Phylogenetic relationships and neotenic evolution of Metzgeriopsis (Lejeuneaceae) based on chloroplast DNA sequences and morphology

The systematic position of the enigmatic Asiatic liverwort Metzgeriopsis pusilla (Marchantiophyta), the single thalloid member of the largest family of leafy liverworts, Lejeuneaceae, was investigated based on sequences of the rbc L gene and the trn L-F region of chloroplast DNA, and morphology. Metzgeriopsis is not phylogenetically isolated from other taxa of Lejeuneaceae, as previously suggested, but is nested within Cololejeunea . The two genera share many morphological features, especially in the sporophyte generation, which is newly described. Metzgeriopsis and Cololejeunea are members of the Tuyamaella – Cololejeunea clade, characterized by neotenic features that probably evolved as adaptations to ephemeral habitats in tropical rain forests. Three different types of neoteny occur in this clade, which apparently originated by multiple heterochronic events. Within Lejeuneaceae, protonemal neoteny is unique to Metzgeriopsis and apparently resulted by evolution from a primary neotenic ancestor. The occurrence of protonemal neoteny in bryophytes is briefly reviewed and two types are recognized: species having a persistent thalloid protonema and species having a persistent filamentous protonema. The thalloid type is very rare and exclusive to epiphyllous liverworts, whereas the filamentous type is much more common and is characteristic of many terrestrial bryophytes, including the rare genus Protocephalozia . The new combination Cololejeunea metzgeriopsis (K.I.Goebel) comb. nov. is proposed.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 293–308.     

A preliminary account of colura (Hepaticae,Lejeuneaceae) in Australia

Four species of Colura not previously reported from Australia are: C. acroloba (Mont.) Jovet-Ast, C. ari Steph., C. conica (Sande Lac.) Goeb., and Colura queenslandica sp. nov. A key to all ten species occurring in Australia is provided. Saccate lobules of C. simplicior are described and illustrated for the first time. The name Colura sect. Lingua Jovet-Ast ex Thiers is validated.     

苔纲(Hepaticae)植物雌性生殖苞的形态学研究

以比较形态解剖学方法,对苔纲 (Hepaticae) 植物雌性生殖苞的结构及其不同成分起源进行了研究。首次把苔纲雌性生殖苞（Perichaetium）分为蒴帽型（Calyptra type）、蒴萼型（Perianth type）、鞘萼型（Coelocaule-perianth type）、茎鞘型（Coelocaule type）、蒴囊型（Perigynium type）、总苞型（Involucre type）和托生总苞型（Carpocephalum involucre type）等雌苞类型,并指明其中蒴帽型雌苞（Perichaetium of calyptra type of Hepaticae）最简单,为演化原始类群,托生总苞型雌苞（Perichaetium of carpocephalum involucre type of Hepaticae）演化最高级。还首次提出了雌苞演化示意图和“苞图式”表示苔类的雌苞结构。     

Pictolejeunea reginac, a new species of Lejeuneaceae (Hepaticae) from Venezuela

Pictolejeunea reginae, a new species from Venezuela is described and illustrated. It is the type of the new subgenusNeopitolejeunea Ilkiu-Borges, which is characterized by entire underleaves. A key to the two subgenera and four known species ofPictolejeunea is provided.

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Ilkiu-Borges, A. L. (Museu Paraense Emílio Goeldi, C. P. 399, Belém, Pará, CEP 66040-170. Brasil: email: ilkiuborges@hotmail.com).Pictolejeunea reginae, a new species of Lejeuneaceae (Hepaticae) from Venezuela. Brittonia 54: 318–321. 2002.—Pictolejeunea reginae, uma nova espécie encontrada na Venezuela, é descrita e ilustrada. A espécie é tratada como tipo do novo subgêneroNeopictolejeunea Ilkiu-Borges, caracterizada pelos anfigastros inteiros. é apresentada uma chave para os dois subgêneros e as quatro espécies dePictolejeunea.

     

A phylogenetic analysis of Sciomyzidae (Diptera) and some related genera

Sciomyzidae is a family of acalyptrate flies with 546 species in 61 genera that is among the most extensively studied groups of higher Diptera. Most of the known larvae are obligate enemies of Gastropoda. Hundreds of studies published over the past 50 years have resulted in detailed information concerning morphology of adults and immature stages, biology, development, behaviour, phenology and distribution. However, studies of phylogenetic relationships are based almost exclusively on morphological characters of adults, and no comprehensive molecular analysis across the family has been published. Here we fill this void by generating and analysing molecular data for 54 species of Sciomyzidae (22 genera), including Phaeomyiidae (one genus), and seven representative species of five other families of Sciomyzoidea (Coelopidae, Dryomyzidae, Helcomyzidae, Heteromyzidae and Huttoninidae) as outgroups. The reconstruction is based on morphological characters as well as nucleotide sequences for genes from the mitochondrial (12S, 16S, COI, COII, Cytb) and nuclear genome (28S, EF1α). The results are compared with recent morphological analyses. Our analyses support the monophyly of Sciomyzidae + Phaeomyiidae, and place Phaeomyiinae as a unique lineage within Sciomyzidae. A modified classification comprising three subfamilies is proposed. The major subfamily, Sciomyzinae, consists of two monophyletic and well separated groups, the tribes Sciomyzini and Tetanocerini.     

A phylogenetic analysis of the monogenomic Triticeae (Poaceae) based on morphology

A cladistic analysis, primarily based on morphology, is presented from 40 diploid taxa representing the 24 monogenomic genera of the Triticeae. General problems related to the treatment of hybrids and supposedly allopolyploid heterogenomic taxa are highlighted. Special emphasis is given to taxa not traditionally included in Aegilops s.J. Most of the 33 characters used in the analysis are coded as binary. The only four multistate characters in the matrix are treated as unordered. Three diploid species of Bromus are used as outgroup. The number of equally parsimonious trees found is very large (approx. 170000; length = 107, ci = 0.36, ri = 0.75) and the strict consensus tree has an expectedly low level of resolution. However, most of the equally parsimonious trees owe their existence to an unresolved Aegilops clade. If this clade is replaced by its hypothetical ancestor, the number of equally parsimonious trees drops dramatically (48; length = 78, ci = 0.45, ri = 0.76). When trees for which more highly resolved compatible trees exist are excluded, only two trees remain. Bremer support is used as a measure of branch support. The trees based on morphology and on molecular data are largely incongruent.     

Phylogenetic relationships in the Lejeuneaceae (Hepaticae) inferred using ITS sequences of nuclear ribosomal DNA

Sequences of the ITS1–5.8S–ITS2 region of nuclear ribosomal DNA were generated for 12 species from 9 genera of Lejeuneaceae and a single species of Jubulaceae (outgroup). The taxon sampling of Lejeuneaceae included representatives of the two widely recognized subfamilies, Lejeuneoideae and Ptychanthoideae. The molecular dataset was analysed independently and in combination with a morphological dataset. The nrITS dataset and the combined dataset resulted in identical topologies. The genus Bryopteris, sometimes treated as a separate family Bryopteridaceae, is nested within the Lejeuneaceae subfamily Ptychanthoideae. Lejeuneaceae subfamily Lejeuneoideae proved to be paraphyletic with the tribe Lejeuneeae sister to Ptychanthoideae, albeit without significant bootstrap support. The tribes Brachiolejeuneeae and Cheilolejeuneeae of Lejeuneoideae, established recently based on morphological evidence, are well supported in bootstrap analyses both of the ITS and the combined molecular–morphological datasets. The results support classifications of Lejeuneaceae based on morphological data and demonstrate the usefulness of the ITS region for phylogenetic studies within or among closely related genera of Lejeuneaceae.     

A phylogenetic supertree of the bats (Mammalia: Chiroptera)

We present the first estimate of the phylogenetic relationships among all 916 extant and nine recently extinct species of bats Mammalia: Chiroptera), a group that accounts for almost one-quarter of extant mammalian diversity. This phylogeny was derived by combining 105 estimates of bat phylogenetic relationships published since 1970 using the supertree construction technique of Matrix Representation with Parsimony (MRP). Despite the explosive growth in the number of phylogenetic studies of bats since 1990, phylogenetic relationships in the order have been studied non-randomly. For example, over one-third of all bat systematic studies to date have locused on relationships within Phyllostomidae, whereas relationships within clades such as Kerivoulinae and Murinae have never been studied using cladistic methods. Resolution in the supertree similarly differs among clades: overall resolution is poor (46.4%, of a fully bifurcating solution) but reaches 100% in some groups (e.g. relationships within Mormoopidae). The supertree analysis does not support a recent proposal that Microchiroptera is paraphyletic with respect to Megachiroptera, as the majority of source topologies support microbat monophyly. Although it is not a substitute for comprehensive phylogenetic analyses of primary molecular and morphological data, the bat supertree provides a useful tool for future phylogenetic comparative and macroevolutionary studies. Additionally, it identifies clades that have been little studied, highlights groups within which relationships are controversial, and like all phylogenetic studies, provides preliminary hypotheses that can form starting points for future phylogenetic studies of bats.     

Intercontinental distribution of Plagiochila corrugata (Plagiochilaceae, Hepaticae) inferred from nrDNA ITS sequences and morphology

Plagiochila sect. Vagae is a large pantropical clade that is characterized morphologically by frequent terminal branching, vegetative distribution by propagules on the ventral surface of the leaves and a capsule wall with thickenings in all layers. Plagiochila corrugata from Brazil is characterized by strongly undulate, toothed leaf margins and represents the only known neotropical species of sect. Vagae with unispiral elaters. Plagiochila cambuena from Madagascar is distinguished by the same features. Maximum likelihood and parsimony analyses of 38 nrDNA ITS sequences of Plagiochila reveal P. corrugata and P. cambuena in a weakly (ML) to well (MP) supported monophyletic lineage within P.  sect.  Vagae . As an outcome of the morphological and molecular investigation, P. cambuena is relegated to the synonymy of P. corrugata. Plagiochila corrugata is placed in a Vagae -subclade with 11 further American species. The range of P. corrugata can be ascribed to long-range dispersal from the Neotropics rather than a Gondwanan distribution. Species from tropical Asia and Africa are placed at the base of the Vagae clade. Branch length within P.  sect.  Vagae points to a sudden radiation.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 469–481.     

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.     

Review of Erigonine spider genera in the Neotropics (Araneae: Linyphiidae,Erigoninae)

The Neotropical genera of the linyphiid spider subfamily Erigoninae are revised at the genus level. Emphasis was placed on genera endemic to the Neotropics and species with dubious relationships to their nominal genera, especially species from the older literature. This work recognizes 50 genera in the Neotropics, of which 39 genera are strictly endemic to the Neotropics, three are represented outside the Neotropics by one species, and eight genera have significant representation both in and beyond the Neotropics. Three additional genera, Ceraticelus Simon, 1884, Idionella Banks, 1893, and Eulaira Chamberlin & Ivie, 1933, are represented in northern Mexico and/or the West Indies, but are best classified as having a Nearctic or Holarctic distribution. Species previously placed in the typically northern hemisphere genera Gongylidiellum Simon, 1884, Leptorhoptrum Kulczynski, 1894, Macrargus Dahl, 1886, Minyriolus Simon, 1884, Oedothorax Bertkau, 1883, Phanetta Keyserling, 1886, and Tmeticus Menge, 1868 are found to be misplaced or nomina dubia; two genera endemic to the Neotropics, Clitistes Simon, 1902 and Zilephus Simon, 1902 are nomina dubia. The genus Beauchenia Usher, 1983 is an erigonine, not a mynoglenine; there are no known representatives of the Mynogleninae in the Neotropics. One hundred and forty new combinations are established; 19 genera are synonymized including Micromaso Tambs‐Lyche, 1954, revalidation rejected; 34 species are synonymized. The following new genera are established: Gigapassus gen. nov. , Intecymbium gen. nov. , Moyosi gen. nov. , Orfeo gen. nov. and Toltecaria gen. nov. Malkinella Millidge, 1991 and Valdiviella Millidge, 1985 are preoccupied; Malkinola nom. nov. and Valdiviola nom. nov. are established as replacement names. The following new species are described: Asemostera daedalus sp. nov. , Asemostera enkidu sp. nov. , Asemostera janetae sp. nov. , Fissiscapus attercop sp. nov. , Gonatoraphis lysistrata sp. nov. , Gravipalpus standifer sp. nov. , Microplanus odin sp. nov. , Moyosi chumota sp. nov. , Myrmecomelix leucippus sp. nov. , Neomaso damocles sp. nov. , Notiomaso exonychus sp. nov. , Paraletes pogo sp. nov. , Psilocymbium acanthodes sp. nov. , Smermisia holdridgi sp. nov. and Smermisia parvoris sp. nov. The following species remain misplaced in inappropriate genera: Erigone fellita Keyserling, 1886, Erigone zabluta Keyserling, 1886, and Oedothorax fuegianus (Simon, 1902). For 23 species, type specimens could not be located and the species could not be unambiguously identified; the type of Macrargus pacificus Berland, 1924 could not be located, but it is transferred to Laminacauda Millidge, 1985. The female of Onychembolus subalpinus Millidge, 1985 described by Millidge in 1991 is mismatched; this female is Notiomaso exonychus sp. nov. ; the true female of Onychembolus subalpinus was described as both Neomaso bidentatus Millidge, 1991 syn. nov. and Neomaso tridentatus Millidge, 1991 syn. nov. The male and female of Asemonetes[now Asemostera]arcana (Millidge, 1991) are not conspecific; a male thought to be conspecific with the female of A. arcana is newly described; the true female of A. arcana is unknown. The transfer of Emenista dentichelis Berland, 1913 to Laminacauda comb. nov. renders Laminacauda dentichelis Millidge, 1991 a junior homonym; the replacement name Laminacauda baerti nom. nov. is provided for Laminacauda dentichelis Millidge. The following species were erroneously placed in erigonine genera: Oedothorax bisignatus Mello‐Leitão, 1945 is synonymized with Theridion calcynatum Holmberg, 1876 syn. nov. (Theridiidae); Liger incomta O. Pickard‐Cambridge, 1896 is transferred to Theridion Walckenaer, 1805 (Theridiidae) [Theridion incomtum comb. nov. ]; Erigone ectrapela Keyserling, 1886 is transferred to Dictyna Sundevall, 1833 (Dictynidae) [Dictyna ectrapela comb. nov. ]; Erigone peruana Keyserling, 1886 is transferred to Thymoites Keyserling, 1884 (Theridiidae) [Thymoites peruanus comb. nov. ]; Adelonetria dubiosa Millidge, 1991 is not a linyphiid and will be dealt with elsewhere. Lomaita darlingtoni Bryant, 1948 is confirmed as a linyphiid, not a mysmenid. © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 149 (Suppl. 1), 1–263.     

Two new species of Lejeunea Lib. (Hepaticae: Lejeuneaceae) from Sikkim,India

Abstract

Two new species of Lejeunea Lib., L. kashyapii sp. nov. and L. mehrana sp. nov. are described from Sikkim, India. The former is characterized by sub-orbicular leaves; first tooth of leaf lobule, 1–3 cells long 1–2 cells wide at base; reniform underleaves slightly wider than long; monoicous sexuality; 2–3-paired male bracts; obcordate perianth with 4–5 smooth keels (2 lateral, auriculate; 2 ventral; 0–1 dorsal, indistinct). Whereas, the latter is characterized by oblong – ovate leaves; lobule 1/6–1/5 as long as lobe; distant – contiguous underleaves with 1–3 teeth on each lobe; dioicous (?) sexuality; 2–4-paired male bracts and 1–2 male bracteoles present near the base of androecia. A key to the Indian species of the genus are provided.     

Reinerantha foliicola,a new genus and species of Lejeuneaceae subtribe Cololejeuneinae (Marchantiophyta) from Ecuador

Cololejeuneinae is the largest subtribe of the liverwort family Lejeuneaceae and the crown group of the family. The species often grow on the surfaces of living leaves and twigs or in swiftly running waters and many of them possess neotenic gametophytes, considered an adaptation to these extreme habitats. We describe a new, epiphyllous genus and species in Cololejeuneinae from the Western Cordillera of Ecuador, Reinerantha foliicola Gradst. & R.L.Zhu. Analysis of combined molecular datasets with parsimony, maximum likelihood and Bayesian methods revealed Reinerantha in a sister relationship to the large pantropical genus Diplasiolejeunea. Reinerantha differs from Diplasiolejeunea in leaf segmentation, epidermal wall thickening, stylus, male bracts and perianth keels, and is morphologically more similar to the genus Tuyamaella. In the molecular analysis, however, Reinerantha and Tuyamaella were resolved in different clades and not closely related. Tuyamaella is shown to be restricted to Asia; the report of Tuyamaella from South America (Peru) was erroneous and a case of mislabeling.     

A comprehensive phylogenetic analysis of termites (Isoptera) illuminates key aspects of their evolutionary biology

The first comprehensive combined molecular and morphological phylogenetic analysis of the major groups of termites is presented. This was based on the analysis of three genes (cytochrome oxidase II, 12S and 28S) and worker characters for approximately 250 species of termites. Parsimony analysis of the aligned dataset showed that the monophyly of Hodotermitidae, Kalotermitidae and Termitidae were well supported, while Termopsidae and Rhinotermitidae were both paraphyletic on the estimated cladogram. Within Termitidae, the most diverse and ecologically most important family, the monophyly of Macrotermitinae, Foraminitermitinae, Apicotermitinae, Syntermitinae and Nasutitermitinae were all broadly supported, but Termitinae was paraphyletic. The pantropical genera Termes, Amitermes and Nasutitermes were all paraphyletic on the estimated cladogram, with at least 17 genera nested within Nasutitermes, given the presently accepted generic limits. Key biological features were mapped onto the cladogram. It was not possible to reconstruct the evolution of true workers unambiguously, as it was as parsimonious to assume a basal evolution of true workers and subsequent evolution of pseudergates, as to assume a basal condition of pseudergates and subsequent evolution of true workers. However, true workers were only found in species with either separate- or intermediate-type nests, so that the mapping of nest habit and worker type onto the cladogram were perfectly correlated. Feeding group evolution, however, showed a much more complex pattern, particularly within the Termitidae, where it proved impossible to estimate unambiguously the ancestral state within the family (which is associated with the loss of worker gut flagellates). However, one biologically plausible optimization implies an initial evolution from wood-feeding to fungus-growing, proposed as the ancestral condition within the Termitidae, followed by the very early evolution of soil-feeding and subsequent re-evolution of wood-feeding in numerous lineages.     

A new african fossil caprin and a combined molecular and morphological bayesian phylogenetic analysis of caprini (Mammalia: Bovidae)

Given that most species that have ever existed on Earth are extinct, no evolutionary history can ever be complete without the inclusion of fossil taxa. Bovids (antelopes and relatives) are one of the most diverse clades of large mammals alive today, with over a hundred living species and hundreds of documented fossil species. With the advent of molecular phylogenetics, major advances have been made in the phylogeny of this clade; however, there has been little attempt to integrate the fossil record into the developing phylogenetic picture. We here describe a new large fossil caprin species from ca. 1.9-Ma deposits from the Middle Awash, Ethiopia. To place the new species phylogenetically, we perform a Bayesian analysis of a combined molecular (cytochrome b) and morphological (osteological) character supermatrix. We include all living species of Caprini, the new fossil species, a fossil takin from the Pliocene of Ethiopia (Budorcas churcheri), and the insular subfossil Myotragus balearicus. The combined analysis demonstrates successful incorporation of both living and fossil species within a single phylogeny based on both molecular and morphological evidence. Analysis of the combined supermatrix produces superior resolution than with either the molecular or morphological data sets considered alone. Parsimony and Bayesian analyses of the data set are also compared and shown to produce similar results. The combined phylogenetic analysis indicates that the new fossil species is nested within Capra, making it one of the earliest representatives of this clade, with implications for molecular clock calibration. Geographical optimization indicates no less than four independent dispersals into Africa by caprins since the Pliocene.     

Comparative and phylogenetic analysis of the echo clicks and social vocalizations of swiftlets (Aves: Apodidae)

Several swiftlet species echolocate to navigate in the darkness of their nesting caves. We investigated whether swiftlets' echo clicks and social vocalizations differed among species and whether they could be used in phylogenetic inference. We incorporated echo clicks of eight swiftlet species and social vocalizations of 27 echolocating and nonecholocating swifts and swiftlets. Characters in the frequency and time domain were used and analysed using nonparametric multivariate analysis of variance and stepwise discriminant analysis. We performed maximum parsimony analyses and character mapping to investigate phylogenetic signal and evolutionary patterns of swiftlet vocalizations. Echo clicks were species-specific. As well as in navigation, they may be used in recognition of conspecifics. Social vocalizations were species-specific also. However, the vocalizations contained no phylogenetic interpretable information. Maximum parsimony trees of echo clicks and of social vocalizations were almost completely unresolved. Moreover, resolved branches showed no congruence to a DNA-based tree that had been obtained previously. Mapping of the vocalization characters on the DNA-based tree revealed no consistent phylogenetic patterns. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 631–643.     

Proopiomelanocortin (POMC) and testing the phylogenetic position of turtles (Testudines)

Morphological and molecular studies have inferred multiple hypotheses for the phylogenetic relationships of Testudines. The hypothesis that Testudines are the only extant anapsid amniotes and the sister taxon of diapsid amniotes is corroborated by morphological studies, while the hypothesis that Testudines are diapsid amniotes is corroborated by more recent molecular and morphological studies. In this study, the placement of Testudines is tested using the full length cDNA sequence of the polypeptide hormone precursor proopiomelanocortin (POMC). Because only extant taxa have been used, the hypotheses being tested are limited to the following (1) Testudines as the sister taxon of Archosauria, (2) Testudines included in Archosauria and the sister taxon of Crocodilia, (3) Testudines as the sister taxon of Lepidosauria, (4) Testudines as the sister taxon of Sauria, and (5) Testudines as the sister taxon of a monophyletic Mammalia–Sauria clade. Neither Maximum likelihood, Bayesian, or maximum parsimony analyses are able to falsify the hypothesis of (Archosauria (Lepidosauria, Testudines)) and as such is the preferred inference from the POMC data.