Available online at www.sciencedirect.com R Review of Palaeobotany and Palynology 123 (2003) 237^246 www.elsevier.com/locate/revpalbo Proustia and Lophopappus (Asteraceae, Mutisieae): generic and subtribal relationships based on pollen morphology Mar|¤a Cristina Teller|¤a a; , Estrella Urtubey b , Liliana Katinas b b a Divisio¤n Paleobota¤nica, Facultad de Ciencias Naturales y Museo, Paseo del Bosque, 1900 La Plata, Argentina Divisio¤n Plantas Vasculares, Facultad de Ciencias Naturales y Museo, Paseo del Bosque, 1900 La Plata, Argentina Received 8 January 2002; received in revised form 30 May 2002; accepted 16 October 2002 Abstract Pollen of Proustia (10 specimens examined) and Lophopappus (12 specimens examined) (Asteraceae, Mutisieae) are described as part of an analysis of the classification of these genera. Their exine structure is determined and compared with structures found in other genera of Mutisieae (25 specimens examined and data of other authors), in order to clarify the subtribal position of Proustia and Lophopappus. Pollen of Proustia and Lophopappus have the following characters in common: Grains radially symmetrical and isopolar; tricolporate, long colpi with a microgranulate membrane, endoaperture lalongate with acute or bifurcate equatorial ends; subprolate or prolatespheroidal; elliptic or spheroidal; polar caps conspicuous or not so distinctive, and exine microechinate. Two types of sexine structure are observed: (1) Trixis type, with two columellate ramified sublayers, the inner sublayer thicker than the outer one (found in Lophopappus berberidifolius, L. tarapacanus, Proustia cuneifolia var. cuneifolia, P. cuneifolia var. mendocina, P. cuneifolia var. mollis, P. ilicifolia var. baccharoides, and P. pyrifolia); and (2) Proustia type, with two equally thick sublayers of ramified columella (found in Lophopappus blakei, L. cuneatus, and L. foliosus). Proustia ilicifolia var. ilicifolia and Lophopappus peruvianus have both types of sexine structure. Pollen analysis of other genera of Mutisieae subtribes Nassauviinae, Mutisiinae and Gochnatiinae demonstrate two new exine types: (1) Ainsliaea type, with two sublayers poorly distinguishable, the exine appearing compact, the outer surface microechinate; and (2) Wunderlichia type, with two well-differentiated sublayers, the outer surface echinate, the spines with apical channels. The results show that: (1) Proustia and Lophopappus cannot be differentiated from each other by pollen characters; (2) the genera of subtribe Nassauviinae can be differentiated from the other two subtribes by having sexine sublayers with similar structure; (3) Proustia and Lophopappus have the pollen exine structural types that are typical of pollen of subtribe Nassauviinae; (4) subtribes Gochnatiinae and Mutisiinae are characterized by having sexine sublayers with different structures; but (5) those two subtribes cannot be differentiated based on pollen characters. : 2003 Elsevier Science B.V. All rights reserved. Keywords: pollen; Proustia; Lophopappus; Mutisieae; Asteraceae 1. Introduction * Corresponding author. Fax: +54-221-425-7527. E-mail addresses: telleria@netverk.com.ar (M.C. Teller|¤a), urtubey@museo.fcnym.unlp.edu.ar (E. Urtubey), katinas@museo.fcnym.unlp.edu.ar (L. Katinas). Proustia Lagasca and Lophopappus Rusby (Asteraceae, Mutisieae) are distributed from Peru to Northern Argentina and Chile. Proustia comprises three species and seven infraspeci¢c taxa 0034-6667 / 03 / $ ^ see front matter : 2003 Elsevier Science B.V. All rights reserved. PII: S 0 0 3 4 - 6 6 6 7 ( 0 2 ) 0 0 2 2 7 - 0 238 M.C. Teller|¤a et al. / Review of Palaeobotany and Palynology 123 (2003) 237^246 of shrubs and vines, and Lophopappus six species of shrubs. Within the subtribal classi¢cation of Mutisieae sensu Cabrera (1977), various authors have placed these two genera alternatively into subtribe Gochnatiinae (Cabrera, 1953, 1961), subtribe Mutisiinae (Fabris, 1968; Cabrera, 1961), and subtribe Nassauviinae (Crisci, 1974a), primarily using features of the corollas and styles. Both genera have style branches with rounded tips (characteristic of Mutisiinae and Gochnatiinae) and bilabiate corollas (characteristic of Nassauvinae), and Lophopappus has occasionally 5-partite corollas (characteristic of Gochnatiinae). Most recently, in a morphological study of Mutisieae, Hansen (1991a) suggested that these two genera may be misplaced in tribe Mutisieae. Following the taxonomic treatments of Lophopappus by Cabrera (1953), and Proustia by Fabris (1968), most recent authors agree that they are distinct but closely relationed. Their close resemblance led Ferreyra (1995) to consider Proustia and Lophopappus as congeneric, treating them all as species of Proustia. In the ¢rst palynological studies in Proustia, Wodehouse (1929) examined pollen of Proustia cuneifolia var. cuneifolia (sub P. pungens) and P. pyrifolia with light microscopy. Later, Parra and Marticorena (1972) established several pollen groups in their study of the Chilean Mutisieae. They analyzed the pollen grains of Proustia cuneifolia var. cuneifolia, P. ilicifolia, P. pyrifolia and Lophopappus tarapacanus, and established for them the ‘Proustia pollen group’, characterized by a bi-layered columellate sexine, with the two sublayers of similar thickness. They suggested a close relationship between the ‘Proustia pollen group’ and the ‘Trixis pollen group’ of the Nassauviinae. The ‘Trixis pollen group’, characteristic of Leunisia, Macrachaenium, Pleocarphus, and Trixis, also has bi-layered columellate sexine, but the inner sublayer is thicker than the outer sublayer. The ‘Proustia pollen group’ was related to a lesser degree with the ‘Perezia pollen subgroup’, found in Nassauvia, Triptilion, and Perezia, which is characterized by two sublayers of the same thickness and separated by a thin non-parallel (‘zigzag’) nexine layer. Crisci (1974a) summarized the palynological data for subtribe Nassauviinae, recognizing six types of exine patterns, one being the Proustia type found in Proustia and Lophopappus, which resembles the Trixis type, as noted by Parra and Marticorena (1972). Crisci (1974a) emphasized the importance of the pollen exine structure in Proustia and Lophopappus for their inclusion in Nassauviinae. Hansen (1991b) examined general pollen characteristics (size, sculpture and structure) of some species of Mutisieae, including Proustia cuneifolia and Lophopappus foliosus. Hansen observed that Gochnatiinae and Mutisiinae have a poorly di¡erentiated ectosexine (as tectum), with the endosexine (as infratectum) di¡erentiated into stout, spaced columellae, whereas the Nassauviinae have both sublayers di¡erentiated into stout, spaced columellae. The aims of this study are : (1) To describe and to illustrate the pollen morphology of all species of Proustia and Lophopappus; (2) to determine the pollen types for all of these taxa; (3) to determine whether Proustia and Lophopappus can be distinguished based on pollen data; and (4) to determine whether the exine structure of Proustia and Lophopappus can provide support for the placement of these genera within the subtribal classi¢cation of tribe Mutisieae sensu Cabrera. 2. Materials and methods Pollen samples were collected from herbarium specimens from AAU, BKF, C, CM, LP, MO, UC, and W (Holmgren et al., 1990 ; see Appendix A). For light microscopy (LM), pollen was acetolyzed (Erdtman, 1960), mounted in glycerol jelly and sealed with para⁄n. Measurements of the polar axis (P), equatorial diameter (E) and exine thickness (measured in equatorial view), are based on 25 grains. The P/E ratio was calculated for each specimen. Table 1 lists species examined and pollen data. For scanning electron microscopy (SEM), unacetolyzed grains were suspended in 90% ethanol, mounted on stubs, sputter-coated with gold palladium, and examined with a Jeol JSM T-100 microscope. Terminology in general follows Punt et al. (1994). Table 2 lists the exine structural type M.C. Teller|¤a et al. / Review of Palaeobotany and Palynology 123 (2003) 237^246 239 Table 1 List of species and varieties of Lophopappus and Proustia examineda Taxa a. P (Wm) b. E (Wm) c. P/E d. Exine (Wm) e. Polar caps f. Exine structure 1. 2. 3. 4. 5. 6. 7a. 45^50 42.5^45 40^45 45^50 42^54 44^46 32^39.5 37 31^36 36^38 37^45 35^40 40^44 28^34 Subprolate Subprolate Subprolate Prolatespheroidal Subprolate Prolatespheroidal Subprolate ca. 7 6^7 4^5 5^6 ca.6 4^6 4.0^5.5 + +(3) +(3) +(3) + +(3) +(3) Trixis Proustia Proustia Proustia Proustia+Trixis Trixis Trixis 31^40 29^37 28^33 23^30 30^33 28^37 25^35 23^29 19^21 23^27 Subprolate Subprolate Subprolate Subprolate Subprolate 4.0^6.0 4.5^6.0 2.0^4.5 2.5^3.5 3.5 +(3) +(3) +(3) +(3) + Trixis Trixis Trixis Proustia+Trixis Trixis Lophopappus berberidifolius L. blakei L. cuneatus L. foliosus L. peruvianus L. tarapacanus Proustia cuneifolia var. cuneifolia 7b. P. cuneifolia var. mendocina 7c. P. cuneifolia var. mollis 8a. P. ilicifolia var. baccharoides 8b. P. ilicifolia var. ilicifolia 9. P. pyrifolia var. pyrifolia a The measurements of polar diameter (P, Wm), equatorial diameter (E, Wm); shape (P/E), exine thickness (Wm), polar caps, and structure types are indicated; + = presence; 3 = absence. Table 2 Exine types in genera of subtribes Nassauviinae, Gochnatiinae and Mutisiinae (tribe Mutisieae sensu Cabrera, 1977)a a. Exine types 1. 2. 3. Calopappus Cephalopappus Oxyphyllum 4. 5. Proustia Trixis 6. 7. 8. 9. Ainsliaea Erythrocephalum Dicoma Mutisia 10. Wunderlichia a b. Nassauviinae c. Gochnatiinae d. Mutisiinae Ainsliaea* Erythrocephalum* Dicoma8 Aphyllocladus6 , Chimantaea*, Cnicothamnus*, Gochnatia5 , Gypothamnium6 , Hecastocleis*, Hyalis*, Lycoseris*, Onoseris7 , Plazia6 , Stenopadus* Sti¡tia*, Urmenetea6 Wunderlichia* Brachyclados6 , Chaetanthera6 , Chaptalia6 , Glossarion7 , Hyaloseris*, Mutisia1 *, Lulia*, Pachylaena6 , Trichocline9 2 Calopappus Cephalopappus* Leucheria4 , Moscharia6 , Nassauvia6 , Oxyphyllum*, Perezia6 , Polyachyrus6 , Triptilion6 Lophopappus*, Proustia* Acourtia2 , Ameghinoa2 , Burkartia*, Criscia2 , Dolichlasium2 , Holocheilus2 , Jungia2 , Leunisia6 , Lophopappus*, Macrachaenium*, Marticorenia3 , Pleocarphus6 , Proustia*, Trixis2 Gongylolepis* Taxa analyzed in this study indicated by asterisk. Species of Proustia and Lophopappus are cited in Table 1, the other taxa are cited in Appendix A. Numbers refer to the literature where the data were extracted: (1) Cabrera, 1965; (2) Crisci, 1974a; (3) Crisci, 1974b; (4) Crisci, 1976; (5) Morbelli, unpubl.; (6) Parra and Marticorena, 1972; (7) Skvarla et al., 1977; (8) Stix, 1960; (9) Zardini, 1975. 240 M.C. Teller|¤a et al. / Review of Palaeobotany and Palynology 123 (2003) 237^246 for genera of subtribes Nassauviinae, Mutisiinae and Gochnatiinae, based on our own observations or from the literature (see Appendix A). 3. Results 3.1. Proustia (Plate I) and Lophopappus (Plate II) pollen description General description (Table 1). Pollen radially 2 1 7 3 5 4 8 symmetrical and isopolar; subprolate or prolate^ spheroidal, spheroidal or elliptic in equatorial view, circular in polar view. Tricolporate, long colpi with a microgranulate membrane. Endoaperture lalongate, with bifurcate (Plate I, ¢g. 6) or acute equatorial ends (Plate II, ¢g. 15). Exine tectate, microechinate. Sexine about 2 or 3 times as thick as nexine; thickened at polar areas (Plate I, ¢g. 6), equatorial areas (Plate I, ¢g. 1), at both equatorial and polar areas (Plate II, ¢g. 10), or not; consisting of two rami¢ed columellate sub- 6 9 M.C. Teller|¤a et al. / Review of Palaeobotany and Palynology 123 (2003) 237^246 layers; the sublayers equal in thickness (Proustia type) (Plate II, ¢g. 12; Fig. 1) or the endosexine thicker than the ectosexine (Trixis type) (Plate I, ¢gs. 1 and 5; Fig. 1). Internal tectum parallel to the nexine. Nexine thickened towards apertures, forming costae. Size (Table 1, a,b). Pollen grains of Lophopappus are larger (40^50U31^45 Wm) than those of Proustia (23^40U19^37 Wm), with overlap. Thus, it is not possible to distinguish species within Proustia and Lophopappus based on pollen size. Shape in equatorial view. Most species have a mixture of elliptic and spheroidal pollen grains, except for Lophopappus berberidifolius, L. peruvianus, Proustia cuneifolia var. mollis, P. ilicifolia var. baccharoides, and P. pyrifolia var. pyrifolia, which have exclusively elliptic grains. Shape (ratio P/E) (Table 1, c). Pollen grains of all species of Proustia and most species of Lophopappus are subprolate, except for Lophopappus foliosus and L. tarapacanus, which have prolate^spheroidal pollen. Pollen of these genera were described as prolate by Parra and Marticorena (1972) and Hansen (1991b) or prolate^spheroidal to prolate by Crisci (1974a). Exine thickness (Table 1, d). The exine of Lophopappus pollen (4^7 Wm) is generally thicker than that of Proustia (2^6 Wm), with considerable overlap in size. Polar caps (Table 1, e). These structures are conspicuous in Lophopappus berberidifolius (Plate II, ¢g. 10), L. peruvianus, and Proustia pyrifolia var. pyrifolia (Plate I, ¢g. 6), but are indistinct in the other taxa. Hansen (1991b) noted that the caps may be related to the shape of the pollen grains, i.e. polar caps are well-developed on elliptic grains, and poorly developed on spheroidal grains. Exine structure (Table 1, f; Fig. 1). All of the taxa of Proustia and Lophopappus are characterized by either the Trixis or Proustia exine structural type. Lophopappus berberidifolius, L. tarapacanus, Proustia cuneifolia var. cuneifolia, P. cuneifolia var. mendocina, P. cuneifolia var. mollis, P. ilicifolia var. baccharoides and P. pyrifolia var. pyrifolia have the Trixis type (ectosexine thinner than endosexine, both sublayers columellate; microechinate). Lophopappus blakei, L. cuneatus and L. foliosus have the Proustia type (ectosexine of same thickness as endosexine, both sublayers columellate; microechinate). Lophopappus peruvianus and Proustia ilicifolia var. ilicifolia have both types of exine structure. 3.2. Exine types found in other genera of Mutisieae We recognize eight types of exine of Mutisieae in addition to the Proustia and Trixis types described above. Six of these types coincide with the exine types previously reported, and two new types are reported here. These are based on the number of sexine sublayers, the relative thickness of the two sublayers, the distinction of columellae (clearly di¡erentiated or not), and the orientation of the internal tectum (parallel to the nexine or zigzag; Crisci, 1974a). We added an additional character: type of surface (echinate or microechi- Plate I. Proustia 1^3. 1. 2. 3. 4^5. 4. 5. 6^9. 6. 7. 8. 9. 241 P. cuneifolia var. cuneifolia (Okada 3036 LP). LM, equatorial view in optical section. SEM, equatorial view. SEM, polar view. P. ilicifolia var. ilicifolia (Ricardi 5553 LP). SEM, equatorial and polar views. LM, equatorial view in optical section. P. pyrifolia var. pyrifolia (Friedrich LP 071038). LM, equatorial view, showing the endoaperture with bifurcate ends. LM, polar view in optical section. SEM, details of aperture showing granular membrane. SEM, polar view. 242 M.C. Teller|¤a et al. / Review of Palaeobotany and Palynology 123 (2003) 237^246 nate). These eight exine types are as follows (Fig. 1): Ainsliaea type (established here; Table 2, a.6): Ectosexine and the endosexine poorly di¡erentiated; both compact. Microechinate. 14 10 11 Calopappus type (Crisci, 1974a ; Table 2, a.1): Ectosexine thinner than the endosexine; both layers columellate and separated by a zigzag (non-parallel to nexine) internal tectum. Microechinate. 13 15 17 18 12 16 M.C. Teller|¤a et al. / Review of Palaeobotany and Palynology 123 (2003) 237^246 243 Fig. 1. Exine types of the three subtribes (Nassauviinae, Gochnatiinae and Mutisiinae) of Mutisieae sensu Cabrera (1977), showing the degree of overlapping. The Ainsliaea and Wunderlichia exine types are described here. Cephalopappus type (Crisci, 1974a; Table 2, a.2): Ectosexine thicker than endosexine; both layers columellate and separated by an internal tectum more or less parallel to nexine. Microechinate. Dicoma type (Stix, 1960; Table 2, a.8): Ectosexine slightly thinner than endosexine; ectosexine compact or scarcely columellate, endosexine with stout, unrami¢ed columellae. Microechinate. Erythrocephalum type (Stix, 1960; Table 2, a.7): Ectosexine (bi-layered) thicker than endosexine; both layers columellate. Echinate, spines with apical channels. Mutisia type (Stix, 1960 ; Table 2, a.9): Ectosexine slightly thinner than endosexine; ectosexine compact or scarcely columellate, endosexine with stout, rami¢ed columellae. Microechinate. Oxyphyllum type (Crisci, 1974a ; Table 2, a.3): Ectosexine and endosexine equally thick, separated from each other by zigzag internal tectum. Microechinate. Wunderlichia type (established here; Table 2, a.10): Ectosexine slightly thinner than endosexine ; ectosexine with compact inner layer and thinner scarcely columellate outer layer, endosexine with stout rami¢ed columellae. Echinate or microechinate, spines with apical channels. 4. Discussion 4.1. Generic status and relationships between Proustia and Lophopappus The exine structure of the pollen of Lophopap- Plate II. Lophopappus. 10. 11. 12. 13. 14^18. 14. 15. 16. 17. 18. L. berberidifolius (Tovar 961 LP), LM, equatorial view in optical section. L. cuneatus (Meyer et Bianchi 33175 LP), SEM, equatorial view. L. blakei (Cabrera et Fabris 13462 LP), LM, equatorial view in optical section. L. foliosus (Cabrera 25270 LP), SEM, equatorial view. L. tarapacanus (Ricardi et Marticorena 25604 LP) LM, equatorial view in optical section. LM, equatorial view, high focus showing aperture. LM, polar view in optical section. SEM, polar view. SEM, detail of apertural membrane and sculpture. 244 M.C. Teller|¤a et al. / Review of Palaeobotany and Palynology 123 (2003) 237^246 pus and Proustia clearly belongs to the Anthemoid pattern established by Skvarla et al. (1977). This pattern is characterized by the absence of a cavus and the presence of thick columellae that support a level of shorter columellae with an internal tectum. Based on pollen data, Proustia and Lophopappus di¡er in several ways: (1) Lophopappus pollen grains (polar diameter 40^54 Wm) are larger than those of Proustia (polar diameter 23^40 Wm); (2) in general, the exine is thicker in Lophopappus (4^ 7 Wm) than in Proustia (2^6 Wm), although they overlap considerably; and (3) most taxa of Lophopappus have Proustia type (except L. berberidifolius and L. tarapacanus with Trixis type, and L. peruvianus with both Proustia and Trixis types), whereas most taxa of Proustia have the Trixis type (except Proustia ilicifolia var. ilicifolia with both Proustia and Trixis types). The Proustia type is found exclusively in Proustia and Lophopappus, whereas the Trixis type occurs in some species of Proustia and Lophopappus as well as other several genera of Nassauviinae (Table 2). Prior to this study, the Proustia type was the only one known for Proustia and Lophoppapus (Parra and Marticorena, 1972; Crisci, 1974a), but we also found the Trixis type to be present in both genera. This analysis reinforces the general taxonomic consensus (Parra and Marticorena, 1972; Crisci, 1974a; Ferreyra, 1995) that these two genera are very closely related. Moreover, we should emphasize that they are the only genera of Mutisieae with the Proustia type. It may be signi¢cant to note that the three species of Lophopappus with Proustia type (L. blakei, L. cuneatus and L. foliosus) are also the only species with exclusively bilabiate £orets, the other three species of Lophopappus having mainly or uniquely tubular £orets. 4.2. Subtribal relationships and position of Proustia and Lophopappus We found, like Hansen (1991b), that pollen of Nassauviinae di¡er from those of Gochnatiinae and Mutisiinae. The exine of Nassauviinae pollen is distinctly bilayered, with both ectosexine and endosexine clearly columellate (i.e. Calopappus, Cephalopappus, Oxyphyllum, Proustia, and Trixis types). This pattern is consistent throughout the subtribe, the sublayers di¡ering only in relative thickness and in the orientation of the internal tectum (parallel to the nexine or zigzag). Proustia and Lophopappus appear strongly and clearly related to the pollen of the genera of Nassauviinae, supporting Crisci’s (1974a) statement that both genera belong to this subtribe. The types of exine structure found in Gochnatiinae and Mutisiinae are heterogeneous, with most having a compact ectosexine constituted by columellae very thickly disposed, and an endosexine with the columellae clearly distinguishable, stout and rami¢ed. Exine types do not di¡erentiate these two subtribes from one another, since the Mutisia and Wunderlichia types are found in both subtribes (Table 2; Fig. 1). Because the pollen data demonstrate a broad overlap in exine features among the genera of Mutisiinae and Gochnatiinae, these data support the hypothesis of other authors (Robinson, 1991; Hansen, 1991a; Bremer, 1994) that the subtribes Gochnatiinae and Mutisiinae are arti¢cially delimited, and that the tribe Mutisieae is comprised of only two subtribes, that is, Nassauviinae and Mutisiinae sensu lato. We should note that several distinctive exine structural types are to be found, especially in tribe Gochnatiinae. The species of Ainsliaea (Gochnatiinae) have a unique exine type, characterized by a compact structure with poorly di¡erentiated sublayers. Likewise, the echinate pollen sculpturing of Erythrocephalum, Gongylolepis, and Wunderlichia is unique in Mutisieae, but is quite very similar to exine features found in some members of the Cardueae (Tormo Molina and Ubera Jime¤nez, 1995), a cichorioid tribe closely related to Mutisieae. We hope to expand our analysis of pollen features, including size, shape, exine thickness, and apertural characteristics, to a more complete sampling of genera and species groups of Mutisieae and its relatives, in order to contribute to our understanding of the diversity of pollen types and of the relationships within Mutisieae and among Mutisieae and other tribes of Cichorioideae. M.C. Teller|¤a et al. / Review of Palaeobotany and Palynology 123 (2003) 237^246 Acknowledgements We thank J.V. Crisci, S.E. Freire, and P.C. Hoch for critical reviews of the manuscript. Special thanks are due to W. Punt and an anonymous reviewer for helpful comments. We are also grateful to the curators of the listed herbaria for loan of specimens and permission to sample pollen, and to Hugo Calvetti for the illustrations. This work was supported by the Consejo Nacional de Investigaciones Cient|¤¢cas y Te¤cnicas (CONICET), Argentina, and by the National Geographic Society (Grant 5776^96 to L.K.). Appendix A Specimens examined and voucher data. Subtribe Nassauviinae Burkartia lanigera (Hooker et Arnott) Crisci. Argentina. Prov. Chubut. Kreibohm 290 (LP). Prov. Santa Cruz. Ruiz Leal 25930 (LP). Cephalopappus sonchifolius Nees et Martius. Brasil. Edo. R|¤o de Janeiro. Cabrera 12243 (LP). Lophopappus berberidifolius Cabrera. Peru¤. Dpto. Huancavelica. Tovar 961 (LP). L. blakei Cabrera. Peru¤. Dpto. Cusco. Cabrera et Fabris 13462 (LP). L. cuneatus R.E. Fries. Argentina. Prov. Jujuy. Cabrera 9496, 15330, 19023 (LP), Cabrera et al., 15425, 21480, 21494 (LP). Meyer et Bianchi 33175 (LP). L. foliosus Rusby. Argentina. Prov. San Juan. Cabrera 25270 (LP). L. peruvianus Cabrera. Peru¤. Dpto. Ancash. Cerrate 1552 (LP). L. tarapacanus (Phil.) Cabrera. Chile. Prov. Tarapaca¤. Ricardi et Marticorena 25604 (LP). Macrachaenium gracile Hooker f. var. gracile. Argentina. Prov. Tierra del Fuego. Correa et Pe¤rez Moreau 1978 (LP). Oxyphyllum ulicinum Philippi. Chile. Prov. Antofagasta. Zo«llner 3892 (LP). Proustia cuneifolia D. Don var. cuneifolia. Argentina. Prov. Jujuy. Cabrera 12009 (LP), Cabrera et al., 13322 (LP), Okada 3036 (LP). P. cuneifolia D. Don var. mendocina (Philippi) Ariza. Argentina. Prov. Mendoza. Dawson 1085 (LP). 245 P. cuneifolia D. Don var. mollis (Kuntze) Cabrera. Argentina. Prov. Jujuy. Cabrera et Fabris 21042 (LP). Bolivia. Dpto. Cochabamba. Herzog 2021 (LP). P. ilicifolia Hooker et Arnott. var. baccharoides (D. Don) Fabris. Chile. Prov. Aconcagua. Garaventa 332 (LP). P. ilicifolia Hooker et Arnott var. ilicifolia. Chile. Prov. Coquimbo. Ricardi 5553 (LP). Argentina. Prov. La Rioja. Hunziker 1821 (LP). P. pyrifolia de Candolle var. pyrifolia. Chile. Prov. Caut|¤n. Friedrich s.n. (LP 071038). Subtribe Gochnatiinae sensu Cabrera (1977) Ainsliaea acerifolia Schultz var. subapoda Nakai. Japan. Honshu, Pref. Miyagi. Bou¡ord 19871 (CM). A. latifolia (D. Don) Schultz var. latifolia. China. Prov. Yunnan. Handel et Mazzetti 8501 (W). A. pertyoides Franchet var. pertyoides. China. Prov. Yunnan. Delavay s.n. (UC). A. okinawensis Hayata. Japan. Ryukyu. Pref. Okinawa. Murata 17066 (AAU). A. spanocephala Y.C.Tseng. Thailand. Loei. Nuijomdha et Vidal 438 (C); Smithland 077323 (BKF). Chimantaea cinerea (Gleason et S.F. Blake) Maguire, Steyermark et Wurdack subsp. cinerea. Venezuela. Edo. Bol|¤var. Steyermark et al., 116028 (MO). Cnicothamnus lorentzii Grisebach. Argentina. Prov. Salta. Maldonado 408 (LP). Erythrocephalum zambesianum Oliver et Hiern. Tanzania. Lovett et Congdon 1097 (MO). Hecastocleis shockleyi A. Gray. U.S.A. Nevada State. Train 3973 (LP). Hyalis argentea Hooker et Arnott. Argentina. Prov. Buenos Aires. Pertusi 259 (LP). Lycoseris trinervis (D. Don) S.F. Blake. Peru¤. Dpto. Chanchamayo. Diers 1249 (LP). Stenopadus talaumifolius S.F. Blake. Venezuela. Territorio Federal Amazonas. Liesner 18346 (MO). Sti¡tia uni£ora Ducke. Brasil. Edo. Amazonas. Ducke s.n. (LP). Wunderlichia mirabilis Riedel ex Baker. Brasil. Edo. Minas Gerais. Hatschbach 26992 (LP). Subtribe Mutisiinae sensu Cabrera (1977) Gongylolepis huachamacari Maguire. Venezuela. 246 M.C. Teller|¤a et al. / Review of Palaeobotany and Palynology 123 (2003) 237^246 Dpto. R|¤o Negro. Liesner et Delascio 21960 (MO). Hyaloseris cinerea (Grisebach) Grisebach var. tomentella Grisebach. Argentina. Prov. Co¤rdoba. Nicora 1589 (LP). Lulia nervosa (Lessing) Zardini. Brasil. Edo. Santa Catarina. Smith et Klein 1^576 (LP). Mutisia kurtzii R.E. Fries var. kurtzii. Argentina. Prov. Jujuy. Budin s.n. (LP). Mutisia sinuata Cavanilles. Argentina. Prov. Mendoza. Pe¤rez Moreau 12751 (LP). References Bremer, K., 1994. 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