Using palynomorphological characteristics for the identification of species of Alsinoideae (Caryophyllaceae): a systematic approach

Palynological characters of 18 species belonging to seven genera of the traditional subfamily Alsinoideae including Arenaria, Cerastium, Eremogone, Lepyrodiclis, Minuartia, Sabulina and Stellaria were studied in detail using light and scanning electron microscopy. Pollen grains of subfamily Alsinoideae are subspheroidal or prolate, pantoporate and 20.04 to 51.4 µm in size, prominent and sunken apertures uniformly distributed on the pollen surface. Two types of pollen grain ornamentation were observed, i.e. microechinate-punctate or microechinate-perforate. Echini are present on the surface of the pollen of all investigated species with medium, dense, or sparse echinodensity. These species exhibit variation in polar view, equatorial diameter, number of apertures, exine thickness, diameter of pore, appendages per pore, pore ornamentation, echini arrangement, echinidensity and shape of pollen. Based on qualitative characters, a dichotomous key has been developed for quick and easy identification. The present investigation contributes to the systematic approach using palynological characteristics and correct identification of species for members of subfamily Alsinoideae (Caryophyllaceae).     

Circumscription of Malvaceae (Malvales) as determined by a preliminary cladistic analysis of morphological, anatomical, palynological, and chemical characters

We report a phylogenetic analysis of “core” Malvales (Tiliaceae, Sterculiaceae, Bombacaceae, and Malvaceae) based on morphological, anatomical, palynological, and chemical features. The results of the analyses lead to the conclusion that Tiliaceae, Sterculiaceae, and Bombacaceae, as variously delimited, are paraphyletic; only the Malvaceae are likely monophyletic. The genera of “core” Malvales form a well-defined clade. Genera of “Tiliaceae” constitute the basal complex within “core” Malvales. The “Sterculiaceae” (most genera)+ “Bombacaceae” + Malvaceae form a clade on the basis of a monadelphous androecium; “Bombacaceae”+ Malvaceae also form a clade, which is diagnosable on the basis of monoloculate anthers. It is clear that the traditional classification, with its arbitrarily delimited evolutionary grades, is unsatisfactory, especially if one seeks to reflect phylogeny accurately. Thus, Malvaceae is redefined to refer to the most recent common ancestor of plants previously considered to be “Tiliaceae,” “Sterculiaceae,” “Bombacaceae,” and Malvaceae, and all of the descendants of that ancestor. This broadly circumscribed Malvaceae can be diagnosed by several presumed synapomorphies, but we draw special attention to the unusual floral nectaries that are composed of densely packed, multicellular, glandular hairs on the sepals (or less commonly on the petals or androgynophore).     

Systematic position of Gyrocheilos and some odd species of Didymocarpus (Gesneriaceae) inferred from molecular data,with reference to pollen and other morphological characters

A phylogenetic analysis was carried out to clarify the systematic position of Gyrocheilos and Didymocarpus, particularly the species placed in Didymocarpus sect.Heteroboea by Wang et al. Based on sequencing the internal transcribed spacer and the chloroplast spacer trnL-F, parsimony and Bayesian inference analyses were carried out using separate nuclear and chloroplast datasets, as well as a combined dataset. Our results showed that the two sections of Didymocarpus in China andGyrocheilos did not form separate monophyletic subclades, but turned up in three different places in the phylogenetic trees. In the frame of the present study, the pollen morphology of the species included in the analysis was studied. It proved inconsistent with the delimitation between Didymocarpus and Gyrocheilos. Furthermore, pollen and other morphological characters indicate that Gyrocheilos and some taxa of Didymocarpus should be placed within Didymocarpus.     

New morphological characters for classifying Phoridae (Diptera) from the structure of the thorax

Forty‐six characters, mostly of the thoracic pleuron, are proposed for the reconstruction of the phylogenetic relationships among the major groups of the family Phoridae. Analysis of these characters, in conjunction with the pre‐existing suite of morphological characters from other body parts, provides a basis for a strongly supported new classification of the Phoridae, in which Sciadocerinae is the sister group of ((Chonocephalus Wandolleck & Cyphocephalus Borgmeier) + (Termitoxeniinae + (Metopininae + Phorinae s.l.))). A new subfamily, Chonocephalinae subfamily nov. , is proposed for Chonocephalus and Cyphocephalus, and a new genus, Hirotophora gen. nov. , is proposed for Chaetopleurophora multiseriata (Aldrich) ( comb. nov. ). © 2015 The Linnean Society of London     

Dioecy and the evolution of pollination systems inSchiedea and Alsinidendron (Caryophyllaceae:Alsinoideae) in the Hawaiian Islands

The transition from biotic to wind pollination and the consequencesof this transition for the evolution of dioecious breeding systems wereinvestigated in Schiedea and Alsinidendron(Caryophyllaceae: Alsinoideae), genera endemic to the Hawaiian Islands. The potential for wind pollination was studied for five species ofSchiedea using a wind tunnel. Morphological correlates of windpollination for these species were then used to infer the presence orabsence of wind pollination in the remaining Schiedea species. Hermaphroditic Alsinidendron and Schiedea species,which occur in mesic to wet forests, or less commonly in dry habitats,show little or no indication of wind pollination. These species had lowpollen:ovule ratios, large relative pollen size, diffuse inflorescences,substantial nectar production in several cases, and appear to bebiotically pollinated or autogamous. Sexually dimorphic species, whichall occur in dry habitats, are wind pollinated, based on wind tunnelresults or morphological adaptations indicating the potential for windpollination. These adaptations include high pollen:ovule ratios, smallpollen size, moderately to highly condensed inflorescences, and reducednectaries and nectar production. Shifts to wind pollination anddimorphism are strongly correlated in Schiedea, suggesting theclose functional relationship of the twophenomena.     

Systematic relevance of pollen and orbicule characters in the tribe Hillieae (Rubiaceae)

Pollen and orbicule morphology of 26 species in the tribe Hillieae is described based on observations by light, scanning and transmission electron microscopy. Pollen and orbicule characters are critically evaluated and discussed in the context of existing hypotheses of systematic relationships within the tribe. Pollen is 3-zonocolporate with a perforate, microreticulate, reticulate or eutectate sexine. In the two species of Blepharidium , however, the pollen has one, four or five apertures. These pollen morphological data were incorporated into an existing macromorphological matrix of the group and cladistically analysed. The resulting phylogenies indicate that it may be appropriate to reduce Cosmibuena to a subgenus of the genus Hillia , while Blepharidium should be removed from Hillieae because of the deviating pollen type that is unique to the Rubiaceae. All species investigated produce orbicules, which are mostly spherical and possess an electron-lucent core that is sometimes characteristically flattened.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 303−321.     

Suprageneric relationships of galliform birds (Aves, Galliformes): a cladistic analysis of morphological characters

Of the basal clades of extant birds (Neornithes) the 'landfowl' or galliforms (Aves, Galliformes) are the most speciose. Cladistic analysis of more than 100 morphological characters coded at the generic level for most putative galliform genera confirms that the megapodes ('mound builders'; Megapodiidae) are the most basal clade within the order. They are followed successively by the curassows, guans and chachalacas (Cracidae), which comprise the sister-group to all other extant Galliformes (i.e. Phasianoidea). Within this large 'phasianoid' clade, analyses suggest that the guineafowl (Numididae) are the most basal taxon, although monophyly of this 'family' is not strictly supported on the basis of the morphological characters employed. An additional major clade within the phasianoid Galliformes is recovered by this analysis, comprising the traditional groupings of New World quails (Odontophoridae) and Old World quails ('Perdicini'), yet only monophyly of the former is supported unambiguously by morphological characters. Relationships within the remainder of the phasianoid taxa, including the grouse (Tetraonidae), turkeys (i.e. Meleagris / Agriocharus spp.) as well as other 'pavonine' galliforms (i.e. peafowl; Pavo , Afropavo , Rheinardia , Argusianus and Polyplectron spp.) remain largely unresolved on the basis of morphological characters, yet monophyly of the major subdivisions is supported here. Although there are a number of important differences, especially with regard to relationships within the nonquail phasianoids, the results of this morphological phylogenetic (cladistic) analysis are broadly congruent both with traditional classifications and existing molecular hypotheses of galliform phylogenetic relationships.     

Monophyly and phylogenetic placement of the spider genus Labulla Simon, 1884 (Araneae, Linyphiidae) and description of the new genus Pecado

The genus Labulla Simon is circumscribed in phylogenetic terms to include the species Labulla thoracica (Wider), L. flahaulti Simon and L. machadoi sp. nov. The genital anatomy of the genus is described in detail and the taxonomy of the genus is reviewed. The monophyly of Labulla is supported by numerous morphological apomorphies of the male palp and female epigynum. Based on a cladistic analysis, a new genus, Pecado gen. nov. , is erected to place Labulla impudica Denis, from Northern Africa. Lepthyphantes insularis Saito and ' Labulla ' nepula Tikader, both formerly included in Labulla , are not congeneric with the type species of Labulla .  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 143 , 359–404.     

Relationships within the family Actiniidae (Cnidaria,Actiniaria) based on molecular characters

Current sea anemone systematics is based on relatively few morphological characters, and potentially could benefit from the use of molecular characters. In this paper, the phylogenetic relationships of 12 species from 6 genera in the family Actiniidae have been investigated using electrophoretically separated isozymes. A numerical cladistic analysis has produced an estimated phylogeny. The implications of this phylogeny for the taxonomic use of certain morphological characters are discussed.     

Phylogeny, evolution and systematics of Moehringia (Caryophyllaceae) as inferred from molecular and morphological data: a case of homology reassessment

The phylogeny of the genus Moehringia (Caryophyllaceae) is investigated by means of analyzing nuclear (ITS) and chloroplast (matK) sequence data in combination with morphological characters. Parsimony and Bayesian methods yield consistent results, and a common phylogenetic signal is shared by the nuclear and chloroplast data. Morphological characters are affected by a high level of homoplasy, but they provide valuable information when analyzed in combination with the molecular data. Moehringia is paraphyletic to Arenaria with the Iberian taxa belonging to Moehringia sect. Pseudomoehringia McNeill more closely related to Arenaria. This cladistic evidence led us to reinterpret the homology of the key character used in most, if not all, floras, to separate Moehringia from Arenaria, i.e., the seed strophiole. Thorough anatomical studies were carried out to elucidate the ontogeny of the strophiole, which proved different in Moehringia s. str. and the Iberian taxa. Within Moehringia s. str., two sister clades are recognized (i.e., Moehringia sect. Moehringia and M. sect. Latifoliae much as recognized by McNeill, whereas representatives of M. sect. Diversifoliae are assigned to either groups), and biogeographical events related to the Würm glaciation are considered to play a fundamental role in the evolution and present distribution of the genus. The variation of the strophiole is regarded as adaptability to ecological conditions and dispersal agents. We also propose two new combinations and two replacement names: Arenaria glochidisperma (J.M. Mont.) Fior et P.O. Karis, comb. nov., Arenaria tejedensis (Willk.) Fior et P.O. Karis, comb. nov., Arenaria suffruticosa Fior et P.O. Karis, nom. nov. for Moehringia intricata Willk., and Arenaria funiculata Fior et P.O. Karis, nom. nov. for Moehringia fontqueri Pau. © The Willi Hennig Society 2007.     

Multivariate analysis of morphological and anatomical characters of Calophyllum (Calophyllaceae) in South America

Calophyllum (Calophyllaceae), previously placed in Clusiaceae, is easily recognizable by its opposite entire leaves with close parallel venation alternating with resin canals. However, distinction between species has been difficult, because of infraspecific variation in tepal and stamen number and resemblance among species that share similar habitats. Here, I report the results of multivariate analyses of morphological and anatomical characters for Calophyllum in South America, and provide a taxonomic treatment for the genus in South America, the first since that of Vesque in 1893. Thirteen preliminary morphogroups were identified. Thirty‐two morphological characters of the leaf, flower and fruit from 401 specimens, and 17 anatomical leaf characters from 45 specimens were measured and analysed using principal component analyses (PCAs) and discriminant analyses (DAs). PCAs were used to find groups and DAs were used to validate those PCAs that were potential groups. Two main subgroups were identified in the general analysis. Subgroup M1 has terete stems and smaller leaves and flowers than subgroup M2, which, instead, has quadrangular stems. Only subgroup M2 showed distinctive clusters in regional and local analyses. Distinctive clusters and morphological and anatomical characters helped us to recognize four species in South America, including a new species, Calophyllum pubescens sp. nov. . In addition, a new species, Calophyllum mesoamericanum sp. nov. , is described from Central America. © 2013 The Linnean Society of London,     

Relationships between floral characters, pollination mechanisms, life forms, and habitats in Araceae

The floral traits of the inflorescences of angiosperms have coevolved to ensure and maximize pollination success. Other factors believed to influence floral architecture are external (for example, ecological) to the inflorescence. In order to understand the relationships between such factors and floral characters, 12 floral traits were measured in 54 species of Araceae. An analysis was performed to determine how these traits are linked to the following: (1) self-pollination capacity; (2) life form (evergreen versus seasonally dormant); (3) climatic conditions; and (4) type of pollinator (i.e. flies, bees, or beetles). A significant difference was found between the pollen to ovule ratio of the species able to self-pollinate and those unable to self-pollinate. Evergreen and tropical aroids produced a larger number of gametes than did seasonally dormant and temperate taxa. Finally, several floral traits, such as pollen volume and number, number of female flowers, and flower sexual type (unisexual or bisexual), showed clear differences between the three pollinator types. Variations in floral traits between the different life forms and climatic conditions are discussed with respect to pollination efficiency and properties of the growing season. The pollen to ovule ratio cannot be considered as an accurate indicator of breeding systems in aroids because of the particular pollination ecology of the family.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 29–42.     

Composition and phylogenetic affinities of vangas (Vangidae, Oscines, Passeriformes) based on morphological characters

The first cladistic analysis of the relationship of Malagasy vangas (Vangidae) based on morphological characters is presented here. The comparative morphological analysis provides new arguments for the exclusion of several taxa traditionally or recently assigned to Vangidae such as Tylas 'Vanga' ( Tylas eduardi ), Crossley's Babbler ( Mystacornis crossleyi ) and newtonias ( Newtonia spp.), but also for the exclusion of the Red-tailed 'Vanga' ( Calicalicus madagascariensis ) from the Vangidae. It leads also to the hypothesis that vangas sensu stricto , the Australasian woodswallows (Artamidae) and the Australo-Papuan butcherbirds (Cracticidae) form a monophyletic group, with the New Guinean peltops ( Peltops spp.) as its probable sister taxon. Such a relationship implies that the last common ancestor of vangas reached Madagascar from the east by means of a dispersal event across the Indian Ocean. This hypothesis is partly in accordance with that of William Pycraft (1907) published a century ago, but in conflict with contemporary hypotheses based on DNA sequence analysis which favour helmet-shrikes, bush shrikes and relatives as the closest relatives of vangas and therefore an African origin of this group.     

RAPD patterns,nrDNA ITS sequences and morphological patterns inSilene sectionSedoideae (Caryophyllaceae)

Hierarchical patterns inSilene sect.Sedoideae were investigated using random amplified polymorphic DNA (RAPD), nucleotide sequences of the internal transcribed spacer (ITS) regions of the 18S–28S nuclear ribosomal DNA, and discrete morphological characters. All data sets firmly supported the species recognized. The RAPD data offered the best resolution at the intraspecific level, supporting the current intraspecific classifications ofS. sedoides andS. integripetala. The ITS sequences and the morphological data gave poor resolution within species, and the three data sets disagreed about the relationships among species. The signal from the RAPD data was strongest and remained when the total data set was analysed. The three data sets all support an amphiploid origin ofS. aegaea, with the strongest evidence from the ITS sequences. Incongruences among data sets as well as merits and shortcomings of each are discussed. The robustness of the results can be evaluated using perturbations of data, i.e., bootstrap and jackknife of taxa and characters. These methods should not be taken as methods of statistical inference at the taxonomic level, because unbiased sampling appears impossible. RAPD data, however, come close to being suitable for statistical estimation of hierarchies at the genome level, but several methodological problems have to be solved.     

The relationships of Heberdenia bahamensis and H. penduliflora (Myrsinaceae)

A morphological and anatomical investigation is presented of the two species of Heberdenia, H. bahamensis (Macaronesia) and H. penduliflora (Mexico). A cladistic analysis including 27 taxa of the Myrsinaceae and using Jacquinia (Theophrastaceae) and Manilkara (Sapotaceae) as outgroup was performed to provide a hypothesis of the relationships of the two species of Heberdenia. It was concluded that H. bahamensis and H. penduliflora are not more closely related to each other than either is to many other species of the Myrsinaceae, and that they should not be referred to the same genus. Heberdenia bahamensis appears to be most closely related to the Old World genera Pleiomeris, Embelia , and Grenacheria , whereas H. penduliflora is nested within Ardisia s. L , possibly being most closely related to the segregate genus Gentlea. It is suggested that H. penduliflora is probably best referred to a genus of its own.     

Phylogeny and morphological evolution of tribe Menispermeae (Menispermaceae) inferred from chloroplast and nuclear sequences

The Menispermaceae family contains ca. 72 genera with 450 species that are almost entirely tropical. Its phylogeny at the tribal level has never been examined using molecular data. Here we used DNA sequences of the chloroplast matK gene and trnL-F regions, and the nuclear ITS region to study the delimitation and position of the tribe Menispermeae within the family and its subtribal monophyletic groups. Family-wide phylogenetic analyses of the chloroplast data produced two strongly supported clades. The first clade contains two subclades: Coscinieae including Arcangelisia and Anamirta, and Tinosporeae sensu lato including Fibraureae, supported by morphological characters, such as traits of the cotyledon, stylar scar and embryo. The second clade consists of the tribes Menispermeae sensu DC. and Tiliacoreae Miers. All our analyses surprisingly recognized that tribe Menispermeae is not monophyletic unless tribe Tiliacoreae is included, suggesting that characters of cotyledon and stylar scar are very important for the infrafamilial classification, and that endosperm presence vs. absence was over-emphasized in traditionally tribal division of the family. Our topologies indicate a secondary loss of endosperm. The monophyly of two subtribes of the tribe Menispermeae, Stephaniinae and Cissampelinae, is supported by the cpDNA and ITS data, as well as by morphological characters, including aperture types and shapes, and colpal membrane features of pollen grains, and sepal number of male flowers. The Cocculinae was recognized as a paraphyletic group containing the remaining genera of the tribe Menispermeae.     

菝葜科基于形态学证据的系统发育分析

对全世界范围分布的菝葜科Smilacaceae的79个代表种(包括了全部的属和组), 以分布于南美洲的Philesia Comm. ex Juss.和Lapageria Ruiz &; Pav.属为外类群, 选取包括花粉和染色体性状在内的47个广义的形态学性状进行了分支分类系统发育分析, 同时以表征分类的方法构建了距离树(NJ)辅助分析, 首次对世界分布的菝葜科各属间及属内的系统发育关系作了探讨。(1)Ripogonum与菝葜属Smilax +肖菝葜属Heterosmilax互为姐妹群, 但是距离较远, 支持将类菝葜属(新拟中文名)Ripogonum独立为科的观点; (2)肖菝葜属在菝葜科内处于较为进化的分支上, 并与菝葜属土茯苓组sect. Coilanthus的部分种组成一个具较高支持率(88%)的单系分支, 分析表明肖菝葜属并非是一个好属, 应归入菝葜属; (3)菝葜属6个组的划分大都没有得到支持, 只有东亚北美间断分布的草本菝葜组sect. Nemexia的单系得到很好的支持(93%); (4)分布于南美洲巴西的种类聚为一个单系类群, 表明它们可能有共同的起源, 但由于取样局限, 南美洲种类的系统地位有待进一步研究。     

The relative lability of floral vs. non-floral characters and a morphological phylogenetic analysis of Cobaea (Polemoniaceae)

The relative levels of lability in floral vs. vegetative characters have been suggested to give insight into the mechanisms of adaptation and speciation. Cobaea (Polemoniaceae) exhibits a remarkable diversity in floral form. A morphology-based phylogeny was constructed and is congruent in most regards with a previously published molecular phylogeny, but significant incongruence was found in the placement of two taxa. A combined analysis was performed, excluding C. aquatoriensis and C. lutea, and was nearly identical to the molecular analysis. The phylogenies are compatible with a recently published classification. Incongruencies between the phylogenies within section Rosenbergia may have profound implications for floral evolution. A simple method for testing the levels of homoplasy, as an indicator of lability between two classes of data, is developed and used to test for differences between floral and non-floral classes of characters. The method is based on repeated randomization of the data into partitions and recalculation of a test statistic defined as the difference between consistency index (CI) values of the data in each partition over a given tree. The null hypothesis that the floral characters are no more homoplasious than the non-floral characters cannot be rejected (P=0.09).     

Polygyrid relations: a phylogenetic analysis of 17 subfamilies of land snails (Mollusca: Gastropoda: Stylommatophora)

The family of polygyrid land snails in North America is significant for its sympatric shell convergences, diversity of mating systems and complex zoogeography; its monophyly and its relation to other families has long been questionable. Cladistic analysis has been performed on one representative each of 17 subfamilies, including all three polygyrid subfamilies and one or more subfamilies each from all ten of the families that have been proposed as the polygyrid sister groups. Eighteen anatomical synapomorphies are used, of which eight are newly discovered, four are differently assessed from previous studies, and six are traditional. The resulting strict consensus tree of alternative maximum-parsimony cladograms is: (Acavidae (Ammonitellidae Corillidae ((Discidae Oreohelicidae) (Helminthoglyptidae Bradybaenidae Polygyridae (Thysanophoridae (Camaenidae Sagdidae)))))).
According to this working hypothesis, the Polygyridae are monophyletic, and their sister group remains unresolved, although the Acavidae, Ammonitellidae, Corillidae, Discidae and Oreohelicidae can be ruled out. Of the five classifications of stylommatophoran families that have been proposed in the past 12 years, the consensus tree is closest to that of Nordsieck. For future morphological work, three regions are recommended as potentially rich in unused phylogenetic information: the fertilization pouch-seminal receptacle complex, the ureter at the pneumostome and the ventral-chain ganglia. Simultaneous dissection, wkh side-by-side comparisons, is recommended over other methods for detecting homologies in land snails. Molecular characters should be exploited, because of the scarcity and the frequent homoplasy of morphological characters.     

A combined morphological and molecular phylogenetic analysis of Parthenocissus (Vitaceae) and taxonomic implications

Parthenocissus (the Virginia creeper genus, Vitaceae) consists of 13 species and shows a disjunct distribution between Asia and North America. We investigated the inflorescence structure, calyx morphology, appendages on the inner side of petals, leaf epidermis, pollen and seed characters throughout the genus. A combined phylogenetic analysis with 27 morphological and 4137 molecular characters was conducted and the result was largely congruent with that of the previous molecular work, but with higher resolution. The combined analysis identified two clades corresponding to the Asian and North American taxa. Parthenocissus feddei was resolved as closely related to the clade containing P. cuspidifera, P. heterophylla and P. semicordata. The four species share synapomorphies of having conspicuously raised veinlets, an obscurely five‐ (to eight‐) lobed calyx, appendages on the inner side of petals covering the entire length of anthers and foveolate pollen exine ornamentation. Within the Old World clade, the pentafoliolate species were weakly supported as more closely related to species with both simple and trifoliolate leaves. Furthermore, the ancestral states of tendril apices, inflorescence structure, appendages on the inner side of petals and exine ornamentation of pollen grains were reconstructed on the molecular strict consensus tree. The appendages on the inner side of petals and exine ornamentation of pollen grains were suggested to be important characters in the taxonomy of Parthenocissus, especially for species with trifoliolate leaves. Finally, the previous classifications of Parthenocissus were evaluated within the phylogenetic framework. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, ?? , ??–??.