On the systematic status of the desert plated lizard (Angolosaurus skoogi): phylogenetic inference from DNA sequence analysis of the African Gerrhosauridae

The desert plated lizard ( Angolosaurus skoogi ), a 'sand sea' endemic of the northern Namib Desert, exhibits remarkable morphological convergence with other dune-dwelling lizards worldwide. This distinct ecomorphic condition sets Angolosaurus apart from the remaining genera in the family Gerrhosauridae. Indeed, a morphological phylogeny addressing generic relationships within the Cordyliformes (Cordylidae + Gerrhosauridae) identified Angolosaurus as the earliest diverging taxon among African gerrhosaurids. We re-evaluated the basal status of Angolosaurus , conducting a molecular phylogenetic analysis of the African and Madagascan gerrhosaurid genera. Our survey involved a comprehensive species-level comparison among the four nominal genera of mainland Africa ( Angolosaurus , Cordylosaurus , Tetradactylus and Gerrhosaurus ). Mitochondrial DNA sequence data from the cytochrome b , ND2, 12S and 16S rRNA genes were combined for analysis using both parsimony and maximum likelihood procedures. In contrast to the morphological hypothesis, our results do not depict Angolosaurus as the sister taxon to other African gerrhosaurids. Rather, the molecular analyses consistently place Angolosaurus within Gerrhosaurus , rendering the latter genus paraphyletic.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 78 , 253–261.     

Simultaneous analysis of basal Hymenoptera (Insecta): introducing robust-choice sensitivity analysis

Molecular characters are analysed on their own and in combination with morphological data to examine the phylogenetic relationships of the basal lineages of Hymenoptera ('Symphyta'). This study covers 47 sawfly genera and nine apocritan families and includes molecular sequences from five genes − 12S, 16S, 18S and 28S ribosomal genes and cytochrome oxidase 1 − as well as 343 morphological characters. A robust-choice sensitivity analysis is performed with the data. First, the simultaneous analysis is repeated three times, each time employing a different step matrix for weighting the transformations of the molecular characters. Then, the results of all three simultaneous analyses are summarized in a strict consensus in order to avoid basing the conclusions on a narrow set of assumptions. This methodology is discussed in the paper. The relationships among superfamilies largely confirm previous hypotheses, being (Xyeloidea (Tenthredinoidea s.l. (Pamphilioidea (Cephoidea (Siricoidea (Xiphydrioidea (Orussoidea Apocrita))))))), where Siricoidea is understood as Siricidae+Anaxyelidae. However, the relationships within Tenthredinoidea s.s. proposed here are novel: ({Argidae Pergidae}[ Athalia {(Diprionidae Cimbicidae) Tenthredinidae minus Athalia }]).  © 2003 The Linnean Society of London . Biological Journal of the Linnean Society , 2003, 79, 245–275.     

Systematic relationships of theLactoridaceae,an endemic family of the Juan Fernandez Islands,Chile

The systematic relationships ofLactoridaceae are problematical, with alternative assignments toMagnoliales, Laurales, andPiperales. Phenetic analyses suggest thatLactoridaceae are best accomodated withinMagnoliales and are most closely related toAnnonaceae. Cladistic analyses indicate that the family is a relatively derived member of theMagnoliales, with affinities toAnnonaceae, Eupomatiaceae, Himantandraceae, andMyristicaceae. These analyses together with fossil pollen data are used to interpret the phylogeny ofLactoridaceae and its relatives.     

Phylogenetic analysis of non-stereotyped behavioural sequences with a successive event-pairing method

A new method is proposed which uses transitions among acts in non-stereotyped behavioural sequences as phylogenetic characters. This method is derived from the event-pairing method designed for the phylogenetic study of developmental sequences and from ethological analyses of transition matrices. It is applied to study the phylogenetic relationships within a well-known group, the presocial Zetoborinae cockroaches. The analysis is carried out with three data sets: a behavioural data set with transitions among acts in behavioural dyadic sequences, together with morphological and molecular data sets. Non-stereotyped behaviour proved to be phylogenetically informative and to display low homoplasy. This new method opens an avenue for studying the evolution of behaviour in the framework of phylogenetic analysis, which was restricted until now to the study of stereotyped sequences and/or isolated features involved in courting or building activities.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 853–867.     

Systematic revision of Myristicaceae (Magnoliales) in Madagascar, with four new species of Mauloutchia

The systematics of Malagasy Myristicaceae is revised to take into account new collections made since the work by Capuron in the 1970s, as well as improved knowledge from fieldwork and two scanning electron microscopy studies by the author. Four new species are described: Mauloutchia annickiae Sauquet, M. capuronii Sauquet, M. echinocarpa Capuron ex Sauquet, and M. sambiranensis (Capuron) Sauquet. In addition, basic information is given for each remaining Malagasy species of the family (synonymy, type specimen, updated distribution and main distinctive features). According to this treatment, Malagasy Myristicaceae now consist of four endemic genera and 15 species: Brochoneura Warb. (three species), Doyleanthus Sauquet (one species), Haematodendron Capuron (one species) and Mauloutchia (Baill.) Warb. (ten species). Two identification keys to these species are provided: one based primarily on fruit characters and one based primarily on male flower and inflorescence characters. Putative phylogenetic relationships among these species are also indicated, based on a previous combined morphological and molecular study by the author.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 351–368.     

Genetic discontinuities among populations of Cleistes (Orchidaceae, Vanilloideae) in North America

Genetic patterns were examined in five populations of North American Cleistes with three sources of molecular data: amplified fragment length polymorphisms (AFLP), DNA sequencing, and plastid microsatellites. Populations of C. bifaria were sampled in four areas of the south-eastern US: the coastal plains of Florida and North Carolina and the mountains of North Carolina and West Virginia. A population of C. divaricata sympatric with the North Carolina coastal plain C. bifaria was also sampled. Analysis of the three types of molecular data resulted in the same relationships among the five sampled populations. The coastal plain population of C. divaricata was consistently grouped with the C. bifaria populations from the mountains of West Virginia and North Carolina, and the two coastal plain populations of C. bifaria formed a separate group, results not supporting the existing concepts of species relationships. For future studies, greater sampling of C. divaricata populations and more detailed morphological and phenological studies are recommended for better characterization of the diversity within North American Cleistes . © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 87–95.     

Gynoecium diversity and systematics of the Magnoliales and winteroids

Carpel and ovule structure was compared in representatives of all 11 families of the Magnoliales (Annonaceae, Canellaceae, Degeneriaceae, Eupomatiaccae, Himantandraceae, Magnoliaceae, Myristicaceae) and winteroids (Austrobaileyaceae, Illiciaceae, Sehisandraceae, Winteraceae). Special attention was paid to features that are constant at family level. Bisexual flowers are always protogynous. In all representatives studied the carpels are closed at anthesis. Caipel closure is attained in three different ways: (1) postgenital fusion of inner surfaces (Degeneriaceae, Eupomatiaccae. Winteraceae), or (2) occlusion by secretion (Austrobaileyaceae, Sehisandraceae), or (3) a combination of (1) and (2): in Annonaceae, Canellaceae, Myristicaceae there is a conspicuous secretory canal in the innermost part of the ventral slit; in Illiciaceae and Magnoliaceae there is a narrow canal in the innermost part of the ventral slit; and in Himantandraceae the ventral slit is postgenilally fused in the style but completely open in the ovary. In most families the carpels have a double stigmalic crest or they have two tips in the transversal symmetry plane (i.e. at right angles to the median plane). Stigmas are unicellular papillate in most families but the papillae are bi-to multicellular (uniseriate) in Degeneriaceae and Eupomatiaceae. An unusual cryptic exlracarpellary compitum was found in Himantandraceae and Sehisandraceae. Intrusive oil cells were found in the carpel epidermis of Illiciaceae and Sehisandraceae. Mature ovules vary in length between 0.15 and 1.1 mm. The outer integument is fully annular (not semiannular) in Degeneriaceae, Himantandraceae, Canellaceae, Myristicaceae, and Illiciaceae. A rudimentary aril occurs in Canellaceae, and originates at the same site as in arillate Annonaceae and Myristicaceae. The results most strongly support an Annonaceae-Myristicaceae-Canellaceae alliance, to some degree also an Eupomatiaccac-Degeneriaceae-Himantandraceae-Magnoliaceae alliance, and an Illiciaceae-Schisandraceae-Winteraccae-Austrobaileyaceae alliance.     

Molecular systematics of North American phoxinin genera (Actinopterygii: Cyprinidae) inferred from mitochondrial 12S and 16S ribosomal RNA sequences

The cyprinid fish fauna of North America is relatively large, with approximately 300 species, and all but one of these are considered phoxinins. The phylogenetic relationships of the North American phoxinins continue to pose difficulties for systematists. Results of morphological analyses are not consistent owing to differences interpreting and coding characters. Herein, we present phylogenetic analyses of mitochondrial 12S and 16S ribosomal RNA sequence data for representatives of nearly all genera of North American phoxinins. The data were analysed using parsimony, weighted parsimony, maximum likelihood and bayesian analyses. Results from weighted parsimony, likelihood and the bayesian analysis are largely consistent as they all account for differing substitution rates between transitions and transversions. Several major clades within the fauna can be recognized and are strongly supported by all analyses. These include the western clade, creek chub–plagopterin clade and the open posterior myodome clade. The shiner clade is nested in the open posterior myodome clade and is the most species-rich clade of North American phoxinins. Relationships within this clade were not well resolved by our analyses. This may reflect the inability of the mitochondrial RNA genes to resolve recent speciation events or taxon sampling within the shiner clade.  © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society , 2003, 139 , 63–80.     

Evolutionary systematics of the Indian mouse Mus famulus Bonhote, 1898: molecular (DNA/DNA hybridization and 12S rRNA sequences) and morphological evidence

The genus Mus encompasses 38 species of mice divided into four subgenera: Mus , Pyromys , Nannomys and Coelomys . Each of these four taxa is characterized by discrete morphological as well as biochemical traits. We used two different molecular approaches to determine the relationships between these subgenera: DNA/DNA hybridization and 12S rRNA mitochondrial sequences. We compared the resulting phylogenies from each method and with phylogenies derived from morphological data. The degree of resolution of each molecular approach is discussed. The two molecular studies indicate that Mus , Pyromys , Nannomys and Coelomys are clearly distinct monophyletic groups, as previously indicated by morphological data and other biochemical and molecular approaches. There is one divergence between previous morphological and the molecular and morphological studies presented here: the position of the Indian species Mus famulus . This taxon, which was formerly included in the subgenus Coelomys , is demonstrated here to belong to the subgenus Mus. We also propose the following relationships within Mus sensu lato : Mus and Pyromys are the closest relatives, followed by Nannomys and Coelomys , whose relationships are still unclear. This arrangement is more robustly supported by DNA/DNA hybridization than by 12S rRNA data. A molecular time scale for the evolution within Mus sensu lato is proposed, using as a reference the Mus/Rattus divergence estimated by the fossil record at around 12 mya.  © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society , 2003, 137 , 385–401.     

Systematics of the tribe Podalyrieae (Fabaceae) based on DNA, morphological and chemical data

Phylogenetic relationships among nine genera and 28 species of the southern African tribe Podalyrieae were estimated from sequences of the internal transcribed spacer (ITS) of nuclear ribosomal DNA as well as morphological and chemical data. Morphological and ITS sequence data produced cladograms with similar topologies, both supporting the monophyly of Podalyrieae (excluding Hypocalyptus ). The combined data sets indicate that subtribe Xiphothecinae are monophyletic, but embedded within Podalyriinae. The high degree of congruence between previous taxonomic hypotheses and those based on DNA data provides further evidence for the utility of ITS sequences in studying phylogeny.  © 2002 The Linnean Society of London , Botanical Journal of the Linnean Society , 2002, 139 , 159–170.     

The systematic relationships of glucosinolate-producing plants and related families: a cladistic investigation based on morphological and molecular characters

A cladistic analysis is performed using 94 morphological and biochemical characters for 42 genera to compare a phylogeny based on morphological data with those obtained using different genes ( rbc L, atp B, 18S RNA, mat K) or their combination with morphological data, and to understand the floral evolution within the expanded Brassicales (Capparales) relative to Sapindales and Malvales. The tree produced with morphological data is congruent with those obtained from macromolecular studies in obtaining a well-supported glucosinolate-producing clade and an expanded Sapindales. The combined analysis of the morphological and molecular characters is generally well resolved with support for many of the relationships. The inclusion of the fossil taxon Dressiantha demonstrates the value of inserting fossil evidence in phylogenetic analyses. However, the fossil appears to be related to the Anacardiaceae and not to the Brassicales. The core Brassicales are well supported by a number of synapomorphies, although the internal position of Tovariaceae and Pentadiplandraceae is not well resolved. Emblingiaceae appears to be related to Bataceae and Salvadoraceae. Several significant morphological characters are mapped on the combined trees and their evolutionary significance is discussed. Within Brassicales and Sapindales several well supported clades can be recognized which merit ordinal or subordinal status, putting the present orders at a higher level; these include: Tropaeolales, Setchellanthales, Batidales, Brassicales (Brassiciflorae), Burserales, Sapindales and Rutales (Sapindiflorae). The present scheme of affinities within the Brassicales corresponds well with a gradual morphological evolution in the order.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 453–494.     

A morphological phylogeny for four families of amblyceran lice (Phthiraptera: Amblycera: Menoponidae, Boopiidae, Laemobothriidae, Ricinidae)

The suborder Amblycera (Insecta: Phthiraptera) comprises seven recognized families of parasitic lice. Three of these families (the Menoponidae, Laemobothriidae and Ricinidae) are present on a wide range of avian hosts. The four remaining families are restricted to a small section of mammals (the Boopiidae are parasites of Australian and New Guinean marsupials, and the Gyropidae, Trimenoponidae and Abrocomophagidae parasitize South and Central American rodents). This study uses a morphological approach to examine the evolutionary relationships between the genera from four amblyceran families: the Menoponidae, Boopiidae, Laemobothriidae and Ricinidae. Genera are represented by exemplars and a total of 44 louse taxa and one outgroup taxon were included. A cladistic analysis of 147 unordered characters recovered six equally parsimonious trees. Bootstrap, jackknife and Bremer support analyses were undertaken to assess the level of support for each resolved node in the strict consensus topology. Strong support was found for deep branch relationships between the families and in some cases for supra-generic groupings within families. The clades present in the strict consensus tree are discussed with reference to supra-generic and interfamily relationships, character choice, morphological convergence and host distribution. This study is the first phylogeny presented solely for amblyceran genera.  © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society , 2003, 138 , 39–82.     

Molecules,morphology, missing data and the phylogenetic position of a recently extinct madtom catfish (Actinopterygii: Ictaluridae)

Taxa missing large amounts of data pose challenges that may hinder the recovery of a well‐resolved, accurate phylogeny and leave questions surrounding their phylogenetic position. Systematists commonly have to contend with one or two species in a group for which there is little or no material available suitable for recovering molecular data. It is unclear whether these taxa can be better placed using analyses based on morphological data only, or should be included in broader analyses based on both morphological and molecular data. The extinct madtom catfish Noturus trautmani is known from few specimens for which molecular data are unavailable. We included this taxon in parsimony and Bayesian analyses of relationships of madtom catfishes based on a combination of morphological and molecular data. Results indicate that using a combination of morphological and molecular data does a better job at providing a phylogenetic placement for N. trautmani than morphology alone, even though it is missing all of its molecular characters. We provide a novel hypothesis of relationships among Noturus species and recommendations for classification within the group. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 60–75.     

The utility of morphological characters in gastropod phylogenetics: an example from the Calyptraeidae

Organismal taxonomy is often based on a single or a small number of morphological characters. When they are morphologically simple or known to be plastic, we may not have great confidence in the taxonomic conclusions of analyses based on these characters. For example, calyptraeid gastropod shells are well known for their simplicity and plasticity, and appear to be subject to frequent evolutionary convergences, but are nevertheless the basis for calyptraeid taxonomy. In a case like this, knowing how the pattern of relationships inferred from morphological features used in traditional taxonomy compares to the patterns of relationships inferred from other morphological characters or DNA sequence data would be useful. In this paper, I examine the relative utility of traditional taxonomic characters (shell characters), anatomical characters and molecular characters for reconstructing the phylogeny of calyptraeid gastropods. The results of an ILD test and comparisons of the recovered tree topologies suggest that there is conflict between the DNA sequence data and the morphological data. Very few of the nodes recovered by the morphological data were recovered by any other dataset. Despite this conflict, the inclusion of morphological data increased the resolution and support of nodes in the topology recovered from a combined dataset. The RIs and CIs of the morphological data on the best estimate topology were not any worse than these indices for the other datasets. This analysis demonstrates that although analyses can be misled by these convergences if morphological characters are used alone, these characters contribute significantly to the combined dataset.  © 2003 The Linnean Society of London . Biological Journal of the Linnean Society , 2003, 78 , 541–593.     

Phylogenetic systematics of Erythronium (Liliaceae): morphological and molecular analyses

For a nearly complete set of species of Erythronium (Liliaceae), we examined two plastid loci (the rps16 intron and the 5′ trnK intron, excluding the matK exon), one nuclear locus (nrITS) and morphology to evaluate species relationships and that of Erythronium to Amana, the putatively most closely allied genus. A matrix of morphological characters was developed through observation of around 900 living and herbarium specimens; evolution of these was examined using character optimization on the combined (total‐evidence) tree. Parsimony methods were used to examine the morphological and molecular data sets produced, both separately and in combination, with Bayesian methods also used on the molecular data sets. These established that the genus is probably sister to Amana (although most analyses placed Amana inside Erythronium) and that Tulipa is sister to the pair of Erythronium and Amana. Within Erythronium, there are three strongly supported geographically distinct clades: (1) Eurasian and (2) eastern and (3) western North American. Separation of species in these three clades is less clear, particularly among the western North American taxa. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, ●●, ●●–●●.     

Sclerotesta morphology and its systematic implications in magnoliaceous seeds

Seeds of 101 species from 14 genera were observed using stereoscopic and scanning electron microscopy. Sclerotesta morphology is stable within the genera of Magnoliaceae. Two different morphological types are described according to features of the chalazal region, which have great value in classification and have been found only in Magnoliaceae. One is the pore type, characterized by being simple, observed in the relatively primitive taxa of this family, including Manglietia, Pachylarnax, Magnolia (19 species), Aromadendron, Talauma (eight species), Parakmeria (one species), Kmeria (one species), Elmerrillia and Liriodendron . The other one is the tube type, which is characterized by having a more complex structure consisting of a central hollow tube contained within a hole. This type was observed in relatively advanced taxa, including Manglietiastrum, Magnolia (15 species), Talauma (three species), Parakmeria (four species), Kmeria (one species), Alcimandra, Michelia, Paramichelia and Tsoongiodendron . Transitional types between these two were observed in some species of Magnolia . Chalazal region morphology, together with other useful sclerotesta characters, including seed size, shape, the raphal sinus and the external surface of the sclerotesta, may be used as diagnostic characteristics of genera, and even species in Magnoliaceae. A key to identify the different genera is supplied.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 407–424.     

Phylogenetic analysis of Nymphaeales using fast-evolving and noncoding chloroplast markers

The Nymphaeales (water-lilies and relatives) represent one of the earliest branching lineages of angiosperms and comprise about 70 aquatic species. Here, we present a comprehensive study of phylogenetic relationships within the Nymphaeales from a dataset containing 24 representatives of the order, including all currently recognized genera and all subgenera of the genus Nymphaea , plus 5 outgroup taxa. Nine different regions of the chloroplast genome − comprising spacers, group II introns, a group I intron, and a protein coding gene − were analysed. This resulted in a character matrix of 6597 positions and an additional 369 characters obtained from coded length mutations. Maximum parsimony and Bayesian analyses of the complete dataset yielded congruent, fully resolved and well-supported trees. Our data confirm the monophyly of the Cabombaceae but do not provide convincing support for the monophyly of Nymphaeaceae with respect to Nuphar . Moreover, the genus Nymphaea is inferred to be paraphyletic with respect to Ondinea , Victoria and Euryale . In fact, the Australian endemic Ondinea forms a highly supported clade with members of the Australian Nymphaea subgenus Anecphya . In addition, Victoria and Euryale are inferred to be closely related to a clade comprising all night-blooming water-lilies ( Nymphaea subgenera Hydrocallis and Lotos ). An experimental approach showed taxon sampling to be of influence on the nodes reconstructed in core Nymphaeaceae. The results underscore that more diverse genera, if not clearly known to be monophyletic, should be represented by all major lineages.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 154 , 141–163.     

Comparative osteology of the Danio (Cyprinidae: Ostariophysi) axial skeleton with comments on Danio relationships based on molecules and morphology

Danio is a diverse group of small, colourful and easily bred freshwater fishes native to Southern Asia. Biological interest in danios has increased in recent years because the zebrafish, Danio rerio , has become an important model organism, particularly for studies of vertebrate developmental biology and genetics. Though several phylogenetic studies of Danio have been done on a subset of Danio species, the resulting phylogenies conflict in detail. To examine the utility of osteology for systematics of this group at the species level, we studied the axial skeleton for 11 species of Danio. We analyse our morphological data alone and combined with DNA sequence data for five gene sequences generated in earlier phylogenetic studies. The axial skeleton of Danio exhibits 14 characters that prove useful in phylogenetic analysis. Both molecular and morphological data support the monophyly of the danios included in our analysis and both data sets support the monophyly of two subclades: a deep-bodied group and a slender-bodied group. Separate analysis of molecular and morphological data sets show that the molecular data resolves relationships within the slender subclade whereas the topology of the deep-bodied subclade is determined by morphological data.  © 2002 The Linnean Society of London. Zoological Journal of the Linnean Society , 2002, 135 , 529–546.     

Canarian land snail diversity: conflict between anatomical and molecular data on the phylogenetic placement of five new species of Napaeus (Gastropoda, Pulmonata, Enidae)

Five new species of land snail (family Enidae) are described from La Gomera (Canary Islands) of which the majority, on the basis of anatomy alone, could be incorporated within a new supraspecific taxon. In addition to the morphological study of these new species, a region of the 16S mitochondrial gene is sequenced from three of the new species and a range of species of Napaeus from within its two subgenera ( Napaeinus and Napaeus ) . There is a disparity between the morphological and preliminary molecular phylogenetic data. Possible explanations for this conflict are discussed, as well as the evolutionary relationships among these different taxa, and it is suggested that this group may be an excellent model for further studies of adaptation and diversification.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 169–187.     

Systematics and evolutionary relationships of the mountain lizard Liolaemus monticola (Liolaemini): how morphological and molecular evidence contributes to reveal hidden species diversity

The delimitation of species is a major issue in systematic biology and has been a re-emerging discipline in the last decade. A number of studies have shown that the use of multiple data sets is critical for the identification of cryptic species, particularly in groups with complex evolutionary histories. Liolaemus monticola is a montane lizard species distributed in central Chile (32°–42°S), with four described subspecies in a latitudinal gradient from north to south: L. m. monticola , L. m. chillanensis , L. monticola ssp. and L. m. villaricensis . In order to test the systematic status and phylogenetic relationships of the taxa included in the L. monticola group, we analysed morphological (morphometric and meristic) and molecular (allozyme and mitochondrial DNA) data sets. The results of the morphological analyses showed that meristic variables correctly assigned individuals with higher accuracy than did morphometric characters. The results of the analyses of allozyme data revealed eight diagnostic loci that are evidence for significant differences among the four L. monticola subspecies. Phylogenetic analyses with mitochondrial DNA data, including additional species, showed that the L. monticola group is polyphyletic. We postulate that the four current subspecies represent independent evolutionary lineages and must be raised to the specific level as L. monticola , L. chillanensis and L. villaricensis . The taxonomic status of the unnamed L. monticola ssp. remains unresolved, although we provide a preliminary proposal.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 635–650.