Critical Reexamination of Palynological Characters Used to Delimit Asclepiadaceae in Comparison to the Molecular Phylogeny Obtained from PlastidmatK Sequences

The family Asclepiadaceae (Dicotyledones) was created by Brown in 1810 by splitting in two the family Apocynaceae of Jussieu established in 1789. The morphological characters used to make this distinction were mainly palynological, such as presence of tetrads or pollinia and number and orientation of pollinia. Those characters, still used in higher taxonomic delimitation (families, subfamilies, and tribes), are here critically reexamined and compared to a molecular phylogeny obtained with one of the more variable plastid genes (matK) of 46 species in the order Gentianales. In this molecular phylogeny, Asclepiadaceae form a monophyletic group derived from within Apocynaceae. Each of the subfamilies of Asclepiadaceae is monophyletic and based on reliable palynological characters, but palynological characters are not useful to delimit tribes of the subfamily Asclepiadoideae. Based on the molecular data, these tribes have undergone parallelisms in several reproductive traits.     

The taxonomy and systematics of Apocynaceae: where we stand in 2012

In their most recent classification of Apocynaceae in 2000, Endress and Bruyns recognized five subfamilies of Apocynaceae (Rauvolfioideae, Apocynoideae, Periplocoideae, Secamonoideae and Asclepiadoideae). Subsequently, through various studies using molecular data, it has been shown that most tribes and subtribes of Rauvolfioideae were not monophyletic, and new tribes and subtribes have been erected to reflect improved phylogenetic understanding of the family: Aspidospermeae in Rauvolfioideae; Nerieae, Odontadenieae and Baisseeae in Apocynoideae; Fockeeae in Asclepiadoideae; and Orthosiinae in Asclepiadeae. Several genera in Rauvolfioideae have been reassigned to different tribes in order to improve the monophyly of these tribes. The sister group of Asclepiadoideae plus Secamonoideae is not Periplocoideae, as formerly assumed, but tribe Baisseeae. Periplocoideae are nested in Apocynoideae. However, tribal composition remains unclear in some parts of the family. Clade structure in Apocynaceae is now generally well understood. The principal challenges now lie in identifying characters that can reflect and articulate these clades in a formal classification. Species‐rich, recent radiations such as core Asclepiadinae in Africa and the Metastematinae in Latin America present particular problems in this regard. © 2013 The Linnean Society of London     

中国特有种苦绳(广义夹竹桃科)的大小孢子发生和雌雄配子体发育及其分类学意义

近年来的分子系统学把狭义萝藦科和狭义夹竹桃科合并为广义夹竹桃科,包括5个亚科和25个族,但亚科和族间的亲缘关系较为复杂,亟待多学科证据澄清。本文利用常规石蜡切片技术观察了马利筋亚科南山藤属中的中国特有植物苦绳(Dregea sinensis var. sinensis)的孢子发生和配子体发育,结合已有资料比较了5个亚科的胚胎学特征。结果表明:(1)苦绳的花药由一对侧生并列药室组成,各有一个花粉团。(2)花药壁有6层,由外至内分别为表皮、2层药室内壁、中层和2层绒毡层,花药壁发育模式属于多层型。(3)绒毡层细胞单核,排成2列,为腺质型; 在小孢子四分体形成时期,药室内壁发生明显纤维状加厚; 花药成熟时,位于药室远轴最外侧处的花药壁发生断裂,准备散粉。(4)小孢子母细胞减数分裂中,胞质分裂方式为连续型,小孢子四分体排列方式为左右对称; 成熟花粉粒为3-细胞型,排列紧密,形成花粉团。(5)雌蕊含有两枚离生心皮,具边缘胎座,胚珠倒生,单珠被,薄珠心,蓼型胚囊。本文观察到的这些胚胎学特征为牛奶菜族提供了新资料。同时,胚胎学特征在5个亚科间的区别和联系,支持广义夹竹桃科的成立。     

Morphological and ultrastructural diversity of orbicules in relation to evolutionary tendencies in apocynaceae s.L

Minute granules of sporopollenin, called orbicules, can be observed on the innermost tangential and/or radial walls of secretory tapetum cells. Orbicules were investigated in 62 species (50 genera) of Apocynaceae s.l. using light microscopy, scanning electron microscopy and transmission electron microscopy. Orbicules were found in 43 species (34 genera) distributed amongst the subfamilies Rauvolfioideae, Apocynoideae, Periplocoideae, and in the genus Riocreuxia (Asclepiadoideae). Absence of orbicules is apparent in Secamonoideae and Asclepiadoideae (except Riocreuxia). The orbicule types described are based on observed morphological and ultrastructural variation. Of the six orbicule types previously described, Type I and Type II orbicules are lacking. In the majority of species, Type III orbicules were recorded in addition to Types IV, V and VI. In this study we suggest that embedded Type VI orbicules are more derived. A correlation between orbicule typology and evolutionary tendencies in Apocynaceae s.l. palynology was found. A trend was observed from the presence of Type III orbicules in the majority of species belonging to the basal group of genera characterized by colporate to porate single pollen grains, or 3-6-porate tetrads, towards the more derived embedded Type VI orbicules in the more advanced Periplocoideae genera with multiporate tetrads or pollinia. Orbicule data have proven not to be useful for evaluating tribal delimitation within the Apocynaceae s.l. contrary to the Rubiaceae and Loganiaceae s.l.     

Phylogenetic relationships within the Gentianales based on NDHF and RBCL sequences, with particular reference to the Loganiaceae

Phylogenetic relationships in the Gentianales with focus on Loganiaceae sensu lato are evaluated using parsimony analyses of nucleotide sequence data from the plastid genes rbcL and ndhF. Inter- and intrafamilial relationships in the Gentianales, which consist of the families Apocynaceae (including Asclepiadaceae), Gelsemiaceae, Gentianaceae, Loganiaceae, and Rubiaceae, are studied and receive increased support from the combination of rbcL and ndhF data, which indicate that the family Rubiaceae forms the sister group to the successively nested Gentianaceae, Apocynaceae, and Loganiaceae, all of which are well supported. The family Gelsemiaceae forms a distinct, supported group sister to Apocynaceae. The Loganiaceae sensu stricto form a strongly supported group consisting of 13 genera: Antonia, Bonyunia, Gardneria, Geniostoma, Labordia, Logania, Mitrasacme, Mitreola, Neuburgia, Norrisia, Spigelia, Strychnos, and Usteria. These genera form two well-supported lineages. Several members of Loganiaceae sensu Leeuwenberg and Leenhouts, i.e., Androya, Peltanthera, Plocosperma, Polypremum, and Sanango are clearly not members of the Gentianales. The earlier exclusion of Buddlejaceae (including Buddleja, Emorya, Gomphostigma, and Nicodemia) as well as the reclassification of the genera Nuxia and Retzia to Stilbaceae of the Lamiales are all well supported.     

CLADISTICS AND FAMILY LEVEL CLASSIFICATION OF THE GENTIANALES

Abstract— The most recent classification of the angiosperm order Gentianales (Thorne, 1992) includes four principal families: Apocynaceae, Gentianaceae, Loganiaceae, and Rubiaceae. Ever since Bentham (1857) the status of Loganiaceae has been questioned, and several segregates of that family have been proposed both before and after his treatment. In this study we present cladistic results that show Loganiaceae, sensu lato, to be a paraphyletic group definable only by plesiomorphies, with members showing closest relationships to other families of the order. As the impact of different character-state representations of polymorphic terminals remains largely untested, our morphological and phytochemical data were analysed both with restricted polymorphism coding as well as with the monomorphic "subtaxon" recoding method of Nixon and Davis (1991). Both approaches yield highly compatible results, and we here discuss a new classification of the Gentianales based on (i) monophyletic groups identified by outgroup analysis, and (ii) the maximal portrayal of evidence provided by subtaxon polymorphism recoding. Most prominently, the Loganiaceae sensu lato are divided into four segregate families, two previously named (Loganiaceae sensu stricto and Strychnaceae), and two defined as a result of this study (Gelsemiaceae, L. Struwe & V. A. Albert, stat. nov. and Geniostomaceae, L. Struwe & V. A. Albert, fam. nov.). Apocynaceae (incl. Asclepiadaceae), Gentianaceae (incl. Loganiaceae—Potalieae), and Rubiaceae remain as monophyletic families. Outgroup analysis supports both the monophyly of the Gentianales as well as the exclusion from the order of Buddleja, Desfontainia, Plocosperma, Polypremum , and Retzia (all Loganiaceae sensu Leeuwenberg and Leenhouts).     

Changes in Structure of Phytomonas elmassiani in Experimental Infections of Apocynaceae, a Presumably Foreign Plant Host

SYNOPSIS. Allamanda nereifolia and Plumeria rubra, plants belonging to the family Apocynaceae were infected experimentally with the trypanosomatid flagellate, Phytomonas elmassiani. In these presumably exotic hosts, P. elmassiani, ordinarily infecting plants belonging to the family Asclepiadaceae, underwent marked structural changes. The presence of numerous choanomastigote like organisms along with a certain proportion of promastigotes and amastigotes, would, under ordinary circumstances, result in classification of the infecting organisms as Crithidia. Obviously, the change in environment from Asclepiadaceae to the closely related Apocynaceae, exerts a profound influence on structure of the organism.     

PHYLOGENY OF THE RUBIACEAE AND THE LOGANIACEAE: CONGRUENCE OR CONFLICT BETWEEN MORPHOLOGICAL AND MOLECULAR DATA?

Phylogenetic analyses of 33 genera of Rubiaceae were performed using morphological and a few chemical characters. Parsimony analysis based on 29 characters resulted in eight equally parsimonious trees, with a consistency index of 0.40 and a retention index of 0.69. These results were compared to a phylogenetic analysis of the same genera based on chloroplast DNA restriction site data. There are discrepancies between the two analyses, but if we consider groupings reflected in the present classification there is much congruency. With the exception of four genera, all the genera are positioned in the same group of taxa in the two analyses. Clades of taxa representing three of the four subfamilies (~the Antirheoideae, ~the Rubioideae, and the ~Ixoroideae) are monophyletic, while the fourth subfamily Cinchonoideae is shown to be paraphyletic. Both analyses support a widened tribe Chiococceae, including the former subtribe Portlandiinae (Condamineeae). Furthermore, in both analyses the tribe Hamelieae is placed outside the subfamily Rubioideae where it is now housed. In search for the most plausible sister group to the Rubiaceae, the genus Cinchona (Rubiaceae) was analyzed together with 13 genera of the Loganiaceae, Nerium (Apocynaceae), and Exacum (Gentianaceae). Cornus (Comaceae), Olea (Oleaceae), and these two genera together were used as outgroups. The analysis, including 25 characters, 16 taxa, and with Cornus and Olea together as an outgroup, resulted in four equally parsimonious trees, with a consistency index of 0.53 and a retention index of 0.62. The non-Loganiaceae taxa Cinchona (Rubiaceae), Nerium (Apocynaceae), and Exacum (Gentianaceae) were all found to have their closest relatives within the Loganiaceae indicating that the Loganiaceae are paraphyletic and ought to be reclassified. As a result of the morphological data the most plausible sister group to the Rubiaceae is the tribe Gelsemieae of the Loganiaceae.     

Classification of apocynaceae s.l. according to a new approach combining Linnaean and phylogenetic taxonomy

A new approach to a nomenclatural system, including elements from both Linnaean and phylogenetic nomenclature, is proposed. It is compatible with the existing Linnaean system, including "standard names" corresponding to principal and secondary ranks, and uses a variant of the definitions from the Phylocode system. A new infrafamilial classification, using this nomenclatural approach, of the Apocynaceae s.l. (i.e., including the Asclepiadaceae) based mainly on analyses of rbcL and ndhF data is discussed. Twenty-one tribes and four rankless taxa are defined.     

浙江姬小蜂科分类学研究(膜翅目:小蜂总科)

在最近提交的天目山姬小蜂科的报告基础上,本文再次对浙江该科昆虫进行了分类学研究。根据从天目山采集的标本和中国科学院动物研究所动物标本馆馆藏标本。研究了分属于姬小蜂科4个亚科的20属35种。文中提供了亚科,属及物种的检索表,同时还提供了关键性状的扫描电镜照片,每个物种均附有最新的寄主记录和分布记录。根据现有材料,对浙江姬小蜂科昆虫进行了初步的分析。     

Phylogeny and classification of Leptophlebiidae (Ephemeroptera) with an emphasis on Neotropical fauna

Mayflies from the family Leptophlebiidae are cosmopolitan and highly diverse morphologically; they are also the largest family in numbers of genera and the second in number of species in the order Ephemeroptera. In spite of their broad diversity and the efforts employed to understand the evolution of this group, the internal classification of Leptophlebiidae remains controversial at all levels. More recently, important changes have been incorporated into the systematics of the family, increasing the number of subfamilies (from two to six) and recognizing several tribes. We present a phylogeny of the family based on 153 taxa (53 genera) and two molecular markers, representing 1655 bp, and verify the taxonomic status of the subfamilies, tribes and complexes. Based on these results, the number of subfamilies has been increased from six to eight and one new tribes and two new subtribes have been added. In addition, new ranks are proposed and the concept of Atalophlebiinae revised, including genera with distributions in the Australasian and Neotropical regions.     

Phylogenetic relationships, character evolution, and taxonomic implications within the slipper lobsters (Crustacea: Decapoda: Scyllaridae)

The slipper lobsters belong to the family Scyllaridae which contains a total of 20 genera and 89 species distributed across four subfamilies (Arctidinae, Ibacinae, Scyllarinae, and Theninae). We have collected nucleotide sequence data from regions of five different genes (16S, 18S, COI, 28S, H3) to estimate phylogenetic relationships among 54 species from the Scyllaridae with a focus on the species rich subfamily Scyllarinae. We have included in our analyses at least one representative from all 20 genera in the Scyllaridae and 35 of the 52 species within the Scyllarinae. Our resulting phylogenetic estimate shows the subfamilies are monophyletic, except for Ibacinae, which has paraphyletic relationships among genera. Many of the genera within the Scyllarinae form non-monophyletic groups, while the genera from all other subfamilies form well supported clades. We discuss the implications of this history on the evolution of morphological characters and ecological transitions (nearshore vs. offshore) within the slipper lobsters. Finally, we identify, through ancestral state character reconstructions, key morphological features diagnostic of the major clades of diversity within the Scyllaridae and relate this character evolution to current taxonomy and classification.     

Phylogeny, diversity, and classification of the Accipitridae based on DNA sequences of the RAG-1 exon

The avian family Accipitridae has historically been divided into subfamilies or tribes based on features such as general resemblance, feeding ecology, and behavior. Consequently, the monophyly of those groups has been questionable. Recently, three phylogenetic analyses of a majority of the genera have appeared, one based on osteology, one on DNA sequences from a single mitochondrial gene, and the third on mitochondrial plus nuclear DNA sequences, and the resulting phylogenies were in substantial disagreement concerning the composition and basal branching patterns of the clades and hence require further analysis and confirmation. Here we use DNA sequences from the large nuclear RAG-1 exon to investigate the phylogenetic relationships of these birds. Our results largely corroborated the prior study that included nuclear genes. We found strong support for a monophyletic clade comprising the secretarybird Sagittarius serpentarius , the osprey Pandion haliaetus , and the traditional accipitrids. However, every one of the traditionally recognized subfamilies of accipitrids was found to be polyphyletic. The most basal nodes in the phylogeny separate small clades of insectivorous and scavenger species, such as kites and Old World vultures, from the rest of the family. The speciose genera of bird and mammal predators are all relatively derived (terminal) in the phylogeny. Many of the basal clades are cosmopolitan in their distributions, consistent with the great mobility of these raptors. A new classification is proposed that eliminates the problem of polyphyletic intrafamilial taxa.     

Implications of the changing phylogenetic relationships of Acacia s.l. on the biological control of Vachellia nilotica ssp. indica in Australia

Plant relationships have implications for many fields including weed biological control. The use of DNA sequencing and new tree building algorithms since the late 1980s and early 1990s have revolutionised plant classification and has resulted in many changes to previously accepted taxonomic relationships. It is critical that biological control researchers stay abreast of changes to plant phylogenies. One of the largest plant genera, Acacia, has undergone great change over the past 20 years and these changes have ramifications for weed biological control projects in a number of countries. Vachellia nilotica (prickly acacia) is a major weed in Australia, originating from the Indian subcontinent and Asia, and it has been a target for biological control since 1980. Once a member of Acacia, a large (>1,000 spp.) and iconic group in Australia, prickly acacia is now part of the genus Vachellia. Current knowledge suggests that Vachellia is more closely related to mimosoid genera than it is to Acacia s.s. There has also been a recent reclassification of legume subfamilies with subfamily Mimosoideae now part of subfamily Caesalpinioideae, and four new subfamilies. In this paper we review the changes that have occurred to this group since the prickly acacia biological control project began and discuss the implications for the project. A new host test list for quarantine testing is proposed. Developed following the modernisation of the centrifugal‐phylogenetic method, it is shorter than past lists, containing 46 species, although still lengthy because of the expectations of regulatory bodies, which are slower to accept advances in scientific knowledge. The list includes five Vachellia species, six “Mimoseae” species and 26 Acacia species. The number species from legume subfamilies other than the new Caesalpinioideae is greatly reduced.     

The Chemotaxonomy of Cynanchum and Its Allied Genera

This paper is a continuation of “Chemotaxonomy of Cynanchum and Its Allied Genera with Notes on the Generic Characteristics of Vincetoxicum”, published in Acta Botanica Yunnanica (11: 41-50, 1989). There are other three sections of Cynanchum in “Flora Reipublicae Popularis Sinicae” (Vol. 63), which were considered as separate genera by some authors. These sections are Cynanchum Sect. Rhodostegiella, Sect. Seutera and Sect. Cyathella, also of a single corona. They were not given generic rank as chemical data of Sect. Rhodostegiella were inavailable. It is for this reason that we have studied the chemical constituents of this section. A very marvellous result was obtained. No C₂₁ steroidal compounds but flavonols were identified from the section Rhodostegiella. Some flavones were isolated from Hoya bella of Asclepiadaceae (Bass et al., 1979), but no report has been given from Cynanchum and the tribe Asclepiadeae up to now. The family Asclepiadaceae mainly contains C21 steroids, but some genera contain cardenolides or terpenoids, and only a few genera, such as Cryptolepis and Tylophora, have alkaloids. The genus Cynanchum sensu Tsiang et P. T. Li, like most members of Asclepiadaceae, has C₂₁ steroids except the section Rhodostegiella. Flavonols, therefore, may be the characteristic of the section. Considering the corona characteristics and geographical distribution, we suggest that the section Rhodostegiella be given generic rank. Because of the separation of Vincetoxicum Wolf and Rhodostegiella (Pobed.) C. Y. Wu et D. Z. Li from Cynanchum sensu Tsiang et P.T. Li, the rank of the sections Seutera and Cyathella should be reconsidered. Obviously, the problem of evolution of the sections of Cynanchum sensu Tsiang et P. T. Li is complicated. The corona, chemical constituents, habit, root-type are not congruent with one another in evolution. The authors support the establishment of the genera Seutera and Cyathella by Reichenbach (1828) and Decne (1838) respectively. Taxonomic treatments, including a new status, Rhodostegiella (Pobed.) C.Y. Wu et D. Z. Li, and 12 new combinations, are given.Key words Cynanchum Linn.; Rhodostegiella (Pobed.) C. Y. Wu et D. Z. Li; Seutera Reichb.; Cyathella Decne.; Flavonols; Chemotaxonomy; New combination     

Deviating chromosome numbers in Asclepiadaceae

The most common base chromosome number in the Asclepiadaceae is x = 11. Deviating base chromosome numbers have been found in the genera Cynanchum, Microloma , and Sarcostemma . An account is also given of previously published deviating chromosome numbers in the Asclepiadaceae.     

The Drosophilidae (Diptera) of the Scattered Islands,with the description of a novel association with Leptadenia madagascariensis Decne. (Apocynaceae)

Thirteen drosophilid species belonging to seven genera and two subfamilies are reported from three coral islands (namely Europa, Juan de Nova and Glorioso) that belong to the Scattered Islands in the Indian Ocean. Five species are cosmopolitan and five are African. Three are endemic to the insular Western Indian Ocean, including a presumably new Scaptodrosophila species. On the island of Juan de Nova, most captured flies had pollinia attached to the bases of their proboscis. DNA analysis using the rbcl gene revealed that these pollinia belong to the genus Leptadenia (Apocynaceae), of which a single species L. madagascariensis, endemic in Madagascar and Comoros, is present in this island. This is the first reported association between this plant and drosophilids.     

西双版纳植物区系的特点与亲缘

西双版纳植物区系经初步资料整理统计有种子植物 3336种 ,隶属于 1 1 4 0个属 ,大约 1 97个科。植物区系成分构成是热带分布的科和属分别占总科和属数的 60 .1 %和 83.5% ,在热带分布属中 ,又以热带亚洲成分占优势 (占 32 .8% )。通过植物区系组成和地理成分分析 ,认为该植物区系是热带性质的 ,属于热带亚洲区系的一部分 ,并带有明显热带边缘性质和几种地理成分交汇的特点。西双版纳植物区系与广西热带植物区系和海南热带植物区系在发生上同源 ,主要来自于古南大陆与古北大陆区系成分的融合 ,在发展上受热带亚洲植物区系的渗透和强烈影响。     

A reconsideration of the generic circumscription of Heterostemma Wight & Arn. (Asclepiadaceae) and a new species from India

Heterostemma Wight & Arn. (Asclepiadaceae) and a new species from India. Heterostemma vasudemni Swarupanandan & Mangaly, a new species belonging to Asclepiadaceae is described and illustrated. The discoid-urceoloate corolla and the staminal coronal scales with a ventral appendage show this species to be intermediate between the two genera Heterostemma and Oianthus. Basically, the concepts of the two genera differ only in a pair of characters which, in the light of the new species seem not to be particularly diagnostic. On this basis it is proposed to merge the two genera together, by reducing Oianthus as a section under Heterostemma. Consequently, three new combinations are proposed. Examination of recent collections extends the distribution of what was 0. decanense , earlier considered to be endemic to Maharashtra, further south to Kerala.