Glands, muscles and genitalia. Morphological and phylogenetic implications of histological characters in the male genitalia of Fritillary butterflies (Lepidoptera: Nymphalidae: Argynnini)

The histology and internal morphology of the male valve in a broad sample of Fritillary butterflies are examined. A large internal ridge to which the extensor of the valve muscle fastens is probably an autapomorphy for the tribe Argynnini. A large glandular pocket is found in the valves of all examined Heliconiinae, but is missing in other examined Nymphalidae. The pocket shows considerable variation between the higher taxa. In the higher Fritillaries (subtribe Argynnina) the pocket probably has an effective closing mechanism allowing the male to store a considerable amount of glandular secretion. It has earlier been suggested that the secreta from the pocket glands are transferred to the female during copulation, where they probably have an antiaphrodisiac effect, preventing other males from mating a once mated female. All examined male Fritillaries also have glands in the anellus, the terminal membrane between the valves. In Boloria these glands are extremely well developed and abundant, a condition that is probably an autapomorphy for that genus. The obtained characters are tested against a known phylogeny of the tribe Argynnini.     

Evolution and phylogeny of the genus Natrix (Serpentes: Colubridae)

Some aspects of the natural history of snakes of the colubrid genus Natrix have been well studied. With their extensive European distribution and relative abundance, their ecology, reproduction and behaviour are well known. Yet other facets of their biology remain poorly understood. These include knowledge of Natrix phylogeny, hypotheses explaining the current distribution of the three extant members of the genus, and their evolution and relationships. In this study we used molecular data, the nucleotide sequences of four protein-coding mitochondrial genes (3806 bp total), to provide a well-supported phylogeny for the genus Natrix . With these molecular data, evidence from the fossil record, and knowledge of palaeogeological events, we used two approaches in designing a time scale which we used to date the major events in Natrix speciation and intraspecific variation. Our data strongly support a phylogeny for the genus in which N. maura is basal with N. natrix and N. tessellata being sister species. The calibrated molecular clock suggests that N. maura diverged from the common ancestor of the three species 18–27 mya and that N. natrix and N. tessellata diverged 13–22 mya. Although the ranges of these estimates are large they support an early Miocene to late Oligocene origin for the three species. Intraspecific divergence is estimated to have commenced 5.3, 6.0 and 6.7 mya with evolutionary rates of 1 : 1.25 : 1.35% per million years for N. maura, N. natrix and N. tessellata , respectively.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 127–143.     

A new species of the genus Lightiella: the first record of Cephalocarida (Crustacea) in Europe

A new species of Cephalocarida belonging to the genus Lightiella is described. Like all known species of Lightiella , the new species is characterized by reduction of trunk segment 8, which also lacks both pleura and thoracopods. The diagnostic characters of the species are: (1) one seta on the inner distal corner of the penultimate endopodal segment of second maxilla and thoracopods 1–5; (2) only one claw on the distal segment of the endopod of thoracopod 6. A cladistic analysis of 27 morphological characters was used to estimate the phylogeny of all species of Lightiella , with all other cephalocarid species used as outgroups. The discovery of this species in the Mediterranean fills a gap in the distribution of the genus and of the entire class.  © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society , 2006, 148 , 209–220.     

A new species of Hybanthus (Violaceae) from north-eastern Brazil

A new species of Hybanthus (Violaceae) from north-eastern Brazil is described and illustrated. Hybanthus verrucosus Paula-Souza is endemic to 'caatingas' of Bahia and Alagoas states and is very distinct from all other Brazilian species of the genus, for its anterior petal, which is covered with small wart-like outgrowths. Other distinctive characteristics of the new species and its relationship with a closely related species ( H. atropurpureus (A.St.-Hil.) Taub.) are discussed.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 141 , 503–506.     

A distinctive new species of Viola (Violaceae) from Yunnan, China

A new species of Viola (Violaceae) from north-western Yunnan, China, is described and illustrated. Viola dimorphophylla Y. S. Chen & Q. E. Yang sp. nov. is endemic to Zhongdian County, north-western Yunnan, and is very easily distinguishable from all other Chinese species of the genus by having obviously dimorphic leaves, with the basal ones being long petiolate, undivided and widely cordate, and the cauline ones sessile, linear and verticillate.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 149 , 115–119.     

A revision of Cerdia (Caryophyllaceae)

Cerdia is a poorly understood genus of minute cushiony plants endemic to Mexico that is traditionally placed in subfamily Paronychioideae, tribe Polycarpeae (Caryophyllaceae). Morphological observations and phylogenetic analysis of ITS DNA sequences were carried out to determine the number of species, heretofore controversial, that constitute the genus as well as its position within Caryophyllaceae. Samples of Cerdia were collected through its entire range of distribution and together with previous specimens were used in morphological analyses. Nineteen terminal taxa were considered in a phylogenetic analysis with representatives of the different lineages in Caryophyllaceae. Results indicated that Cerdia is a monotypic genus, including only one variable species, C .  virescens , but its phylogenetic affinities remain doubtful.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 152 , 1–13.     

Mitochondrial DNA phylogeny of the woodpecker genus Veniliornis (Picidae, Picinae) and related genera implies convergent evolution of plumage patterns

The woodpecker genus Veniliornis comprises 12 species, all restricted to the New World tropics. The seemingly distantly related genus Picoides is broadly distributed in Eurasia and North America with two putative species, P. lignarius and P. mixtus , occurring in South America. The two genera are clearly distinct with respect to general plumage colouration and patterning as well as habitat utilization and thus traditionally have been placed in different tribes. Phylogenetic analyses of mtDNA sequences from the COI and cyt b genes indicated that both genera are reciprocally paraphyletic. The two South American species of Picoides belong to a clade comprising most species of Veniliornis , but V. fumigatus of Central and north-western South America belongs to a clade comprising species of Picoides . The mtDNA tree also indicated that Veniliornis is not closely related to the genus Piculus, as is implicit in current classifications. Misclassifications involving Veniliornis at both the generic and tribal levels appear to result from convergent evolution of plumage traits in specific forest types. We infer that the common ancestor of Veniliornis entered South America approximately at the time the Isthmus of Panama was formed, and diversification within South America was rapid.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 611–624.     

Flies on thistles: support for synchronous speciation?

Synchronous speciation of hosts and herbivorous insects predicts a congruent topology of host and insect phylogenies and similar evolutionary ages of host and insect taxa. To test these predictions for the specialized herbivorous fly genus Urophora (Diptera: Tephritidae), we used three different approaches. (i) We generated a phylogenetic tree of 11 European Urophora species from allozyme data and constructed a phylogeny of their hosts from published sources. Superimposing the Urophora tree on the host-plant tree we found no evidence for general congruence. (ii) We correlated genetic distances (Nei distances) of the host plants vs. the genetic distances of associated Urophora species. Overall, the relationship was not positive. Nevertheless, for some pairs of Urophora species and host plants genetic distances were in the same order of magnitude. (iii) We collected allozyme data for pairs of thistle taxa and pairs of herbivores on thistles together with independent time estimates. With these data we calibrated a molecular clock. There was a non-linear relationship between phylogenetic age and genetic distance, rendering the dating of deep events in thistle–insect evolution difficult. Nevertheless the derived molecular clock showed that the split of insect taxa lagged behind the split of hosts.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 775–783.     

Phylogenetic analysis of Zygaenoidea small-subunit rRNA structural variation implies initial oligophagy on cyanogenic host plants in larvae of the moth genus Zygaena (Insecta: Lepidoptera)

Cyanogenesis, the release of toxic cyanide from living cells, plays an important role in the defence system of certain plant (e.g. Fabaceae) and animal (e.g. Zygaenidae) taxa. The larvae of a significant number of Zygaena moth species (Zygaenidae) preferentially feed on cyanogenic Fabaceae and some of them are able to sequester cyanogenic compounds of their host plants. Using secondary structure variation of the small-subunit rRNA, we tested the currently accepted evolutionary hypothesis explaining species diversification in the genus Zygaena . We derived secondary structures considering evidence from covariation patterns and thermodynamic folding and applied structural information in a phylogenetic analysis. Contrary to previous assumptions, our results suggest that the use of cyanogenic larval host plants is an ancient trait and that the ability to feed on cyanogenic plants was probably already present in the most recent common ancestor of Zygaena . The utilization of acyanogenic plants in Zygaena species appears to be the result of a single secondary, reverse, larval host-plant shift. © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society , 2006, 147 , 367–381.     

Phyllanthus section Choretropsis (Euphorbiaceae) in Brazil

Section Choretropsis genus Phyllanthus (Euphorbiaceae) is formed by plants whose branches are modified into phylloclades. Despite the suitable characteristics of phylloclades within the Phyllanthus genus, the systematics of these species are poorly understood. Morphological data are presented here to allow future revision of this taxonomic group. The section is represented by nine species distributed in South America, eight of them endemic to Brazil. Our taxonomic studies distinguish the species and propose the synonymization of nine varieties of P. klotzschianus and two varieties of P. flagelliformis based on vegetative and reproductive characters. The diversity of branching and the morphology of phylloclades are discussed.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 150 , 131–164.     

Phylogeny, biogeography, and recurrent evolution of divergent bill types in the nectar-stealing flowerpiercers (Thraupini: Diglossa and Diglossopis)

Neotropical tanagers known as flowerpiercers ( Diglossa and Diglossopis ) have a novel feeding adaptation, comprising a downward curved hook on the maxilla that allows these species to obtain floral nectar without pollination. Using mitochondrial DNA sequences, the phylogenetic relationships of all 18 species of flowerpiercers were studied for the first time. Strong support was found for the monophyly of flowerpiercers and for the monophyly of four superspecies within flowerpiercers. However, previously described species-groups, as well as the genus Diglossopis , are not monophyletic. The biogeographic origin of flowerpiercers was identified as Andean, with a single dispersal event from the northern Andes to Central America and a single dispersal event from the northern Andes to the tepuis. The first principal component, representing a contrast between hook size and bill size, was mapped onto the phylogeny to examine the evolution of relative hook size in the group. Across the phylogeny, a relatively large hook and a relatively small hook evolved multiple times in unrelated lineages, indicating lability in bill morphology. Differences in hook size among sympatric species, together with habitat partitioning and behavioural differences, can explain the coexistence of multiple species of flowerpiercers at the same locality.   © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 14–28.     

Molecular phylogeny of shrimps from the genus Lysmata (Caridea: Hippolytidae): the evolutionary origins of protandric simultaneous hermaphroditism and social monogamy

Shrimps from the genus Lysmata are known because of their wide diversity of lifestyles, mating systems, symbiotic partnerships, and conspicuous coloration. They can occur in crowds (large aggregations), in small groups, or as socially monogamous pairs. Shrimps from this genus are rare, if not unique among crustaceans, because of their unusual sexual system. To date, the sexual system of all species investigated comprises a protandric simultaneous hermaphroditism: shrimps initially mature and reproduce as males and later in life turn into functional simultaneous hermaphrodites. The evolutionary relationships of the species within the genus are unsettled. A molecular phylogeny of the group may shed light on the evolutionary origins of the peculiar sexual and social systems of these shrimps and help resolve standing taxonomic questions long overdue. Using a 647-bp alignment of the 16S rRNA mitochondrial DNA, we examined the phylogenetic relationship of 21 species of shrimps from the genus Lysmata from several biogeographical regions; the Atlantic, Pacific, and Indo-Pacific. The resulting phylogeny indicates that the genus is paraphyletic and includes the genus Exhippolysmata . The constituent species are subdivided into three well supported clades: one group exclusively composed of neotropical species; a second clade comprising the Indo-Pacific and Atlantic symbiotic fish cleaner shrimps; and a third clade including tropical and temperate species from the Atlantic and Pacific. The molecular phylogeny presented here does not support a historical contingency hypothesis, previously proposed to explain the origins of protandric simultaneous hermaphroditism within the genus. Furthermore, the present study shows that monogamous pair-living is restricted to one monophyletic group of shrimps and therefore probably evolved only once.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 415–424.     

Phylogeny and species boundaries in the gobiid genus Gnatholepis (Teleostei: Perciformes)

Species of the goby genus Gnatholepis Bleeker, 1874, are common inhabitants of shallow tropical seas worldwide. In this study, mitochondrial DNA sequence (ND2 gene), from 349 Gnatholepis individuals sampled from across the South and Central Pacific and Caribbean, is used to infer phylogeny and determine species boundaries. Seven species of Gnatholepis are recognized: the Indo-Pacific G. anjerensis (Bleeker, 1851) [ G. cauerensis (Bleeker, 1853) is a synonym]; G. scapulostigma Herre, 1953; G. davaoensis Seale, 1910; G. knighti Jordan & Evermann, 1903; G. gymnocara Randall & Greenfield, 2001; G.  sp. Randall & Greenfield, 2001; and the Atlantic/Caribbean G. thompsoni Jordan, 1904. Results from the molecular phylogeny are compared with a previous morphology-based revision of the genus in order to establish which morphological characters diagnose species in correspondence with the molecular phylogeny.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 142 , 573–582.     

Chromosomes of four species in three genera of Commelinaceae from China and their systematic implications

Chromosomes of four species in three genera of Commelinaceae from China were examined. Spatholirion longifolium had 2 n  = 20. This small genus is considered to be of a possible polyploid origin based on x  = 5, a basic number shared by its closest ally, the monospecific genus Streptolirion , with its only species having 2 n  = 10. The two genera were found to be very different from each other in their karyotypic constitution. The differences in both chromosome number (2 n  = 20 vs. 10) and karyotypic constitution support their separation as two independent genera. The remaining three species, Amischotolype hispida , Porandra ramosa and P. scandens , all had 2 n  = 36. In view of the occasional occurrence of 2 n  = 18 in the African Coleotrype , a genus very close to Amischotolype and Porandra in gross morphology, these three species are all very probably polyploid based on x  = 9. The high degree of karyotypic similarity of the three genera also strongly indicates their close affinity and supports their placement in the subtribe Coleotrypinae. © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 152 , 399–403.     

Checklist of the Caesalpinioideae (Leguminosae) of Equatorial Guinea (Annobón, Bioko and Río Muni)

This study provides a checklist of the Caesalpinioideae (Leguminosae) present in Equatorial Guinea, comprising 52 genera and 124 taxa. Seven species are known from Annobón, 33 from Bioko and 109 from Río Muni. The best represented genus is Senna with eight species. In addition, bibliographic references for Caesalpinioideae (Leguminosae) from Equatorial Guinea have been gathered and checked. Fourteen species are included based on literature records, because their distribution ranges suggest they may occur in Equatorial Guinea, 11 introduced species could be naturalized, and 45 taxa are recorded for the first time from the country. This represents an increase of over 35% in the floristic knowledge of Caesalpinioideae from Equatorial Guinea. A statistical summary is presented at the end of the checklist.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 541–562.     

Island-like radiation of Saussurea (Asteraceae: Cardueae) triggered by uplifts of the Qinghai–Tibetan Plateau

Increasing evidence suggests that geological or climatic events in the past triggered the radiative diversification of both animals and plants on islands as well as continents. The Qinghai–Tibetan Plateau (QTP) has been extensively uplifted since the Miocene, but there is little information on possible links between these events and biological diversification in this and adjacent regions. Partly to explore such links, we have examined the diversification of Saussurea (Asteraceae: Cardueae), a species-rich genus that is mostly endemic to QTP, but also occurs in arid highlands elsewhere in the Northern Hemisphere. The phylogenetic analyses were conducted on the basis of the nuclear (internal transcribed spacer, ITS) and plastid ( trnL-F and psbA-trnH ) sequences from 55 species, representing 19 sections from all six subgenera of Saussurea , and species from 15 genera of the Cardueae. The results suggest that the currently circumscribed genus Saussurea ( s.l. ) is a polyphyletic group and that five sections should be excluded from the genus. Samples from the other 14 sections (representing five subgenera) clustered as a monophyletic group (here designated the Saussurea s.s. lineage, SSSL) with high statistical support. However, none of the analyses (nuclear, plastid or combined) resolved SSSL's infrageneric phylogeny, and the parallel clades of the lineage indicate that island-like adaptive radiation occurred. Furthermore, this radiation appears to have occurred 14–7 Mya, during the period of the major uplift events of QTP. Thus, our results support the hypothesis that geological events may play important roles in driving biological diversification through continental radiation.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 893–903.     

High photobiont diversity associated with the euryoecious lichen-forming ascomycete Lecanora rupicola (Lecanoraceae, Ascomycota)

The diversity and phylogenetic position of photobionts in the widespread saxicolous, crustose lichen-forming ascomycete Lecanora rupicola s.l. is presented. The algal partners of this lichen species complex belong to diverse and unrelated lineages in the genus Trebouxia . Specimens were sampled from different habitats and geographical origins. Either whole thallus DNA extractions or minute fragments of the algal layer of the lichen thallus were subjected to polymerase chain reaction, using primers that specifically amplify internal transcribed spacer rDNA of the photobionts. No correlations between different chemical races of L. rupicola with particular lineages of Trebouxia spp. were found. Irrespective of the different algal partners, all lichen thalli abundantly developed ascomata. L. rupicola apparently maintains full fecundity with a low degree of selectivity for photobionts, which promotes the occurrence of this lichen-forming species in ample ecological situations.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 283–293.     

The biogeography of the austral, subalpine genus Ourisia (Plantaginaceae) based on molecular phylogenetic evidence: South American origin and dispersal to New Zealand and Tasmania

Molecular phylogenetic analyses of 26 of the 28 species of Ourisia , including eight of ten subspecies and two purported natural hybrids, are presented and used to examine the biogeography of the genus, which is distributed in subalpine to alpine habitats of South America, New Zealand and Tasmania. Gondwanan vicariance, often cited as the cause of this classic austral biogeographical pattern, was rejected by parametric bootstrapping of our combined dataset. Alternatively, various lines of evidence are presented in favour of a South American origin of Ourisia and subsequent dispersal to Australasia. Specifically, the genus likely arose in the Andes of central Chile and spread to southern Chile and Argentina, to the north-central Andes, and finally to Tasmania and New Zealand. The ancestor of the New Zealand species probably first arrived on the South Island, where the New Zealand species of Ourisia are most diverse, and migrated to the North and Stewart Islands. Because the Tasmanian and New Zealand species are sister to one another, the direction of dispersal between these two areas is equivocal. These results agree with other molecular phylogenetic studies that show that past dispersal between southern hemisphere continents has played an important role in the evolutionary history of many high-elevation austral plants. Our data also show that within South America, many of the geographical barriers (with the exception of the Atacama Desert) that have played a role in the evolution of other plant groups have not affected Ourisia species. Within New Zealand, the phylogeny and biogeography of species of Ourisia coincide with the geological history of the country and patterns of other alpine plants. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 479–513.     

The phylogeny and taxonomy of Hippopotamidae (Mammalia: Artiodactyla): a review based on morphology and cladistic analysis

The phylogeny and taxonomy of the whole family Hippopotamidae is in need of reconsideration, the present confusion obstructing palaeoecology and palaeobiogeography studies of these Neogene mammals. The revision of the Hippopotamidae initiated here deals with the last 8 Myr of African and Asian species. The first thorough cladistic analysis of the family is presented here. The outcome of this analysis, including 37 morphological characters coded for 15 extant and fossil taxa, as well as non-coded features of mandibular morphology, was used to reconstruct broad outlines of hippo phylogeny. Distinct lineages within the paraphyletic genus Hexaprotodon are recognized and characterized. In order to harmonize taxonomy and phylogeny, two new genera are created. The genus name Choeropsis is re-validated for the extant Liberian hippo. The nomen Hexaprotodon is restricted to the fossil lineage mostly known in Asia, but also including at least one African species. The genus Hippopotamus is confirmed. These changes represent substantial advances for understanding the evolutionary history of the Hippopotamidae, and provide a new framework for future studies.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 143 , 1–26.