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Consensus tree of 10,000 shortest trees (each of 553 steps) resulting from matrix representation with parsimony (MRP) analysis of the data matrix (461 characters and 212 terminals, including the all-zero outgroup) based on 11 inter-or infrafamilial source trees (see text). To highlight relationships among the major clades of Apiales and their close relatives, the majority-rule (50%) tree is presented in abbreviated form; dashed lines (with majority rule percentages) indicate those branches not found in all shortest-length trees; the strict consensus can be derived by collapsing these dashed branches. Bracketed numbers in the clade labels, adjacent to abbreviated (triangular) clades, indicate the number of terminals represented. Taxa labelled 'woody south African clade I' include the apioids Anginon Raf., Heteromorpha Cham. & Schltdl., Dracosciadium Hilliard & B. L. Burtt, Polemannia Eckl. & Zeyh., and Glia Sond.; the 'woody south African clade II' comprises the apioids Polemanniopsis B. L. Burtt and Steganotaenia Hoscht. (see Downie & Katz-Downie, 1999); taxa in the Mackinlayeae clade includes the araliads Mackinlaya and Apiopetalum, and four hydrocotyloids (Centella, Micropleura, Actinotus and Xanthosia Rudge); the Myodocarpeae clade comprises Delarbrea plus Pseudosciadium (scored as a single terminal ) and Myodocarpus, plus the hydrocotyloid Spananthe Jacq. The Pittosporaceae clade includes Pittosporum Gaertn., Hymenosporum F. Muell., and Sollya Lindl. Relationships within the core Apiaceae clade are further detailed in Downie et al. (2001), and those in the core Araliaceae clade by Lowry et al. (2001), Wen et al. (2001) and Plunkett & Lowry (2001).
Source publication
Phylogenetic relationships involving the angiosperm order Apiales (Apiaceae and Araliaceae) are troublesome at nearly every taxonomic level and have eluded several generations of botanists. Because of difficulties in interpreting and polarizing morphological character states at deeper phylogenetic levels, most studies in Apiales have focused on rel...
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Citations
... The systematics of the family has undergone a radical transformation during the past quarter-century, resulting from extensive field work, collections-based morphological studies, and molecular phylogenetics. Major changes include a re-assessment of the relationship of Araliaceae to the rest of the eudicot order Apiales (e.g., Plunkett & al., , 2004bPlunkett, 2001;Wen & al., 2001;Chandler & Plunkett, 2004;Nicolas & Plunkett, 2009, a re-circumscription of the family (Plunkett & al., 2004a), a re-ordering of its genera (Lowry & al., 2004;Plunkett & al., 2004b, and the discovery of poly-and paraphyly in several genera (Plunkett & al., 2001. Phylogenetic studies have also yielded insights into relationships within genera such as Aralia Wen, 2000), Brassaiopsis (Mitchell & Wen, 2005), Dendropanax (Li & Wen, 2013), Hedera (Valcárcel & al., 2014), Meryta (Tronchet & al., 2005), Panax (Wen & Zimmer, 1996;Lee & Wen, 2004), Plerandra (Plunkett & Lowry, 2012), Polyscias (Eibl & al., 2001;Plunkett & al., 2001;, Pseudopanax (Perrie & Shepherd, 2009), Raukaua (Mitchell & al., 2012), and Schefflera Fiaschi & Plunkett, 2011;Li & Wen, 2014). ...
... The systematics of the family has undergone a radical transformation during the past quarter-century, resulting from extensive field work, collections-based morphological studies, and molecular phylogenetics. Major changes include a re-assessment of the relationship of Araliaceae to the rest of the eudicot order Apiales (e.g., Plunkett & al., , 2004bPlunkett, 2001;Wen & al., 2001;Chandler & Plunkett, 2004;Nicolas & Plunkett, 2009, a re-circumscription of the family (Plunkett & al., 2004a), a re-ordering of its genera (Lowry & al., 2004;Plunkett & al., 2004b, and the discovery of poly-and paraphyly in several genera (Plunkett & al., 2001. Phylogenetic studies have also yielded insights into relationships within genera such as Aralia Wen, 2000), Brassaiopsis (Mitchell & Wen, 2005), Dendropanax (Li & Wen, 2013), Hedera (Valcárcel & al., 2014), Meryta (Tronchet & al., 2005), Panax (Wen & Zimmer, 1996;Lee & Wen, 2004), Plerandra (Plunkett & Lowry, 2012), Polyscias (Eibl & al., 2001;Plunkett & al., 2001;, Pseudopanax (Perrie & Shepherd, 2009), Raukaua (Mitchell & al., 2012), and Schefflera Fiaschi & Plunkett, 2011;Li & Wen, 2014). ...
... Major changes include a re-assessment of the relationship of Araliaceae to the rest of the eudicot order Apiales (e.g., Plunkett & al., , 2004bPlunkett, 2001;Wen & al., 2001;Chandler & Plunkett, 2004;Nicolas & Plunkett, 2009, a re-circumscription of the family (Plunkett & al., 2004a), a re-ordering of its genera (Lowry & al., 2004;Plunkett & al., 2004b, and the discovery of poly-and paraphyly in several genera (Plunkett & al., 2001. Phylogenetic studies have also yielded insights into relationships within genera such as Aralia Wen, 2000), Brassaiopsis (Mitchell & Wen, 2005), Dendropanax (Li & Wen, 2013), Hedera (Valcárcel & al., 2014), Meryta (Tronchet & al., 2005), Panax (Wen & Zimmer, 1996;Lee & Wen, 2004), Plerandra (Plunkett & Lowry, 2012), Polyscias (Eibl & al., 2001;Plunkett & al., 2001;, Pseudopanax (Perrie & Shepherd, 2009), Raukaua (Mitchell & al., 2012), and Schefflera Fiaschi & Plunkett, 2011;Li & Wen, 2014). These advances have benefited from and contributed to major improvements to the taxonomy of the family, including a recent treatment for The families and genera of vascular plants series , generic revisions and synopses (e.g., Shang, 1983;Wen, 1993Wen, , 2002Wen, , 2004Wen, , 2011Mitchell & al., 1997;Jebb, 1998;Shang & al., 2000;Lowry & Plunkett, 2011;Lowry & al., 2013, floristic treatments (e.g., Allan, 1961;Bamps, 1974a,b;Philipson, 1979;Maguire & al., 1984;Smith, 1985;Lowry, 1989Lowry, , 1990; Marais, 1990;Xiang & Lowry, 2007;Cannon & Cannon, 2009;Lowry & Plunkett, 2011), regional generic treatments (Smith & Stone, 1965;Bernardi, 1969Bernardi, , 1971Bernardi, , 1980Shang, 1984;Cox, 1985;Friedmann, 1986;Borchsenius, 1997), and descriptions of new species (e.g., Wen & Lowry, 2002, 2006Fiaschi, 2004;Fiaschi & Pirani, 2005;Fiaschi & Frodin, 2006;Lowry & Shang, 2006;Fiaschi & al., 2008;Callmander & al., 2009;Callmander & Lowry, 2011;Tronchet & Lowry, 2011;Idárraga & al., 2015;Costion & Plunkett, 2016;Jiménez-Montoya & Idárraga-Piedrahíta, 2018). ...
Schefflera is the largest and most complex genus of Araliaceae, with ~600 described species (and many additional species awaiting formal description), but recent studies indicate that it is polyphyletic, forming five geographically centered clades spread across the major lineages of the family. Significant progress has been made in revising the three smallest clades, but the two largest groups, centered in Asia and the Neotropics, remain poorly understood. To advance our knowledge of these groups, a taxon set of 211 Schefflera samples and 38 samples representing other genera of Araliaceae (including all genera of the “Asian Palmate clade”) was assembled, from which sequences were obtained for six DNA spacer regions, including two nuclear (ITS, ETS) and four plastid markers (rpl32‐trnL, trnF‐rpl32, trnK‐rpl16, psbA‐trnH). Results were compared to an informal system of classification based on morphology. They confirm the monophyly of both the Asian and Neotropical groups, and their inclusion within the Asian Palmate clade, but not as sister groups. In the Neotropical clade, five major subclades can be recognized, four corresponding closely to morphological groupings (Cephalopanax, Crepinella, Didymopanax, Sciodaphyllum) and one that is geographically coherent (Gleasonia). Within Asian Schefflera, there are four major subclades, corresponding closely to currently recognized informal groups (Agalma, Heptaphyllum, Heptapleurum, Hypoleucoi). Biogeographic analyses suggest that both the Asian and Neotropical groups can trace their origins to SE Asia, where Asian Schefflera is still well represented and where Chengiopanax, the possible sister group of Neotropical Schefflera, is native. Trait‐evolution analyses of a remarkable series of convergences in habit, foliar, and reproductive characters suggest multiple independent origins (and some reversals), both within and between the Neotropical and Asian clades of Schefflera.
... Phylogenetic studies based on molecular data now suggest that these three genera form a single clade most closely related to Araliaceae, Apiaceae, Myodocarpaceae, and Pittosporaceae (Plunkett et al. 1996(Plunkett et al. , 1997Plunkett 2001;Chandler and Plunkett 2004;Nicolas and Plunkett 2009). To reflect these relationships, Plunkett et al. (2004) recircumscribed Torricelliaceae to include Aralidium and Melanophylla as well as Torricellia, and placed this family in an expanded order Apiales, together with the monogeneric Griseliniaceae. ...
Small trees or shrubs, evergreen or deciduous; plants dioecious or hermaphroditic, lacking secretory canals. Leaves spiral and simple; margins often dentate, serrate, crenate to pinnatifid, more rarely entire; leaf venation pinnate or palmate; petioles exstipulate and distinctly sheathing; blades chartaceous to subcoriaceous or somewhat succulent, glabrous or trichomes unicellular or multicellular, unbranched, glandular and sometimes also nonglandular. Inflorescence terminal (occasionally in the upper leaf axils) and paniculate with racemules or cymules as the ultimate inflorescence unit (or bifid of two racemes), erect or pendulous. Flowers minute to medium-sized, perfect or imperfect, actinomorphic, epigynous, pedicel articulated or not, bracteolate; calyx a low rim or tube with (3–)5 small teeth or subequal lobes; corolla imbricate or induplicate-valvate; petals free 5 (sometimes lacking), broad at base, plane or inflexed at apex, ovate to long-elliptical; stamens 5 (lacking or present as staminodia in carpellate flowers), alternipetalous; anthers tetrasporangiate, basi- or dorsifixed, with longitudinal dehiscence; pollen tricolporate; carpels 3, connate; ovary inferior; epigynous disc absent to flat or forming a gibbous stylopodium; styles 3, free, erect to recurving (sometimes bifid); locules 3, sometimes all but one aborting; a single functional ovule per ovary (others abortive); ovules pendant and anatropous, placentation apical or axile. Fruits drupaceous, single-seeded crowned by the persistent calyx and styles; exocarp smooth, mesocarp fleshy, endocarp sclerified or lignified. Seeds with copious endosperm (ruminate or uniform) and a small embryo.
... Malaysia). Plunkett (2001) and Lundberg (2001) both suggested that Torricelliaceae and Griseliniaceae are successive clades near the base of Apiales (Soltis et al. 2011). Clokie (2001) investigated infrageneric relationships with an array of molecular markers (rpoA, trnL-F, trnH-K and ITS) and confirmed the monophyly of the genus. ...
The Griseliniaceae is a monophyletic group of dioecious, evergreen trees, shrubs, or trailing habits that currently comprises one genus, Griselinia, and seven species. It exhibits a trans-Pacific disjunct distribution between New Zealand (2 spp.) and Chile, (5 spp.); one species in Argentina; and, one variety of a Chilean species disjunct to south-eastern Brazil. Species occupy a variety of habitats from littoral sites to montane rainforests environments, and within desert habitats (lomas) to Araucaria forests (sea-level to 3200 m). All members of the family are woody, from small trees or large shrubs reaching a height of 20 m. The recognition of the family has been supported by molecular data and its removal from the Cornaceae where it had traditional been placed; relationships within the Apiales have been confirmed. The New Zealand species, Griselinia littoralis and G. lucida, have been introduced into the horticultural trade.
... Among campanulid angiosperms, the family Torricelliaceae was proposed relatively recently, following the results of molecular phylogenetic investigations demonstrating the monophyly of a clade composed of Torricellia DC, Melanophylla Baker, and Aralidium Miq. (e.g., Plunkett 2001;Kårehed 2003;APG III 2009). Previously, these three genera had been placed in separate families and were thought to have cornalean affinities (Eyde 1988;Takhtajan 2009). ...
Recognition of Torricellia DC, Melanophylla Baker, and Aralidium Miq. as members of the same angiosperm family, Torricelliaceae, has come relatively recently, bolstered by analyses of molecular sequence data. Fruits of all three genera, endemic to eastern Asia, Madagascar, and Malesia, respectively, were compared morphologically and anatomically as a basis for evaluating systematic relationships among extant and fossil representatives. Application of X-ray tomography to fossil and extant fruits has augmented traditional approaches of physical sectioning and LM to facilitate more thorough systematic comparisons. The fruits vary from subglobose (Torricellia) to boat shaped (Melanophylla) to elongate-ellipsoidal (Aralidium) but are consistent in being tricarpellate and trilocular but with only one fertile locule. In Torricellia and Melanophylla the sterile lateral locules become larger than the central seed-bearing locule, but in Aralidium the pair of sterile locules becomes enveloped within the greatly enlarged fertile locule. In all three genera, the sterile lateral carpels each contain a prominent circular to elliptical aperture in the endocarp wall. A germination valve is located near the apex of the fertile locule in Torricellia and runs the length of the fertile locule in Melanophylla and Aralidium fruits. This work shows that the fruits of these three genera are distinctive in their morphology and anatomy, allowing for identification of fossils to the generic level, and supports the previous recognition of Torricellia from the middle Eocene of North America and from the middle Eocene to middle Miocene of Europe.
... Many phylogenetic studies based on molecular sequence data have demonstrated that the concept of Drude's (1897Drude's ( -1898 and Pimenov and Leonov's (1993) Apiaceae subfamily Hydrocotyloideae is highly artificial (Plunkett et al. 1996(Plunkett et al. , 1997Plunkett and Downie 1999;Plunkett 2001;Plunkett and Lowry 2001;Chandler and Plunkett 2004;Nicolas and Plunkett 2009). Based on these studies, four genera (Hydrocotyle L., Neosciadium Domin, Trachymene Rudge, and Uldinia J. M. Black) were moved to Araliaceae and most of the remaining genera of the now defunct Hydrocotyloideae were placed in two new subfamilies, Mackinlayoideae and Azorelloideae Nicolas and Plunkett 2009). ...
Fruit and trichome structures of Apiaceae subfamily Mackinlayoideae were studied in detail using light microscopy to identify morphological features that might be useful in delimiting this subfamily and to identify structural characters that can be used to define subclades identified by molecular phylogenetic studies (e.g. the Centella and Xanthosia clades). Three types of trichomes are present in all genera except Mackinlaya and Schoenolaena: equisetiform (Actinotus, Centella, and Chlaenosciadium), digitiform (Apiopetalum, Micropleura, and Pentapeltis), and dendritic (Xanthosia). Fruits usually have laterally compressed mericarps. Dorsal bundles and rib ducts are usually branching (or branching and anastomosing) in most taxa studied. Branching and anastomosing vittae only occur in Apiopetalum, which is also characterized by sclereids in the mesocarp. Two types of crystals were found, rhomboidal and druse. Carpophores are entirely absent from the subfamily, but homologous features are present in the form of ventral bundles. Within Mackinlayoideae, the Actinotus-piopetalum and Mackinlaya clades can be distinguished from the Centella clade (Centella, Micropleura, Pentapeltis, and Schoenolaena) and Xanthosia clade (Xanthosia and Chlaenosciadium) in having fleshy fruits, specialized trichomes, and sclereids in the fruit mesocarp. Despite its phylogenetic position, Mackinlayoideae are more similar to Araliaceae than to Azorelloideae or the rest of Apiaceae, suggesting the retention of plesiomorphic character states, such as digitiform trichomes, laterally compressed mericarps, and sclereids. The equisetiform and dendritic trichomes are synapomorphies of Centella and Xanthosia clades and the Xanthosia clade differs from the Centella clade in lacking digitiform trichomes. Overall, trichome and fruit characters provide useful structural features in defining ackinlayoideae and in differentiating its subclades.
... obs.) and Dasispermum often have ten unequal wings and show great variability within and between populations (Tilney & Van Wyk, 1995). The taxa with heteromorphic fruits are all from Africa and they are placed in a basal position in the Apioideae (Plunkett, 2001;Plunkett et al., 2004;Calviño et al., 2006). Lateral-winged mericarps are present in many genera of Azorelloideae (Asteriscium Cham. ...
Fruit structure of Apiaceae was studied in 19 species representing the 10 genera of the tribe Heteromorpheae. Our results indicate this group has a woody habit, simple leaves, heteromorphic mericarps with lateral wings. fruits with bottleshaped or bulging epidermal cells which have thickened and cutinized outer wall, regular vittae (one in furrow and two in commissure) and irregular vittae (short, dwarf, or branching and anatosmosing), and dispersed druse crystals. However, lateral winged mericarps, bottle-shaped epidermal cells, and branching and anatosmosing vittae are peculiar in the tribe Heteromorpheae of Apioideae sub family. Although many features share with other early-diverging groups of Apiaceae, including Annesorhiza clade, Saniculoideae sensu lato, Azorelloideae, Mackinlayoideae, as well as with Araliaceae. Our study shows that fruit anatomy can be used to define the tribe by molecular phylogenetic studies and support that Heteromorpheae are close to Annesorhiza clade and both are placed in the basal position of Apioideae.
... Among these deviations, high floral merism has traditionally been considered as a primitive condition for the family (Bentham, 1867;Clarke, 1879;Harms, 1898;Viguier, 1906;Li, 1942;Tikhomirov, 1961;Melchior, 1964;Takhtajan, 1966Takhtajan, , 1987Philipson, 1970;Eyde & Tseng, 1971;Grushvitzky & Skvortsova, 1973;Grushvitzky, 1981). According to another view (Cronquist, 1968(Cronquist, , 1981(Cronquist, , 1988, which is currently commonly accepted (Plunkett, 2001;Plunkett et al., 2005;Jabbour, Damerval & Nadot, 2008;Takhtajan, 2009), polymerous flowers are not primitive in Araliaceae. Rather, multiple gains of floral polymery took place in the family (Nuraliev et al., 2010). ...
Gross morphology and the development of flowers in Schefflera subintegra (Araliaceae) are examined. The floral groundplan of this species is found to be very similar to that of Tupidanthus calyptratus representing a case of most extreme floral polymery within Araliaceae. Schefflera subintegra differs from T. calyptratus with respect to a lower floral merism (19–43 versus 60–172 stamens and 15–33 versus 60–138 carpels respectively) and by transformation from polysymmetry to disymmetry of flower in the course of its development. Close relationships between S. subintegra, T. calyptratus, and Schefflera hemiepiphytica have been confirmed by phylogenetic analysis based on nuclear ribosomal internal transcribed spacer sequences. These species form a subclade within the Asian Schefflera clade, with T. calyptratus as a sister taxon to two other species. Apart from more or less pronounced floral polymery, the species of this subclade share calyx and corolla without any traits of individual sepals and petals, and also a massive calyptra. As these data suggest, the extremely polymerous flowers of Tupidanthus apparently evolved in two steps: (1) the saltational multiplication of floral elements together with a loss of individuality of sepals in the calyx and petals in the corolla and (2) further polymerization of androecium and gynoecium. Mutation(s) in CLAVATA-like gene(s) are suggested as a possible mechanism of the saltation event. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, ●●, ●●–●●.
... Despite the persistence of Drude's classification and its most recent update, that of Pimenov and Leonov (1993), this system is highly artificial, as demonstrated by numerous phylogenetic studies based on molecular sequence data, which have shown the polyphyly or paraphyly of nearly every traditional subfamily and tribe (e.g. Plunkett et al., 1996Plunkett et al., , 1997Plunkett et al., , 2004Downie and Katz-Downie, 1999;Downie et al., 2001;Plunkett, 2001;Valiejo-Roman et al., 2002;Chandler and Plunkett, 2004). Plunkett et al. (2004) transferred the genus Hydrocotyle and several close relatives to Araliaceae, and described two new subfamilies, Azorelloideae and Mackinlayoideae, into which the other hydrocotyloid genera had historically been placed. ...
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.
... Con lo studio dell"ecologia della germinazione si è cercato inoltre di capire quali sono le cause che hanno portato le popolazioni di F. arrigonii ad avere una distribuzione molto ridotta ed isolata rispetto a F. communis, che non presenta apparentemente nessun tipo di difficoltà a livello ecologico e riproduttivo in Sardegna. Mattana E., Grillo O., Venora G., Bacchetta G., 2008 -Germplasm Plunkett (2001) ...
The Ferula L. genus (Apiaceae), has the highest concentration of endemic species in Central Asia, but it presents another important distribution area in the Western Mediterranean. It is represented in Sardinia (Italy) by F. arrigonii Bocchieri, endemic species of Sardinia and Corsica and F. communis L., a species widespread in the Mediterranean basin. In this graduation thesis project, the two species have been compared by
studying seeds germination ecology and morphocolorimetric analysis. To perform the germination ecology study, seeds from three different populations of both species were incubated in Petri dishes for 50 days on 1% agar water medium and subjected to constant temperature regimes (10°C, 15°C and 20°C), alternating temperature (20/10°C) and a 12 h light/12 h dark photoperiod. The same number of seeds was subjected to a regime of 4°C in the dark (pre-chilling) for 90 days before being transferred to the mentioned above temperature conditions; treatments in 1% agar supplemented with GA3 (120 ppm) and KNO3 (0.20%) were also tried. Seeds of both species have achieved high rates of viability and germination, but F. arrigonii showed, in both cases, higher percentages than F. communis. For the morphocolorimetric analysis, the scanned images of seeds from
seven populations of F. communis and five populations of F. arrigonii were processed with the system KS-400 V. 3.0 (Carl Zeiss Vision, Oberkochen, Germany) and data obtained were statistically analyzed with the stepwise Linear Discriminant Analysis (LDA) algorithm and the cross validation procedure. This method allowed the correct classification of 92.2% of the seeds of the two species and proved the existence of phenotypic seed differences between them. This graduation thesis project has provided new data on the germination ecology and taxonomic distances of the genus Ferula in Sardinia and Corsica.
... Within the subfamily Azorelloideae, the count of n = 7 found in Hermas, is unique. Traditionally Hermas had been included in the tribe of Mulineae, but modern molecular analyses (Nicolas and Plunkett, 2009;Plunkett, 2001) showed an isolated position for the tribe on the border between Umbelliferae and Araliaceae. Morphological analyses by Liu (2004) and Liu et al. (2009), as well as molecular systematic evidence indicated that the group should be recognized as a separate subfamily, the Azorelloideae. ...
Chromosome numbers are reported for 12 species from nine genera of South African Umbelliferae, of which seven species and one genus (Itasina) are recorded for the first time. A detailed list of all published chromosome counts for southern African species is also presented, together with a review of the literature. The new data obtained are briefly discussed in the context of the taxonomy and relationships of local Umbelliferae. The counts agree with previous reports except that Annesorhiza appears to have 2n=22, with or without one additional B-chromosome, and not 2n=24 as reported in the literature. The number for Itasina (2n=24) is of considerable interest and indicates that a detailed chromosome study of the South African genera Annesorhiza and Chamarea may yield valuable taxonomic information.
