Gochnatia (Asteraceae, Mutisieae) and the Gochnatia Complex: Taxonomic Implications from Morphology

Susana E. Freire; Liliana Katinas; Gisela Sancho

The new genus Moquiniastrum, the result of recent phylogenetic analyses, is described. Although these analyses are based on cpDNA and nDNA, they also involve documentation of the distinctive morphological characters supporting this new genus. The recognition of Moquiniastrum is necessary to accurately reflect the relationships of the taxa found in the tribe Gochnatieae. Moquiniastrum includes twenty-one species that are usually gynodioecious and found mainly in Brazil but with some species elsewhere in South America. A description of Moquiniastrum, together with the corresponding new combinations, new lectotypifications of three names and one new neotypification is here provided. Key words: classification, Compositae, nomenclature, taxonomy

GOCHNATIA (ASTERACEAE, MUTISIEAE) AND THE GOCHNATIA COMPLEX: TAXONOMIC IMPLICATIONS FROM MORPHOLOGY 1 Susana E. Freire 2 , Liliana Katinas 2 , and Gisela Sancho 2 ABSTRACT Gochnatia is one of the largest genera of the tribe Mutisieae (Asteraceae) and has been traditionally characterized by its homogamous capitula with isomorphic corollas. A morphological study of Gochnatia and associated genera, i.e., Actinoseris, Chucoa, Cnicothamnus, Cyclolepis, Hyalis, Ianthopappus, Nouelia, Pleiotaxis, and Wunderlichia, was carried out to evaluate the circumscription of Gochnatia and its sections, and the affinities of this complex of genera. The characters studied involve habit, leaf features (consistency, pubescence type, shape, margin, and venation), types of capitulescence, involucre features (shape, size, phyllary series, pubescence, and shape), paleae, floret features (morphology, sex, number, color of corolla, anthers, style shapes, achenial pubescence, and pappus). Analysis of these features revealed: (1) although Gochnatia is highly variable in most of the characters studied, it can be defined by this suite of features: isomorphic to subdimorphic corollas, apiculate anther appendages, and smooth style branches; (2) sections of Gochnatia needed to be re-evaluated. As result of this, two sections, i.e., sect. Discoseris and sect. Pentaphorus, are confirmed; two sections are proposed, i.e., sect. Glomerata sect. nov. and sect. Rotundifolia sect. nov.; three sections are redefined, i.e., sect. Hedraiophyllum, sect. Leucomeris, and sect. Gochnatia, while sect. Anastraphioides is formally published; and (3) the combination of apiculate anther appendages and smooth style branches is unique to Actinoseris, Cnicothamnus, Cyclolepis, Gochnatia, Hyalis, Ianthopappus, and Nouelia within the Mutisieae. This group of genera is recognized here as the Gochnatia complex, with Gochnatia as the basal genus of this complex. Chucoa, Pleiotaxis, and Wunderlichia do not belong to the Gochnatia complex. Key words: Asteraceae, Compositae, Gochnatia, Gochnatia complex, infrageneric classification, morphology, Mutisieae. Gochnatia Kunth is one of the largest genera of the tribe Mutisieae, subtribe Mutisiinae sensu lato (including Gochnatiinae; Robinson, 1991; Bremer, 1994). It comprises 68 species, nearly all Neotropical and 2 endemic to the mountains of southeastern Asia. All the species of Gochnatia have been traditionally described as discoid with actinomorphic, deeply 5-lobed corollas, features that are plesiomorphic within Mutisieae (Bremer, 1994). The only apomorphic character suggested for the genus is the acuminate to apiculate apical anther appendage of the stamens (Bremer, 1994). This character, however, is shared by other genera of Mutisieae. The infrageneric taxonomy of Gochnatia has been much discussed (Lessing, 1832; de Candolle, 1838; Jervis, 1954; Cabrera, 1971) since the genus was established by Kunth (1818). Cabrera (1971), in his monograph of the genus, divided it into six sections: sect. Discoseris, sect. Gochnatia, sect. Hedraiophyllum, sect. Leucomeris, sect. Moquiniastrum, and sect. Pentaphorus. He suggested the artificial delimitation of some sections, such as Hedraiophyllum, and in other cases emphasized geographical distribution in distinguishing groups of species, such as in section Gochnatia. Jeffrey (1967), when relating Gochnatia to other taxa, established the Stifftia-series of genera, which all have short, rounded, glabrous style arms, and commonly glabrous corollas. The Stifftia-series was then divided into four subseries by slight differences in the shape of the style arms, although these 1 We thank Vicki Funk, Per Ola Karis, Richard Keating, Harold Robinson, Peter Stevens, and Tod Stuessy for helpful suggestions on an earlier draft of this manuscript, and one anonymous reviewer for comments. We are especially grateful to John Pruski, who enthusiastically improved the manuscript with his suggestions. We are also grateful to the curators of G, K, LP, MO, NY, S, SI, and US for the loan of specimens, and Nadia Roque for access to her prepublication manuscript on Ianthopappus (now published in Novon). Finally, we thank Victoria Hollowell because her editorial work was extremely helpful, and Vı́ctor H. Calvetti for inking our original pencil illustrations. Support for this study by the Consejo Nacional de Investigaciones Cientı́ficas y Técnicas (CONICET), Argentina, the Smithsonian Institution, Washington, D.C. (for G.S.), and the National Geographic Society (grant 5776-96) is gratefully acknowledged. 2 División Plantas Vasculares, Museo de La Plata, Paseo del Bosque, 1900 La Plata, Argentina. freire@museo. fcnym.unlp.edu.ar (author for correspondence). ANN. MISSOURI BOT. GARD. 89: 524–550. 2002. Volume 89, Number 4 2002 Freire et al. Gochnatia Morphology differences were not described for each subseries. Of these subseries, the Gochnatia-subseries included Achnopogon, Cnicothamnus, Gochnatia, Nouelia, and Oldenburgia. Cabrera (1971) considered Actinoseris, Cyclolepis, and Pleiotaxis closely related to Gochnatia by their apiculate anthers. Later, he also associated Chucoa with Gochnatia (Cabrera, 1977). Hansen (1991) considered only Actinoseris and Cyclolepis of Cabrera’s group to be close to Gochnatia, excluding Pleiotaxis and Chucoa, and added Hyalis and Nouelia to what he called the ‘‘Gochnatia-group.’’ Hansen argued the presence of cone-like involucres, i.e., with light brownish bracts imbricately arranged and resembling the cone of a spruce as a probable synapomorphy of the group. Recently, Roque and Hind (2001) created the new genus Ianthopappus for the species Actinoseris corymbosa. In this work, the authors grouped the genera Actinoseris, Chucoa, Cnicothamnus, Cyclolepis, Gochnatia, Hyalis, Ianthopappus, and Nouelia by their apiculate to acuminate apical anther appendages. Furthermore, Roque (2001) and Roque and Pirani (2001) re-circumscribed the genus Richterago Kuntze according to the genus concept first employed by Lessing (1830) to include the species of Actinoseris and the species of Gochnatia sect. Discoseris. We consider, however, that the characters used to distinguish Richterago are widespread in Gochnatia or are shared with other genera, such as herbaceous to subshrubby habit (shared among Gochnatia sect. Discoseris, Actinoseris, and other such related genera as Hyalis and Ianthopappus), leaves rosulate to alternate (alternate leaves are present in most species of Gochnatia), venation pinnate (present in most species of Gochnatia), capitula homogamous and discoid (present in most species of Gochnatia) or heterogamous and radiate (present in some species of Actinoseris), as well as pappus features, i.e., pappus uniseriate, with 25 to 42 bristles, united into a fleshy ring (present in species of Gochnatia sect. Gochnatia). For these reasons, we affirm here the traditional concept of Gochnatia and Actinoseris (excluding A. corymbosa 5 Ianthopappus corymbosus) as was established by Cabrera (1970, 1971). Some phylogenetic studies relate Gochnatia with other genera of Mutisieae; for example, Jansen and Palmer (1987) related Gochnatia with Ainsliaea, Onoseris, and Stifftia, and Karis et al. (2001) related it with Mutisia and Trixis. However, because of the few taxa sampled in the tribe, these analyses are not considered here to address relationships at the generic level. In other analyses (Karis et al., 1992; Jansen & Kim, 1996), Gochnatia was found to be an isolated taxon within Cichorioideae. The cladogram of Karis et al. (1992), based on morphological characters, shows Gochnatia as sister to most Asteraceae, excluding Barnadesioideae and four genera of Gochnatiinae. The ndhF tree of Jansen and Kim (1996) also shows Gochnatia as an independent lineage positioned basal to most Asteraceae, excluding Barnadesioideae and the core of Mutisieae examined. There are other genera within Mutisieae such as Quelchia and Neblinaea from the Guayana Highland with apiculate anther appendages, but morphological features of these genera (Pruski, 1991; Bremer, 1994) mark a departure from Gochnatia and associated taxa. On the other hand, the planaltive Brazilian genus Wunderlichia, with shortly apiculate anther appendages, was either placed in the ‘‘Stenopadus group’’ (Pruski, 1991) or it was considered isolated within Mutisieae (Hansen, 1991; Karis et al., 1992; Bremer, 1994). The group selected here for the analysis is mainly represented by the genera with apiculate anther appendages suggested by Jeffrey (in part, 1967), Cabrera (1971, 1977), Hansen (1991), and Roque and Hind (2001). It is comprised of Actinoseris sensu Cabrera, excluding A. corymbosa (7 species), Chucoa (1 species), Cnicothamnus (2 species), Cyclolepis (1 species), Gochnatia (68 species), Hyalis (2 species), Ianthopappus (1 species), Nouelia (1 species), Pleiotaxis (ca. 25 species), and the controversial Wunderlichia (6 species). There are several potential diagnostic features in Gochnatia and relatives such as the habit, leaf morphology, types of capitulescence, involucre, florets, trichomes, and pappus that have never been studied comparatively or in detail. A detailed morphological study of Gochnatia and its associated taxa is needed as a first step to provide a robust base for discussion. On the basis of morphological evidence, the goals of this study are: (1) to evaluate the circumscription of Gochnatia and its sections, and (2) to group Gochnatia with other genera based on their similarities. MATERIAL AND 525 METHODS Herbarium specimens of the studied taxa, i.e., Actinoseris, Chucoa, Cnicothamnus, Cyclolepis, Gochnatia (64 of its 68 species), Hyalis, Ianthopappus, Nouelia, Pleiotaxis, and Wunderlichia (Appendix 1), were examined to assess characters used previously to distinguish taxa in this group of genera, and to search for additional characters. The data were augmented by literature studies (Fran- 526 Annals of the Missouri Botanical Garden chet, 1888; Cabrera, 1950, 1951, 1955, 1970, 1971; Jeffrey, 1967; Barroso & Maguire, 1973; Zardini, 1975; Sancho, 1997, 2000; Roque, 1997; Roque & Hind, 2001; Roque & Pirani, 1997, 2001). Vegetative and floral parts were dissected and observed after boiling in water and stained with 2% safranin. Freehand sections of leaves were performed and these sections examined to determine the presence of hypodermis, and then stained with safranin. Drawings were made by the authors using a stereomicroscope Wild M5 and a microscope Leitz SM Lux with the camera lucida technique. Apical anther appendages were described and drawn to include the portion ranging from the thecae apex to the apex of the stamen. Descriptive terminology for the trichomes follows Ramayya (1962). Whenever possible, additional synonyms of trichome terminology were added (Payne, 1978; Font Quer, 1979; Moreno, 1984; Harris & Harris, 1994; Metcalfe & Chalk, 1950). Some authors use the term ‘‘trichome,’’ whereas others apply the term ‘‘hair’’ in their pubescence classifications. Both terms are cited here following each author classification. G. palosanto, G. purpusii, G. rotundifolia, G. vernonioides), Actinoseris, Cnicothamnus, Cyclolepis, Hyalis, Ianthopappus, and in some species of Pleiotaxis (e.g., P. eximia, P. huillensis). Occasionally, the hypodermis is discontinuous (e.g., Gochnatia argentina, G. discolor, G. orbiculata, G. ramboi). Leaf shape: The leaves are always simple in the taxa under study, but show great variation in shape, from linear to suborbicular. Most species of Gochnatia sect. Gochnatia and Nouelia have leaves ovate to ovate-elliptic, obtuse to subobtuse at the apex; occasionally ovate leaves are acute at the apex (e.g., G. polymorpha, G. vernonioides), or cordate at the base (e.g., G. cordata, G. haumaniana) (Fig. 1A). Most of the Caribbean species of Gochnatia sect. Gochnatia, Actinoseris (A. hatschbachii, A. polymorpha, A. radiata), Chucoa, Ianthopappus, and Wunderlichia have obovate leaves (Fig. 1B). Elliptic leaves (Fig. 1C) are rounded in the base and apex in G. discoidea, attenuate at the base and apex in G. magna, G. sect. Leucomeris, Actinoseris (A. polyphylla), Cnicothamnus, and some species of Pleiotaxis (e.g., P. huillensis, P. newtonii, P. selina, P. subscaposa). Gochnatia sect. Pentaphorus, G. argyrea, Actinoseris (A. angustifolia, A. stenophylla), Cyclolepis, Hyalis, and most species of Pleiotaxis (e.g., P. ambigua, P. decipiens, P. dewevrei, P. linearifolia, P. rogersii) have linear leaves (Fig. 1D). Leaf margin: Only the Caribbean species of Gochnatia sect. Gochnatia (except G. crassifolia and G. picardae) and Chucoa have spinose margins (Fig. 1E). In the remaining taxa the leaf margin is predominantly entire or denticulate (Fig. 1F). Leaf venation: The venation is predominantly pinnate, although in taxa such as Gochnatia arequipensis, G. glutinosa, G. rotundifolia, Hyalis, and Ianthopappus the leaves are three-veined (actinodromous). RESULTS For comparison with our results, sections of Gochnatia with their corresponding species according to Cabrera (1971) are listed in Table 1. VEGETATIVE CHARACTERS Habit. Taxa under study display four types of habit: small trees, shrubs, subshrubs, and perennial herbs. Several species of Gochnatia (e.g., G. arborescens, G. decora, G. elliptica, G. ilicifolia, G. magna, G. oligocephala, G. palosanto, G. polymorpha, G. spectabilis, G. tortuensis), Cnicothamnus, and Wunderlichia are small trees. Most species of Gochnatia, Chucoa, Cyclolepis, and Nouelia are shrubs. Gochnatia sect. Discoseris, Hyalis, and Ianthopappus are subshrubs. Actinoseris and Pleiotaxis are perennial herbs. Cyclolepis is the only spinose genus and is aphyllous at maturity. Leaves. All taxa analyzed have alternate, occasionally rosulate, subsessile to shortly petiolate leaves. Leaf consistency: The leaves are coriaceous or subcoriaceous, but only some species have leaves with an adaxial hypodermis. It is absent in some species of Gochnatia (e.g., G. amplexifolia, G. arequipensis, G. cardenasii, G. discoidea, G. foliolosa, G. glutinosa, G. hypoleuca, G. intertexta, G. magna, CAPITULESCENCE Capitula are borne singly or 2 or 3 at the end of the branches or may be clustered in pseudocorymbs, pseudoracemes, or pseudopanicles, in open to condensed or glomerulose synflorescences depending upon the length of the peduncles. Solitary or few capitula (2 or 3) are short-pedunculate to glomerulose in Gochnatia sect. Gochnatia, Cnicothamnus, Nouelia, and some species of Wunderlichia (W. azulensis, W. crulsiana, W. mirabilis) (Fig. 2A). They are scapiform, long-pedunculate in Gochnatia suffrutescens, some species of Actinoseris (e.g., A. angustifolia, A. hatschbachii, A. polymorpha, A. radiata), Chucoa, and some species of Pleiotaxis (e.g., P. subscaposa) (Fig. 2B). Sections sensu Cabrera (1971) Sections according to this study Gochnatia sect. Discoseris G. amplexifolia (Gardner) Cabrera *G. discoidea (Less.) Cabrera G. suffrutescens Cabrera Gochnatia sect. Discoseris G. amplexifolia G. discoidea G. suffrutescens Gochnatia sect. Gochnatia South American species: G. arequipensis Sandw. G. boliviana S. F. Blake G. cardenasii S. F. Blake G. curviflora (Griseb.) O. Hoffm. G. patazina Cabrera G. vargasii Cabrera *G. vernonioides Kunth Gochnatia sect. Gochnatia arequipensis boliviana cardenasii curviflora patazina vargasii vernonioides Gochnatia sect. Rotundifolia *G. rotundifolia Subshrubs; capitula solitary, long-pedunculate, or numerous in scapose pseudocorymbs; appendage abruptly apiculate, laciniate tails; pappus type A Peru Bolivia Bolivia S Bolivia, NW Argentina Peru Peru N Peru Leaves pinnately (subthree-) veined; solitary capitula (2 or 3); anther appendage caudate (abruptly apicualte), laciniate tails; pappus type B S Brazil Leaves three-veined, glabrous; solitary capitula; involucre with subglabrous phyllaries, ciliolate at the margins; corollas white; anther appendage attenuate, laciniate tails; pappus type B Cuba Caribbean Cuba Cuba Cuba Leaves with spinose (entire) margin; solitary capitula (2 or 3); corollas deeply to very deeply 5lobed; anther appendage caudate (abruptly apiculate), smooth tails (laciniate); capitate glandular hairs in achenes; pappus type A (type B) Cuba Cuba Cuba Dominican Republic Cuba Caribbean 527 Caribbean species: Gochnatia sect. Anastraphioides G. attenuata (Britton) Jervis & Alain G. attenuata G. buchii (Urb.) Jiménez G. buchii G. calcicola (Britton) Jervis & Alain G. calcicola G. cowellii (Britton) Jervis & Alain G. cowellii G. crassifolia (Britton) Jervis & G. crassifolia Alain G. cubensis (Carabia) Jervis & Alain G. cubensis G. ekmanii (Urb.) Jervis & Alain G. ekmanii G. elliptica (León) Alain G. elliptica G. enneantha (S. F. Blake) Alain G. enneantha G. gomezii (León) Jervis & Alain G. gomezii G. ilicifolia Less. *G. ilicifolia Brazil Brazil Brazil Diagnostic characters Freire et al. Gochnatia Morphology G. rotundifolia Less. G. G. G. G. G. G. G. Distribution Volume 89, Number 4 2002 Table 1. Sections within Gochnatia established in this study on the basis of the morphology, compared with the sections delimited by Cabrera (1971), with the addition of G. hatschbachii (Cabrera, 1974) and G. densicephala (Sancho, 2000). The geographic distribution and diagnostic characters for each section are provided, with uncommon features appearing in parentheses. * 5 type species of each section. 528 Table 1. Continued. Sections sensu Cabrera (1971) Gochnatia sect. Hedraiophyllum G. arborescens T. S. Brandegee *G. cordata Less. G. magna Cabrera G. purpusii T. S. Brandegee G. hypoleuca (DC.) A. Gray G. palosanto Cabrera G. smithii Robinson & Greenm. Distribution G. intertexta G. maisiana G. mantuensis Cuba Cuba Cuba G. microcephala Cuba G. montana G. obtusifolia Cuba Cuba G. G. G. G. G. G. G. G. G. Caribbean Cuba Bahamas Haiti Cuba Cuba Cuba Haiti Cuba oligantha parvifolia pauciflosculosa picardae recurva sagraeana shaferi tortuensis wilsonii Gochnatia sect. Glomerata *G. arborescens (moved to another section) G. magna G. purpusii Gochnatia sect. Leucomeris G. hypoleuca G. palosanto G. smithii Mexico Mexico Mexico S U.S.A., Mexico N Argentina, S Bolivia Mexico Diagnostic characters Capitula few together in glomerulose pseudocorymbs; florets 12 to 20(50); anther appendage abruptly apiculate (attenuate), smooth tails; pappus type B (type A) Numerous capitula in glomerulose pseudocorymbs, pseudoracemes or pseudopanicles; involucre with glabrous, ciliolate at the margin phyllaries extending into the peduncle; florets 4 to 6(8 to 12), white (yellow); anther appendage attenuate (abruptly apiculate), laciniate tails (smooth); pappus type B Annals of the Missouri Botanical Garden G. intertexta (Griseb.) Jervis & Alain G. maisiana (León) Jervis & Alain G. mantuensis (Griseb.) Jervis & Alain G. microcephala (Griseb.) Jervis & Alain G. montana (Britton) Jervis & Alain G. obtusifolia (Britton) Jervis & Alain G. oligantha (Urb.) Howard G. parvifolia (Britton) Jervis & Alain G. pauciflosculosa (Hitchc.) Cabrera G. picardae (Urb.) Jiménez G. recurva (Britton) Jervis & Alain G. sagraeana Jervis & Alain G. shaferi (Britton) Jervis & Alain G. tortuensis (Urb.) Jiménez G. wilsonii (Britton) Jervis & Alain Sections according to this study Continued. Sections sensu Cabrera (1971) Gochnatia sect. Leucomeris G. decora (Kurz) Cabrera *G. spectabilis (D. Don) Less Gochnatia sect. Moquiniastrum G. argentina (Cabrera) Cabrera G. argyrea (Malme) Cabrera G. barrosii Cabrera G. blanchetiana (DC.) Cabrera G. discolor Baker G. floribunda Cabrera G. gardneri (Baker) Cabrera haumaniana Cabrera mollissima (Malme) Cabrera oligocephala (Gardner) Cabrera orbiculata (Malme) Cabrera paniculata (Less.) Cabrera polymorpha (Less.) Cabrera pulchra Cabrera ramboi Cabrera rusbyana Cabrera sordida (Less.) Cabrera velutina (Bong.) Cabrera Gochnatia sect. Pentaphorus *G. foliolosa (D. Don) Hook. & Arn. G. glutinosa (D. Don) Hook & Arn. G. decora G. spectabilis Distribution Diagnostic characters Asia Asia Gochnatia sect. Hedraiophyllum G. argentina G. argyrea G. barrosii G. blanchetiana *G. cordata Less. G. densicephala (Cabrera) G. Sancho G. discolor G. floribunda G. gardneri G. hatschbachii Cabrera G. haumaniana G. mollissima G. oligocephala G. orbiculata G. paniculata G. polymorpha G. pulchra G. ramboi G. rusbyana G. sordida G. velutina N Argentina S Brazil S Brazil, Paraguay E Brazil Paraguay, Brazil, Uruguay, Argentina E Brazil Brazil Brazil Brazil Brazil Paraguay, Brazil S Brazil E Brazil E Brazil SE Brazil Brazil, Paraguay, Uruguay, Argentina S Brazil, Paraguay S Brazil Andes of Peru, Bolivia S Brazil S Brazil Gochnatia sect. Pentaphorus G. foliolosa G. glutinosa Central Chile W Argentina Trees or shrubs, gynodioecious, polygamous (monoecious); leaves with 2-, 3- to 5-armed (flagellate) hairs; numerous capitula in glomerulose to loose pseudopanicles; capitula heterogamous (homogamous), disciform (discoid); corollas subdimorphic (isomorphic); anther appendage abruptly apiculate, laciniate, smooth tails; pappus type C Freire et al. Gochnatia Morphology G. G. G. G. G. G. G. G. G. G. G. Sections according to this study Volume 89, Number 4 2002 Table 1. Linear leaves, pinnately, three-veined, glandulate; numerous capitula in glomerulose, leafy pseudoracemes; florets 5 (3, 7, or 10 to 20), white or lilac; anther appendage abruptly apiculate, smooth, laciniate tails; pappus type D 529 530 Annals of the Missouri Botanical Garden Figure 1. Leaf shape. —A. Ovate (left to right): Gochnatia rotundifolia (Handro 157, LP), G. ramboi (Rambo 50005, S), G. vernonioides (López & Sagástegui 3354, LP), G. cordata (Serrano 6, LP), Nouelia insignis (l’Abbé Delavay 2498, US). —B. Obovate (left to right): Wunderlichia crulsiana (Ratter et al. 2615, MO), Gochnatia intertexta (Alain A-1680, NY), G. pauciflosculosa (Wilson 7428, K), G. crassifolia (Ekman 4023, S), Ianthopappus corymbosus (Palacios & Cuezzo 2304, LP), Actinoseris radiata (Hatschbach 690, LP), Chucoa ilicifolia (López Miranda 1090, LP). —C. Elliptic (left to right): Gochnatia discoidea (Blanchet 3345, LP), G. magna (Cronquist 11277, NY), G. spectabilis (Galrola 32, LP), Volume 89, Number 4 2002 Freire et al. Gochnatia Morphology Few to several capitula (more than 4) arranged in short-pedunculate to glomerulose pseudocorymbs are found in Gochnatia arborescens, G. magna, G. purpusii, Gochnatia sect. Leucomeris, Hyalis lancifolia, and Wunderlichia (W. bahiensis, W. insignis) (Fig. 2C). Numerous capitula arranged in scapiform, long-pedunculate pseudocorymbs are found in Gochnatia sect. Discoseris (except G. suffrutescens), Actinoseris (A. stenophylla), Hyalis argentea, Ianthopappus, and Wunderlichia senaeii (Fig. 2D). Gochnatia sect. Pentaphorus and Cyclolepis have numerous capitula borne in short-pedunculate to glomerulose clusters, that are in turn arranged in leafy pseudoracemes (Fig. 2E). Pseudoracemes in compact or glomerulose clusters, which lack leaves, at the tip of the branches, are characteristic of Gochnatia palosanto (Fig. 2F). Loose pseudoracemes are present only in some species of Pleiotaxis (e.g., P. angusterugosa, P. gombensis, P. oxylepis, P. racemosa) (Fig. 2G). Numerous capitula arranged in glomerulose pseudopanicles are present in most species of Gochnatia sect. Moquiniastrum, in G. cordata, G. hypoleuca, and G. smithii (Fig. 2H). Short-pedunculate (Fig. 2I) or loose pseudopanicles (Fig. 2J) are present in species of Gochnatia sect. Moquiniastrum (G. argentina, G. argyrea, respectively). (A) The smallest involucres [(2–)4–7(–8) mm] are present in Gochnatia sect. Pentaphorus, Gochnatia sect. Moquiniastrum (except G. argyrea with 9–10 mm), G. hypoleuca, G. microcephala, G. palosanto, G. smithii, Cyclolepis, and Hyalis. (B) Intermediate involucres (10–18 mm high) are present in some species of Gochnatia (e.g., G. arborescens, G. cordata, G. discoidea, G. ilicifolia, G. magna, G. patazina, G. purpusii, G. recurva, G. rotundifolia, G. vernonioides), Actinoseris, Chucoa, and Ianthopappus. (C) The biggest involucres (20–45 mm) are displayed by a few Caribbean species of Gochnatia sect. Gochnatia (e.g., G. cowellii, G. cubensis, G. ekmanii, G. picardae, G. sagraeana), Cnicothamnus, Nouelia, Pleiotaxis, and Wunderlichia. INVOLUCRE Shape. The involucre shape is either oblong to campanulate or turbinate. Oblong to campanulate involucres are characteristic of most species of Gochnatia (Fig. 3A), Chucoa (Fig. 3B), Cnicothamnus (Fig. 3C), Cyclolepis (Fig. 3D), Hyalis (Fig. 3E), Nouelia (Fig. 3F), Pleiotaxis (Fig. 3G), and Wunderlichia (Fig. 3H). Turbinate involucres, on the other hand, occur in a few Caribbean species of Gochnatia sect. Gochnatia (e.g., G. cubensis, G. intertexta, G. pauciflosculosa), in some species of G. sect. Hedraiophyllum (G. hypoleuca, G. palosanto, G. smithii) (Fig. 3I), G. suffrutescens, Actinoseris, and Ianthopappus (Fig. 3J). Size. The involucre size ranges from 2 to 45 mm high, with three main categories: (2–)4–7(–8) mm, 10–18 mm, and 20–45 mm high. 531 Series of phyllaries. The phyllaries are arranged in several imbricate series. The highest number is (6)7 to 10 series, and this is found in a few Caribbean species of Gochnatia sect. Gochnatia (e.g., G. cowellii, G. cubensis, G. picardae, G. recurva), G. arborescens, Cnicothamnus (Fig. 3C), Nouelia (Fig. 3F), Pleiotaxis (Fig. 3G), and Wunderlichia (Fig. 3H). Three- to six-seriate involucres occur in most species of Gochnatia (Fig. 3A), and Actinoseris, Chucoa (Fig. 3B), Cyclolepis (Fig. 3D), Hyalis (Fig. 3E), and Ianthopappus (Fig. 3J). Gochnatia sect. Leucomeris and three species of G. sect. Hedraiophyllum (G. hypoleuca, G. palosanto, G. smithii) show a peculiar feature in their involucres. The capitula are 3- to 4-seriate, but the peduncle bracts extend into the involucre giving a 7- to 12-seriate condition (Fig. 3I). Phyllary pubescence and shape. In almost all the taxa studied the phyllaries are dorsally pubescent or subglabrous. However, Gochnatia sect. Leucomeris, G. hypoleuca, G. palosanto, G. smithii, and G. rotundifolia have dorsally glabrous phyllaries with ciliate margins. Most genera have linear to oblong or obovate phyllaries with entire margins. Cnicothamnus has phyllaries with an apical appendage rounded or rhombic, with lacerate margins (Fig. 3C), and Wunderlichia has phyllaries with scarious, colored, and occasionally fimbriate margins (Fig. 3H). ← Actinoseris polyphylla (Hatschbach 35304, LP), Cnicothamnus lorentzii (Padaci 84, LP), Pleiotaxis huillensis (Gossweiler 10780, US). —D. Linear (left to right): Gochnatia foliolosa (Cabrera 3451, LP), G. argyrea (Hatschbach 9578, LP), Hyalis lancifolia (Schinini 16098, LP), Actinoseris angustifolia (Hatschbach 29986, LP), Pleiotaxis rogersii (Rolyns 1568, US), P. dewevrei (de Hitte 288, US), Cyclolepis genistoides (Rojas 7104, LP). Leaf margin. —E. Spinose (left to right): Chucoa ilicifolia (López Miranda 1090, LP), Gochnatia montana (Ekman 18725, S), G. enneantha (Ekman H-15498, S), G. cubensis (Ekman 9632, S), G. intertexta (Alain A-1680, NY). —F. Denticulate to entire (left to right): G. polymorpha (Pereira 8609 & Pabst 7984, LP), G. rotundifolia (Handro 157, LP), G. foliolosa (Cabrera 3451, LP). 532 Annals of the Missouri Botanical Garden Figure 2. Types of capitulescence. —A. Solitary or few capitula short-pedunculate. —B. Solitary or few capitula longpedunculate and scapiform. —C. Pseudocorymbs short-pedunculate. —D. Pseudocorymbs long-pedunculate and scapiform. —E. Pseudoracemes glomerulose and leafy. —F. Pseudoracemes apically glomerulose. —G. Loose pseudoracemes. —H. Pseudopanicles glomerulose. —I. Pseudopanicles short-pedunculate. —J. Loose pseudopanicles. PALEAE Wunderlichia is the only genus in the group with receptacular paleae. For this reason, Pruski (1991) included it in the ‘‘Stenopadus group.’’ Cabrera (1971) described Gochnatia as typically epaleate, and only exceptionally with some paleae, but mentioned no species. However, we Volume 89, Number 4 2002 Freire et al. Gochnatia Morphology 533 Figure 3. Involucre. A–H. Oblong to campanulate. —A. Gochnatia floribunda (Hatschbach 43151, NY). —B. Chucoa ilicifolia (López Miranda 1090, LP). —C. Cnicothamnus lorentzii (Padaci 84, LP). —D. Cyclolepis genistoides (Tinto 2038, LP). —E. Hyalis lancifolia (Schinini 16098, LP). —F. Nouelia insignis (Maire 2516, NY). —G. Pleiotaxis rogersii (Rolyns 1568, US). —H. Wunderlichia crulsiana (Ratter et al. 2615, MO). I, J. Turbinate. —I. Gochnatia palosanto (Schreiter in 1925, LP). —J. Ianthopappus corymbosus (Palacios & Cuezzo 2304, LP). found no evidence of paleae in any species of Gochnatia examined. FLORETS Corolla morphology and sex arrangement. Corollas in Gochnatia and allies may be isomorphic, subdimorphic, or clearly dimorphic. All florets of the capitula may be hermaphroditic or female (homogamous capitula), or the capitula may have central hermaphroditic florets and marginal female florets (heterogamous capitula). These features are combined as follows: (A) Corollas isomorphic. Plants with discoid and homogamous capitula made up of hermaphroditic florets with isomorphic tubular corollas are char- 534 Annals of the Missouri Botanical Garden acteristic of most species of Gochnatia (Fig. 4A, B), Chucoa (Fig. 4C), Pleiotaxis (Fig. 4D), and Wunderlichia (Fig. 4E). Only G. argyrea of section Moquiniastrum has isomorphic tubulose corollas and heterogamous capitula (Sancho, 2000). Very occasionally marginal female florets (with staminodes) are found in Chucoa (Cabrera, 1955, and our own observations). (B) Corollas subdimorphic. Such corollas are present only in Gochnatia sect. Moquiniastrum (except G. argyrea and G. gardneri) (Fig. 4F) and Cyclolepis (Fig. 4G). Gochnatia sect. Moquiniastrum includes monoecious, polygamous, and gynodioecious plants (Sancho, 2000). Polygamous and gynodioecious species have disciform capitula made up of subdimorphic corollas, i.e., central hermaphroditic florets with tubular actinomorphic corollas and coiled lobes (Fig. 4F1), and outer, female florets with tubular subzygomorphic corollas and erect or coiled lobes of different lengths (Fig. 4F2). In the gynodioecious genus Cyclolepis the female florets have tubular-filiform corollas (Fig. 4G1), and the hermaphroditic florets have tubular corollas (Fig. 4G2). (C) Corollas dimorphic. Plants with radiate, homogamous capitula comprising dimorphic, hermaphroditic florets, with outer bilabiate or subbilabiate and central tubular corollas, are present in some species of Actinoseris (e.g., A. polyphylla, A. revoluta, A. stenophylla) (Fig. 4H), Cnicothamnus (Fig. 4I), Hyalis (Fig. 4J), and Nouelia (Fig. 4L). Plants with heterogamous capitula and the type of dimorphic corollas previously mentioned are found in Ianthopappus (Fig. 4K) and some species of Actinoseris (e.g., A. arenaria, A. hatschbachii, A. radiata); the outer florets are female and the central florets hermaphroditic. These truly dimorphic corollas are absent in Gochnatia. It may be noted that tubular, deeply lobed corollas with coiled lobes (reaching up to 2/3 of the corolla) are present in most species of Gochnatia (Fig. 4A), and in Actinoseris, Chucoa (Fig. 4C), Cnicothamnus, Cyclolepis, Hyalis, Ianthopappus, Nouelia, and Pleiotaxis (Fig. 4D). Tubular, very deeply lobed corollas with straight lobes (more than ⅔ the length of the corolla, almost reaching the base) are present in most Caribbean species of Gochnatia sect. Gochnatia (e.g., G. calcicola, G. cubensis, G. gomezii, G. oligantha) (Fig. 4B). Bilabiate corollas have an external 3-dentate lip and an internal 2cleft lip (312 arrangement, e.g., Actinoseris revoluta, Hyalis, Ianthopappus, Nouelia) (Fig. 4J–L), and subbilabiate corollas have an external 3-dentate and one entire, internal lip (311, e.g., Acti- noseris hatschbachii, A. polyphylla, Cnicothamnus) (Fig. 4H, I). Number of florets per capitulum. The number of florets per capitulum ranges from 4 to 300. A few species of Gochnatia (G. crassifolia, G. magna, G. recurva, G. rotundifolia, G. sagraeana) and most species of Actinoseris, Cnicothamnus, and Wunderlichia have 50 to 300 florets per capitulum. Some species of Gochnatia (e.g., G. amplexifolia, G. boliviana, G. cordata, G. cowellii, G. patazina, G. vernonioides), some species of Actinoseris (e.g., A. hatschbachii, A. revoluta), Ianthopappus, and Nouelia have capitula with 30 to 50 florets. Some species of Gochnatia (e.g., G. arequipensis, G. cardenasii, G. ilicifolia, G. purpusii), G. sect. Moquiniastrum, Chucoa, and Cyclolepis have 7 to 30 florets per capitulum, whereas some species of Gochnatia (e.g., G. hypoleuca, G. smithii), G. sects. Leucomeris and Pentaphorus, a few Caribbean species of G. sect. Gochnatia (e.g., G. calcicola, G. cubensis, G. gomezii, G. oligantha), and Hyalis have capitula with 4 to 6 florets. Corolla color. The corollas in Gochnatia are predominantly yellow or cream. In some species (e.g., G. rotundifolia, G. decora, G. spectabilis, G. foliolosa) they are white and/or lilac. In others (e.g., G. cowellii, G. ilicifolia, G. intertexta, G. mantuensis, G. sagraeana) they are orange. The corollas are white, pink, or purple in Actinoseris; in Cnicothamnus they are orange; in Chucoa and Cyclolepis, yellow; in Hyalis lilac; in Nouelia white; in Pleiotaxis deep red, pink, white, or cream; in Wunderlichia white or yellow. Ianthopappus shows white marginal corollas and purple disc corollas. ANTHERS As well as most species of the tribe Mutisieae, all the taxa in this study have an apical anther appendage and basal tails. The apical appendage can be viewed as an adaptation to protect the pollen in the anther tube from moisture and insect predators until the stigma and style push upward through it for pollen presentation (Stuessy et al., 1996). With the exception of Chucoa, with acute anther appendages (Fig. 5A), anther appendages in Gochnatia and related taxa have usually been described as apiculate. Pleiotaxis differs in its thickened and bulbous apical appendage (Fig. 5B). Analysis of this feature in all the genera under study revealed further variation in its shape that is particularly useful in grouping species within Gochnatia. The appendage can be short (anthers Volume 89, Number 4 2002 Freire et al. Gochnatia Morphology 535 Figure 4. A–E. Isomorphic corolla. —A. Gochnatia magna (Cronquist 11277, NY). —B. G. pauciflosculosa (Eggers 3866, K). —C. Chucoa ilicifolia (López Miranda 1090, LP). —D. Pleiotaxis dewevrei (de Hitte 288, US). —E. Wunderlichia mirabilis (Martinelli & Stutts 999, NY). F, G. Subdimorphic corolla. —F. Gochnatia polymorpha (Hashimoto 624, LP): F1, disc corolla, F2, marginal corolla. —G. Cyclolepis genistoides (Cabrera 3782, LP): G1, marginal corolla, G2, disc corolla. H–L. Dimorphic corolla. —H. Actinoseris polyphylla (Hatschbach 35304, LP): H1, disc corolla, H2, marginal corolla. —I. Cnicothamnus lorentzii (Cabrera et al. 22576, LP): I1, marginal corolla, I2, disc corolla. —J. Hyalis lancifolia (Schinini 16098, LP): J1, marginal corolla, J2, disc corolla. —K. Ianthopappus corymbosus (Palacios & Cuezzo 2304, LP): K1, marginal corolla, K2, disc corolla. —L. Nouelia insignis (Maire 2516, NY): L1, marginal corolla, L2, disc corolla. 536 Annals of the Missouri Botanical Garden Figure 5. Apical anther appendage. —A. Not apiculate: Chucoa ilicifolia (López Miranda 1090, LP). —B. Apiculate bulbous: Pleiotaxis eximia (Rolyns 1836, US). C–F. Caudate anther appendage. —C. Gochnatia sect. Gochnatia sensu Cabrera (from left to right): Gochnatia cardenasii (Cordo & Ferrer 88-B-17, SI), G. cowellii (Britton & Cowell 10183, NY), G. curviflora (Jeréz et al. 49120, LP), G. ilicifolia (Small & Carter 8526, K), G. montana (Ekman 18725, S), G. parvifolia (Shafer 2938, NY), G. patazina (López & Sagástegui 3409, LP), G. recurva (León 20946, LP). —D. Actinoseris Volume 89, Number 4 2002 Freire et al. Gochnatia Morphology apiculate) to very long (anthers apically caudate). In addition, it can abruptly terminate in a sharp point, or gradually taper above into an attenuate point. The following three combinations were observed: (1) Apically caudate (Fig. 5C–E): in most species of Gochnatia sect. Gochnatia (except G. boliviana, G. buchii, G. microcephala, G. tortuensis, and G. vernonioides with abruptly apiculate apices) (Fig. 5C), Actinoseris (Fig. 5D), and Cnicothamnus (Fig. 5E). (2) Attenuate apiculate (Fig. 5G): in Gochnatia sect. Leucomeris, G. hypoleuca, G. palosanto, G. magna, and G. rotundifolia. (3) Abruptly apiculate (Fig. 5J–L): in Gochnatia sect. Discoseris, sect. Moquiniastrum (short appendage in G. argentina, G. densicephala, and G. floribunda), and sect. Pentaphorus, Gochnatia arborescens, G. cordata, G. purpusii (Fig. 5J), Nouelia (Fig. 5K), and Wunderlichia (very short) (Fig. 5L). Although useful in distinguishing taxa, some overlap exists between these broad categories such as between apically caudate-attenuate (e.g., Ianthopappus, Fig. 5F), attenuate-abruptly apiculate (e.g., Cyclolepis, Fig. 5H; Hyalis, Fig. 5I), and abruptly apiculate-caudate (e.g., G. velutina, Fig. 5J). Anther tails are free and can be smooth or laciniate. They are smooth in the Caribbean species of Gochnatia sect. Gochnatia (except G. attenuata, G. ilicifolia, and G. microcephala), some species of Gochnatia sect. Moquiniastrum (e.g., G. barrosii, G. densicephala, G. floribunda, G. paniculata), some species of Gochnatia sect. Hedraiophyllum (e.g., G. arborescens, G. magna, G. purpusii), Chucoa, Cnicothamnus, and Wunderlichia. The tails are laciniate, at least in one side, in most species of Gochnatia (e.g., G. sect. Discoseris, G. sect. Leucomeris, the South American species of G. sect. Gochnatia, and G. glutinosa), and in Actinoseris, Cyclolepis, Hyalis, Ianthopappus, Nouelia, and Pleiotaxis. STYLE 537 Most taxa have smooth styles except for Wunderlichia (Fig. 6A) and Chucoa (Fig. 6B) with dorsally papillose styles, i.e., with the two branches (rounded and acute at the apex, respectively) covered by short sweeping hairs, and Pleiotaxis (Fig. 6C) with a subapical crown of short hairs. All species of Gochnatia (Fig. 6D, E), Actinoseris (Fig. 6F), Cnicothamnus (Fig. 6G), Cyclolepis (Fig. 6H), Hyalis (Fig. 6I), Ianthopappus (Fig. 6J), and Nouelia (Fig. 6K) have smooth, apically rounded styles. Most have the inner surface of the branches covered by stigmatic papillae prolonged into the outer surface of the style constituting a ridge of cells, which is less evident in Cnicothamnus. TRICHOMES Leaf pubescence. Excluding some species such as Gochnatia rotundifolia, Actinoseris hatschbachii, A. stenophylla, and Hyalis lancifolia, which have glabrous leaves, at least when mature, there are five different types of trichomes in Gochnatia and its relatives. (1) Oblique septate flagellate hairs: one or two foot cells, one- or more-celled stalks or stipes, and unicellular, very long, flagellate, tubular heads (Fig. 7A). This trichome type is present in most species of Gochnatia, Chucoa, Cnicothamnus, Pleiotaxis, and Wunderlichia. (2) Two-armed hairs (or T-shaped, malpighiaceous, anvil, dolabriform hairs): one or two foot cells, uniseriate, generally two- to more-celled stalks, and unicellular heads. The apical cell, which constitutes the head, initially assumes the shape of a hammer and later becomes T-shaped by further outgrowth of the two ends (Fig. 7B, C). This trichome type is present in Gochnatia sect. Moquiniastrum (Fig. 7B) and G. cordata, Cyclolepis (Fig. ← radiata (Hatschbach 690, LP). —E. Cnicothamnus lorentzii (Ruiz Leal 14162, LP). —F. Ianthopappus corymbosus (Palacios & Cuezzo 2304, LP). —G–I. Attenuate-apiculate anther appendage. G1, Gochnatia sect. Leucomeris sensu Cabrera (from left to right): Gochnatia decora (Maung Mya 5309, LP), G. spectabilis (Galrola 32, LP). G2, G. hypoleuca (González Quintero 3215, LP), G. palosanto (Schreiter in 1925, LP), G. magna (Fernández 3666, NY), G. rotundifolia (Joly 596, LP). —H. Cyclolepis genistoides (Correa 3172 & Nicora, LP). —I. Hyalis lancifolia (Cabrera 4083, LP). — J–L. Abruptly apiculate anther appendage. J1, Gochnatia sect. Discoseris sensu Cabrera (from left to right): Gochnatia amplexifolia (Hatschbach 35312, LP), G. discoidea (Blanchet 3345, LP). J2, Gochnatia sect. Moquiniastrum sensu Cabrera (from left to right): G. argyrea (Hatschbach 9578, LP), G. densicephala (Sheperd 5771 et al., UEC), G. haumaniana (Rojas, herb. Hassler 9752, LP), G. oligocephala (Blanchet 3288, US), G. orbiculata (Brade 5523, US), G. paniculata (Gardner 4810, US), G. polymorpha (Glaziou in 1876, LP), G. velutina (Smith & Klein 14885, LP), G. vernonioides (López et al. 4364, LP). J3, Gochnatia sect. Pentaphorus sensu Cabrera (from left to right): G. foliolosa (Cabrera 3451, LP), G. glutinosa (King 183, LP). J4 (from left to right): G. arborescens (Johnston 4023, LP), G. cordata (Rambo 545, LP), G. purpusii (Purpus 4248, NY). —K. Nouelia insignis (Maire 2516, NY). —L. Wunderlichia azulensis (Harley et al. 25209, MO) (very shortly apiculate). 538 Annals of the Missouri Botanical Garden Figure 6. Style branches. A–C. Papillose style branches. —A. Wunderlichia azulensis (Harley et al. 25209, MO). —B. Chucoa ilicifolia (López Miranda 1090, LP). —C. Pleiotaxis eximia (Rolyns 1836, US). D–K. Smooth style branches. —D. Gochnatia discoidea (Blanchet 3345, LP). —E. G. cordata (Rambo 545, LP). —F. Actinoseris angustifolia (Hatschbach 28756, LP). —G. Cnicothamnus lorentzii (Ruiz Leal 14162, LP). —H. Cyclolepis genistioides (Zardini & Kiesling 114, LP). —I. Hyalis argentea (Ruiz Leal 3701, LP). —J. Ianthopappus corymbosus (Palacios & Cuezzo 2304, LP). —K. Nouelia insignis (Maire 2516, NY). 7C), and Hyalis argentea. In Nouelia one end of the apical cell is very short. (3) Three- to 5-armed hairs (or 3- to 5-branched, stellate hairs sensu Cabrera, 1971): similar to the 2-armed hairs, but the apical cell has 3 to 5 branches (Fig. 7D). The apical cell does not divide and thus the head remains one-celled. This type is found only in Gochnatia barrosii and G. rusbyana of section Moquiniastrum. (4) Multistoried T-shaped hairs: similar to the 2-armed hairs, but further periclinal divisions take place in the apical cell. The head is thus comprised of 3 or 4 one-celled layers, all oriented transversally and parallel, but of different lengths (Fig. 7E). This type of trichome, not very common in Asteraceae, has been reported in the tribe Senecioneae (Robinson, 1989). Ianthopappus is the only genus with this type of trichome. (5) Biseriate glandular hairs: comprised of 2 rows of cells in the body, with two to many cells in each row, enclosed by a persistent or collapsed cuticular vesicle (Fig. 7F, G). Glandular hairs are widespread in all the taxa studied, and especially in species of Gochnatia sect. Pentaphorus (i.e., Gochnatia foliolosa, G. glutinosa) where they cover almost the entire surface of the leaf, with the flagellate hairs restricted to the margins. Achenial pubescence. Glabrous or slightly papillose achenes occur in Chucoa, in a few species of Pleiotaxis (e.g., P. decipiens, P. huillensis, P. linearifolia, P. welwitschii), and Wunderlichia (e.g., W. bahiensis, W. crulsiana, W. insignis, W. senaei). The remaining taxa have villose achenes. Three types of achenial trichomes were found: Duplex or twin hairs, leaf-like hairs (2-armed, oblique-septate flagellate), and glandular hairs. (1) Duplex or twin hairs: Twin hairs are the common type in Asteraceae, comprised of two triangular or rectangular basal cells, one sometimes reduced, and two cylindrical or elliptical hair cells, equal or subequal in length, generally in contact up to their tips (Hess, 1938; Freire & Katinas, 1995) (Fig. 7H, I). All the taxa studied, in- Volume 89, Number 4 2002 Freire et al. Gochnatia Morphology 539 Figure 7. Trichomes. A–G. Leaf hairs. —A. Oblique septate flagellate hair: Gochnatia tortuensis (Ekman H-3553, S). B, C. 2-armed hair. —B. Gochnatia polymorpha (Pedersen 8587, LP). —C. Cyclolepis genistoides (Correa 3172 & Nicora, LP). —D. 3- to 5-armed hair: Gochnatia barrosii (Hatschbach 16945, LP). —E. Multistoried T-shaped hair: Ianthopappus corymbosus (Palacios & Cuezzo 2304, LP). F, G. Biseriate glandular hairs. —F. Gochnatia discoidea (Blanchet 3345, LP). —G. Gochnatia glutinosa (Fabris 1343, LP). H–L. Achenial hairs. H–K. Duplex hairs: Wunderlichia azulensis (Harley et al. 25209, MO). H–I. Duplex hairs with one of the hair cells shorter. —J. Septate duplex hair. —K. Duplex hair with only one hair cell. —L. Capitate glandular biseriate hair: Gochnatia cowellii (Britton & Cowell 10183, NY). cluding most species of Gochnatia, have achenes with twin hairs, usually very long and filiform. In some cases twin hairs have one hair cell very short (e.g., Gochnatia purpusii, G. recurva, G. tortuensis, Cyclolepis, Wunderlichia azulensis, W. mirabilis) (Fig. 7I), they are septate (e.g., Gochnatia hatschbachii, G. oligocephala, Hyalis, Pleiotaxis eximia, Wunderlichia azulensis, W. mirabilis) (Fig. 7J), or have only one hair cell (Wunderlichia azulensis) (Fig. 7K). (2) Leaf-like hairs: Two-armed hairs in the achenes similar in morphology to the leaf hairs, as described above, were found in Gochnatia orbiculata. Only a few species of Gochnatia (e.g., G. cubensis, G. magna, G. ramboi) have oblique-septate flagellate hairs. (3) Achenial biseriate glandular hairs: These are similar to those on the leaves (Fig. 7F, G) and are very widespread in the group under study, occurring with the other types. A modification of the typical biseriate glandular hair with a very enlarged head, i.e., capitate glandular hair (Metcalfe & Chalk, 1950) (Fig. 7L), was found in the Caribbean species of Gochnatia sect. Gochnatia. PAPPUS In all taxa analyzed the pappus is comprised of rigid and scabrid bristles. However, there is interesting variation in bristle length and width, modifications of the lateral cells at the apex of the bristles, and in the number of series of the bristles. Five pappus types were found. Type A. All the bristles have the same length 540 Annals of the Missouri Botanical Garden Figure 8. Pappus. General aspect (on the left) and detail of bristle apex (on the right). —A. Type A: Gochnatia recurva (León 20946, LP). —B. Type B: Gochnatia decora (Maung Mya 5309, LP). —C. Type C: Gochnatia cordata (Serrano 6, LP). —D. Type D: Gochnatia glutinosa (Fabris 1343, LP). —E. Type E: Wunderlichia mirabilis (Irwin et al. 9913, NY). and width (Fig. 8A). This is present in the Caribbean species of Gochnatia sect. Gochnatia (except G. cubensis, G. oligantha, G. sagraeana, G. tortuensis), G. sect. Discoseris, and Actinoseris. Type B. All the bristles have the same width, but about half are shorter than the others (Fig. 8B). This type is present in Gochnatia sect. Leucomeris, the South American species of section Gochnatia, Volume 89, Number 4 2002 Freire et al. Gochnatia Morphology most species of section Hedraiophyllum, and Chucoa, Hyalis, and Nouelia. L1) present in the taxa studied show the great variation of this character in Mutisieae. Cabrera (1961, 1977) characterized Gochnatiinae and Barnadesiinae (the latter subtribe currently constitutes the subfamily Barnadesioideae; Bremer & Jansen, 1992) in the first comprehensive key to subtribes of Mutisieae by having more or less actinomorphic, deeply 5-lobed disc corollas, with equal or unequal segments, but never truly bilabiate, and bilabiate or subligulate ray corollas. The Mutisiinae and Nassauviinae, on the other hand, have clearly bilabiate (the disc florets exceptionally actinomorphic) or ligulate disc and ray florets. According to this key all the taxa studied, although variable in their corollas, would correspond to the subtribe Gochnatiinae (sensu Cabrera, 1977). Other authors (e.g., Robinson, 1991; Bremer, 1994) regard the distinction between Gochnatiinae and Mutisiinae, based only on the actinomorphic versus bilabiate disc florets, to be artificial and recognize only Mutisiinae sensu lato. Apiculate anther appendages have been strongly considered to be an advanced character within Mutisieae (Cabrera, 1977; Hansen, 1991; Karis et al., 1992; Bremer, 1994). The shape of the anther appendage, i.e., caudate, attenuate, and abruptly apiculate (Fig. 5C–L), led to distinction within Gochnatia and also among genera. Indeed, Gochnatia can be associated with Actinoseris, Cnicothamnus, Cyclolepis, Hyalis, Ianthopappus, Nouelia, and Wunderlichia by the common possession of flat (not bulbous), apiculate anther appendages. Chucoa and Pleiotaxis (Fig. 5A, B), based on their acute and bulbous anther appendages, respectively, are very different. The current tribal position of Pleiotaxis is controversial. Some authors regard Pleiotaxis as forming part of a mutisiean ‘‘Dicomagroup’’ (Bremer, 1994; Ortiz, 2000; Ortiz & Coutinho, 2001). According to Hansen (1991), this group of genera (including Pleiotaxis) should be excluded from Mutisieae by features of style branches, anthers, and pollen type. Smooth styles are revealed as another informative feature in this group of taxa, although the value of this character is controversial. Some authors (Bremer, 1987) consider the glabrous styles to be plesiomorphic within Asteraceae (although the Lobeliaceae, characterized by hairy style branches, were used as an outgroup), while others (Stuessy et al., 1996) postulate the smooth condition to be derived with Calyceraceae as the outgroup. At present the hypothesis that Calyceraceae are the sister group of Asteraceae (e.g., Albach et al., 2001; Urtubey & Stuessy, 2001) is widely accepted. Since the style branches in Calyceraceae are papillose Type C. All the bristles have the same width, half of them are shorter, and the longest are plumose at the apex (Fig. 8C). This pappus is present in Gochnatia cordata, G. sect. Moquiniastrum, and Ianthopappus. Type D. Half of the bristles are long and relatively wide and flat (somewhat paleaceous) and the other half are short and thin (Fig. 8D). This type is present in Gochnatia sect. Pentaphorus and some species of Pleiotaxis (e.g., P. dewevrei, P. eximia, P. pulcherrima), all with the long bristles acute at the apex, and in Cnicothamnus, which has the long bristles clavate at the apex. Type E. Half of the bristles are long and relatively wide and flat (somewhat paleaceous) and half are short and thin; all are scabrid, but the longer are plumose at the apex (Fig. 8E). This kind of pappus is found in Cyclolepis, some species of Pleiotaxis (e.g., P. huillersii, P. rogersii), and Wunderlichia. The type A pappus, i.e., with all the bristles of the same length, is 1-seriate with 30 to 40 bristles. Types B–E, i.e., with the bristles of different length, are 2- to 3-seriate with more than 50 bristles. DISCUSSION CHARACTER VALUE Some characters such as habit, most leaf features, and pappus vary within Gochnatia. Other characters, although constant and common to all species of Gochnatia, are not exclusive to it, such as multiseriate involucres, tailed anthers with apiculate anther appendages, and smooth style branches. Table 2 shows this and the main morphological characters that allow the genera associated with Gochnatia to be distinguished. Some of the characters studied merit a brief discussion. The involucre of Gochnatia, resembling a spruce cone, suggested by Hansen (1991) as a distinguishing character for the group of Gochnatia and its associated genera, also occurs in other Mutisieae such as Aphyllocladus, Dinoseris, Hyaloseris, and Stenopadus, which are not very closely related to Gochnatia. We do not therefore consider that this feature has diagnostic value. Corolla morphology has been used as a key diagnostic character in Mutisieae. The tubular actinomorphic (Fig. 4A–E), tubular-filiform (Fig. 4G1), tubular subzygomorphic (Fig. 4F2), subbilabiate (Fig. 4H2, I1), and bilabiate corollas (Fig. 4J1, K1, 541 542 Table 2. Main morphological characters that allow the distinction of Gochnatia and associated genera. The exclusive characters are in boldface. Uncommon features are in parentheses. Characters that distinguish Gochnatia from other genera are in italics. Gochnatia Habit trees, shrubs, (subshrubs) Non-glandular foliar trichomes Pappus Types A, B, C, D Chucoa Cnicothamnus herbs shrubs trees flagellate flagellate epaleate homogamous, heterogamous absent epaleate homogamous (heterogamous) absent dimorphic Cyclolepis Hyalis Ianthopappus Nouelia Pleiotaxis Wunderlichia subshrubs subshrubs shrubs herbs trees flagellate spinose, aphyllous shrubs 2-armed 2-armed multistoried T-shaped 2-armed flagellate flagellate epaleate homogamous epaleate homogamous epaleate homogamous epaleate heterogamous epaleate homogamous epaleate homogamous paleate homogamous absent absent absent absent absent absent isomorphic rhombic, lacerate dimorphic subdimorphic dimorphic dimorphic dimorphic isomorphic isomorphic white, pink, purple yellow orange yellow lilac white and purple white red, pink, white, cream white, yellow apiculate: caudate acute apiculate: caudate apiculate: abruptlyattenuate apiculate: abruptlyattenuate apiculate: caudateattenuate apiculate: abruptly apiculate: abruptly laciniate smooth smooth laciniate laciniate laciniate laciniate apiculate: thickened, bulbose laciniate smooth pubescent: dorsally smooth smooth smooth smooth smooth Type A Type B Type D Type E Type B Type C Type B pubescent: subapically Types D, E smooth pubescent: dorsally Type E Annals of the Missouri Botanical Garden flagellate, 2armed, 3to 5armed Receptacle epaleate Capitula homogamous (heterogamous) Phyllaries apical absent appendage Corollas isomorphic (subdimorphic) Corolla color yellow, cream (lilac, orange, white) Anther apical apiculate: appendage abruptly, caudate, attenuate Anther tails smooth, laciniate Style branches smooth Actinoseris Volume 89, Number 4 2002 Freire et al. Gochnatia Morphology (Hansen, 1992, and our own observations) the smooth style branches in Asteraceae become an advanced character for the family. The multistoried T-shaped hair (Fig. 7E), the 2-, and 3- to 5-armed hairs (Fig. 7B–D), the capitate glandular achenial hair (Fig. 7L), and the pappus types established here (Fig. 8) are revealed as new diagnostic characters. In fact: (1) the multistoried T-shaped hair is exclusive to Ianthopappus and becomes another character to distinguish it; (2) the 2-armed hairs are present in Gochnatia cordata, G. sect. Moquiniastrum, Cyclolepis, Hyalis, and Nouelia; (3) the 3- to 5-armed hairs are exclusive to some species of Gochnatia sect. Moquiniastrum; (4) the capitate glandular achenial hairs, with a very enlarged head, distinguish most of the Caribbean species of Gochnatia sect. Gochnatia; and (5) pappus types A, B, C, D are present in Gochnatia and allow the distinction of sections within the genus (see below), while type E is present only in Cyclolepis, Pleiotaxis, and Wunderlichia. sect. Gochnatia, sect. Hedraiophyllum, and sect. Leucomeris, are redefined resulting in the establishment of five sections: sect. Anastraphioides, sect. Glomerata, sect. Gochnatia, sect. Leucomeris, and sect. Rotundifolia. Cabrera (1971) established two groups in his key to section Gochnatia, the ‘‘South American species’’ and the ‘‘Caribbean species.’’ Species of this section have some characters in common, such as solitary or 2 to 3 capitula and caudate anther appendages, but other characters suggest that it should be separated into two sections: sect. Anastraphioides and sect. Gochnatia. The Caribbean species form the new section Anastraphioides clearly differentiated by spiny leaf margins (Fig. 1E), corollas very deeply 5-lobed (Fig. 4B), and pappus type A (Fig. 8A). The South American species, on the other hand, have predominantly ovate leaves with entire margins (Fig. 1A, F), corollas deeply 5-lobed, and pappus type B (Fig. 8B), and correspond to section Gochnatia. The species Gochnatia rotundifolia, included by Cabrera in section Gochnatia, has characters that show a departure from the other sections of the genus such as glabrous and 3-veined leaves, white corollas, and anther appendages attenuate. It would approach Cabrera’s section Leucomeris by its phyllaries glabrous with ciliate margins and conspicuous parallel veins, but G. rotundifolia lacks the involucre extending into the peduncle typical of this section. Consequently, this species is placed in the new, monotypic section Rotundifolia. Gochnatia sect. Hedraiophyllum sensu Cabrera, which was already recognized by Cabrera (1971) to be artificial, is split off. Three species of this section, G. arborescens, G. magna, and G. purpusii, are placed in the new section Glomerata characterized by the exclusive presence of numerous capitula arranged in glomerulose pseudocorymbs (Fig. 2C). As mentioned above, the type species of section Hedraiophyllum, G. cordata, was moved to Cabrera’s Gochnatia sect. Moquiniastrum. The remaining species of section Hedraiophyllum, G. hypoleuca, G. palosanto, and G. smithii, were placed in section Leucomeris largely based on their involucre with glabrous, conspicuously veined phyllaries, ciliolate at the margins, extending into the peduncle (Fig. 3I). In summary, we propose the following eight sections for the genus Gochnatia: sect. Anastraphioides, sect. Discoseris, sect. Glomerata, sect. Gochnatia, sect. Hedraiophyllum, sect. Leucomeris, sect. Pentaphorus, and sect. Rotundifolia. This new infrageneric classification, with descriptions and a key to the sections, is shown in Appendix 2. CIRCUMSCRIPTION OF GOCHNATIA AND ITS SECTIONS Our morphological analysis confirms that Gochnatia has no single exclusive feature that distinguishes it from related taxa, but it can be defined by a set of characters, i.e., homogamous (rarely heterogamous) capitula, isomorphic (tubular) to subdimorphic (tubular and tubular subzygomorphic) corollas, apiculate anther appendages, and smooth style branches (Table 2). After analyzing the morphological features of Gochnatia, the sections established by Cabrera (1971) were reviewed and some changes are proposed. The distinctiveness of two of the six Cabrera sections, sect. Discoseris and sect. Pentaphorus, was confirmed and additional characters supporting them were found. For instance, Discoseris has pappus type A and Pentaphorus type D. Gochnatia cordata, placed by Cabrera in section Hedraiophyllum, has characters that link it to section Moquiniastrum such as the 2-armed foliar hairs (Fig. 7B), numerous capitula arranged in glomerulose pseudopanicles (Fig. 2H), and pappus type C (Fig. 8C) and must be included in this section. However, it constitutes the type species of section Hedraiophyllum established by Lessing (as a subgenus) in 1832 and has priority over the name of section Moquiniastrum established by Cabrera in 1971 (with G. polymorpha as the type species). Thus, the name Hedraiophyllum is retained in what was known until now as section Moquiniastrum. The remaining three sections sensu Cabrera, 543 544 Annals of the Missouri Botanical Garden Figure 9. Main morphological characters defining the Gochnatia complex, allowing the distinction of Gochnatia from its related taxa, and also sections within Gochnatia. Volume 89, Number 4 2002 Freire et al. Gochnatia Morphology THE GOCHNATIA COMPLEX terminación de los géneros. Revista Mus. Argent. Ci. Nat., Bernardino Rivadavia Inst. Nac. Invest. Ci. Nat. Bot. 2: 291–362. . 1970. Actinoseris, nuevo género de compuestas. Bol. Soc. Argent. Bot. 13: 45–52. . 1971. Revisión del género Gochnatia (Compositae). Revista Mus. La Plata. Bot. 12(66): 1–160. . 1974. Tres Compositae nuevas de Minas Gerais (Brasil). Bol. Mus. Bot. Munic. 15: 1–4. . 1977. Mutisieae. Systematic review. Pp. 1039– 1066 in V. H. Heywood, J. B. Harborne & B. L. Turner (editors), The Biology and Chemistry of the Compositae 2. Academic Press, London. Candolle, A. P. de. 1838. Prodromus 7(1): 24. Paris. Carlquist, S. 1976. Tribal interrelationships and phylogeny of the Asteraceae. Aliso 8: 465–492. Font Quer, P. 1979. Diccionario de Botánica. Ed. Labor, S. A., Barcelona, España. Franchet, M. A. 1888. Les Mutisiacées du Yun-nan. J. Bot. 5: 65–67. Freire, S. E. & L. Katinas. 1995. Morphology and ontogeny of the cypsela hairs of Nassauviinae (Asteraceae, Mutisieae). Pp. 107–143 in D. J. N. Hind, C. Jeffrey & G. Pope (editors), Advances in Compositae Systematics. The Royal Botanic Gardens, Kew. Hansen, H. V. 1991. Phylogenetic studies in Compositae tribe Mutisieae. Opera Bot. 109: 1–50. . 1992. Studies in the Calyceraceae with a discussion of its relationship to Compositae. Nordic J. Bot. 12: 63–75. Harris, J. G. & M. W. Harris. 1994. Plant Identification Terminology: An Illustrated Glossary. Spring Lake Publishing, Spring Lake, Utah. Hess, R. 1938. Vergleichende Untersuchungen über die Zwillingshaare der Kompositen. Bot. Jahrb. Syst. 68: 435–496. Jansen, R. K. & K.-J. Kim. 1996. Implications of chloroplast DNA data for the classification and phylogeny of the Asteraceae. Pp. 317–339 in D. J. N. Hind & H. J. Beentje (editors), Compositae: Systematics. Proceedings of the International Compositae Conference, Kew 1994, Vol. 1. The Royal Botanic Gardens, Kew. & D. J. Palmer. 1987. A chloroplast DNA inversion marks an ancient evolutionary split in the sunflower family (Asteraceae). Proc. Natl. Acad. U.S.A. 84: 5818–5822. Jeffrey, C. 1967. Notes on Compositae: II. The Mutisieae in East Tropical Africa. Kew Bull. 21: 177–223. Jervis, R. N. 1954. A Summary of the Genus Gochnatia including a Revision of the West Indian Species which Comprise the Section Anastraphioides. Ph.D. Dissertation, University of Michigan, Ann Arbor. Karis, O., M. Källersjö & K. Bremer. 1992. Phylogenetic analysis of the Cichorioideae (Asteraceae), with emphasis on the Mutisieae. Ann. Missouri Bot. Gard. 79: 421– 427. , P. Eldenäs & M. Källersjö. 2001. New evidence for the systematic position of Gundelia L. with notes on delimitation of Arctoteae (Asteraceae). Taxon 50: 105– 114. Kunth, C. S. 1818. In F. W. Humboldt, A. J. A. Bonpland & C. S. Kunth, Nov. gen. sp. 4: 19. Lessing, C. F. 1830. Synanthereis Herbarii Regii Berlinensis, Dissertatio Tertia. Linnaea 5: 237–263. . 1832. Gochnatia. Syn. Gen. Compos.: 102–103. Maguire, B. & J. J. Wurdack. 1957. The botany of the The combination of apiculate anther appendages and smooth style branches is exclusive to Actinoseris, Cnicothamnus, Cyclolepis, Gochnatia, Hyalis, Ianthopappus, and Nouelia within Mutisieae. Consequently, these genera are recognized here as the Gochnatia complex (Fig. 9). Chucoa, Pleiotaxis, and Wunderlichia, as already mentioned, differ in their styles and anthers. Within the Gochnatia complex, the genus most morphologically similar to Gochnatia is Cyclolepis. This genus shares trichomes 2-armed, subdimorphic corollas, and gynodioecy with section Hedraiophyllum, which constitutes a pivotal group among the remaining species of Gochnatia and Cyclolepis, providing a ‘‘link’’ between both genera. Cyclolepis, however, can be distinguished from Gochnatia by its leafless, spiny branches, tubular, filiform female florets (Fig. 4G1), and pappus type E (Fig. 8E). The hypothesis that a shrubby habit, large, homogamous, and solitary or few capitula, with yellow, and actinomorphic corollas represent primitive conditions in Asteraceae has been widely discussed and recognized (Maguire & Wurdack, 1957; Carlquist, 1976; Bremer, 1987, 1994; Pruski, 1991; Stuessy et al., 1996). From this point of view, most species of Gochnatia have a set of plesiomorphic characters when compared with the remaining genera of the complex. This hypothesis is consistent with Cabrera’s (1977) idea that Gochnatia is the basal genus in a complex from which the other genera in the subtribe are derived. Literature Cited Albach, D. C., P. S. Soltis, D. E. Soltis & R. G. Olmstead. 2001. Phylogenetic analysis of Asterids based on sequences of four genes. Ann. Missouri Bot. Gard. 88: 163–212. Barroso, G. M. & B. Maguire. 1973. A review of the genus Wunderlichia (Mutisieae, Compositae). Rev. Brasil. Biol. 33(3): 379–406. Beltrán, H. & R. Ferreyra. 2001. Una especie nueva de Asteraceae para Perú y Bolivia: Gochnatia lanceolata. Compositae Newsl. 36: 26–30. Bremer, K. 1987. Tribal interrelationships of the Asteraceae. Cladistics 3: 210–253. . 1994. Asteraceae. Cladistics and Classification. Timber Press, Portland, Oregon. & R. K. Jansen. 1992. A new subfamily of Asteraceae. Ann. Missouri Bot. Gard. 79: 414–415. Cabrera, A. L. 1950. Observaciones sobre los géneros Gochnatia y Moquinia. Notas Mus. La Plata, Bot. 15(74): 37–48. . 1951. Notas sobre compuestas de América Austral. Darwiniana 9(3–4): 363–386. . 1955. Un nuevo género de Mutisieas del Perú. Bol. Soc. Argent. Bot. 6: 40–44. . 1961. Compuestas argentinas: Clave para la de- 545 546 Annals of the Missouri Botanical Garden Guayana Highland, Part II. Mem. New York Bot. Gard. 9(3): 235–392. Metcalfe, C. R. & L. Chalk. 1950. Anatomy of the Dicotyledons, Leaves, Stem, and Wood in Relation to Taxonomy, with Notes on Economic Uses, Vol. II. Clarendon Press, Oxford. Moreno, N. P. 1984. Glosario Botánico Ilustrado. Compañı́a Editorial Continental, S. A. de C. V. (C.E.C.S.A.), Xalapa, Mexico. Ortiz, S. 2000. A phylogenetic analysis of Dicoma Cass. and related genera (Asteraceae: Cichorioideae: Mutisieae) based on morphological and anatomic characters. Ann. Missouri Bot. Gard. 87: 459–481. & A. P. Coutinho. 2001. Achyrothalamus reduced to Erythrocephalum (Asteraceae, Mutisieae). Taxon 50: 389–403. Payne, W. W. 1978. A glossary of plant hair terminology. Brittonia 30: 239–255. Pruski, J. F. 1991. Compositae of the Guayana Highland—V. The Mutisieae of the Lost World of Brazil, Colombia and Guyana. Bol. Mus. Paraense, ser. Bot. 7: 335–392. Ramayya, N. 1962. Studies in the trichomes of some Compositae. I. General structure. Bull. Bot. Surv. India 4: 177–188. Robinson, H. 1989. A revision of the genus Dresslerothamnus (Asteraceae: Senecioneae). Syst. Bot. 14: 380– 388. . 1991. Two new species of Stifftia with notes on relationships of the genus (Asteraceae: Mutisieae). Syst. Bot. 16: 685–692. Roque, N. 1997. A reassessment of Actinoseris polymorpha (Less.) Cabrera (Compositae: Mutisieae) with a new combination. Kew Bull. 52: 197–204. . 2001. Five new species of Richterago (Compositae, Mutisieae): A genus endemic to Brazil. Novon 11: 341–349. & D. J. N. Hind. 2001. Ianthopappus, a new genus of the tribe Mutisieae (Compositae). Novon 11: 97– 101. & J. R. Pirani. 1997. Flora da Serra do Cipó, Minas Gerais: Compositae–Barnadesieae e Mutisieae. Bol. Bot., Univ. Sâo Paulo 16: 151–185. & . 2001. Reinstatement of the name Richterago Kuntze and recircumscription of the genus to include species formerly treated as Actinoseris (Endl.) Cabrera (Compositae, Mutisieae). Taxon 50: 1155– 1160. Sancho, G. 1997. Revisión Sistemática, Análisis Cladı́stico y Biogeográfico de la Sección Moquiniastrum del Género Gochnatia (Asteraceae, Mutisieae). Ph.D. Dissertation, Universidad Nacional de La Plata, La Plata, Argentina. . 2000. Revisión y filogenia de la sección Moquiniastrum Cabrera del género Gochnatia Kunth (Asteraceae, Mutisieae). Fontqueria 54(5): 61–122. Stuessy, T. F., T. Sang & M. L. Devore. 1996. Phylogeny and biogeography of the Barnadesioideae, with implications for early evolution of Compositae. Pp. 463–490 in D. J. N. Hind & H. J. Beentje (editors), Compositae: Systematics. Proceedings of the International Compositae Conference, Kew 1994, Vol. 1. The Royal Botanic Gardens, Kew. Urtubey, E. & T. F. Stuessy. 2001. New hypotheses of phylogenetic relationships in Barnadesioideae (Asteraceae) based on morphology. Taxon 50: 1043–1063. Zardini, E. M. 1975. A new species of Actinoseris (Compositae). Bol. Mus. Bot. Munic. 23: 1–3. APPENDIX 1. Index to specimens examined, with vouchers. Note: During preparation of this work, the new species Gochnatia lanceolata Beltrán & Ferreyra was published (Beltrán & Ferreyra, 2001). Specimens examined of this species (Carolina 01, isotype, US; Torrico & Peca 336, US) have pubescent styles and acute anther appendages, so this species must be excluded from Gochnatia and does not belong to the Gochnatia complex. Actinoseris A. angustifolia: BRAZIL. Minas Gerais: Jaboticatuba, Hatschbach 28756 (LP); Serra do Cipó, Hatschbach 29986 (LP). A. polymorpha: BRAZIL. Minas Gerais: Serra de Cipó, Brade 2007 (LP). A. polyphylla: BRAZIL. Minas Gerais: Santa Ana do Riacho, Hatschbach 35304 (LP). A. radiata: BRAZIL. Paraná: Campo Largo, Hatschbach 690 (LP). A. stenophylla: BRAZIL. Minas Gerais: Santa Ana do Riacho, Hatschbach 35388 & Koczicki (LP). Cyclolepis C. genistoides: PARAGUAY. Chaco Paraguayo, Rojas 7104 (LP). ARGENTINA. Salta: Quebrada de las Conchas, Cabrera 3782 (LP). Rı́o Negro: Gral. Conessa, Correa 3172 & Nicora (LP). Salta: La Candelaria, Schreiter 6630 (LP). San Juan: La Laja, Tinto 2038 (LP). Neuquén: Plotier, Banda, Zardini & Kiesling 114 (LP). Chucoa C. ilicifolia: PERU. La Libertad: Santiago de Chuco, López Miranda 1090 (LP). Cnicothamnus C. azafran: ARGENTINA. Salta: Serranı́as del Crestón, Bortagaray 226 (LP). Jujuy: Santa Bárbara, Cabrera et al. 26287 (LP). C. lorentzii: ARGENTINA. Salta: Caldera, Cabrera et al. 22576 (LP); Capital, Cerro San Bernardo, Padaci 84 (LP); Abra de Santa Laura, Ruiz Leal 14162 (LP); Capital, Bonavı́a 61 (LP); camino de cornisa SaltaJujuy, Zardini 1292 (LP). Gochnatia G. amplexifolia: BRAZIL. Minas Gerais: Santa Ana do Riacho, Hatschbach 35312 (LP). G. arborescens: MEXICO. Baja California: Isla Cerralvo, Johnston 4023 (LP); entre Santonio y Puerto de Bahı́a de los Muertos, Wiggins 5632 (US); along the Pacific Coast, 14 mi. S of Pescadero, Spjut & Edson 6085 (US). G. arequipensis: PERU. Arequipa: Monte Chiwata, Eyerdam & Beetle 22120 (LP). G. argentina: ARGENTINA. Tucumán: Trancas, Meyer 22421 (LP). Entre Rı́os: Paracao, Paraná, Schulz 173 (LP); Leales, Venturi 703 (LP). G. argyrea: BRAZIL. Paraná: Vila Velha, Dusén 4035 (LP), Dusén 9115 (G), Hatschbach 9578 (LP). G. attenuata: CUBA. Oriente: Sierra de Nipe, Ekman 19174 (LP). Volume 89, Number 4 2002 Freire et al. Gochnatia Morphology 547 G. barrosii: BRAZIL. Paraná: Cianorte, Hatschbach 16945 (LP). Minas Gerais: Ituitaba, Macedo 1138 (LP). PARAGUAY. Amambay: Parque Nacional Cerro Corá, Sancho 8 (LP). G. blanchetiana: BRAZIL. Ceará: Serra do Araripe, Gardner 1735 (K). Goiás: Glaziou 21663 (G). G. boliviana: BOLIVIA. Santa Cruz: Florida, Cabrera & Gutiérrez 33804 (LP). G. buchii: DOMINICAN REPUBLIC. Falda del Morro, Monte Cristo, Jiménez 1598 (LP). G. calcicola: CUBA. Oriente: Guantánamo, Ekman 15764 (S). G. cardenasii: ARGENTINA. Jujuy: Tafna, Cordo & Ferrer 88-B-17 (SI). G. cordata: ARGENTINA. Entre Rı́os: Federación, Burkart 23169 & Crespo (LP); Santa Ana, Serrano 6 (LP). BRAZIL. Rio Grande do Sul: Porto Alegre, Rambo 545 (LP). G. cowellii: CUBA. Santa Clara: Santa Clara, Britton & Cowell 10183 (NY). G. crassifolia: CUBA. Oriente: Baracoa, Ekman 4023 (S). G. cubensis: CUBA. Oriente: Sierra de Nipe, Ekman 4767 (S), 9632 (S). G. curviflora: BOLIVIA. Tarija: Tarija, Fiebrig 2838 (LP). ARGENTINA. Jujuy: Humahuaca, Meyer 21409 (LP). Salta: La Candelaria, Schreiter 9409 (LP); La Candelaria, Jeréz et al. 49120 (LP). G. decora: MYANMAR. Bhamo: Lweji, Maung Mya 5309 (LP). G. densicephala: BRAZIL. Rio de Janeiro: Rı́o de Janeiro, Glaziou 11072 (K). Minas Gerais: Poços de Caldas, Leoncini 88 (LP); Pico da Bandeira, Shepherd 5771 et al. (UEC). G. discoidea: BRAZIL. Bahia: Igregia Velha, Blanchet 3345 (LP). G. discolor: BRAZIL. Minas Gerais: Claussen 1301 (NY), Claussen in 1840 (K). G. ekmanii: CUBA. Sin. loc., Wright 2875 (GH). G. enneantha: DOMINICAN REPUBLIC. Cordillera Central, Samaná, Los Haitises, Ekman H-15498 (S). G. floribunda: BRAZIL. Minas Gerais: Morro do Frı́o, Gardner 4806 (K); Diamantina, Hatschbach 30192 (US). Distrito Federal: Brasilia, Hatschbach 43151 (NY). G. foliolosa: CHILE. Santiago: Cerro Renca, Cabrera 3451 (LP). Valparaı́so: El Quisco, Mahu 10336 (LP). G. gardneri: BRAZIL. Goyaz: Capella da Passe, Gardner 4183 (K). G. glutinosa: ARGENTINA. Tucumán: Tafı́, Fabris 1343 (LP). Mendoza: Cerro de La Gloria, King 183 (LP). G. gomezii: CUBA. Oriente: Región de Moa, Cerro Miraflores, Marie-Victorin et al. 21591 (GH). G. hatschbachii: BRAZIL. Minas Gerais: Jaboticatubas, Hatschbach 29951 (LP); Datas, Hatschbach 30155 (LP). G. haumaniana: BRAZIL. Mato Grosso do Sul: Ponta Porá, Meyer 18770 (LP). PARAGUAY. Amambay: Sierra de Amambay, Rojas 6575 (LP), Rojas [herb. Hassler 9752] (G); Est. Los cinco hermanos, Sancho 41 (LP). G. hypoleuca: MEXICO. Hidalgo: Cañada del Vaquero, González Quintero 3215 (LP). Coahuila: Sierra Gavia, Johnston 7223 (LP). G. ilicifolia: BAHAMAS ISLANDS. Andros Island: Coppice, Small & Carter 8526 (K). G. intertexta: CUBA. Pinar del Rı́o: Cajalbana, Alain A1680 (NY). G. magna: MEXICO. San Luis Potosı́: Cronquist 11277 (NY); Queretaro, 5 km SW Jalpan, Fernández 3666 (NY). G. mantuensis: CUBA. Pinar del Rı́o: Guane, Shafer 11208 (LP). G. microcephala: CUBA. Oriente: Boca Guantánamo a Mantua, Bro. Hioram 4874 (NY). G. mollissima: BRAZIL. Rio Grande do Sul: Malme 648 (S); pr. Santa Marı́a, Malme 1261 (S). G. montana: CUBA. Pinar del Rı́o: Guane, Ekman 18725 (S). G. obtusifolia: CUBA. Oriente: Southern Baracoa region, Mesa de Prada, León 11963 (NY). G. oligantha: REPUBLICA DOMINICANA. Monte Cristi, N of Villa Isabel, Jiménez 3614 (US). G. oligocephala: BRAZIL. Bahia: Serra do Jacobina, Blanchet 3288 (US); Rı́o Branco, Curran 284 (NY), Salzmann s.n. (G). G. orbiculata: BRAZIL. São Paulo: Moóca, Brade 5523 (US). Rio Grande do Sul: Guaiba, Sancho 48 (LP); rı́o Jaquety hills, Tweedie 998 (K). G. palosanto: ARGENTINA. Jujuy: San Pedro, Cabrera & Fabris 21157 (LP). Tucumán: Vipos, Trancas, Venturi 1296 (LP), Schreiter in 1925 (LP). G. paniculata: BRAZIL. Minas Gerais: Massa, Brade 13550 (LP), Gardner 4810 (US). G. parvifolia: CUBA. Oriente: Barren Savannas, Shafer 2938 (NY). G. patazina: PERU. La Libertad: Pataz entre Huaylillas y Tayabamba, López & Sagástegui 3409 (LP). G. pauciflosculosa: BAHAMAS ISLANDS. Fortune Island: Eggers 3866 (K); Mariguana Island, 10 mi. W of Abraham Bay, Wilson 7428 (K). G. picardae: HAITI. Massif de la Selle, croix-des Bauquets, gorge of Grande-Rivière de cul-de sac, Ekman H-5385 (K). G. polymorpha: BRAZIL. Rio de Janeiro: Tijuca, Glaziou in 1876 (LP). São Paulo: Paranaiba do Sul, Hashimoto 624 (LP). Rio Grande do Sul: Pereira 8609 & Pabst 7984 (LP). PARAGUAY. Amambay: Sancho 26, Sancho 32 (LP). San Pedro: Lima, Pedersen 8587 (LP). G. pulchra: BRAZIL. São Paulo: Campos de Emas, Cabrera 12311 (LP); Ityrapina, Gehrt 8296 (LP). G. purpusii: MEXICO. Puebla: Tehuacán, Purpus 4248 (NY). G. ramboi: BRAZIL. Santa Catarina: Xanxeré, Rambo 50005 (S). Rio Grande do Sul: Palmeira, Rambo 51961 (LP). G. recurva: CUBA. Oriente: entre Moa y Punta Andén, León 20946 (LP). G. rotundifolia: BRAZIL. São Paulo: Capital, Jabaquara, Handro 157 (LP); Vila Esperança, Joly 596 (LP). G. rusbyana: BOLIVIA. Yungas: Bang 2252 (LP). PERU. Cusco: Alto Urumbamba, Zamalloa 2015 (LP). G. sagraeana: CUBA. Habana: León 7094 (NY); Vedado, Alain 2532 (NY). G. shaferi: CUBA. Oriente: La Caridad, López Figueiras 1738 (NY). G. smithii: MEXICO. Oaxaca: Cuesta de Coyula, Conzatti 4135 (US). G. sordida: BRAZIL. Paraná: Ponta Grossa, Hatschbach 17422 (LP); Senges, Hatschbach 27167 (K). G. spectabilis: INDIA. Uttar Pradesh: Rajpur, Dehra Dun District, Galrola 32 (LP). 548 Annals of the Missouri Botanical Garden G. tortuensis: HAITI. Presquile du Nord-Ouest, Port-de Paix, Ekman H-3553 (S). G. vargasii: PERU. Apurimac: Abancay, Vargas 16317 (LP). G. velutina: BRAZIL. Paraná: Ponta Grossa, Vila Velha, Hatschbach 23447 (LP), López 4364 (LP), Smith & Klein 14885 (LP). G. vernonioides: PERU. Amazonas: Chachapoyas, Tingo, Ferreyra 7097 (LP); entre Chachapoyas y Leimebamba, López et al. 4364 (LP). La Libertad: Bolı́var, Infantes 1701 (LP), López & Sagástegui 3354 (LP). 29. Hyalis H. argentea: ARGENTINA. Mendoza: Tupungato, Ruiz Leal 3701 (LP). H. lancifolia: ARGENTINA. Chaco: San Fernando, Cabrera 4083 (LP), Schinini 16098 (LP). Ianthopappus I. corymbosus: ARGENTINA. Corrientes: Paso Troncón, Palacios & Cuezzo 2304 (LP). 3(2). 39. 4(3). 49. 5(3). 59. 6(5). 69. 7(1). Nouelia N. insignis: CHINA. Maire 2516 (NY). Yunnan: l’ Abbe Delavay 2498 (US), Rock 11714 (US). Pleiotaxis P. dewevrei: DEMOCRATIC REPUBLIC OF CONGO. Lukulu, de Hitte 288 (US). P. eximia: DEMOCRATIC REPUBLIC OF CONGO. Tshinloingwe (Hout Katanga), Rolyns 1836 (US). ZIMBABWE. 20 km de Mangula, Lavranos 22745 (US). P. huillensis: ANGOLA. Huila, Hampata, Gossweiler 10780 (US). P. pulcherrima: ANGOLA. Distr. do Cuanza Sul, Seles, Gossweiler 9367 (US). P. rogersii: ZAIRE—DEMOCRATIC REPUBLIC OF CONGO. Elisabethville, Rolyns 1568 (US). Wunderlichia W. azulensis: BRAZIL. Minas Gerais: Pedra Azul, Harley et al. 25209 (MO). W. crulsiana: BRAZIL. Goiás: Chapada dos Veadeiros, Ratter et al. 2615 (MO). W. mirabilis: BRAZIL. Goiás: Serra do Cristais, Irwin et al. 9913 (NY); about 52 km W of Alto Paraiso, Martinelli & Stutts 999 (NY). APPENDIX 2. New infrageneric classification of Gochnatia. KEY TO THE SECTIONS OF GOCHNATIA 1. Pappus type A (all bristles are thin and have the same length and width) and/or pappus type B (all bristles are thin and have the same width, but about half are shorter) ---------------------------------- 2 19. Pappus type C (all bristles are thin and have the same width, half of them are shorter, and the longer are plumose at the apex) or pappus type D (half of the bristles are long and wide, and the other half are short and thin) ---------------- 7 2(1). Subshrubs; capitula solitary, very long-pedunculate, or arranged in scapiform pseudocorymbs ---------------------------------------------------- G. sect. Discoseris 79. Trees or shrubs; capitula short-pedunculate or sessile ------------------------------------------------------------------- 3 Involucral phyllaries dorsally glabrous or subglabrous and ciliolate at the margins ----------- 4 Involucral phyllaries dorsally tomentose and not ciliolate at the margins --------------------------------- 5 Capitula arranged in glomerulose pseudocorymbs, pseudoracemes, or pseudopanicles; phyllaries extending into the peduncle ------------------------------------------------------------------ G. sect. Leucomeris Solitary capitula; phyllaries not extending into the peduncle -------------------------- G. sect. Rotundifolia Solitary capitula or 2 or 3 ----------------------------------- 6 Capitula in glomerulose pseudocorymbs ----------------------------------------------------------------- G. sect. Glomerata Leaves spiny; corollas deeply to very deeply lobed --------------------------------- G. sect. Anastraphioides Leaves not spiny; corollas deeply lobed ----------------------------------------------------------------- G. sect. Gochnatia Pappus type C; plants not glandulose, with 2-, and 3- to 5-armed hairs; gynodioecious or polygamous dioecious; corollas subdimorphic (isomorphic), 9–15(50) ---- G. sect. Hedraiophyllum Pappus type D; plants glandulose; monoecious; corollas isomorophic, 5 (3, 7, or 10–20) -------------------------------------------------------------- G. sect. Pentaphorus Gochnatia sect. Anastraphioides Jervis ex S. E. Freire, L. Katinas & G. Sancho, sect. nov. TYPE: Gochnatia ilicifolia Less. Arbores vel frutices, foliis alternis, spinoso-dentatis vel integerrimis. Capitula apicibus ramulorum, solitaria vel 2– 3, sessilia. Involucrum campanulatum vel turbinatum. Flores 4–150, lutei vel aurantiaci, isomorphi, hermaphroditi, corolla tubulosa pentasecta vel profunde pentasecta. Antherae appendicibus connectivalibus caudatis vel abruptis, appendicibus basalibus integerrimis vel laciniatis. Pappus uniseriatus vel biseriatus. Shrubs or small trees, monoecious. Leaves alternate, petiolate or shortly petiolate; obovate, oblong, or elliptic with margins spinose-dentate (rarely entire); pinnately veined; upper surface generally glabrous, lower surface densely tomentose (with flagellate hairs). Capitula homogamous, discoid, sessile, solitary at the tip of the branches (rarely 2 or 3). Involucre campanulate or turbinate. Phyllaries in 4 to 10(15) series, dorsally tomentose. Florets 4 to 150, yellow or orange, hermaphroditic, isomorphic, tubulose, deeply to very deeply 5-lobed. Anthers with appendages caudate (rarely abruptly apiculate), and commonly smooth, occasionally laciniate tails. Style bilobate or shortly bifid; style branches rounded, dorsally glabrous. Cypselas with duplex hairs and commonly capitate glandular hairs (rarely with flagellate hairs). Pappus uniseriate with all bristles of the same length (rarely biseriate with a reduced number of outer short bristles) all thin. Twenty-six species: G. attenuata, G. buchii, G. calcicola, G. cowellii, G. crassifolia, G. cubensis, G. ekmanii, G. elliptica, G. enneantha, G. gomezii, G. ilicifolia (type species), G. intertexta, G. maisiana, G. mantuensis, G. microcephala, G. montana, G. obtusifolia, G. oligantha, G. parvifolia, G. pauciflosculosa, G. picardae, G. recurva, G. sagraeana, G. shaferi, G. tortuensis, G. wilsonii. Distribution. Bahamas Islands, Cuba, Haiti, Dominican Republic, Puerto Rico. Observations. As pointed out by Jervis (1954) and Ca- Volume 89, Number 4 2002 Freire et al. Gochnatia Morphology brera (1971), Anastraphia D. Don (Trans. Linn. Soc. London 16: 295. 1830) is considered an unknown genus because its type (Anastraphia ilicifolia D. Don based on a ‘‘Joannes Fraser’’ specimen in the Lambert herbarium) has never been located. The original diagnosis of Anastraphia does not agree with the later interpretation of the genus by de Candolle (1838: 26). Shrubs, monoecious. Leaves alternate, shortly petiolate; ovate or ovate-elliptic with margins entire (rarely denticulate); pinnately, occasionally subtri-veined; upper surface generally glabrous or tomentulose, lower surface densely tomentose (with flagellate hairs). Capitula homogamous, discoid, sessile or short-pedunculate, solitary (rarely 2 or 3). Involucre oblong to campanulate. Phyllaries in 3 to 7 series, dorsally tomentose or only tomentose above. Florets 7 to 40, yellow, hermaphroditic, isomorphic, tubulose, deeply 5-lobed. Anthers with appendages caudate, more rarely abruptly apiculate, and laciniate tails. Style shortly bifid; style branches rounded, dorsally glabrous. Cypselas with duplex hairs and usually glandular hairs. Pappus biseriate of numerous scabrid bristles, with a reduced number of outer, short bristles, all thin. Gochnatia sect. Discoseris (Endlicher) Cabrera, Revista Mus. La Plata 12, Secc. Bot. 66: 150. 1971. Seris Less., non Willd. 1807, Linnaea 5: 253. 1830. Seris sect. Discoseris Endl., Gen. Pl.: 483. 1838. Richterago Kuntze, Rev. Gen. Pl. 1: 360. 1891. Discoseris (Endl.) T. Post & Kuntze, Lex. Gen. Phan.: 181. 1904, nom. superfl. TYPE: Seris discoidea Less. (5 Gochnatia discoidea (Less.) Cabrera). Subshrubs, monoecious. Leaves alternate, petiolate; ovate to ovate-elliptic with margins entire or denticulate; pinnately veined; glabrous or tomentose on both sides (with flagellate hairs). Capitula homogamous, discoid, long-pedunculate and solitary or arranged in scapose pseudocorymbs. Involucre campanulate to turbinate. Phyllaries in 4 or 5 series, dorsally tomentose. Florets numerous (ca. 30), yellow, hermaphroditic, isomorphic, tubulose, deeply 5-lobed. Anthers with appendages abruptly apiculate and laciniate tails. Style shortly bifid; style branches rounded, dorsally glabrous. Cypselas with duplex and glandular hairs. Pappus uniseriate with all bristles of the same length and thin. Three species: G. amplexifolia, G. discoidea (type species), G. suffrutescens. Distribution. Southeastern Brazil. Gochnatia sect. Glomerata S. E. Freire, L. Katinas & G. Sancho, sect. nov. TYPE: Gochnatia arborescens T. S. Brandegee (selected here). Arbores vel frutices, foliis alternis, integerrimis. Capitula pauca, sessilia vel subsessilia, apicibus ramulorum glomerata. Involucrum campanulatum. Flores 12–50, lutei, isomorphi, hemaphroditi, corollis tubulosis, pentasectis. Antherae appendicibus connectivalibus attenuatis vel abruptis, appendicibus basalibus integerrimis. Pappus biseriatus vel uniseriatus. Small trees or shrubs, monoecious. Leaves alternate, shortly petiolate; ovate to elliptic with margins entire or denticulate; pinnately veined; upper surface glabrous or tomentulose, lower surface usually densely tomentose (with flagellate hairs). Capitula homogamous, discoid, sessile or subsessile, few together in terminal glomerulose pseudocorymbs. Involucre campanulate. Phyllaries in 5 or 6 series (rarely 8 to 10), dorsally tomentose or glabrous. Florets 12 to 20 (rarely ca. 50), yellow, hermaphroditic, isomorphic, tubulose, deeply 5-lobed. Anthers with appendages abruptly apiculate (rarely attenuate) and smooth tails. Style shortly bifid or bilobate; style branches rounded, dorsally glabrous. Cypselas with duplex and glandular hairs. Pappus biseriate of numerous scabrid bristles, with a reduced number of outer short bristles (rarely uniseriate with all bristles of the same length), all thin. Three species: G. arborescens (type species), G. magna, G. purpusii. Distribution. Northern Mexico. Gochnatia Kunth sect. Gochnatia, Nov. Gen. Sp. 4: 15. 1818. TYPE: Gochnatia vernonioides Kunth. 549 Seven species: G. arequipensis, G. boliviana, G. cardenasii, G. curviflora, G. patazina, G. vargasii, G. vernonioides (type species). Distribution. Argentina. Andes of Peru, Bolivia, and northwestern Gochnatia sect. Hedraiophyllum (Lessing) DC., Prodr. 7(1): 24. 1838. Gochnatia subg. Hedraiophyllum Less., Syn. Gen. Compos.: 103. 1832. TYPE: Gochnatia cordata Less. Gochnatia sect. Moquiniastrum Cabrera, Revista Mus. La Plata 12, Secc. Bot. 66: 73. 1971. TYPE: Spadonia polymorpha Less. (5 G. polymorpha (Less.) Cabrera). Shrubs or trees, commonly gynodioecious or polygamous dioecious. Leaves alternate, petiolate or shortly petiolate; ovate, elliptic (rarely linear or cordate) with margins entire, rarely denticulate; pinnately veined; upper surface glabrous (rarely tomentose), lower surface densely tomentose (with 2-, 3- to 5-armed hairs, occasionally flagellate). Capitula heterogamous (homogamous), disciform (rarely discoid), subsessile to pedunculate, numerous arranged in terminal and usually loose leafy pseudopanicles. Involucre oblong to campanulate. Phyllaries in 3 or 4 series, dorsally tomentose. Florets 9 to 15(50), creamy or white, tubulose, deeply 5-lobed, subdimorphic (occasionally isomorphic); functionally female, corollas slightly zygomorphic, with straight lobes; hermaphroditic, actinomorphic, with resupinate lobes. Anthers with appendages abruptly apiculate and laciniate or smooth tails. Style shortly bifid (rarely bilobate); style branches rounded, dorsally glabrous. Cypselas with duplex, glandular, and flagellate hairs. Pappus biseriate of numerous scabrid bristles, with a reduced number of outer short bristles, all thin, and the longest are plumose at the apex. Twenty-one species: G. argentina, G. argyrea, G. barrosii, G. blanchetiana, G. cordata (type species), G. densicephala, G. discolor, G. floribunda, G. gardneri, G. hatschbachii, G. haumaniana, G. mollissima, G. oligocephala, G. orbiculata, G. paniculata, G. polymorpha, G. pulchra, G. ramboi, G. rusbyana, G. sordida, G. velutina. Distribution. Andes of Peru and Bolivia, eastern Brazil, Paraguay, Uruguay, and central-eastern Argentina. Gochnatia sect. Leucomeris (D. Don) Cabrera, Revista Mus. La Plata 12, Secc. 66: 128. 1971. Leucomeris D. Don, Prodr. Fl. Nepal.: 169. 1825. Gochnatia subg. Leucomeris (D. Don) Less., Syn. Gen. Compos.: 103. 1832. TYPE: Gochnatia spectabilis (D. Don) Less. (5 Leucomeris spectabilis D. Don). 550 Annals of the Missouri Botanical Garden Trees or shrubs, monoecious. Leaves alternate, shortly petiolate; elliptic, margins entire or denticulate; pinnately (3-)veined; upper surface generally glabrous to tomentose, lower surface densely tomentose (with flagellate hairs) or glabrous on both sides. Capitula homogamous, discoid, subsessile, many in glomerules, arranged in terminal leafy pseudoracemes, pseudopanicles, or pseudocorymbs. Involucre campanulate, oblong or turbinate. Phyllaries in 3 to 6(7) series, dorsally glabrous with ciliolate margins, extending to the peduncle. Florets 4 to 6 (8 to 12), white (rarely yellow), hermaphroditic, isomorphic, tubulose, deeply 5-lobed. Anthers with appendages attenuate (rarely abruptly apiculate) and laciniate tails (rarely smooth). Style bilobate or shortly bifid, style branches obtuse or rounded, dorsally glabrous. Cypselas with duplex hairs and commonly with glandular hairs. Pappus biseriate of numerous scabrid bristles, with a reduced number of outer short bristles, all thin. or tomentulose with ciliolate margin. Florets 5 (rarely 3, 7, or 10 to 20), white or lilac, hermaphroditic, isomorphic, tubulose, deeply 5-lobed. Anthers with appendages abruptly apiculate and smooth or laciniate tails. Style bilobate or shortly bifid; style branches rounded, dorsally glabrous. Cypselas with duplex and glandular hairs. Pappus biseriate of numerous scabrid bristles, half of them relatively wide and long, and the others short and thin. Five species: G. hypoleuca, G. palosanto, G. smithii, G. decora, G. spectabilis (type species). Distribution. Mexico, Brazil, Andean region of Bolivia and Argentina, and southeastern Asia. Gochnatia sect. Pentaphorus (D. Don) DC., Prodr. 7(1): 24. 1838. Pentaphorus D. Don, Trans. Linn. Soc. London 16: 296. 1830. Gochnatia subg. Pentaphorus (D. Don) Hook. & Arn., Comp. Bot. Mag. 1: 108. 1835. TYPE: Gochnatia foliolosa D. Don ex Hook. & Arn. Shrubs, monoecious. Leaves alternate, sessile; linearovate or linear-obovate with margins entire or upper portion denticulate; pinnately or three-veined; upper and lower surfaces glandulate (with or without flagellate hairs). Capitula homogamous, discoid, sessile or subsessile, numerous in leafy glomerulose pseudoracemes. Involucre campanulate. Phyllaries in 4 to 6 series, dorsally glabrous View publication stats Two species: G. foliolosa, G. glutinosa (type species). Distribution. Western Argentina and central Chile. Gochnatia sect. Rotundifolia S. E. Freire, L. Katinas & G. Sancho, sect. nov. TYPE: Gochnatia rotundifolia Less. (selected here). Frutices, foliis alternis, integerrimis vel denticulatis. Capitula apicibus ramulorum solitaria, sessilia. Involucrum campanulatum; bracteis involucralibus subglabris, margine ciliatis. Flores multi, albi, isomorphi, hermaphroditi, corollis tubulosis, pentasectis. Antherae appendicibus connectivalibus attenuatis apiculatis, appendicibus basalibus laciniatis. Pappus biseriatus, setosus. Shrubs, monoecious. Leaves alternate, shortly petiolate; broadly elliptic with margins entire to denticulate; threeveined, glabrous on both surfaces at maturity (young leaves with flagellate hairs). Capitula homogamous, discoid, sessile, solitary. Involucre campanulate. Phyllaries in 4 or 5 series, dorsally subglabrous with ciliolate margin. Florets ca. 50, white, hermaphroditic, isomorphic, tubulose, deeply 5-lobed. Anthers with appendages attenuate and laciniate tails. Style bilobate; style branches rounded, dorsally glabrous. Cypselas with duplex and glandular hairs. Pappus biseriate of numerous scabrid bristles, with a reduced number of outer, short bristles, all thin. One species: G. rotundifolia. Distribution. Southeastern Brazil.

RELATED PAPERS

RELATED TOPICS

Log In

Gochnatia (Asteraceae, Mutisieae) and the Gochnatia complex: Taxonomic implications from morphology

Gochnatia (Asteraceae, Mutisieae) and the Gochnatia complex: Taxonomic implications from morphology

Related Papers

RELATED PAPERS

RELATED TOPICS