A revised classification of the Apocynaceae s.l.

The Asclepiadaceae, as traditionally defined, have repeatedly been shown to be an apomorphic derivative of the Apocynaceae. It has often been recommended that the Asclepiadaceae be subsumed within the Apocynaceae in order to make the latter monophyletic. To date, however, no comprehensive, unified classification has been established. Here we provide a unified classification for the Apocynaceae, which consists of 424 genera distributed among five subfamilies: Rauvolfioideae, Apocynoideae, Periplocoideae, Secamonoideae, and Asclepiadoideae. Keys to the subfamilies and tribes are provided, with lists of genera that (as far as we have been able to ascertain) are recognized in each tribe.     

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.     

Vessel grouping patterns in subfamilies Apocynoideae and Periplocoideae confirm phylogenetic value of wood structure within Apocynaceae

This study contributes to our understanding of the phylogenetic significance and major evolutionary trends in the wood of the dogbane family (Apocynaceae), one of the largest and economically most important angiosperm families. Based on LM and SEM observations of 56 Apocynoideae species-representing all currently recognized tribes-and eight Periplocoideae, we found striking differences in vessel grouping patterns (radial multiples vs. large clusters) between the mainly nonclimbing apocynoid tribes (Wrightieae, Malouetieae, Nerieae) and the climbing lineages (remaining Apocynoideae and Periplocoideae). The presence of large vessel clusters in combination with fibers in the ground tissue characterizing the climbing Apocynoideae and Periplocoideae clearly contrasts with the climbing anatomy of the rauvolfioids (solitary vessels plus tracheids in ground tissue), supporting the view that (1) the climbing habit has evolved more than once in Apocynaceae, (2) the three nonclimbing apocynoid tribes are basal compared to the climbing apocynoids, and (3) Periplocoideae belong to the crown clade. The wood anatomy within the nonclimbing and climbing lineages is rather homogeneous, although a combination of specific characters (e.g. presence of septate fibers, axial parenchyma distribution, abundance of uniseriate compared to multiseriate rays, and presence and location of prismatic crystals) may be used to identify several tribes.     

The tribal position of Fockea and Cibirhiza (Apocynaceae: Asclepiadoideae): evidence from pollinium structure and cpDNA sequence data

The pollinium morphology of the two members of the Asclepiadoideae, tribe Fockeeae, Fockea Endl. and Cibirhiza Bruyns, has been studied in detail and compared with that of eight genera of Marsdenieae, the tribe in which Fockea and Cibirhiza were previously accommodated and thus their putative closest relatives, as well as nine genera of Asclepiadeae. Both Fockea and Cibirhiza have several morphological characteristics in common, the most important of which is the absence of well-developed caudicula, which distinguishes them from all other genera of Asclepiadoideae known. The pollinium structure of these two genera, however, differs significantly. Whereas the pollinium of Cibirhiza consists of single pollen grains and is covered by a pollinium wall, as is typical for other Asclepiadoideae, the pollinium of Fockea consists of tetrads and is not covered by a pollinium wall, a condition otherwise typical of Secamonoideae. Fockea, however, has only two pollinia per anther, as does Cibirhiza and all other Asclepiadoideae, whereas the Secamonoideae have four pollinia per anther. Sequence data from two intergenic spacers, trnT-L and trnL-F and the trnL intron of cpDNA was analyzed. The ingroup included three species of Fockea and one species of Cibirhiza. The outgroup taxa consisted of three representatives each of Periplocoideae, and Secamonoideae and 24 species of Asclepiadoideae, including representatives of all tribes, of which eight genera belong to Marsdenieae, as outgroups. The results of the DNA analysis provide strong support for Fockeeae as a monophyletic tribe, distinct from Marsdenieae and, to the rest of the Asclepiadoideae. With the exception of pollen data, all morphological and molecular evidence clearly support recognition of the tribe Fockeeae. The occurrence of two such significantly different types of pollinia structure – characters elsewhere in the family used to distinguish subfamilies – within the small tribe Fockeeae was unexpected, and can perhaps best be understood as yet another attestment to the basal position of the Fockeeae in the nascence of the Asclepiadoideae.     

Climate niches of milkweeds with plesiomorphic traits (Secamonoideae; Apocynaceae) and the milkweed sister group link ancient African climates and floral evolution

? Premise of the study: Climate change that increases mortality of plants and pollinators can create mate-finding Allee effects and thus act as a strong selective force on floral morphology. Milkweeds (Secamonoideae and Asclepiadoideae; Apocynaceae) are typically small plants of seasonally dry habitats, with pollinia and high pollen-transfer efficiency. Their sister group (tribe Baisseeae and Dewevrella) is mostly comprised of giant lianas of African rainforests, with pollen in monads. Comparison of the two groups motivated a new hypothesis: milkweeds evolved in the context of African aridification and the shifting of rainforest to dry forest. Pollinia and high pollen-transfer efficiency may have been adaptations that alleviated mate-finding Allee effects generated by high mortality during droughts. We formally tested whether milkweeds have a drier climate niche by comparing milkweeds with plesiomorphic traits (Secamonoideae) and the milkweed sister group in continental Africa. ? Methods: We georeferenced specimens of the milkweed sister group and Secamonoideae in continental Africa, extracted 19 climatic variables from the Worldclim model, conducted factor analysis to identify correlated suites of variables, and compared the frequency distributions of the two lineages relative to each factor. ? Key results: The distributions of Secamonoideae and the milkweed sister group differed significantly relative to four factors, each correlated with a distinct suite of climate parameters: (1) air temperature (Secamonoideae: cooler), (2) total and (3) summer precipitation (Secamonoideae: drier), and (4) temperature seasonality and isothermality (Secamonoideae: more seasonal and less isothermal). ? Conclusions: Secamonoideae in continental Africa inhabit drier, cooler sites than do the milkweed sister group, consistent with a shift from rainforests to dry forests in a cooling climate.     

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.     

Asclepiadoideae and Periplocoideae (Apocynaceae s.l.) of the Thunberg herbarium

The 141 specimens of Asclepiadoideae and Periplocoideae in the Thunberg herbarium, Uppsala, Sweden, are identified according to the most recent classification of the two subfamilies.     

Phylogenetic relationships of the Aurantioideae inferred from chloroplast DNA sequence data

The tribes and subtribes of Aurantioideae, an economically important subfamily of the Rutaceae, have a controversial taxonomic history because of the lack of a phylogenetic framework. The rps16 and trnL-trnF sequences of the chloroplast were analyzed phylogenetically to construct an evolutionary history and evaluate the most recent classification system of Swingle and Reece (The Citrus Industry, volume 1 [1967]). Taxa representing tribes Citreae and Clauseneae and five of the six subtribes were sampled. Conflicts in the positions of some taxa between the rps16 and trnL-trnF trees are poorly supported. In all analyses, the Aurantioideae are monophyletic. The strict consensus tree of the combined analysis indicates that the two tribes along with the subtribes sampled are not monophyletic. The combined topology is not congruent with the widely used classification of Aurantioideae by Swingle and Reece. The tribes and subtribes are in need of revision.     

Phylogenetic relationships of the Aurantioideae (Rutaceae) based on the nuclear ribosomal DNA ITS region and three noncoding chloroplast DNA regions, atpB-rbcL spacer, rps16, and trnL-trnF

The tribes and subtribes of Aurantioideae, an economically important subfamily of the Rutaceae, have a controversial taxonomic history because a phylogenetic framework has been lacking. In order to construct an evolutionary history and evaluate the most recent classification system [Swingle and Reece 1967. The botany of Citrus and its wild relatives, in: The Citrus Industry, vol. 1, History, World Distribution, Botany, and Varieties. University of California, Berkeley, pp. 190-430], one nuclear and three noncoding chloroplast genes were sequenced and analyzed phylogenetically along with selected non-molecular characters. Taxa representing tribes Citreae and Clauseneae and their six subtribes were sampled. In all analyses Aurantioideae is monophyletic. The majority-rule consensus tree from the combined analysis indicates that the two tribes are not monophyletic. The combined topology is not congruent with the widely used classification of Aurantioideae by Swingle and Reece (1967). The tribes and subtribes are in need of revision.     

Wood anatomy of Rauvolfioideae (Apocynaceae): a search for meaningful non-DNA characters at the tribal level

Wood anatomical studies in the economically important Apocynaceae or dogbane family are fragmentary. This study represents a first attempt to unravel the phylogenetic significance and major evolutionary trends in the wood of the family, using existing and new microscopic wood observations within the large subfamily Rauvolfioideae. On the basis of LM and SEM observations of 91 species representing all 10 currently recognized tribes, we found that most of the tribes are characterized by a unique combination of wood characters, such as vessel grouping, vessel element length, fiber type, frequency of uniseriate rays, and fused multiseriate rays. Climbing rauvolfioid taxa can generally be distinguished from erect species by their wider vessels, tendency to form paratracheal axial parenchyma, presence of tracheids, and occurrence of laticifers in rays. With respect to the entire family, there is a general phylogenetic trend toward shorter vessel elements, a higher proportion of vessels in multiples and more vessels per multiple, higher tracheid abundance, more paratracheal parenchyma, and fewer cells per axial parenchyma strand in the more derived Apocynaceae. Most of these evolutionary trends are likely to be triggered by drier environmental conditions and/or shifts from an erect to a climbing habit.     

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.     

Generic delimitations in tuberous Periplocoideae (Apocynaceae) from Africa and Madagascar

BACKGROUND AND AIMS: The number of genera included in Apocynaceae subfamily Periplocoideae is a matter of debate. DNA sequences are used here as an independent dataset to clarify generic relationships and classification of the tuberous periplocoid genera and to address the question of the phylogenetic interpretation of pollinia formation in Schlechterella. METHODS: Representatives of nearly all African and Malagasy genera of Periplocoideae possessing root tubers were analysed using ITS and plastid DNA sequence characters. RESULTS: Sequence data from non-coding molecular markers (ITS of nrDNA and the trnT-L and trnL-F spacers as well as the trnL intron of plastid DNA) give support for a broad taxonomic concept of Raphionacme including Pentagonanthus. Together with Schlechterella, which is sister to Raphionacme, all Raphionacme-like taxa form a derived monophyletic group of somewhat diverse species. Sister to the Schlechterella/Raphionacme clade is a clade comprising Stomatostemma and the not truly tuberous vine Mondia. In the combined analysis, sister to these two clades combined is a clade formed by Petopentia natalensis and Periploca. CONCLUSIONS: The recent inclusion of the monotypic South African Petopentia in the monotypic Malagasy endemic Ischnolepis is to be rejected. The Malagasy Camptocarpus is sister to the remainder of Periplocoideae in the ITS and combined analyses, and a Malagasy origin for the subfamily is discussed.     

Sesquiterpene lactone-based classification of three Asteraceae tribes: a study based on self-organizing neural networks applied to chemosystematics

This work describes an application of artificial neural networks on a small data set of sesquiterpene lactones (STLs) of three tribes of the family Asteraceae. Structurally different types of representative STLs from seven subtribes of the tribes Eupatorieae, Heliantheae and Vernonieae were selected as input data for self-organizing neural networks. Encoding the 3D molecular structures of STLs and their projection onto Kohonen maps allowed the classification of Asteraceae into tribes and subtribes. This approach allowed the evaluation of structural similarities among different sets of 3D structures of sesquiterpene lactones and their correlation with the current taxonomic classification of the family. Predictions of the occurrence of STLs from a plant species according to the taxa they belong to were also performed by the networks. The methodology used in this work can be applied to chemosystematic or chemotaxonomic studies of Asteraceae.     

A worldwide phylogenetic classification of the Poaceae (Gramineae) III: An update

We present an updated worldwide phylogenetic classification of Poaceae with 11 783 species in 12 subfamilies, 7 supertribes, 54 tribes, 5 super subtribes, 109 subtribes, and 789 accepted genera. The subfamilies (in descending order based on the number of species) are Pooideae with 4126 species in 219 genera, 15 tribes, and 34 subtribes; Panicoideae with 3325 species in 242 genera, 14 tribes, and 24 subtribes; Bambusoideae with 1698 species in 136 genera, 3 tribes, and 19 subtribes; Chloridoideae with 1603 species in 121 genera, 5 tribes, and 30 subtribes; Aristidoideae with 367 species in three generaand one tribe; Danthonioideae with 292 species in 19 generaand 1 tribe; Micrairoideae with 192 species in nine generaand three tribes; Oryzoideae with 117 species in 19 genera, 4 tribes, and 2 subtribes; Arundinoideae with 36 species in 14 genera and 3 tribes; Pharoideae with 12 species in three generaand one tribe; Puelioideae with 11 species in two generaand two tribes; and the Anomochlooideae with four species in two generaand two tribes. Two new tribes and 22 new or resurrected subtribes are recognized. Forty-five new (28) and resurrected (17) genera are accepted, and 24 previously accepted genera are placed in synonymy. We also provide an updated list of all accepted genera including common synonyms, genus authors, number of species in each accepted genus, and subfamily affiliation. We propose Locajonoa, a new name and rank with a new combination, L. coerulescens. The following seven new combinations are made in Lorenzochloa: L. bomanii, L. henrardiana, L. mucronata, L. obtusa, L. orurensis, L. rigidiseta, and L. venusta.     

Phylogeny and systematics of the subfamily Bryocorinae based on morphology with emphasis on the tribe Dicyphini sensu Schuh,

The classification of the hyperdiverse true bug family Miridae is far from settled, and is particularly contentious for the cosmopolitan subfamily Bryocorinae. The morphological diversity within the subfamily is pronounced, and a lack of explicit character formulation hampers stability in the classification. Molecular partitions are few and only a handful of taxa have been sequenced. In this study the phylogeny of the subfamily Bryocorinae has been analysed based on morphological data alone, with an emphasis on evaluating the tribe Dicyphina sensu Schuh, 1976, within which distinct groups of taxa exist. A broad sample of taxa was examined from each of the bryocorine tribes. A broad range of outgroup taxa from most of the other mirid subfamilies was also examined to test for bryocorine monophyly, ingroup relationships and to determine character polarity. In total a matrix comprising 44 ingroup, 15 outgroup taxa and 111 morphological characters was constructed. The phylogenetic analysis resulted in a monophyletic subfamily Bryocorinae sensu Schuh (1976, 1995), except for the genus Palaucoris, which is nested within Cylapinae. The tribe Dicyphini sensu Schuh (1976, 1995) has been rejected. The subtribe Odoniellina is synonymized with the subtribe Monaloniina and the subtribes Dicyphina, Monaloniina and Eccritotarsina are now elevated to tribal level, with the Dicyphini now restricted in composition and definition. The genus Felisacus is highly autapomorphic and a new tribe – the Felisacini – is erected for the included taxa. This phylogeny of the tribes of the Bryocorinae comprises the following sister‐group relationships: Dicyphini ((Bryocorini + Eccritotarsini)(Felisicini + Monaloniini)).     

An updated classification of Orchidaceae

Since the last classification of Orchidaceae in 2003, there has been major progress in the determination of relationships, and we present here a revised classification including a list of all 736 currently recognized genera. A number of generic changes have occurred in Orchideae (Orchidoideae), but the majority of changes have occurred in Epidendroideae. In the latter, almost all of the problematic placements recognized in the previous classification 11 years ago have now been resolved. In Epidendroideae, we have recognized three new tribes (relative to the last classification): Thaieae (monogeneric) for Thaia, which was previously considered to be the only taxon incertae sedis; Xerorchideae (monogeneric) for Xerorchis; and Wullschlaegelieae for achlorophyllous Wullschlaegelia, which had tentatively been placed in Calypsoeae. Another genus, Devogelia, takes the place of Thaia as incertae sedis in Epidendroideae. Gastrodieae are clearly placed among the tribes in the neottioid grade, with Neottieae sister to the remainder of Epidendroideae. Arethuseae are sister to the rest of the higher Epidendroideae, which is unsurprising given their mostly soft pollinia. Tribal relationships within Epidendroideae have been much clarified by analyses of multiple plastid DNA regions and the low‐copy nuclear gene Xdh. Four major clades within the remainder of Epidendroideae are recognized: Vandeae/Podochileae/Collabieae, Cymbidieae, Malaxideae and Epidendreae, the last now including Calypsoinae (previously recognized as a tribe on its own) and Agrostophyllinae s.s. Agrostophyllinae and Collabiinae were unplaced subtribes in the 2003 classification. The former are now split between two subtribes, Agrostophyllinae s.s. and Adrorhizinae, the first now included in Epidendreae and the second in Vandeae. Collabiinae, also probably related to Vandeae, are now elevated to a tribe along with Podochileae. Malaxis and relatives are placed in Malaxidinae and included with Dendrobiinae in Malaxideae. The increased resolution and content of larger clades, recognized here as tribes, do not support the ‘phylads’ in Epidendroideae proposed 22 years ago by Dressler. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 177 , 151–174.     

A new classification of Cyperaceae (Poales) supported by phylogenomic data

Cyperaceae (sedges) are the third largest monocot family and are of considerable economic and ecological importance. Sedges represent an ideal model family to study evolutionary biology due to their species richness, global distribution, large discrepancies in lineage diversity, broad range of ecological preferences, and adaptations including multiple origins of C₄ photosynthesis and holocentric chromosomes. Goetghebeur′s seminal work on Cyperaceae published in 1998 provided the most recent complete classification at tribal and generic level, based on a morphological study of Cyperaceae inflorescence, spikelet, flower, and embryo characters, plus anatomical and other information. Since then, several family-level molecular phylogenetic studies using Sanger sequence data have been published. Here, more than 20 years after the last comprehensive classification of the family, we present the first family-wide phylogenomic study of Cyperaceae based on targeted sequencing using the Angiosperms353 probe kit sampling 311 accessions. In addition, 62 accessions available from GenBank were mined for overlapping reads and included in the phylogenomic analyses. Informed by this backbone phylogeny, a new classification for the family at the tribal, subtribal, and generic levels is proposed. The majority of previously recognized suprageneric groups are supported, and for the first time, we establish support for tribe Cryptangieae as a clade including the genus Koyamaea. We provide a taxonomic treatment including identification keys and diagnoses for the 2 subfamilies, 24 tribes, and 10 subtribes, and basic information on the 95 genera. The classification includes five new subtribes in tribe Schoeneae: Anthelepidinae, Caustiinae, Gymnoschoeninae, Lepidospermatinae, and Oreobolinae.     

TRIBAL INTERRELATIONSHIPS OF THE ASTERACEAE

Abstract— A cladistic analysis involving 27 tribes and subtribes of Asteraceae and 81 characters is presented. The terminal taxa are mainly those of present tribal classification, though some apparently poly- and paraphyletic tribes, notably the Mutisieae and the Inuleae, have been represented by sub-tribal taxa. Characters are assembled from all available sources. Corolla types, styles and stamens have provided many characters. The Lobeliaceae are used as an outgroup and are considered as the most probable sister group of the Asteraceae. There is a basal dichotomy in the family, the Mutisieae-Barnadesiinae being the monophyletic sister group of the remaining major, also monophyletic part of the family. The recent family division into two subfamilies about equal in size, the Cichorioideae and the Asteroideae, neither represents a basal dichotomy nor a sister group relationship within the Asteraceae. The Asteroideae are monophyletic and have their sister group within the paraphyletic Cichorioideae. Interrelationships among the cichorioid tribes are still unclear. The Lactuceae, Eremothamneae, Vernonieae and Liabeae may be one monophyletic group, and the Arctoteae, Carlineae, Echinopsideae and Cardueae another. The Mutisieae are a paraphyletic grade at the base of the family. Within the subfamily Asteroideae tribal interrelationships are also rather unclear. The Anthemideae and the Heliantheae sensu lato (including the Helenieae, Tageteae, Coreopsideae and all helenioid/helianthoid representatives sometimes placed in the Senecioneae) may be sister groups. The Heliantheae appear to be monophyletic and there is little support for the hypothesis that other tribes are derived from or have their sister group within the Heliantheae. The Astereae and the Eupatorieae may be sister groups, though a closer relationship between the Eupatorieae and the Heliantheae is possible. The Inuleae are a paraphyletic grade group at the base of the subfamily Asteroideae in the same way as the Mutiseae are a grade group at the base of the family.     

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。