was probably a result of pollinator satiation since Melampyrum became very abundant with time. Ustilago-infected plants received far fewer visits but because of its long flowering time the proportion of open flowers receiving visits was still high. Viscaria flowers received significantly more visits than flowers of other species when bumblebees made heterospecific flower visits from Ustilago-infected plants; thus Ustilago spores were probably effectively dispersed from infected to healthy plants by the pollinators. The mechanism behind competition for pollination in this system was competition through pollinator preference, since the visitation rate to Viscaria actually decreased, but also competition through improper pollen transfer (grains of both species were found on the bodies of bumblebees) since the incidence of switching between the two species increased, probably resulting in an increased misplacement of conspecific pollen grains with time.     

Seed and capsule morphology of Iranian perennial species of Euphorbia (Euphorbiaceae) and its phylogenetic application

Of the 480 species of Euphorbia subgenus Esula, c. 290 occur in the Mediterranean and Irano‐Turanian regions. Turkey and Iran are the most species‐rich countries in Asia with 83 and 74 species, respectively. Following our previous paper on annual species of Iranian Euphorbia, we studied the quantitative and qualitative macro‐ and micromorphological traits of seeds and capsules of 47 perennial species, including E . ferdowsiana sp. nov. , E . sulphurea sp. nov. and E. glareosa, as a first report from Iran. A key for all Iranian perennial Euphorbia spp. based on seed and capsule morphology is provided. The phylogenetic relationships of Iranian species based on internal transcribed spacer (ITS) nuclear and ndhF plastid regions are updated and used for the characterization of the synapomorphies of each clade. Capsule shape, seed shape, seed surface and shape of the caruncle have been found to be homoplastic, whereas the presence or absence of granulate elements on seed surfaces represents a phylogenetically important trait for section delimitation. The capsule surface is synapomorphic for several sections, including Helioscopia (tuberculate‐verrucose), Myrsiniteae (vesiculate) and Esula (granulate), and seed shape is synapomorphic for sections Helioscopia (ellipsoidal), Myrsiniteae (ovoid‐quadrangular) and Herpetorrhizae (pseudo‐hexahedral). Reversals have also taken place in some features, including capsule surface (E. mazandaranica, E. altissima) and seed shape (E. densa, E. aleppica). It seems that ecarunculate seeds are plesiomorphic in sections Helioscopia (E. eriophora) and Herpetorrhizae (E. consanguinea and E. turczaninowii). © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 00 , 000–000.