The phylogenetic significance of the carpophore in Apiaceae

Background and aims

Fruit structural characters have traditionally been important in the taxonomy of the family Apiaceae. Previous investigations using a limited number of taxa have shown that the carpophore may be especially useful in helping to circumscribe subfamily Azorelloideae. The present study examines, for the first time, carpophore structure in 92 species from 43 genera, representing all subfamilies of Apiaceae, and including all genera assigned to subfamily Azorelloideae. Phylogenetic interpretations are made for the first time, using all available information, and a standard terminology is proposed to describe the various character states found in carpophores.

Methods

Carpophore structure was studied in detail using light microscopy.

Key Results

Carpophores, when present, may be categorized into two main groups (B and C) based mainly on the arrangement of the vascular bundles in transverse section, and further divided into six sub-types according to the length of the carpophore (short in B1 and C1) and whether they are entire (B1–B3 and C1) or bifurcate (B4 and C2). Free carpophores are absent in subfamily Mackinlayoideae, and in tribes Lichtensteinieae and Phlyctidocarpeae, which have two opposite vascular bundles (Group A). Entire carpophores with one or two vascular bundles, or bifurcate carpophores with lateral vascular bundles (arranged side by side within the commissural plane), are the main types characterizing Azorelloideae. The short, hygroscopic carpophores found in Choritaenia are unique in Apiaceae and provide additional evidence for the exclusion of this genus from Azorelloideae. Carpophore type C2 is typical for most Apioideae sensu lato (exceptions are, for example, Arctopus and Alepidea, which have type B2).

Conclusions

A single carpophore and ventral vascular bundles not forming free carpophores are proposed to be the ancestral conditions in Apiaceae, while bifurcate carpophores with opposite vascular bundles are the derived state, present in most Apioideae. Secondary reductions seem to have occurred in several unrelated lineages in all major groups, e.g. many Azorelloideae, several protoapioids (including nearly all members of the tribe Saniculeae) and 29 euapioid genera (e.g. some Oenantheae).     

Billburttia, a new genus of Apiaceae (tribe Apieae) endemic to Madagascar

The genus Billburttia is described to include two new species endemic to Madagascar, B. capensoides and B. vaginoides. Both species were tentatively placed within the problematic genus Peucedanum L. (as Peucedanum sp. A and Peucedanum sp. B) based on their dorsally compressed fruits lacking prominent dorsal ribs and with winged marginal ribs. Recently, however, the African members of Peucedanum have been shown to be only distantly related to the type of the genus and have therefore been segregated into six African endemic genera. While the Malagasy species appear superficially similar to members from one of these segregate genera, viz. Notobubon, they differ markedly in their fruit anatomical characters, notably a narrower commissure, six commissural vittae, vascular tissue in the tip of the ribs and sphaerocrystals distributed in and around the epidermis. The latter two characters apparently represent apomorphies for Billburttia. The non-peucedanoid affinity of the genus, as suggested by the fruit anatomical data, was confirmed using ITS and rps16 intron sequences. Both parsimony and Bayesian analyses of these data place Billburttia within the tribe Apieae of subfamily Apioideae and not closely related to either Peucedanum (Selineae) or the African peucedanoid genera (Lefebvrea clade of Tordylieae).     

23种伞形科植物果实形态及其分类学意义

该实验采用徒手切片法对伞形科17属23种植物的果实进行了外部形态和解剖特征的观察,结果表明:伞形科果实有背腹压扁、两侧压扁和不压扁3种类型;侧棱有宽、有窄;油管的分布有棱槽单油管型和棱槽多油管型;花柱基多数为圆锥状,少数种的花柱基为扁平垫状;萼齿明显或不明显。通过进一步对属间和属内果实解剖特征的比较得出:(1)果实表面被钩刺或刚毛及果棱特征在属间差异明显,在属内表现出一致性,可作为伞形科属间分类的依据。(2)果实横切面的形状、胚乳腹面的凹凸以及萼齿的形态特征在属内种间的分类研究中有重要的意义。基于果实形态特征,编制了17属23种植物的分种检索表。     

The taxonomic value of fruit structure in the Chinese endemic genus Dickinsia (Apiaceae)

The monotypic Dickinsia Franch. is misplaced in its current position in the tribe Hydrocotyleae and is transferred to the tribe Mulineae. Its mericarps are strongly dorsally compressed and the lateral ribs of each mericarp are much more prominent than the dorsal rib. The carpophore is free. These characters are typical for genera of the Mulineae but are obviously different from other genera of Hydrocotyleae, which all have laterally compressed mericarps, dorsal ribs which are more expanded than the lateral ribs and an absence of free carpophores.     

Two new genera and species of the subfamily Brosmophycinae (Bythitidae,Ophidiiformes) from Northern Australia

Two new bythitid genera and species of the subfamily Brosmophycinae are described from Northern Territory, Australia.Brosmolus longicaudus, described from a single male specimen, is unique in the tribe Brosmophycini in having the anal fin origin well anterior to the midpoint of the body and thin, transparent skin on the head and body.Beaglichthys macrophthalmus, described from a single female specimen, differs from all other genera in the subfamily by the following combination of characters: eight branchiostegal rays, eye diameter longer than snout length, cheek scaly, anal fin origin at midpoint of body, three developed rakers on the first gill arch, 12 caudal fin rays, and 14 precaudal vertebrae.     

Leaf anatomy as a contribution to the taxonomy of Salacioideae N.Hallé ex Thorne & Reveal (Celastraceae)

The current classification systems recognize Salacioideae as a monophyletic group within Celastraceae. Nonetheless, some divergences exist for genera: in some cases, most species of the subfamily have been included in only two genera; in others, these genera have been subdivided. This study characterizes the leaf anatomy of 31 species of the subfamily Salacioideae as a contribution to identifying them through features that may also help distinguish among genera. Cross-sections of the median region of the leaf blade and of the petiole and dissociated and macerated epidermis were analyzed. Taxonomically relevant anatomical characters include the type of crystals in the parenchymatous tissue (monocrystals in Cheiloclinium and druses in other genera); the presence of laticifers in Cheiloclinium and Tontelea only; the variable form of the petiole vascular system among studied species; the type of stomata (cyclocytic with two concentric circles of subsidiary cells in P. dulcis; anomocytic in T. attenuata, T. fluminensis, and T. leptophylla; laterocytic in C. anomalum and C. hippocrateoides; and ciclocytic in the other species); the sinuosity of the anticlinal walls of the epidermal cells (sinuous in Cheiloclinium and Peritassa, except P. laevigata, and in S. arborea, S. insignis, S. mosenii, S. nemerosa, and S. opacifolia, and straight in all other studied species); the presence of crystalliferous idioblasts in the epidermis of P. dulcis, P. flaviflora, and P. mexiae; and the presence, form, and disposition of sclereids in the leaf blade, which is a highly variable character among the studied species.     

Systematic and phylogenetic implications of the wood anatomy of six Neotropical genera of Primulaceae

Our main goals were to identify diagnostic characters at the species, genus, and subfamily levels, find anatomical features with potential for future morphological and molecular (combined) phylogenetic analyses, and to reconstruct the evolution of wood anatomical characters in two subfamilies of Primulaceae in a molecular phylogenetic framework. We investigated twenty-seven species from the woody Myrsinoideae (4 genera) and Theophrastoideae (2 genera) using scanning electron, light, and epifluorescence microscopy. Samples were prepared using standard protocols. Based on the wood anatomical characters, we were able to identify synapomorphies and to detect evolutionary trends of interest for the genera and subfamilies. Both subfamilies share the presence of diffuse porosity, simple perforation plates, septate fibres, and scanty paratracheal axial parenchyma. Theophrastoideae species have rays?>?10 cells wide and short (<?350 µm) vessel elements, and Myrsinoideae have breakdown areas in rays and longer vessel elements. Ardisia and Stylogyne have scalariform intervessel pits, Myrsine exhibit breakdown areas in rays, and two Cybianthus species from subgenus Weilgetia have distinguishing features (e.g., scalariform perforation plate in C. nemoralis and the absence of rays in C. densiflorus). Overall, when combining characters, we were able to segregate the Neotropical Primulaceae subfamilies and genera from each other and from the subfamily Maesoideae based on wood anatomy.     

Parabrosmolus novaeguineae,a new genus and species of the subfamily Brosmophycinae from Papua New Guinea (Bythitidae,Ophidiiformes)

Parabrosmolus novaeguineae, a new genus and species of the subfamily Brosmophycinae (family Bythitidae) is described, based on a single specimen from Papua New Guinea. The genus is unique in the tribe Brosmophycini in having six branchiostegal rays and ten precaudal vertebrae, and is also similarly distinguished from two brosmophycine genera,Melodichthys andBeaglichthys, tribal allocations of the two latter being uncertain.Parabrosmolus also differs from all other genera in the subfamily by the following combination of characters: head scaly, anal fin origin slightly before midpoint of body, eye diameter shorter than snout length, three developed rakers on first gill arch and 16 (14+2) caudal fin rays.     

Fruit and Seed Anatomy of Chenopodium and Related Genera (Chenopodioideae,Chenopodiaceae/Amaranthaceae): Implications for Evolution and Taxonomy

A comparative carpological study of 96 species of all clades formerly considered as the tribe Chenopodieae has been conducted for the first time. The results show important differences in the anatomical structure of the pericarp and seed coat between representatives of terminal clades including Chenopodium s.str.+Chenopodiastrum and the recently recognized genera Blitum, Oxybasis and Dysphania. Within Chenopodium the most significant changes in fruit and seed structure are found in members of C. sect. Skottsbergia. The genera Rhagodia and Einadia differ insignificantly from Chenopodium. The evolution of heterospermy in Chenopodium is discussed. Almost all representatives of the tribe Dysphanieae are clearly separated from other Chenopodioideae on the basis of a diverse set of characteristics, including the small dimensions of the fruits (especially in Australian taxa), their subglobose shape (excl. Teloxys and Suckleya), and peculiarities of the pericarp indumentum. The set of fruit and seed characters evolved within the subfamily Chenopodioideae is described. A recent phylogenetic hypothesis is employed to examine the evolution of three (out of a total of 21) characters, namely seed color, testa-cell protoplast characteristics and embryo orientation.     

Analysis of the taxonomic structure of the crane fly family Limoniidae (Diptera) based on the larval characters

A morphological study of the larval characters in the genera Elephantomyia, Helius, Microlimonia, Lipsothrix, and Teucholabis was conducted. Significant differences between Elephantomyia and Helius were discovered; Elephantomyia should be placed in the subfamily Hexatominae, while Helius cannot be assigned to any known subfamily. The genus Microlimonia has nothing in common with other representatives of the tribe Limoniini, and the genera Lipsothrix and Teucholabis, with those of Gonomyini; these genera cannot be included in any of the existing tribes. The presently accepted subdivision of the Limoniidae into 4 subfamilies does not reflect its actual diversity.     

Clarification of the relationship beteen Apiaceae and Araliaceae based on matK and rbcL sequence data

Apiaceae and Araliaceae (Apiales) represent a particularly troublesome example of the difficulty in understanding evolutionary relationships between tropical-temperate family pairs. Previous studies based on rbcL sequence data provided insights at higher levels, but were unable to resolve fully the family-pair relationship. In this study, sequence data from a more rapidly evolving gene, matK, was employed to provide greater resolution. In Apiales, matK sequences evolve an average of about two times faster than rbcL sequences. Results of phylogenetic analysis of matK sequences were first compared to those obtained previously from rbcL data; the two data sets were then combined and analyzed together. Molecular analyses confirm the polyphyly of apiaceous subfamily Hydrocotyloideae and suggest that some members of this subfamily are more closely related to Araliaceae than to other Apiaceae. The remainder of Apiaceae forms a monophyletic group with well-defined subclades corresponding to subfamilies Apioideae and Saniculoideae. Both the matK and the combined rbcL-matK analyses suggest that most Araliaceae form a monophyletic group, including all araliads sampled except Delarbrea and Mackinlaya. The unusual combination of morphological characters found in these two genera and the distribution of matK and rbcL indels suggest that these taxa may be the remnants of an ancient group of pro-araliads that gave rise to both Apiaceae and Araliaceae. Molecular data indicate that the evolutionary history of the two families is more complex than simple derivation of Apiaceae from within Araliaceae. Rather, the present study suggests that there are two well-defined "families," both of which may have been derived from a lineage (or lineages) or pro-araliads that may still have extant taxa.     

Phylogeny of the Archiborborinae (Diptera: Sphaeroceridae) Based on Combined Morphological and Molecular Analysis

The Archiborborinae is a diverse Neotropical subfamily of Sphaeroceridae, with many undescribed species. The existing generic classification includes three genera consisting of brachypterous species, with all other species placed in the genus Archiborborus. We present the first phylogenetic hypothesis for the subfamily based on morphological, molecular, and combined datasets. Morphological data include 53 characters and cover all valid described taxa (33 species in 4 genera) in the subfamily, as well as 83 undescribed species. Molecular data for five genes (mitochondrial 12S rDNA, cytochrome c oxidase subunit I, and cytochrome B, and nuclear alanyl-tRNA synthetase and 28S rDNA) were obtained for 21 ingroup taxa. Data support the separation of the Archiborborinae from the Copromyzinae, with which they were formerly combined. Analyses support consistent groups within the subfamily, but relationships between groups are poorly resolved. The validity of the brachypterous genera Penola Richards and Frutillaria Richards is supported. The former genus Archiborborus Duda is paraphyletic, and will be divided into monophyletic genera on the basis of this work. Aptery and brachyptery have evolved multiple times in the subfamily. Antrops Enderlein, previously including a single brachypterous species, is a senior synonym of Archiborborus.     

Molecular systematics of Apiaceae subfamily Apioideae: phylogenetic analyses of nuclear ribosomal DNA internal transcribed spacer and plastid RPO C1 intron sequences

Evolutionary relationships among representatives of Apiaceae (Umbelliferae) subfamily Apioideae have been inferred from phylogenetic analyses of nuclear ribosomal DNA internal transcribed spacer (ITS 1 and ITS 2) and plastid rpoC1 intron sequences. High levels of nucleotide sequence variation preclude the use of the ITS region for examining relationships across subfamilial boundaries in Apiaceae, whereas the rpoC1 intron is more suitably conserved for family-wide phylogenetic study but is too conserved for examining relationships among closely related taxa. In total, 126 ITS sequences from subfamily Apioideae and 100 rpoC1 intron sequences from Apiaceae (all three subfamilies) and outgroups Araliaceae and Pittosporaceae were examined. Phylogenies estimated using parsimony, neighbor-joining, and maximum likelihood methods reveal that: (1) Apiaceae subfamily Apioideae is monophyletic and is sister group to Apiaceae subfamily Saniculoideae; (2) Apiaceae subfamily Hydrocotyloideae is not monophyletic, with some members strongly allied to Araliaceae and others to Apioideae + Saniculoideae; and (3) Apiaceae subfamily Apioideae comprises several well-supported subclades, but none of these coincide with previously recognized tribal divisions based largely on morphological and anatomical characters of the fruit. Four major clades in Apioideae are provisionally recognized and provide the framework for future lower level phylogenetic analyses. A putative secondary structure model of the Daucus carota (carrot) rpoC1 group II intron is presented. Of its six major structural domains, domains II and III are the most, and domains V and VI the least, variable.     

Systematics of the subfamily Haplorchiinae (Trematoda: Heterphyidae), based on nuclear ribosomal DNA genes and ITS2 region

Phylogenetic relationships of 6 species in the trematode subfamily Haplorchiinae were analyzed using small and large subunit of ribosomal DNA genes (18S rDNA and 28S rDNA) and internal transcribed spacer subunit II (ITS2) region as molecular markers. Maximum Likelihood and Bayesian inference analyses of combined rDNAs and ITS2 indicated a close relationship between the genera Haplorchis and Procerovum, while these two genera were distinct from Stellantchasmus falcatus. These phylogenetic relationships were consistent with the number of testes but not with the characters of the modification of the seminal vesicle or of the ventral sucker. Although three Haplorchis spp. were, together with Procerovum, in the same cluster, their mutual topology was incongruent between rDNA and ITS2 trees. Phylogenetic analyses using other molecular markers with more species are necessary to work out solid phylogenetic relationships among the species in this subfamily.