On the monophyly of Dactylorhiza Necker ex Nevski (Orchidaceae): is Coeloglossum viride (L.) Hartman a Dactylorhiza?

Previous phylogenetic analyses of Orchidaceae subtribe Orchidinae resulted in the proposal to classify Coeloglossum viride (L.) Hartman within the genus Dactylorhiza in order to maintain its monophyly. In this paper, we report some results that contradict previous studies regarding the monophyly of the traditional Dactylorhiza and its phylogenetic relationship with Coeloglossum. Our results, which combine sequences of the internal and external transcribed spacers of the nuclear ribosomal DNA, support the monophyly of Dactylorhiza , with Coeloglossum being a sister clade. The position of C. viride in the phylogenetic tree, and the considerable morphological differences with respect to Dactylorhiza , incline us to retain both lineages as distinct genera.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 152 , 261–269.     

睡莲科的核型分析及其分类学位置的探讨

本文对睡莲科6属6种代表植物的核型进行了研究,并探讨了它的分类学位置。结果如下：莲2n＝16=9sm+4m+3st;王莲2n＝24=8sm+8m+8T,蓝睡莲2n＝28,可配成14对,染色体小,第l号染色体上有2条随体;萍蓬草2n＝34＝18m+16sm;芡实2n=58,可配成29对,染色体小,第l号染色体有2条随体,莼菜2n＝72,可配成36对,染色体按大小可分成大,中、小三个类别。除莲外,其它5种植物的核型为首次报道。莼菜的体细胞染色体数目2n＝72和国外报道的2n＝80不相一致。莲的染色体以及形态学特征和其它睡莲科分类群显著不同,可将其从睡莲科中独立出来,并成立莲科和莲目。原归属于睡莲科的分类群仍组成睡莲目,并分别置于莼菜科和睡莲科。     

Phylogenetic relationship among Libellula, Ladona and Plathemis (Odonata: Libellulidae) based on DNA sequence of mitochondrial 16S rRNA gene

Abstract. The type genus for the dragonfly family Libellulidae is Libellula. At present, Libellula s.l. includes twenty-nine species, whose distribution is largely Nearctic. Whether two other libellulid taxa, Ladona and Plathemis, should be considered synonyms of Libellula, subgenera of Libellula, or separate genera, has been a subject of intermittent debate for over a century. Earlier proposals concerning Ladona and Plathemis were based on a limited number of morphological characters and lacked rigorous phylogenetic analyses. Therefore, we used the DNA sequence of a portion of the mitochondrial 16S rRNA gene and parsimony, maximum likelihood and neighbour-joining analyses to explore whether Ladona and Plathemis are monophyletic lineages distinct from Libellula. We obtained ≈ 415 bp of DNA sequence from twenty-three taxa including thirteen species of Libellula s.s., all three recognized species of Ladona, the two species of Plathemis and representatives of four other libellulid genera. Tetragoneuria williamsoni (Odonata: Corduliidae) was included as the outgroup. Parsimony analysis suggested that Ladona and Plathemis are monophyletic lineages distinct from Libellula s.s. with a sister group relationship between Libellula and Ladona. The monophyly of Ladona, Plathemis and Libellula was supported in > 90% of bootstrap replications and in trees five to ten steps longer than the most parsimonious trees. Relationships inferred from maximum likelihood and neighbour-joining analyses also supported the monophyly of Ladona and Plathemis. The four other libellulid genera included in the study formed a monophyletic clade distinct from Libellula, Ladona and Plathemis. Based on our analysis, we propose that Ladona and Plathemis be considered either genera or subgenera within Libellulidae.     

Phylogeny of the Nymphaeaceae inferred from

The Nymphaeaceae are one of the most important taxa for understanding the origin and evolution of primitive angiosperms due to its basal position in the cladograms of the angiosperms recently given based both on several gene sequences and on morphological characters, but phylogenetic relationships within the family have not as yet been resolved. The sequences of nrDNA ITS region of 11 species representing seven genera of the Nymphaeaceae and one outgroup, Ceratophyllum demersum, were used to reconstruct the phylogeny of the family using PAUP4.0b4A. Three most parsimonious trees (Length=1125, CI=0.7618 and RI=0.7214) were obtained. In the consensus tree: (1) Nelumbo was basal to the other genera with a bootstrap value of 100% and occupied an isolated position, so it could be separated from the Nymphaeaceae and placed in its own family, Nelumbonaceae, and its own order, Nelumbonales; (2) Nuphar was monophyletic, nested in the basal position of clade II and was strongly supported to be the sister group of the other genera (excluding Nelumbo) of the Nymphaeaceae, suggesting that its traditional inclusion in the Nymphaeaceae should be maintained; (3) Cabomba and Brasenia were sister group and formed a subclade with a bootstrap value of 99%, indicating their close affinity; (4) Nymphaea was strongly supported to be the sister group of a subclade comprising Euryale and Victoria with a bootstrap value of 94%, indicating that they should be placed in the Nymphaeaceae.     

Phylogenetic relationships in Cyperus L. s.l. (Cyperaceae) inferred from plastid DNA sequence data

The phylogeny of Cyperus and allied genera has been reconstructed using cladistic analysis of plastid rbcL gene, rps16 intron, trnL intron, and trnL-F intergenic spacer sequence data in 40 species of tribe Cypereae. Cyperus s.s. as currently circumscribed is not monophyletic because ten cyperoid genera are embedded within it. Eucyperoid Cyperus species (with a C₃ anatomy, e.g. C. involucratus ) and the genera Courtoisina , Kyllingiella and Oxycaryum form a clade that is sister to a clade comprising chlorocyperoid species (with a C₄ anatomy, e.g. C. papyrus ) and the genera Alinula , Ascolepis , Kyllinga , Lipocarpha , Pycreus , Remirea and Sphaerocyperus . The position of two species is uncertain; C . tenellus is resolved in a clade together with Isolepis although with typical cyperoid spikelets, whereas I. humillima is not resolved near either Isolepis or Cyperus s . l . © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 138 , 145–153.     

Pollen ontogeny in Brasenia (Cabombaceae, Nymphaeales)

Brasenia is a monotypic genus sporadically distributed throughout the Americas, Asia, Australia, and Africa. It is one of eight genera that comprise the two families of Nymphaeales, or water lilies: Cabombaceae (Brasenia, Cabomba) and Nymphaeaceae (Victoria, Euryale, Nymphaea, Ondinea, Barclaya, Nuphar). Evidence from a range of studies indicates that Nymphaeales are among the most primitive angiosperms. Despite their phylogenetic utility, pollen developmental characters are not well known in Brasenia. This paper is the first to describe the complete pollen developmental sequence in Brasenia schreberi. Anthers at the microspore mother cell, tetrad, free microspore, and mature pollen grain stages were studied using combined scanning electron, transmission electron, and light microscopy. Both tetragonal and decussate tetrads have been identified in Brasenia, indicating successive microsporogenesis. The exine is tectate-columellate. The tetrad stage proceeds rapidly, and the infratectal columellae are the first exine elements to form. Development of the tectum and the foot layer is initiated later during the tetrad stage, with the tectum forming discontinuously. The endexine lamellae form during the free microspore stage, and their development varies in the apertural and non-apertural regions of the pollen wall. Degradation of the secretory tapetum also occurs during the free microspore stage. Unlike other water lilies, Brasenia is wind-pollinated, and several pollen characters appear to be correlated with this pollination syndrome. The adaptive significance of these characters, in contrast to those of the fly-pollinated genus Cabomba, has been considered. Brasenia does not produce pollenkitt nor develop tectal microchannels as does Cabomba. Instead, the discontinuity of the tectum reduces the amount of sporopollenin in the wall, which may allow for more effective wind dispersal. The importance of reassessing palynological characters in light of new ontogenetic data and the phylogenetic implications of this reevaluation are also discussed.     

Phylogeny of Labidodemas and the Holothuriidae (Holothuroidea: Aspidochirotida) as inferred from morphology

The Holothuriidae is one of the three established families within the large holothuroid order Aspidochirotida. The approximately 185 recognized species of this family are commonly classified in five nominal genera: Actinopyga , Bohadschia , Holothuria , Pearsonothuria and Labidodemas. Maximum parsimony analyses on morphological characters, as inferred from type and nontype material of the five genera, revealed that Labidodemas comprises highly derived species that arose from within the genus Holothuria . The paraphyletic status of the latter, large (148 assumed valid species) and morphologically diverse genus has recently been recognized and is here confirmed and discussed. Nevertheless, we adopt a Darwinian or eclectic classification for Labidodemas , which we retain at generic level within the Holothuriidae. We compare our phylogeny of the Holothuriidae with previous classifications of its genera and subgenera, and make suggestions concerning possible systematic changes.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 144 , 103−120.     

Comparative Leaf Anatomy of Nymphaeaceae (s. l.)

Leaf anatomy of six genera was investigated. The petiole of Brasenia schreberi possesses a pair of bundles, the other five possess closed, scattered vascular bundles, resembling those of the Monocotyledons. True vessels are absent. Air passages are present in the ground tissue of the petiole except for Nuphar pumilum. Leaves are all dorsiventral. Stomatal apparatuses of the Ranunculaceous type are generally confined to the upper surface of the lamina, but fewer stomata are found on the lower surface of the lamina of Nelumbo nucifera. The stomatal apparatuses of Nelumbo nucifera are of the Haplocheilic type in development, but of the Ranunculaceous type at maturity. Hydathodes and sclerenchymatous idioblasts are present only in Nelumbo nucifera. Glandular hairs and hydropotens occur on the lower surface of the lamina in all the investigated species except Nelumbo nucifera. Astrosclereids are present in all the taxa except Brasenia schreberi and Nelumbo nucifera. According to observations we consider that the genus Brasenia is a primitive one in Nymphaeaceae and it has a close phylogenetic relationship with Cabomba. They may be treated as an independent family, Cabombaceae, which is better included in the order Nymphaeales. Nymphaea, Nuphar, Euryale and Victoria are closely related to one another and should be retained in the family Nymphaeaceae. Nelumbo shows, specialized features distinct from those of the rest of the Nymphaeaceae. It should be segregatednot only as a distinct family but also as a distinct order, Nelumboales.     

Relationships among extant and fossil echimyids (Rodentia: Hystricognathi)

The echimyid rodents are the most diverse group of Neotropical hystricognaths, with approximately 40 extant and fossil genera. Craniodental characters are proposed in order to formulate hypotheses of phylogenetic relationships within the Echimyidae. A data matrix of 54 taxa and 50 characters is constructed and submitted to parsimony analyses using PAUP and WinClada programs. Analysis of the complete data set results in 47 448 most parsimonious trees 107 steps long. These trees are summarized in a strict consensus tree, which is taken as the main phylogenetic hypothesis resulting from this study. The monophyly of several currently recognized supraspecific taxa is not corroborated. These are: the subfamilies Eumysopinae, Echimyinae, Myocastorinae and Adelphomyinae; and the genera Proechimys , Echimys and Makalata . Conversely, the monophyly of Dactylomyinae and Trinomys is supported. New associations are proposed: (1) a clade comprising the extant Carterodon , Clyomys and Euryzygomatomys and the fossil Pampamys and Theridomysops placed at the base of the crown-group Echimyidae; (2) a clade uniting Proechimys , Hoplomys and Trinomys , which is the sister-taxon of (3) a clade including Mesomys , Lonchothrix , Myocastor and a clade with extant dactylomyines and echimyines and associated fossil taxa. Based on this phylogenetic hypothesis, patterns of tooth evolution in Echimyidae are discussed, and minimum ages for the divergence events within the family are estimated.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 142 , 445–477.     

睡莲科的系统发育:核糖体DNA ITS区序列证据

以金鱼藻Ceratophyllum demersum为外类群,使用PAUP4.0b4A软件对睡莲科Nymphaeaceae植物7属11个代表种的ITS区序列进行了系统发育分析。采用最大简约法分析获得了3个最简约树,步长为1125,一致性指数(CI)和维持性指数(RI)值分别为0.7618和0.7214。利用3个最简约树获取严格一致树。结果表明:(1)莲属Nelumbo位于系统树的基部,自展支持率为100%,可从睡莲科中独立出来成立莲科Nelumbonaceae和莲目Nelumbonales;(2)萍蓬草属Nuphar是一单系类群,位于分支Ⅱ的基部,并和睡莲科其他属(不包括莲属)植物聚在一起构成姐妹群,故萍蓬草属仍应置于睡莲科中;(3)水盾草属Cabomba和莼菜属Brasenia聚成一小支并构成姐妹群,自展支持率为99%,说明这两属之间亲缘关系较近;(4)睡莲属和芡实属Euryale、王莲属Victoria聚成一小支并构成姐妹群,自展支持率为94%,说明三者亲缘关系较近,仍应置于睡莲科中。     

Gnidia (Thymelaeaceae) is not monophyletic: taxonomic implications for Thymelaeoideae and a partial new generic taxonomy for Gnidia

We address the generic limits of Gnidia (Thymelaeaceae) through a phylogenetic analysis of nuclear ribosomal DNA internal transcribed spacer (ITS) and plastid rbcL, trnL intron and trnL‐F intergenic spacer regions. Maximum parsimony and Bayesian inference were used to produce trees and assess internal support. The most significant conclusion drawn from the molecular analysis is that Gnidia is polyphyletic as currently circumscribed, comprising at least four distinct lineages that are each related to other genera within Thymelaeoideae. Gnidia pinifolia and G. racemosa are members of a clade within which Struthiola is embedded; a second group of species allies with Drapetes as sister to Passerina; and a third lineage corresponds to the previously recognized genus Lasiosiphon. The remaining species of Gnidia included in this study are allied with the Australian genus Pimelea. The taxonomic implications of these findings are discussed in relation to the principle of monophyly. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 160 , 402–417.     

Pollen tube growth and the pollen‐tube pathway of Nymphaea odorata (Nymphaeaceae)

An important aspect of the evolution of carpel closure, or angiospermy, is the relationship between pollen tube growth patterns and internalization of the pollen‐tube pathway. True carpel closure, involving postgenital fusion of inner carpel margins, is inferred to have arisen once within the ancient order Nymphaeales, in the common ancestor of Nymphaeaceae. We studied pollen tube development, from pollination to fertilization, in a natural population of Nymphaea odorata, using hand pollinations and timed flower collections. Pollen germinates in stigmatic secretions within 15 min and pollen tubes enter subdermal transmitting tissue within an hour, following wide intercellular spaces towards the zone of postgenital fusion. At the zone of fusion they turn downwards to grow in narrow spaces between interlocked cells and then wander freely to ovules within ovarian secretions. The pollen‐tube pathway is 2–6 mm long and upper ovules are first penetrated 2.5 h after pollination. Pollen tubes grow at rates of approximately 1 mm/h whether in stigmatic fluid, transmitting tissues or ovarian secretions. Pollen‐tube pathways are structurally diverse across Nymphaeales, yet their pollen tubes have similar morphologies and rapid growth rates. This pattern suggests pollen tube growth innovations preceded and were essential for the evolution of complete carpel closure. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162 , 581–593.     

Phylogeny and biogeography of African Biomphalaria (Gastropoda: Planorbidae), with emphasis on endemic species of the great East African lakes

The pulmonate snails of the genus Biomphalaria are widely distributed in the tropics, and they are intermediate hosts of the digenean trematode Schistosoma mansoni that causes intestinal schistosomiasis in humans. Recent molecular evidence suggests that Biomphalaria originated in South America, and following a recent transatlantic migration colonized Africa, where it radiated into the currently recognized 12 species. In the present study we further investigate the internal phylogenetic relationships of African Biomphalaria with emphasis on the dispersal and speciation on the continent, especially in the Great Lakes in East Africa. Our results, based on 16S ribosomal DNA, cytochrome oxidase subunit I (COI), and ribosomal internal transcribed spacer I (ITS1), support the monophyly of an African clade with two separate lineages ( Biomphalaria pfeifferi / Biomphalaria camerunensis and the Nilotic species complex/ Biomphalaria angulosa ). Following the initial colonization of Africa, Biomphalaria spread towards the east where a later radiation occurred in the Lake Victoria basin and the Albertine Rift Valley Lakes. With further dispersal along the River Nile, additional speciation took place giving origin to the North-east African species Biomphalaria alexandrina . Our results present almost no support of the species groups of Mandahl-Barth (except for the pfeifferi group), which is in accordance with other molecular appraisals. Our results suggest that Biomphalaria stanleyi , which is endemic to Lake Albert, is not an ecophenotype of the continental B. pfeifferi as previously suggested by other molecular studies. B. angulosa is sequenced for the first time and it is inferred to have an important phylogenetic position as sister group to the Albertine Rift/Lake Victoria basin radiation.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 151 , 337–349.     

Phylogenetic relationships and evolution of Orbiniidae (Annelida, Polychaeta) based on molecular data

The phylogenetic relationships of orbiniid taxa were reconstructed based on sequence data of the mitochondrial 16S rRNA and nuclear 18S rRNA genes. Both genes were analysed separately and in combination using maximum likelihood, Bayesian inference and maximum parsimony. Regardless of the method used, a clade consisting of the investigated Orbiniidae, Methanoaricia dendrobranchiata and Questa was strongly supported by the 18S dataset. The analysis of the combined dataset suggests inclusion of M. dendrobranchiata within the Orbiniidae with close relationships to species of Orbinia and Phylo, rather than as a sister taxon to all other orbiniids. Evidence is given for the paraphyletic status of Leitoscoloplos , Naineris , Orbinia , Phylo and Scoloplos , which represent the most species-rich genera of the Orbiniidae. It is thus reasoned that the morphological characters presently used for genus diagnosis are not informative for cladistic analysis. No support is found for the hypothesis that taxa of the Protoariciinae represent juveniles of Orbiniinae. Instead, in the case of Protoaricia oerstedi , strong support for a progenetic origin is found.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 144 , 59−73.     

The exine ultrastructure of pollen grains in Gnetum (Gnetaceae) from China and its bearing on the relationship with the ANITA Group

Pollen grains of six species of Gnetum , G. parvifolium , G. hainanense , G. luofuense , G. pendulum , G. cleistostachyum and G. montanum , collected from China were examined using light, scanning and transmission electron microscopy. Pollen grains of Gnetum are subspheroidal or irregular-apolar, inaperturate, 11.21–22.44 µm in long axis and 9.34–20.47 µm in short axis. The exine surface is covered with spinules, 0.50(0.30–0.71) µm long spaced on average 1.12(0.81–1.46) µm apart. The exine is about 0.55 µm thick and comprises ectexine and endexine. The ectexine includes a thin tectum and an infratectal granular layer. The tectum protrudes outwards, forming the spinules. The endexine is composed of discontinuous lamellae, with lacunae between lamellae. The pollen grains of Gnetum are compared with those of Ephedra and Welwitschia , and also those of the ANITA Group of angiosperms, including Amborellaceae, Nymphaeales, Illiciales, Trimeniaceae and Austrobaileyaceae. The exine ultrastructures of Gnetum , Ephedra and Welwitschia are quite similar, consisting of tectum, granular layer and lamellated endexine. The exine ultrastructure of Gnetum is also similar to that of Nymphaea colorata (Nymphaeaceae) in the transitional region between the proximal and distal poles, but differs from that of Amborellaceae, Illicium religiosum (Illiciaceae), Schisandra (Schisandraceae), Trimeniaceae and Austrobaileyaceae. This comparison of exine ultrastructure provides new evidence for consideration of the relationship between Gnetum and the ANITA Group.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 415–425.     

Diversification of the Genus Anopheles and a Neotropical Clade from the Late Cretaceous

The Anopheles genus is a member of the Culicidae family and consists of approximately 460 recognized species. The genus is composed of 7 subgenera with diverse geographical distributions. Despite its huge medical importance, a consensus has not been reached on the phylogenetic relationships among Anopheles subgenera. We assembled a comprehensive dataset comprising the COI, COII and 5.8S rRNA genes and used maximum likelihood and Bayesian inference to estimate the phylogeny and divergence times of six out of the seven Anopheles subgenera. Our analysis reveals a monophyletic group composed of the three exclusively Neotropical subgenera, Stethomyia, Kerteszia and Nyssorhynchus, which began to diversify in the Late Cretaceous, at approximately 90 Ma. The inferred age of the last common ancestor of the Anopheles genus was ca. 110 Ma. The monophyly of all Anopheles subgenera was supported, although we failed to recover a significant level of statistical support for the monophyly of the Anopheles genus. The ages of the last common ancestors of the Neotropical clade and the Anopheles and Cellia subgenera were inferred to be at the Late Cretaceous (ca. 90 Ma). Our analysis failed to statistically support the monophyly of the Anopheles genus because of an unresolved polytomy between Bironella and A. squamifemur.     

Phylogeny and taxonomy of thegenera of south-western North American Euctenizinae trapdoor spidersand their relatives (Araneae: Mygalomorphae, Cyrtaucheniidae)

The primary focus of this paper is to evaluate the monophylyand intergeneric relationships of the cyrtaucheniid subfamily Euctenizinaeand to a lesser degree the monophyly of Cyrtaucheniidae. Using 71morphological characters scored for 29 mygalomorph taxa our cladisticanalysis shows that Cyrtaucheniidae is likely paraphyletic withrespect to the Domiothelina, the clade that comprises the Migidae,Actinopodidae, Ctenizidae, and Idiopidae. Together, the Domiothelinaand Cyrtaucheniidae have been treated as the Rastelloidina clade.A strict interpretation of rastelloid classification based on ourcladogram would require the establishment of four additional spiderfamilies. However, we choose to use informal names for these cladesso that these taxa can be validated by subsequent studies of mygalomorphphylogeny before formal names are introduced. The phylogenetic analysisalso serves as a vehicle for examining the patterns of homoplasyobserved in mygalomorphs. The secondary focus of this paper is ataxonomic revision of Euctenizinae genera from the south-westernUnited States that includes a key to its genera. The cyrtaucheniid genera Enrico and Astrosoga areconsidered junior synonyms of Eucteniza. Actinoxia and Nemesoides arejunior ­synonyms of Aptostichus . At present theNorth American Euctenizinae comprises these seven nominal genera: Eucteniza, Neoapachella gen.nov. ( Neoapachella rothi sp. nov. ), Myrmekiaphila,Entychides, Promyrmekiaphila, Apomastus gen. nov. ( Apomastusschlingeri sp. nov. ), and Aptostichus .  © 2002The Linnean Society of London . Zoological Journal of the LinneanSociety, 2002, 136 , 487−534     

Phylogeny and classification of Muscini (Diptera, Muscidae)

Worldwide in distribution, the tribe Muscini comprises 21 accepted genera and about 350 species. In the present study, a cladistic analysis based upon adult morphological characters is carried out in order to discuss the monophyly of the tribe and its genera, the intergeneric relationships and, in some cases, also the intrageneric relationships. As a result, Muscini is supported as a monophyletic tribe sister-group of Stomoxyini. Except for Morellia Robineau-Desvoidy, Curranosia Paterson, and Eudasyphora Townsend, all the remaining genera are monophyletic. The results are dubious for Polietes Rondani, which was then provisionally kept unchanged. Morellia was broadened to include the Neotropical endemic genera Parapyrellia Townsend, Trichomorellia Stein, and Xenomorellia Malloch. Therefore, a new classification is proposed for Morellia in which it is divided into four subgenera: Morellia s.s. , Parapyrellia , Trichomorellia , and Xenomorellia . Furthermore, the previously proposed subgenus Dasysterna Zimin is given new status as a genus; however, as it is preoccupied by Dasysterna Dejean, the new replacement name Ziminellia nom. nov. is proposed herewith. Eudasyphora was found to be a paraphyletic group relative to Dasyphora Robineau-Desvoidy; both genera are hence synonymized, and Dasyphora is classified in three subgenera: Dasyphora s.s. , Eudasyphora , and Rypellia Malloch. The analysis demonstrated that the traditional classification of Musca Linnaeus into subgenera is artificial and, moreover, that the use of characters from male genitalia could be strongly informative for classifying the genus in phylogeny-supported species groups. Finally, the new classification proposal for Muscini recognizes 18 genera and, furthermore, two undescribed genus-ranked taxa are indicated.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 149 , 493–532.