Morphological analysis, phylogenies and classification in Velloziaceae

A morphological (non-molecular) cladistic analysis of the Velloziaceae is presented. The terminal taxa are 47 species of Velloziaceae plus four taxa as outgroups: Acanthochlamys bracteata (Acanthochlamydaceae), Encholirium scrutor (Bromeliaceae), Thoracocarpus bissectus (Cyclanthaceae) and Pandanus racemosus (Pandanaceae). The species of Velloziaceae sampled represent a significant proportion of the morphological diversity of the family, including all recognized genera and sections. The analysis revealed two major groups within Velloziaceae, supported mainly by stomata, vascular bundles in the pedicel, aquiferous tissues, filaments, anthers, pollen, stigma, seeds, ploidy and chemical characters. Comparison between this and the previous phylogenetic hypotheses for Velloziaceae, together with the two conflicting current classifications, suggests that there is incompatibility concerning the support of current genera and sections. The only three groups supported in all phylogenies are Barbacenioideae sensu Menezes (= Barbacenia sensu Smith & Ayensu), Xerophyta sect. Barbacenioides and Xerophyta sect. Xerophyta sensu Smith & Ayensu.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 157–173.     

Phylogeny and classification of the Scopulini moths (Lepidoptera: Geometridae, Sterrhinae)

The phylogenetic relationships of the genera in the geometrid tribe Scopulini (Lepidoptera: Sterrhinae) were examined using 141 characters of adult morphology and ecology. The study material included 92 species, representing all previously recognized genera and covering the morphological variation and full geographical range of the tribe. The cladistic analysis resulted in 20 equally parsimonious trees and a strict consensus cladogram based on these was well resolved. A majority of the recovered synapomorphic characters have been used previously in the taxonomy of the tribe. However, many novel characters were found in the sclerotized structures of the thorax. Many previously recognized genera were found to be nonmonophyletic and based on the present revised, synapomorphy-based classification, the number of recovered genera is reduced considerably. Twenty new generic synonyms and 90 new or revived species combinations are proposed. Seven genera are considered valid, with the large genus Scopula Schrank including over 85% of all species in the tribe. The taxonomic history of the tribe is reviewed and the problems of earlier classifications are discussed. A key to the genera is presented, although an informal diagnosis is preferred. All recognized genera are illustrated and a revised world checklist of the Scopulini is presented.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 143 , 473−530.     

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.     

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.     

Phylogeny of the riodinid butterfly subtribe Theopeina (Lepidoptera: Riodinidae: Nymphidiini)

Abstract The almost exclusively Neotropical butterfly family Riodinidae is poorly represented in both ecological and systematic studies of Lepidoptera. A comparative morphological study of all seventy-five species in subtribe Theopeina (tribe Nymphidiini) yielded 104 characters, predominantly from wing pattern, male and female genitalia, and abdominal structures. All morphological characters and adults representing the range of wing pattern variation are illustrated. Phylogenetic analysis of the data produced a large number of most parsimonious cladograms, but the strict consensus of these, both when using equal weights and after successive weighting, is well resolved and the majority of terminal clades have high character and branch support. Theopeina is found to consist of five monophyletic genera, Protonymphidia , Archaeonympha , Calicosama , Behemothia and Theope (=  Parnes and Dinoplotis ), with the largest genus Theope containing thirteen monophyletic species groups, which are delineated to facilitate a discussion of broad evolutionary patterns in this morphologically diverse subtribe.     

Asymmetric female genitalia and other remarkable morphology in a new genus of cobweb spiders (Theridiidae, Araneae) from Madagascar

Symmetry is such a conspicuous feature of life that asymmetries draw our immediate attention. While not uncommon in bilateral organisms in general, asymmetry in spiders is rare. Here I report the first case of antisymmetry in external female genitalia in spiders, in the new genus Asygyna (Theridiidae: Araneae) from Madagascar. In the nearly 39 000 species of spiders described to date, the external structure of the female genitalia is symmetric. In entelegyne spiders paired external copulatory openings each lead to an internal copulatory duct, whose roughly symmetrical trajectories terminate in paired receptacles, the spermathecae. In Asygyna , here exemplified by two new species, A. huberi and A. coddingtoni , laterality is evident in the internal and external female genitalia. A single copulatory opening leads (either to the left or right depending on the individual) to a single copulatory duct with a distinctly asymmetric trajectory. The duct splits terminally shortly before entering the two spermathecae. The males are symmetric, but possibly only one palp can be used in copulation with each female. If adaptive, the selective forces behind this asymmetry are perplexing, as male access to females seems reduced. However, if males are plentiful, asymmetry may benefit the female by reducing insertion times and thus shortening copulation time, and by tightening her control over which males sire her offspring. Asygyna has a range of other bizarre sex-related morphologies, including prosomal pits and a well developed stridulatory mechanism in both sexes, a male proboscis, and simplified palps. A phylogenetic analysis, including 63 taxa and 242 morphological characters, places Asygyna in Pholcommatinae, sister to the enigmatic genus Carniella .  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 211–232.     

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.     

The secondary copulatory organ in female ground weta (Hemiandrus pallitarsis,Orthoptera: Anostostomatidae): a sexually selected device in females?

Secondary sexual devices in female insects, primarily abdominal modifications, appear to function as a means of thwarting coercive mating attempts by males or are, in rare cases, sexually selected adaptations. Female ground weta, Hemiandrus pallitarsis (Orthoptera: Anostostomatidae), have an elaborate elbowed device on the underside of the mid-abdomen. Experimental removal of this accessory organ shows that it does not increase the probability of mating as predicted by the thwart-copulation hypothesis. Instead, removal prevents copulation, thus demonstrating that it is a secondary copulatory device. The male attaches to the organ both at the beginning of copulation and at the end, when he positions himself to adhere a spermatophylax food gift onto the mid-ventral region of his mate. The female accessory organ does not function to manipulate eggs or larvae (females provide care to their single clutch of offspring) and is unlikely to be a copulatory structure that prevents hybridization. The great extent of the modification of the ventral abdominal segments of H. pallitarsis females compared to other Hemiandrus species is consistent with a history of sexual selection on the accessory organ. Taken together, these results and the finding that the length of the accessory organ of H. pallitarsis correlates with female fecundity, suggest that this structure evolved under sexual selection to acquire nuptial gifts from males.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 85 , 463–469.     

Phylogenetic relationships of the north-eastern Atlantic and Mediterranean blenniids

The phylogenetic relationships of 27 north-eastern Atlantic and Mediterranean blennioids are analysed based on a total of 1001 bp from a combined fragment of the 12S and 16S mitochondrial rDNA. The most relevant results with implications in current blenniid taxonomy are: (1) Lipophrys pholis and Lipophrys (=  Paralipophrys ) trigloides are included in a well-supported clade that by the rule of precedence must be named Lipophrys ; (2) the sister species of this clade are not the remaining species of the genus Lipophrys but instead a monotypic genus comprising Coryphoblennius galerita ; (3) the smaller species of Lipophrys were recovered in another well-supported and independent clade, which we propose to be recognized as Microlipophrys ; (4) although some authors included the genera Salaria and Lipophrys in a single group we have never recovered such a relationship. Instead, Salaria is more closely related to the genera Scartella and Parablennius ; (5) the genus Parablennius , which was never recovered as a monophyletic clade, is very diverse and may include several distinct lineages; (6) the relative position of Aidablennius sphynx casts some doubts on the currently recognized relationships between the different blenniid tribes. Meristic, morphological, behavioural and ecological characters support our results and are also discussed. The possible roles of the tropical West African coast and the Mediterranean in the diversification of blenniids are discussed.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 283–295.     

A morphometric and molecular study in Tortula subulata complex (Pottiaceae, Bryophyta)

Specimens belonging to Tortula subulata complex ( T. inermis , T. mucronifolia and T. subulata ) were analysed using a combination of morphometric methods based on quantitative characters [principal component analysis (PCA), discriminant analysis (DA); 76 samples)] and molecular methods (ITS1 – 5.8S rRNA gene and ITS2; 47 samples) to assess patterns of morphological and molecular differentiation within this complex of taxa. The study shows that four species can be recognized: T. mucronifolia , T. subulata , T. inermis and T. subulata var. angustata with bistratose border, which is elevated to the species rank as T. schimperi nom. nov. The most valuable quantitative characters for identification of these species are the strata number of the marginal laminal cells, the ratio of middle marginal laminal cell width/middle marginal laminal cell length, basal membrane of peristome length, middle laminal cell width and papillae number on the middle laminal cells. The internal transcribed spacer (ITS) data suggest that this group of taxa is not monophyletic, T. mucronifolia being close to Protobryum bryoides (= Tortula protobryoides ). The remaining species seem to be a monophyletic group, T. schimperi being the sister group of the clade composed by T. inermis and T. subulata . T. subulata is considered to be of high morphological variability, for which ITS sequences did not resolve the internal relationships.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 149 , 333–350.     

Phylogenetic taxonomy of the Cercosaurini (Squamata: Gymnophthalmidae), with new genera for species of Neusticurus and Proctoporus

The tribe Cercosaurini is one of the most poorly studied groups of the lizard family Gymnophthalmidae. Recent studies have suggested that two cercosauriine genera, Neusticurus and Proctoporus , are polyphyletic. The aim of the current study was to rectify the polyphyletic relationships and construct a phylogenetic taxonomy of the Cercosaurini that is congruent with evolutionary history. Neusticurus is divided into two genera, one of them new ( Potamites ), based on the clades recovered by molecular studies and previously discussed morphological data. Proctoporus is divided into three genera, one of which is new ( Petracola ), while an older name ( Riama ) is resurrected for another. All five genera are described and defined and taxonomic keys are presented. This study represents an important advance in rectifying the taxonomy of the Cercosaurini. Many other para- and polyphyletic genera remain in the Gymnophthalmidae and much future work on this group is warranted.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 143 , 405–416.     

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.     

Phylogenetic relationships of thorny catfishes (Siluriformes: Doradidae) inferred from molecular and morphological data

Doradidae is a putatively monophyletic group of South American freshwater catfishes containing 30 extant genera and 72 valid species. Only one study to date has attempted to estimate phylogenetic relationships among doradids. This morphological analysis partitioned species into two basal genera ( Wertheimeria and Francisodoras ) and a crown group of three subfamilies (Platydoradinae, Astrodoradinae and Doradinae) whose relationships were unresolved. No subsequent work has been done to resolve the subfamilial trichotomy or to assess whether postulated intergeneric relationships are accurate. We address this problem with complete sequences (2.5 kilobases, kb) of mitochondrial 12S and 16S rRNA genes and partial (1.3 kb) sequences of the nuclear elongation factor-1 alpha (EF1α) gene from representatives of 23 doradid genera (43 species) and 13 outgroups from additional siluriform families. Phylogenetic analysis of these data yields strong support for the monophyly of Doradidae and Astrodoradinae (as well as other relationships), but otherwise shows significant conflict with morphological results. A partial re-examination of published morphological data indicates that many characters may have been incorrectly polarized and many taxa have incorrect state assignments. Our results provide a framework for ongoing efforts to describe the species-level diversity of this poorly understood neotropical family.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 140 , 551–575.     

Phylogenetic relationships of the tribe Operophterini (Lepidoptera, Geometridae): a case study of the evolution of female flightlessness

A molecular phylogenetic analysis was conducted in order to reconstruct the evolution of female flightlessness in the geometrid tribe Operophterini (Lepidoptera, Geometridae, Larentiinae). DNA variation in four nuclear gene regions, segments D1 and D2 of 28S rRNA, elongation factor 1α , and wingless , was examined from 22 species representing seven tribes of Larentiinae and six outgroup species. Direct optimization was used to infer a phylogenetic hypothesis from the combined sequence data set. The results obtained confirmed that Operophterini (including Malacodea ) is a monophyletic group, and Perizomini is its sister group. Within Operophterini, the genus Malacodea is the sister group to the genera Operophtera and Epirrita , which form a monophyletic group. This relationship is also supported by morphological data. The results suggest that female flightlessness has evolved independently twice: first in the lineage of Malacodea and, for the second time, in the lineage of Operophtera after its separation from the lineage of Epirrita . An alternative reconstruction (i.e. recovery of flight ability in an ancestor of Epirrita ) appears unlikely for various reasons. The similarities shared by Epirrita with a basal representative of Perizomini, Perizoma didymatum , allow the proposal of a sequence of evolutionary events that has led to flightlessness. It is likely that the transition to female flightlessness in the two lineages of Operophterini occurred after the colonization of stable forest habitats, followed by the evolution of a specific set of permissive traits, including larval polyphagy, limited importance of adult feeding, and adult flight during the cold months of the season.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 241–252.     

Nanoa, an enigmatic new genus of pimoid spiders from western North America (Pimoidae, Araneae)

The spider genus Nanoa gen. nov. (Araneae, Pimoidae) is described to place Nanoa enana , a new species of pimoids from Western North America. Parsimony analysis of morphological characters provides support for the monophyly of Pimoa plus Nanoa and corroborates the monophyly of Pimoidae and of the clade Linyphiidae plus Pimoidae. © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 145 , 249–262.     

Higher phylogeny of zygaenid moths (Insecta: Lepidoptera) inferred from nuclear and mitochondrial sequence data and the evolution of larval cuticular cavities for chemical defence

Zygaenid moths are capable of releasing hydrogen cyanide in their defense by enzymatic break-down of cyanoglucosides, but only larvae of chalcosiine and zygaenine moths store cyanogenic compounds in cuticular cavities and thus are able to discharge defense droplets, which effectively deter potential predators. A previously proposed phylogeny of Zygaenidae hypothesized a sister group relationship of chalcosiine and zygaenine moths because of their similar larval defense system. Not all chalcosiine taxa possess cuticular cavities, however, and a comparable defense mechanism has been reported in larvae of the zygaenoid family Heterogynidae. Considering sequence data of seven molecular loci, the present study estimates the posterior probability of phylogenetic hypotheses explaining the occurrence of larval cuticular cavities. The molecular data confirm the previous exclusion of Himantopteridae from Zygaenidae and suggest their close affinity to Somabrachyidae. The sequence data also corroborate the recently proposed exclusion of the Phaudinae from the Zygaenidae, because this subfamily is recovered in a reasonably well supported species cluster consisting of members of the families Lacturidae, Limacodidae, Himantopteridae, and Somabrachyidae. We consequently agree to raise Phaudinae to family rank. Within Zygaenidae, the subfamilies Callizygaeninae, Chalcosiinae, and Procridinae most likely constitute a monophyletic group, which is sister to the Zygaeninae. Our results imply that cuticular cavities were probably present in the larvae of the most recent common ancestor of Zygaenidae. Heterogynidae cannot be confirmed as sister taxon to this family, but appear at the very first split of the Zygaenoidea, although with poor support. The specific pattern of taxa in the molecular phylogeny showing larval cuticular cavities opens the possibility that these structures could have been already present in the most recent common ancestor of the Zygaenoidea.     

A phylogenetic analysis of the genera of Lejeuneaceae (Hepaticae)

The Lejeuneaceae are the largest family of the liverworts (Hepaticae), with almost a thousand species in 91 currently accepted genera. We analysed phylogenetic relationships of 69 genera, representing all major subfamilies and tribes recognized in the family, by using 49 informative morphological characters (31 gametophytic, 18 sporophytic), one chemical character, and applying equal and successive weighting of characters and parsimony analysis. In all trees recovered, the Lejeuneaceae were monophyletic with Nipponolejeunea (subfam. Nipponolejeuneoideae) forming the basalmost lineage. The remaining genera clustered in two major groups, the monophyletic Lejeuneoideae (52 genera) and the paraphyletic Ptychanthoideae (16 genera). Within each, several multigeneric lineages corresponding in part to previously described taxa were recovered: the Acrolejeuneinae and Ptychanthinae clades in the Ptychanthoideae, and the Brachiolejeuneinae, Lejeuneeae and Tuyamaella–Cololejeunea clades in the Lejeuneoideae. Bryopteris , a genus sometimes treated as a separate family, was nested in the Ptychanthinae clade. The Tuyamaella–Cololejeunea lineage corresponded with three previously recognized subfamilies (Cololejeuneoideae, Myriocoleoideae and Tuyamaelloideae) and contained genera with neotenic features, in two subclades. These features seemed to have originated by multiple heterochronic events: single origins were detected for 'protonemal neoteny' and 'primary neoteny', whereas 'secondary neoteny' probably evolved twice. Relationships within the large Lejeuneeae clade (43 genera) remained largely unresolved, although several putative lineages were detected in majority rule trees. Additional characters such as DNA sequences may provide better phylogenetic resolution in this group.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 391–410.     

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.