A new phylogenetic classification for the gymnophthalmid genera Cercosaura , Pantodactylus and Prionodactylus (Reptilia: Squamata)

Because of the poor state of knowledge of many of the gymnophthalmid genera, systematic revision is necessary to render the classification consistent with evolutionary history. To that end, I conducted a review of the species of three genera of the Cercosaurinae which appear to form a monophyletic group: Cercosaura , Pantodactylus , and Prionodactylus . Phylogenetic analysis of 61 morphological characters was conducted after specimens of all species were examined to evaluate the composition of each taxon. The phylogenetic reconstruction suggested that the genus Prionodactylus was paraphyletic. A new phylogenetic classification is proposed that synonymizes Pantodactylus and Prionodactylus with Cercosaura. Cercosaura is redefined to include 11 species and seven subspecies. A key is provided to distinguish among species.  © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society, 2003, 137 , 101−115.     

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.     

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.     

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.     

Review of morphological evidence on the phylogeny of basal Hymenoptera (Insecta), with a discussion of the ordering of characters

In a previous study of the phylogeny of basal Hymenoptera, Vilhelmsen (2001; Zool. J. Linn. Soc . 131 : 393–442) compiled an extensive morphological data matrix for a phylogenetic analysis of basal Hymenoptera, comprising 38 hymenopteran genera. In this study, his characters are revised. This results in a cladogram whose relationships largely agree with those proposed by Vilhelmsen, except that the relationships at the base of the Hymenoptera are unresolved. The revised data matrix is expanded by 17 sawfly and three apocritan taxa. Moreover, 112 new morphological characters from different parts of the larval and adult morphology are also added to the data matrix, including 82 from a recent study of the terminal abdominal segments of male Hymenoptera. The addition of the new characters leads to Xyelidae, again, being the sister-group of all other Hymenoptera. The relationships among the sawfly families as proposed by Vilhelmsen are confirmed, except that the relationships among Syntexis , Siricidae and Xiphydriidae + Vespina are unresolved and that the monophyly of Apocrita is not convincingly supported. A separate analysis is performed which includes all extant genera of Xyelidae. The internal phylogeny of Xyelidae is determined as (( Macroxyela Megaxyela ) Xyelecia ( Xyela Pleroneura )).  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79 , 209–243.     

Phylogenetic relationships of the enigmatic land snail genus Prestonella: the missing African element in the Gondwanan superfamily Orthalicoidea (Mollusca: Stylommatophora)

The land snail superfamily Orthalicoidea, although generally assumed to be of Gondwanan origin, is considered by the majority of recent authors to be absent from the African continent. However, two poorly-known African genera, Aillya and Prestonella , have historically been referred to the orthalicoid family Bulimulidae s.l. Anatomical study of Aillya has subsequently shown it to be morphologically distinct from the Bulimulidae and referable to a family of its own, outside the Orthalicoidea, but Prestonella has remained an enigmatic taxon of unknown affinity. Using molecular and morphological evidence, we demonstrate conclusively that Prestonella is indeed a member of the Bulimulidae s.l. We thus confirm that this family is represented in Africa, and that it has a classical disjunct, tri-continental southern distribution. Thus, either the origin of the family must at the least predate the separation of Africa and South America in the Mid Cretaceous (under a vicariance scenario) or there must have been subsequent dispersal between the isolated Gondwanan fragments. In view of the limited dispersal ability of terrestrial snails, we consider the former more likely. Anatomically, Prestonella exhibits many character states thought to be plesiomorphic, suggesting a relationship with the subfamily Bulimulinae. Bayesian analysis of nuclear DNA sequence data places it as sister group (posterior probability = 1.0) to an Australasian clade comprising Bothriembryon and Placostylus . However, taxon sampling within the Orthalicoidea is currently inadequate to permit meaningful resolution of subfamilial affinity using molecular data. Similarly, although those orthalicoid taxa for which molecular data are available comprise a well-supported clade, the relationships of this clade to other stylommatophoran clades remain unresolved.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 203–221.     

Phylogenetic analysis of Magnoliales and Myristicaceae based on multiple data sets: implications for character evolution

Magnoliales, consisting of six families of tropical to warm-temperate woody angiosperms, were long considered the most archaic order of flowering plants, but molecular analyses nest them among other eumagnoliids. Based on separate and combined analyses of a morphological matrix (115 characters) and multiple molecular data sets (seven variable chloroplast loci and five more conserved genes; 14 536 aligned nucleotides), phylogenetic relationships were investigated simultaneously within Magnoliales and Myristicaceae, using Laurales, Winterales, and Piperales as outgroups. Despite apparent conflicts among data sets, parsimony and maximum likelihood analyses of combined data converged towards a fully resolved and well-supported topology, consistent with higher-level molecular analyses except for the position of Magnoliaceae: Myristicaceae + (Magnoliaceae + (( Degeneria + Galbulimima ) + ( Eupomatia + Annonaceae))). Based on these results, we discuss morphological evolution in Magnoliales and show that several supposedly plesiomorphic traits are synapomorphies of Magnoliineae, the sister group of Myristicaceae (e.g. laminar stamens). Relationships within Annonaceae are also resolved with strong support ( Anaxagorea basal, then ambavioids). In contrast, resolution of relationships within Myristicaceae is difficult and still incomplete, due to a very low level of molecular divergence within the family and a long stem lineage. However, our data provide good evidence that Mauloutchia is nested among other Afro-Malagasy genera, contradicting the view that its androecium and pollen are plesiomorphic  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 125–186.     

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.     

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 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.     

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.     

Comparative morphology of the leaf epidermis in Schisandra (Schisandraceae)

The leaf epidermis of 127 samples representing ten species within Schisandra Michaux. and one species of the related genus, Kadsura Kaempf. ex Juss., was investigated using light and scanning electron microscopy. Many characters of the leaf epidermis in Schisandra , such as pattern of epidermal cells, type of stomata, shape of guard cell pairs and cuticular ornamentation, are usually constant within species and thus make good characters for studying the relationship between and within genera. A new character, rim number, of the outer stomatal rim in the genus is introduced. It is shown that double outer stomatal rims occur only in evergreen Schisandra species, whereas a single rim occurs in deciduous species. This character supports the classification of Schisandra into two subgenera based on habit and androecial organs. This classification is also supported by additional morphological and molecular taxonomic characters. Kadsura coccinea (Lem.) Smith A. C. is the most primitive taxon in the related genus Kadsura . The outer stomatal rim of this species also has double rims. Combined with morphological and molecular evidence, this suggests that Schisandra and Kadsura are closely related and may share a recent common ancestor.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 39–56.     

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.     

Isoetes fluitans sp. nov.: the identity of Spanish plants of 'I. longissimum'

Recent revision of North African specimens of Isoetes velata A. Braun and the closely related taxon I. longissimum Bory, together with Spanish material conventionally designated I. longissimum , suggests that the Spanish specimens constitute a new species, I. fluitans . This is described and illustrated. The North African taxon I. longissimum is probably not specifically distinct from I. velata .  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 231–236.     

A revision and phylogenetic analysis of Stapeliopsis (Apocynaceae)

A survey of morphological characters is carried out for Stapeliopsis . The information obtained from this is combined with molecular data from the plastid trn L-F DNA region and ITS1 of the nuclear encoded 18S−26S rRNA cistron, to obtain a hypothesis of the evolutionary relationships among the species. It is shown that Stapeliopsis is monophyletic in a combined molecular and morphological analysis. Stapeliopsis is sister to a clade containing Huernia , Orbea and Tromotriche . The species of Stapeliopsis group into two clades. One contains S. khamiesbergensis , S. neronis and S. urniflora , and this is highly supported. The remaining species fall into an unsupported clade in which S. exasperata is sister to the others. The genera Hermanschwartzia Plowes and Neopectinaria Plowes are rejected. It is shown that a synapomorphy for Stapeliopsis is the laterally flattened inner corona-lobes, which touch the anthers only at their bases. Eight species of Stapeliopsis are recognized, with no subgeneric divisions.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 125–155.     

What is the subtribe Glossonematinae (Apocynaceae: Asclepiadoideae)? A phylogenetic study based on cpDNA spacer

Evidence from the chloroplast trn T-L spacer, trn L intron and trn L-F spacer shows the subtribe Glossonematinae of the tribe Asclepiadeae, hitherto composed of the Arabian and North African genera Glossonema , Odontanthera and Solenostemma , not to be monophyletic. While the affinities of Solenostemma cannot be determined with certainty at present, molecular, karyological and morphological evidence suggests that Glossonema and Odontanthera are closely allied to Pentarrhinum , an African genus of five species, belonging to the Cynanchinae.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 139 , 145–158.     

Stockwellia quadrifida (Myrtaceae), a new Australian genus and species in the eucalypt group

A new genus and species in the eucalypt group of the Myrtaceae is described. Stockwellia quadrifida D.J. Carr, S.G.M. Carr & B.Hyland gen. et sp. nov. is a rainforest tree of restricted distribution on the Atherton Tableland, North Queensland, Australia. Molecular data suggest that it is the sister taxon to Eucalyptopsis and this is supported by morphological characters. The prolonged hypanthium and reduced perianth appear to be synapomorphies for these two genera and the circumscissile hypanthium a synapomorphy shared with Allosyncarpia . Stockwellia differs from Eucalyptopsis by the distinct, albeit reduced, perianth and the hypanthium splitting into four segments at anthesis. The relationship of these two genera indicates an historical biogeographical link between New Guinea and the Queensland wet tropics region.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 139 , 415–421.     

Morphological and molecular characterization ofAquaticheirospora and phylogenetics of Massarinaceae (Pleosporales)

A morphologically interesting hyphomycete was collected from submerged wood in a stream in Doi Suthep-Pui National Park, Thailand. It is described as Aquaticheirospora lignicola gen. and sp. nov., and is characterized by euseptate conidia with divergent arms, which are vertically inserted in different planes to a basal cell. The genus differs from other chirosporous genera in having synnematous conidioma and conidia that are produced on conidiogenous cells borne at the apices of synnemata. The morphological characterization of this new fungus is reported and compared with similar chirosporous genera. To investigate the teleomorphic and phylogenetic relationships of this new taxon, three different regions of the ribosomal gene [18S rDNA, 28S rDNA, and internal transcribed spacer (ITS) including 5.8S] were sequenced and analysed. The results of phylogenetic analyses based on 18S, 28S, and partial ITS including 5.8S rDNA, employing different tree-making methods, indicate that Aquaticheirospora lignicola is closely related to the ascomycetes family Massarinaceae (Order: Pleosporales). The Massarinaceae as currently circumscribed is monophyletic. Massarina australiensis and M. bipolaris , however, appear to belong to the Lophiostomataceae.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 283–296.