Epidermal Features of Leaves and Their Taxonomic Significance in Subfamily Ophiopogonoideae (Liliaceae)

This paper reports epidermal features of leaves in Ophiopogonoideae. Thirty-nine species and one variety (29 species, 1 variety in Ophiopogon, 6 species in Liriope, 4 species in Peliosanthes)were examined under scanning electron microscope. In addition, transections of stomatal apparatuses of six species (Ophiopogon: 3 species; Liriope: 2 species; Peliosanthes: 1 species) were made and examined under light microscope. The stomatal apparatus in Liriope, Ophiopogon and Peliosanthes is of the anomocytic type. These types of epidermal features of leaves in these genera are recognized: Cuticular processes type, No cuticular processes type and No stomatal band type. The cuticular processes type can be further divided into three patterns: Fibrillose, Massive and Wrinkled-massive. The taxonomic value of the epidermal features of leaves in Ophiopogonoideae is rather evident. (1)These epidermal features can be used to distinguish among those species of Ophiopogon, Liriope and Peliosanthes, even in their vegetative state; (2) The different patterns of cuticular processes are helpful to reasonable classification of some species in Ophiopogon, (3)They are of great value for recognizing some sections, (4) These epidermal features of leaves also provide evidence for further discussion on relationships among Ophiopogon, Liriope, and Peliosanthes. The evolutionary trend of the epidermal features of leaves in Ophiopogonoideae is No stomatal band type→No cuticular process type(stomatal band)→Cuticular process type (stomatal band). According to the epidermal features of leaves, flowers and fruits, Ophiopogon, Liriope and Peliosanthes are closely related, forming a subfamily Ophiopogonoideae. Ophiopogon is more close to Liriope than to Peliosanthes, and they should be grouped into the same tribe-Ophiopogoneae. Liriope seems to be more primitive than Ophiopogon. Peliosanthes, which constitutes another tribe of its ownPeliosantheae, is more advanced than Ophiopogon and Liriope, and it might have beenderived from its ancestor early.     

Pollen Morphology and Generic Phylogenetic Relationships in Ophiopogonoideae (Liliaceae)

In the present work, pollen grains of 3 species of Liriope, 24 species of Ophiopogon and 2 species of Peliosanthes were examined under scanning electron microscope, Among them 4 species were also observed under transmission electron microscope. The observation (Table 3)shows that Liriope and Ophiopogon distinctly differ from Peliosanthes in the exine ornamentation and structure, They may be divided into two types: 1. Rugulate-perforate, ektexine with perforate tectum in Liriope and Ophiopogon. 2. Verrucate, Verracae unequal in size, ektexine intectate in Peliosanthes. Pollen morphology shows the close affinity between Liriope and Ophiopogon, but they are very far from Peliosanthes. The correlation between pollen and gross morphology in Liriope, Ophiopogon and Peliosanthes are stated and their evolutionaly trends are discussed in this paper. The pollen characters support the placement of Liriope and Ophiopogon in one tribe—Ophiopogoneae, and Peliosanthes in another tribe—Peliosantheae. Peliosanthes is more advancedthan Liriope and Ophiopogon.     

Molecular phylogenetics of Ruscaceae sensu lato and related families (Asparagales) based on plastid and nuclear DNA sequences

Background

Previous phylogenetics studies of Asparagales, although extensive and generally well supported, have left several sets of taxa unclearly placed and have not addressed all relationships within certain clades thoroughly (some clades were relatively sparsely sampled). One of the most important of these is sampling within and placement of Nolinoideae (Ruscaceae s.l.) of Asparagaceae sensu Angiosperm Phylogeny Group (APG) III, which subfamily includes taxa previously referred to Convallariaceae, Dracaenaaceae, Eriospermaceae, Nolinaceae and Ruscaceae.

Methods

A phylogenetic analysis of a combined data set for 126 taxa of Ruscaceae s.l. and related groups in Asparagales based on three nuclear and plastid DNA coding genes, 18S rDNA (1796 bp), rbcL (1338 bp) and matK (1668 bp), representing a total of approx. 4·8 kb is presented. Parsimony and Bayesian inference analyses were conducted to elucidate relationships of Ruscaceae s.l. and related groups, and parsimony bootstrap analysis was performed to assess support of clades.

Key Results

The combination of the three genes results in the most highly resolved and strongly supported topology yet obtained for Asparagales including Ruscaceae s.l. Asparagales relationships are nearly congruent with previous combined gene analyses, which were reflected in the APG III classification. Parsimony and Bayesian analyses yield identical relationships except for some slight variation among the core asparagoid families, which nevertheless form a strongly supported group in both types of analyses. In core asparagoids, five major clades are identified: (1) Alliaceae s.l. (sensu APG III, Amarylidaceae–Agapanthaceae–Alliaceae); (2) Asparagaceae–Laxmanniaceae–Ruscaceae s.l.; (3) Themidaceae; (4) Hyacinthaceae; (5) Anemarrhenaceae–Behniaceae–Herreriaceae–Agavaceae (clades 2–5 collectively Asparagaceae s.l. sensu APG III). The position of Aphyllanthes is labile, but it is sister to Themidaceae in the combined maximum-parsimony tree and sister to Anemarrhenaceae in the Bayesian analysis. The highly supported clade of Xanthorrhoeaceae s.l. (sensu APG III, including Asphodelaceae and Hemerocallidaceae) is sister to the core asparagoids. Ruscaceae s.l. are a well-supported group. Asparagaceae s.s. are sister to Ruscaceae s.l., even though the clade of the two families is weakly supported; Laxmanniaceae are strongly supported as sister to Ruscaceae s.l. and Asparagaceae. Ruscaceae s.l. include six principal clades that often reflect previously named groups: (1) tribe Polygonateae (excluding Disporopsis); (2) tribe Ophiopogoneae; (3) tribe Convallarieae (excluding Theropogon); (4) Ruscaceae s.s. + Dracaenaceae + Theropogon + Disporopsis + Comospermum; (5) Nolinaceae, (6) Eriospermum.

Conclusions

The analyses here were largely conducted with new data collected for the same loci as in previous studies, but in this case from different species/DNA accessions and greater sampling in many cases than in previously published analyses; nonetheless, the results largely mirror those of previously conducted studies. This demonstrates the robustness of these results and answers questions often raised about reproducibility of DNA results, given the often sparse sampling of taxa in some studies, particularly the earliest ones. The results also provide a clear set of patterns on which to base a new classification of the subfamilies of Asparagaceae s.l., particularly Ruscaceae s.l. (= Nolinoideae of Asparagaceae s.l.), and examine other putatively important characters of Asparagales.     

Vegetative anatomy of Ophiopogoneae (Convallariaceae)

CUTLER, D. F., 1992. Vegetative anatomy of Ophiopogoneae (Convallariaceae). The vegetative anatomy (particularly leaf) of species of Ophiopogon, Liriope and Peliosanthes is described from observations with light and scanning electron microscopy. A syndrome of leaf characters is present, including epidermal features; hypodermal fibre-like cells; raphides and unusual short, square-ended prismatic crystals arranged in plates; phloem with abundant sclerenchyma and frequent individual strands each composed of a sieve tube element and its associated companion cell; and vascular bundles with unusual orientation, which shows the very close inter-relationship between Ophiopogon and Liriope. Peliosanthes shares the phloem type, hypodermal fibre-like cells and raphides, but is less similar in epidermal characters and vascular bundle orientation. The significance of the unusual phloem type is considered in relation to similar types in other members of the Liliiflorae.     

Phylogenetic analyses and chromosome evolution in Convallarieae (Ruscaceae sensu lato), with some taxonomic treatments

The tribe Convallarieae, comprising 10 genera and 95 species, has recently been transferred from its own family to Ruscaceae sensu lato. In this study, sequence data from trnK and rbcL were analyzed for 19 species in 8 genera, and chromosome morphology was analyzed for 17 species in 7 genera. The parsimony analysis of trnK and rbcL sequences showed that Convallarieae are monophyletic. Although early branches did not receive strong bootstrap support, Convallaria diverged at the first branch, followed by Speirantha. The rest of the tribe was split into three, well-supported clades: one with Reineckea, the second with Campylandra and Rohdea, and the third with Tupistra, Tricalistra, and Aspidistra. Two monotypic genera, Rohdea and Tricalistra, were embedded in a clade of Campylandra and of Tupistra, respectively. Three karyotypes were distinguished in the tribe on the basis of the basic number and morphology of metaphase chromosomes: Convallaria type (with x=19 and unimodal chromosome length), Tupistra type (with x=19 and trimodal chromosome length), and Aspidistra-elatior type (with x=18 and trimodal chromosome length). The character-state distribution in the molecular tree showed that the Convallaria type is plesiomorphic, from which was derived the Tupistra type and subsequently the Aspidistra-elatior type. Taxonomic treatments to transfer Campylandra to Rohdea and Tricalistra to Tupistra are also given.     

SCAR markers for discriminating species of two genera of medicinal plants, Liriope and Ophiopogon

The development of DNA markers that can closely discriminate between Liriope and Ophiopogon species is vital for efficient and accurate identification of these species, and to ensure the quality, safety, and efficacy of medicines made from these plants. We developed species-specific molecular markers for these two genera. Forty RAPD primers were tested to detect polymorphism; species-specific RAPD bands were gel-purified, cloned, and sequenced. Primers for sequence-characterized amplified regions (SCARs) were then designed, based on nucleotide sequences of specific RAPD primers. SCAR markers SA06 and SB05, specific to Ophiopogon japonicus, amplified 460- and 553-bp DNA fragments, respectively. The marker SA12 amplified a 485-bp fragment specific to Liriope platyphylla. This is the first report of a species-specific SCAR marker for this group. These markers will be useful for rapid identification of closely related Liriope and Ophiopogon species.     

Phylogeny and biogeographic diversification of Maianthemum (Ruscaceae: Polygonatae)

Maianthemum (Ruscaceae) comprises 28-38 species and includes the two traditionally recognized genera: Maianthemum sensu stricto and Smilacina. Thirty-seven samples representing 22 species of Maianthemum and six closely related outgroup taxa were sequenced for eight chloroplast and nuclear markers (trnL-F, rps16, rpl16, psbA-trnH, rbcL, ndhF, trnK, and ITS) with a total length of nearly 10,000 base pairs. Phylogenetic analyses supported the monophyly of Maianthemum with Maianthemum sensu stricto nested within Smilacina. Almost all species from the eastern Himalayan region in SW China except for Maianthemum tatsienense and M. stenolobum form a well supported clade. This clade is characterized morphologically by short filaments and large anthers, relatively large flowers, and pubescent stems and leaves. Maianthemum tatsienense and M. stenolobum from SW to central China form another clade. The other species from eastern Asia (central to NE China and Japan) and the New World fall into several clades. The intercontinental disjunction between eastern Asia and North America in Maianthemum sensu stricto is estimated to be at 1.68 million years ago (mya) with the Bayesian relaxed clock relying on uncorrelated rates. A recent radiation at about 2.04mya is suggested in the high mountains of SW China, corresponding to the geographical heterogeneity in that region after the uplift of the Himalayas. Long distance dispersal by birds may have facilitated the evolution of their intercontinental disjunction and their biogeographic diversifications in SW China.     

Phylogeny of Agavaceae based on ITS rDNA sequence variation

Several systems of classification have been proposed for the family Agavaceae. A distinctive bimodal karyotype and similarities of fruits and seeds strongly support close relationships among Yucca, Hesperaloë, Beschorneria, Furcraea, Agave, Manfreda, Polianthes, Prochnyanthes, and perhaps Hosta. However, Dasylirion, Beaucamea, Nolina, Calibanus, Dracaena, and Sansevieria differ in so many cytological and morphological features that many have concluded they should be excluded from Agavaceae and separated into two families, Nolinaceae and Dracaenaceae. Chloroplast DNA restriction site data support these separations and indicate that Nolinaceae and Dracaenaceae are very close to Convallariaceae (Maianthemum, Convallaria, Aspidistra, Liriope, etc.). In this paper we report the results of an ITS rDNA sequencing study of 40 taxa in Agavaceae sensu lato and related groups in the order Asparagales. Sequence alignments were optimized using the Consistency Index, Retention Index, and Rescaled Consistency Index to find the alignment that exhibited the least amount of homoplasy. The results of our study are congruent with the conclusions drawn from cytological, immunological, cpDNA, and rbcL studies, which support a narrow interpretation of Agavaceae and a close relationship among Convallariaceae, Dracaenaceae, and Nolinaceae. In addition, the ITS sequence data provide evidence for some interesting relationships within these families.     

New species of Ophiopogon,Peliosanthes and Tupistra (Asparagaceae s.l.) in the flora of Vietnam

Five new species named Peliosanthes aperta, P. elegans, P. kenhillii, Tupistra densiflora and T. patula are described and illustrated. These species are very local in distribution and endemic to northern or southern Vietnam. Two other species, Ophiopogon ogisui and Peliosanthes griffithii, are recorded as new to the flora of Vietnam. A key to the species of Tupistra occurring in Indochina and its neighboring regions is also provided.     

Molecular phylogenetic analysis of Passifloraceae sensu lato (Malpighiales) based on plastid and nuclear DNA sequences

A phylogenetic analysis of Passifloraceae sensu lato was performed using rbcL, atpB, matK, and 18S rDNA sequences from 25 genera and 42 species. Parsimony analyses of combined data sets resulted in a single most parsimonious tree, which was very similar to the 50% majority consensus tree from the Bayesian analysis. All nodes except three were supported by more than 50% bootstrap. The monophyly of Passifloraceae s.l. as well as the former families, Malesherbiaceae, Passifloraceae sensu stricto, and Turneraceae were strongly supported. Passifloraceae s.s. and the Turneraceae are sisters, and form a strongly supported clade. Within Passifloraceae s.s., the tribes Passifloreae and Paropsieae are both monophyletic. The intergeneric relationships within Passifloraceae s.s. and Turneraceae are roughly correlated with previous classification systems. The morphological character of an androgynophore/gynophore is better used for characterizing genera grouping within Passifloraceae s.s. Other morphological characters such as the corona and aril are discussed.     

Pollen Morphology and Epidermal Characters of Leaves in Convallarieae (s. l.)

The tribe Convallarieae (sensu Krause 1930) consists of 7 genera, i.e. Convallaria, Speirantha, Reineckia, Theropogon, Tupistra, Rohdea and Aspidistra, but now generally recognized as two tribes, Convallarieae (the former 4 genera) and Aspidistreae (the rest). Observed in this work were pollen morphology of 17 species and epidermal characters of leaves of 12 species. All the 7 genera are covered in observations. Pollen grains in Convallarieae (s. str.) are all monosulcate and boat-shaped (Plate 1: A-F). The exine is rather uniformly microperforate (Plate 1: A-F); only Theropogon is exceptional in this respect: it has rugulate exine (Plate 1: O, P). Tang and Zhang (1985) have pointed out the heterogeneity of Theropogon in this tribe. Pollen morphology in the tribe Aspidistreae is widely variable. The genera Tupistra and Rohdea were shown to have monosulcate and boat-shaped pollen grains. Their exine is perforate or reticulate (Plate 1: G-N). Pollen grains in the genus Aspidistra, however, are nonaperturate and spheroidal. The exine in the genus varies from crass-rugulate, variously gemmate to tuberculate-baculate (Plate 2; A-H). The pollen morphology of Aspidistra is therefore distinctly different from that of Tupistra and Rohdea, which supports the Nakai's (1936) establishment of the tribe Rohdeae for Tupistra and Rohdea. Therefore, Krause's Convallarieae is reasonably divided into at least three tribes, Convallarieae (Speirantha, Convallaria, Reineckia and Theropogon), Aspidistreae (Aspidistra) and Rohdeae (Rohdea and Tupistra). The pollen characters of all the 7 genera are shown in Table 1. The evolutionary trends of pollen morphology (aperture and exine) in the three tribes are discussed and our major view-points are shown in Fig. 1. Observations on epidermal characters of leaves show that in the Convallarieae (s. 1.) stomatal apparatuses are all anomocytic; cuticular layer on the upper epidermis is mainly striatethickened or rather uniformly thickened (Plate 2: J--P; Plate 3: A-C, F-N), whereas in the genus Convallaria the cuticular layer is squamosely thickened (Plate 2: I; Plate 3: D, E).The epidermal characters of leaves in the 7 genera are summarized in Table 2.     

Molecular and morphological phylogenetics of weevils (coleoptera,curculionoidea): do niche shifts accompany diversification?

The main goals of this study were to provide a robust phylogeny for the families of the superfamily Curculionoidea, to discover relationships and major natural groups within the family Curculionidae, and to clarify the evolution of larval habits and host-plant associations in weevils to analyze their role in weevil diversification. Phylogenetic relationships among the weevils (Curculionoidea) were inferred from analysis of nucleotide sequences of 18S ribosomal DNA (rDNA; approximately 2,000 bases) and 115 morphological characters of larval and adult stages. A worldwide sample of 100 species was compiled to maximize representation of weevil morphological and ecological diversity. All families and the main subfamilies of Curculionoidea were represented. The family Curculionidae sensu lato was represented by about 80 species in 30 "subfamilies" of traditional classifications. Phylogenetic reconstruction was accomplished by parsimony analysis of separate and combined molecular and morphological data matrices and Bayesian analysis of the molecular data; tree topology support was evaluated. Results of the combined analysis of 18S rDNA and morphological data indicate that monophyly of and relationships among each of the weevil families are well supported with the topology ((Nemonychidae, Anthribidae) (Belidae (Attelabidae (Caridae (Brentidae, Curculionidae))))). Within the clade Curculionidae sensu lato, the basal positions are occupied by mostly monocot-associated taxa with the primitive type of male genitalia followed by the Curculionidae sensu stricto, which is made up of groups with the derived type of male genitalia. High support values were found for the monophyly of some distinct curculionid groups such as Dryophthorinae (several tribes represented) and Platypodinae (Tesserocerini plus Platypodini), among others. However, the subfamilial relationships in Curculionidae are unresolved or weakly supported. The phylogeny estimate based on combined 18S rDNA and morphological data suggests that diversification in weevils was accompanied by niche shifts in host-plant associations and larval habits. Pronounced conservatism is evident in larval feeding habits, particularly in the host tissue consumed. Multiple shifts to use of angiosperms in Curculionoidea were identified, each time associated with increases in weevil diversity and subsequent shifts back to gymnosperms, particularly in the Curculionidae.     

Morphological analysis, phylogenies and classification in Velloziaceae

A morphological (non-molecular) cladistic analysis of the Velloziaceae is presented. The terminal taxa are 47 species of Velloziaceae plus four taxa as outgroups: Acanthochlamys bracteata (Acanthochlamydaceae), Encholirium scrutor (Bromeliaceae), Thoracocarpus bissectus (Cyclanthaceae) and Pandanus racemosus (Pandanaceae). The species of Velloziaceae sampled represent a significant proportion of the morphological diversity of the family, including all recognized genera and sections. The analysis revealed two major groups within Velloziaceae, supported mainly by stomata, vascular bundles in the pedicel, aquiferous tissues, filaments, anthers, pollen, stigma, seeds, ploidy and chemical characters. Comparison between this and the previous phylogenetic hypotheses for Velloziaceae, together with the two conflicting current classifications, suggests that there is incompatibility concerning the support of current genera and sections. The only three groups supported in all phylogenies are Barbacenioideae sensu Menezes (= Barbacenia sensu Smith & Ayensu), Xerophyta sect. Barbacenioides and Xerophyta sect. Xerophyta sensu Smith & Ayensu.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 157–173.     

Developing EST-SSR markers to study molecular diversity in Liriope and Ophiopogon

Twenty-two EST-SSR loci for further genetic analysis of Liriope and Ophiopogon species were developed by analyzing of 545 ESTs in Ophiopogon japonicus cDNA library and used to assess genetic diversity and population structure in 68 Liriope and Ophiopogon accessions (Liriope platyphylla, Liriope spicata, O. japonicus, and Ophiopogon jaburan). Among the 22 SSR loci, the di-nucleotide motif was the most prevalent, with a ratio of 56%, and AG/GA was the most common among di-nucleotide repeats, accounting for 42%. The polymorphic analysis showed a broad genetic background with an average genetic diversity index of 0.46. A model-based STRUCTURE analysis revealed the presence of three subpopulations, which was basically consistent with clustering based on the genetic distance. These newly developed EST-SSR markers will be useful for further studies such as taxonomy and molecular breeding studies and will further enhance our scientific understanding of the population genetic and conservation of Liriope and Ophiopogon.     

Critical notes on the classification of the Liriope muscari complex

The presence of stolons is taxonomically significant in the genus Liriope Linn. However, probably due to a lapse of attention, F. T. Wang and T. Tang failed to recognize Ophiopogon muscari Decne., type of Liriope muscari (Decne.) Bailey, a nonstoloniferous taxon, and erroneously referred it as a synonym to the stoloniferous L. spicata Lour. At the same time, however, they described another non-stoloniferous species-L. platyphylla Wang et Tang (L. muscari Bailey) as new. What, then, is the difference between L. platyphylla Wang et Tang and L. muscari (Decne.) Bailey? A comparison of the type photo of Ophiopogon muscari Decne. with specimens of L. platyphylla Wang et Tang shows that the two forms are much alike, but the latter has longer scapes usually overtopping the foliage, as well as longer spikes and broader leaves. L. H. Bailey must had combined the two forms into one species. Furthermore. there are another two non-stoloniferous forms, both having scapes shorter than their foliage, but differing from L. muscari (Decne.) Bailey in their much narrower leaves. One of them has very short leaves and looks like L. minor Makino. Finally, what are the correlations among the above four forms? By using methods of quantitative analysis, such as pictorialized scatter diagram and histogram, on herbarium material, it has been found that the L. muscari complex can be separated into two parts: 1) muscari (M), and 2) platyphylla (P). But since these two parts display a continuous variation pattern as a whole, it seems advisable to treat both as varieties of the same species, i. e., L. muscari (Decne.) Bailey var. muscari, and L. muscari var. communis (Maxim.) Hsu et L. C. Li. (Ophiopogon spicatus Ker-Cawl. var. communis Maxim.) As to the other two narrow-leaved forms, they can hardly be regarded as sufficient for two distinct infraspecific units, for they are found to be inseparablefrom each other as well as from muscari.     

A new genus of phreatoicidean isopod (Crustacea) from the north Kimberley region, Western Australia

A new genus and species of phreatoicidean isopod, Crenisopus acinifer, has been collected from a freshwater spring in the northern Kimberley region of Western Australia. Empirical cladistic analysis of 10 exemplars of phreatoicidean genera found a single cladogram. The new genus and species assumed a basal position in the Phreatoicidea, placing it within the family Amphisopodidae sensu lata. This family, however, was not monophyletic in the preliminary cladogram, suggesting that the taxonomic structure of the suborder must be revised. The cladogram provided evidence for the monophyly of the Phreatoicidae and its New Zealand clade. The analysis suggested that clades of modern phreatoicideans diverged from one another during the Mesozoic Era after they entered fresh water, but prior to the fragmentation of East Gondwana.     

Phylogeny of Podocarpaceae: comparison of evidence from morphology and 18S rDNA

Maximum parsimony analyses of the genera of Podocarpaceae were conducted using sequence data from 18S ribosomal DNA. Trees from sequence, morphological, and combined data differ in taxon arrangement, but are similar in that Podocarpus sensu lato and Dacrydium s.l. are unnatural, while Podocarpaceae (including Phyllocladus) are monophyletic. The clade Microcachrys + Microstrobos is recognized in all analyses, but its placement differs, i.e., nested among other scale-leaved taxa in the morphological analysis, but associated with Nageia and other tropical genera in the sequence analyses. Trees from combined data reflect this ambiguity. Podocarpus sensu stricto is paraphyletic according to most trees. Inferences of plesiomorphic character states within the family are largely consistent between analyses and support the view that prototypical podocarps had bifacial leaves, cones with several fertile cone scales, and large epimatia (cone scales) that covered the inverted ovules.     

Sapindales: molecular delimitation and infraordinal groups

An analysis of rbcL sequence data for representatives of families of putative sapindalean/rutalean affinity identified a robust clade of core “sapindalean” taxa that is sister to representatives of Malvales. The constitution of this clade approximates the broad concept of Sapindales (sensu Cronquist). Five lineages within the order are recognized: a “rutaceae” clade (Rutaceae, Cneoraceae, Ptaeroxylaceae, Simaroubaceae sensu stricto, and Meliaceae); a “sapindaceae” clade (Sapindaceae, Aceraceae, and Hippocastenaceae); Anacardiaceae plus Burseraceae; Kirkiaceae; and Zygophyllaceae pro parte. Relationships among these groups were only weakly resolved, but there was no support for the recognition of the two more narrowly defined orders, Rutales and Sapindales sensu stricto. Several families that have previously been allied to Sapindales or Rutales show no affinity to the core sapindalean taxa identified with the molecular data, and are excluded from the order: viz. Akaniaceae, Bretschneideraceae, Conneraceae, Coriariaceae, Melianthaceae, Meliosmaceae, Physenaceae, Rhabdodrendraceae, Sabiaceae, Staphyleaceae, Stylobasiaceae, Surianaceae, and Zygophyllaceae sensu stricto.