A review of the genus Critoniopsis in Ecuador (Vernonieae: Asteraceae) Author(s): Xavier Haro-Carrión and Harold Robinson Source: Proceedings of the Biological Society of Washington, 121(1):1-18. 2008. Published By: Biological Society of Washington DOI: http://dx.doi.org/10.2988/07-18.1 URL: http://www.bioone.org/doi/full/10.2988/07-18.1 BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/ terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 121(1):1–18. 2008. A review of the genus Critoniopsis in Ecuador (Vernonieae: Asteraceae) Xavier Haro-Carrión and Harold Robinson* (XH-C) Herbario QCA, Escuela de Ciencias Biológicas, Pontificia Universidad Católica del Ecuador, Avenida 12 de Octubre 1076 y Roca, Apartado 17-01-2184, Quito, Ecuador, e-mail: xavierhc@gmail.com; (HR) Department of Botany, National Museum of Natural History, MRC-166, P.O. Box 37012, Smithsonian Institution, Washington, D.C. 20013-7012, e-mail: robinsoh@si.edu Abstract.—The genus Critoniopsis is found to have 18 species in Ecuador. The treatment includes the removal of a previously cited species for the country, C. bogotana, the reduction of C. sevillana to a variety of C. floribunda, and the description of three new species: Critoniopsis cerulosa, C. persetosa and C. zamorensis. A key for identification of the currently credited species is provided and notes on the distribution and taxonomic problems are presented. The genus Critoniopsis Sch. Bip., typified by C. lindenii Sch. Bip. from Colombia, was often treated as a subgenus of Vernonia Schreb. (Bentham 1873, Cuatrecasas 1956) but has recently been restored to generic status (Robinson 1980, 1993). The genus is a member of neotropical Vernonieae subtribe Piptocarphinae, notable for the easily deciduous inner involucral bracts of the flowering head. The species of the restricted concept of the genus, excluding Eremosis (DC.) Gleason and Tephrothamnus Sch. Bip. (Robinson 2007), occur mostly in the Andes from Colombia south to Bolivia, with a few species in southern Brazil. The genus is composed of shrubs or trees to 13–15 m tall with stems and leaf undersurfaces usually densely pilosulous to tomentose, rarely nearly glabrous. The leaves are always simple, alternate or opposite. There are 1–3 to over 20 florets in the heads. The related genus Piptocarpha R. Br., which consists of mostly scandent plants, differs most importantly by having tails on the anther thecae that are sclerified. * Corresponding author. The first reasonably complete treatment of Critoniopsis for Ecuador was done by Cuatrecasas (1956) as part of his study of mostly Colombian and Ecuadorian species. Of these species, ten were from Ecuador. New species were described separately in subsequent years raising the number to 17 in the last listing done by Robinson in the Catalogue of Vascular Plants for Ecuador (Robinson in Jørgensen & León-Yánez 1999). As is the case with many taxonomic groups of trees and shrubs, the members of the genus are often poorly represented in herbaria, and most of the species outside of the Quito area are represented by few specimens. Many of the species have a superficially similar appearance, although distinctions are obvious once the significant characters are recognized. The new examination of Critoniopsis by the present authors has established which characters are significant, and has greatly clarified species concepts. In the process, the study has revealed three previously undescribed species. Among the characters proving most useful in Critoniopsis are the prominence or lack of prominence of venation on the 2 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON abaxial surface of the leaf, a character used sparingly by Cuatrecasas (1956) in his study. Even more helpful are details of the trichomes. The trichomes are most often irregularly branched to stellate but are also sometimes simple and vermiform, spurred at the base, peltate, or gobletformed as seen in SEM micrographs. The goblet-shaped trichomes can be seen in the Colombian type species of the genus, C. lindenii Sch. Bip. and three species from Ecuador (C. sodiroi, C. tungurahuae, C. yambonensis, Figs. 9E, F, 10B–D). Trichomes that are mostly simple but vermiform are well developed in two Ecuadorian species (C. huairacajana, C. suaveolens, Figs. 7E, F, 10A). Trichomes that seem simple but have spurred bases can be seen in two Ecuadorian species (C. cotopaxensis, C. zamorensis, Figs. 5A–C; 10E, F), and the most nearly glabrous form of pubescence in the genus can be seen in the Ecuadorian, Peruvian and Bolivian C. boliviana (Britton) H. Rob. (Fig. 4A, B). A species that seems glabrous abaxially under the dissecting microscope, C. cerulosa, actually has compacted mass of trichomes embedded in wax (Fig. 4C–F). The Ecuadorian species with the most obvious simple trichomes, C. harlingii (H. Rob.) H. Rob. (Fig. 7C, D), may eventually be excluded from the genus. This revision of Critoniopsis in Ecuador seems to have resolved problems that have appeared up to the present, but due to incomplete collecting of the genus, new additions or modifications might be expected in the future. The occurrence within Ecuador and adjacent territories in neighboring countries might change with more data. The species of the genus in adjacent Peru are still comparatively poorly known. Also, the altitudinal ranges are cited from specimens, but citations are scarce in some cases. One species from the list of species cited in the Catalogue of Vascular Plants for Ecuador (Robinson in Jørgensen & León- Yánez 1999), Critoniopsis bogotana (Cuatrec.) H. Rob. is withdrawn. The species was cited for the country on the basis of a specimen (Palacios & Freire 4885) that has been re-identified as C. floribunda. Consequently, we find no evidence that C. bogotana, well distributed in central and northern Colombia, is present in Ecuador. Three new species are described and one of the previously recognized species is reduced to a variety. The list of the 18 species presently accepted in Critoniopsis in Ecuador can be distinguished by the following key. Key to Species of Critoniopsis 1a. Leaves opposite or mostly opposite 2a 1b. Leaves alternate . . . . . . . . . . . . . . . . 6a 2a. Leaves sessile; heads with ca. 20 florets; involucral bracts with prominent scarious margins . . . C. harlingii 2b. Leaves petiolate; heads with 12 or fewer florets; involucral bracts without prominent scarious margins . . 3a 3a. Leaf blades usually 13–24 cm long; densely villous or tomentose, hairs about 0.55–1.00 mm long on the undersurface and on the costa on the upper surface; stems lanate, and with abundant glands, ca. 0.15 mm long . . . . . . . . . . . . . . . . . . C. palaciosii 3b. Leaf blades mostly 6–13 cm long; lepidote with scales ca. 120 mm long on the costa and on the undersurface; stems with peltate scales, occasionally with glands 20 mm long or less . . . . . . . . . . . . . . . . . . . . . . . . . . . 4a 4a. Leaf margin slightly dentate towards the apex; tertiary nerves weakly prominulous; ca. 4 florets in the head; stellate hairs concave with a goblet shape . . . . . . . . . . . . . C. sodiroi 4b. Leaf margin entire; tertiary nerves strongly prominulous and reticulated; ca. 9–12 florets in the head; stellate hairs not concave and lacking goblet shape . . . . . . . . . . . . . . . . . . . . . . . . . 5a 5a. Leaf bases acute or subacute with secondary veins evenly spaced; inner involucral bracts 5.0–5.5 mm long, VOLUME 121, NUMBER 1 pilose on the outer surface with simple hairs up to 0.3 mm long or glabrous . . . . . . . . . . . . . . . . . C. dorrii 5b. Leaf bases rounded to subtruncate with lower secondary veins congested; inner involucral bracts less than 4.0 mm long, densely pilose on outer surface with simple hairs up to 0.6 mm length . . . . . . . . . C. persetosa 6a. Leaf abaxial surface with a glabrous appearance, without evident trichomes . . . . . . . . . . . . . . . . . . . . . . . . . . 7a 6b. Leaf abaxial surface with evident pubescence, with stellate or gobletshaped hairs . . . . . . . . . . . . . . . . . . . 8a 7a. Leaf surfaces concolorous, occasionally with few simple hairs, 30 mm long or less, scattered on the undersurface; petiole commonly with very small lateral lobes on petiole; less than 6 florets in the head . . . . . . . C. boliviana 7b. Leaf surfaces bicolorous or occasionally concolorous; undersurface with mat of hairs embedded in wax in a homogeneous layer; petiole without lateral lobes; more than 9 florets in the head . . . . . . . . . . . . . . . C. cerulosa 8a. Leaves rugose on the upper surface . . 9a 8b. Leaves essentially smooth on the upper surface . . . . . . . . . . . . . . . . . 10a 9a. Leaves oblong to elliptic, blade 4–7 cm long, 0.8–2.4 cm wide, typically in ratio 5:1 to 3:1 length-width; apex acuminate or occasionally obtuse . . C. huairacajana 9b. Leaves essentially elliptic, blade 6– 16 cm long, 3.0–5.2 cm wide, typically in ratio 3:1 to 2:1 length-width; apex acute to acuminate . . . . . C. suaveolens 10a. Leaves and stems villous or lanate; long hairs on the undersurface of the leaf and much of the stem, 350– 1000 mm long, simple appearing but with spurs at the base, porrect stellate type . . . . . . . . . . . . . . . . . 11a 10b. Leaf undersurface lepidote or tomentellous, stem with no evident pubescense; hairs stellate, not porrect, 350 mm or less in length . . 12a 11a. Florets in the head fewer than 10 . . . . . . . . . . . . . . . C. cotopaxensis 11b. Florets 19 or 20 in the head . . . . . . . . . . . . . . . . . . . . . . C. zamorensis 12a. Florets in the head 7 or fewer . . . 13a 3 12b. Florets in the head 8 or more . . . 17a 13a. Leaf texture papyraceous; achene intercostae with small straight setulae; capitulum with conspicuous pedicel . . . . . . . . . . . . C. yamboyensis 13b. Leaf texture coriaceous or subcoriaceous; achene sides and intercostae glabrous; capitulum sessile or subsessile . . . . . . . . . . . . . . . . . . . 14a 14a. Pubescence and venation of leaf undersurface appearing flat, with tertiary veins very weakly prominulous; scales on the leaf undersurface peltate . . . . . . . . . . . . . C. tungurahuae 14b. Venation prominent on leaf undersurface, with tertiary veins strongly prominulous, forming an irregular surface with pubescence; trichomes not evidently peltate . . . . . . . . . . 15a 15a. Leaves broadly ovate; stellate hairs, ca. 450 mm long, scattered on the stem; pappus darkened; heads in dense glomerules in the inflorescence . . . . . . . . . . . . . . . . . C. occidentalis 15b. Leaves essentially elliptic; peltate scales on the stem surface; pappus white; heads loosely arranged on short cymose branches in the inflorescence . . . . . . . . . . . . . . . . . . 16a 16a. Leaf undersurface tomentellous with hairs stellate multiangulate; veins prominulous to the quaternary venation with evident strong reticulation . . . . . . . . . . C. floribunda 16b. Leaf undersurface lepidote with triangular scale-like hairs; veins prominulous to tertiary venation reticulated or weakly reticulated . . . . . . . . . . . . . . . . . . . . . C. elbertiana 17a. Bracteoles present on peduncle below base of the capitulum, involucral bracts in 7 series; leaf blade elliptic to oblong, apex acute, undersurface of leaf blade tomentellous with broad hairs . . C. jaramilloi 17b. Bracteoles absent at the base of the capitulum, involucral bracts in 4–5 series; leaf blade elliptic, apex acute to acuminate, undersurface of leaf blade pubescent with slender or broad hairs . . . . . . . . . . . . . . . . . 18a 18a. Secondary nerves more congested at the base of the leaf, becoming more 4 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON separated towards the middle of the leaf; veins of leaf undersurface prominulous to the secondary order of venation and weakly reticulated . . . . . . . . . . . . . . . . . C. pycnantha 18b. Secondary leaf nerves equally spaced, veins of leaf undersurface prominulous to the tertiary or quaternary order of venation and strongly reticulated . . . . . . . . . . . . . . . . . . . C. floribunda var. sevillana Critoniopsis boliviana (Britton) H. Rob. Fig. 4A, B Proc. Biol. Soc. Wash. 106(3):609–610. 1993. Vernonia boliviana Britton, Bull. Torrey Bot. Club. 18:332. 1891. (Bolivia. La Paz: Yungas, 4000 ft, Rusby 1729, NY). Vernonia paucisquamata Rusby, Bull. New York Bot. Gard. 4:376. 1907. (Bolivia. La Paz: Yungas, Coroico, 4 Sep 1894, Bang 2400, NY). Baccharis dudleyi Cuatrec., Phytologia 49:70. 1981. (Peru. Huanuco: Southeastern slope of Rı́o Llulla Pichis watershed on the ascent of Cerros del Sira, about halfway between Laguna and Peligrosso, 1450 m, 23 Jul 1969, T.R. Dudley 13183, NA). Distribution.—Zamora (Peru, Bolivia). Altitudinal range.—1000–2300 m. The species was collected in ZamoraChinchipe in 1990 (Neill & Palacios 9523, MO, US) and two other specimens (Parque Nacional Podocarpus, Palacios & Ticado 13056, MO, QCNE, US; Cordillera del Cóndor, Miranda et al. 157, MO, US) confirm its presence in this province. Some of the Ecuadorian specimens do not show the lobed petiole that is generally characteristic of the species. Critoniopsis cerulosa Haro-C. & H. Rob., sp. nov. Figs. 1, 4C–F Type locality.—Ecuador. Tungurahua: Baños, Zona de amortiguamiento del Parque Nacional Llanganates. Rı́o Verde, cañón a las cascadas de Machay, bosque húmedo montano bajo, vegetación secundaria, suelo aluvial, 01u239S, 78u179W, 1630 m, árbol de 20 m, envés de las hojas blanquecino, capı́tulos blancos, 1999, Vargas, Narváez & Sánchez 3628 (holotype US; isotype MO) (Fig. 1). A speciebus congeneribus omnibus in foliis abaxialiter confertim dense obscure pubescentes sed in ceris inclusis et ut videtur glabris distincta. Trees to 30 m tall; Leaves alternate, petioles slender, 2.0–3.5 cm long; blades narrowly to broadly elliptic, 10–21 cm long, 4–9 cm wide, base obtuse to narrowly acute, slightly acuminate at petiole, margins entire, apex short acuminate, secondary veins 7–9 on each side, rather evenly spaced, spreading from midrib at 60–75u angles, upper surface glabrous on lamina and veins, with slightly prominulous reticulum of tertiary and quaternary veinlets, undersurface of the leaf, densely puberulous but trichomes embedded in wax and surface appearing glabrous. Inflorescence a terminal panicle of erectspreading branches and seriate-cymose branchlets, surfaces minutely puberulous and gland-dotted. Heads sessile to subsessile, pedicels to 2 mm long; involucres to 8 mm long, ca. 5 mm wide; bracts ca. 30 in 5–6 series, glabrous outside, broadly ovate to oblong-ovate in many crowded series, 0.5–3.0 mm long, 1.0–1.2 mm wide, persistent, basal bracts spreading, inner bracts erect, rather deciduous, narrowly lanceolate to linear-lanceolate, 5–7 mm long, 0.9–1.1 mm wide near middle, narrowed at base with narrow basal wings, apices obtuse to narrowly rounded. Florets 8–10 in a head; corollas submature, whitish at anthesis (?), 5 mm or more long; anther thecae ca. 1.5 mm long; apical appendages ca. 0.35 mm long. Achenes usually 8–10-ribbed, ca. 3 mm long, glabrous, with scattered idioblasts; pappus of ca. 30 whitish capillary bristles ca. 5.5 mm long, slightly VOLUME 121, NUMBER 1 Fig. 1. 5 Critoniopsis cerulosa Haro-C. & H. Rob. (Vargas, Narváez & Sánchez 3628, holotype, US). broadened at tips; outer series of narrow squamae ca. 0.7 mm long. Paratypes.—Ecuador. Napo: Canto El Chaco, margen derecho del Rı́o Quijos, Finca ‘‘La Ave Brava’’ de Segundo Pacheco, bosque pluvial premontano, bosque primario, sobre suelos saturados, 00u129S, 77u399W, 1800–1900 m, 7–10 Sept 1990, Palacios 5369 (MO, US); Cantón Tena, Parque Nacional Llanganates, Via Salcedo – Tena, km 74, ribera del Rı́o Mulatos, bosque muy húmedo montano bajo roca metamórfica, 01u019S, 78u129W, 2020 m, 10 Sep 1998, Vargas, Narváez & Orellana 2178 (MO, US). Distribution.—Napo. Altitudinal range.—1650–2000 m. The collections are described variously as trees 20–30 m tall with trunks to 50 cm DAP, leaves coriaceous with the underside whitish, ‘‘plateado’’; inflorescences white. 6 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Critoniopsis persetosa Haro-C. & H. Rob. (Jørgensen, Ulloa, Gavilanes, Mena & Suarez L. 92074, holotype, US). Critoniopsis cerulosa is superficially similar to various other species, and specimens have been previously named as C. elbertiana and C. pycnantha. Close examination of the undersurface of the leaves, however, reveals a distinctive glabrous appearance that is actually formed by a dense mat of hairs in a waxy coating. The pores that can be seen are spaces between the hairs, and the true stomata of the leaf surface seem to be surrounded by a waxy collar (Fig. 4F). Critoniopsis cotopaxensis H. Rob. Fig. 5A–C Proc. Biol. Soc. Wash. 106(3):610–611. 1993 (Ecuador. Cotopaxi: Carretera Latacunga-Pilaló-Quevado, 5–15 km al este de Pilaló, 00u539S, 79u019W, 2700–3350 m, 22 May 1988, Cerón, Neill & Palacios 3804, US). Distribution.—Cotopaxi Altitudinal range.—2700–3350 m. VOLUME 121, NUMBER 1 Fig. 3. 7 Critonopsis zamorensis Haro-C. & H. Rob. (Homeier 589, holotype, MO). Known only from the type collection. The species is distinguished from the related C. palaciosii which has alternate leaves and more vermiform trichomes abaxially. Critoniopsis dorrii H. Rob Fig. 5D, E Proc. Biol. Soc. Wash. 106(3):612–613. 1993. (Ecuador. Azuay: Cuenca-Solda- 8 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Critoniopsis. SEM micrographs of abaxial leaf surfaces. A, B, C. boliviana (Solomon 9038), surface mostly glabrous with sparse, appressed, scale-like trichomes; C–F, C. cerulosa (Vargas et al. 3628, holotype US); C, D, Surface showing smooth covering of wax-embedded trichomes; D, Surface showing scattered pores in compacted mass of trichomes; E, Surface showing broken areas in waxy surface; F, Surface with cover of trichomes partially removed showing bases of trichomes and some stomata with waxy collars. dos road (following N bank of Rı́o Yanuncay), 19–20 km W of San Joaquin, 2u559S, 79u059W, 22 Jun 1989, Dorr & Valdespino 6404, US). Distribution.—Azuay, Loja. Altitudinal range.—2400–3200 m. The original range was restricted to Azuay (Dorr & Valdespino 6404) but another specimen has been seen from adjacent northern Loja (Jørgensen & Ulloa 92007). The strongly prominulous dense reticulation of veinlets on the under surface of the leaf is the most useful character to distinguish C. dorrii from the similarly opposite-leaved C. sodiroi. The species also differs from C. sodiroi by its stellate rather than goblet-formed trichomes. Critoniopsis elbertiana (Cuatrec.) H. Rob. Figs. 5F, 6A, B Phytologia 46:440. 1980. VOLUME 121, NUMBER 1 9 Fig. 5. Critoniopsis. SEM micrographs of trichomes on stems and leaves and involucral bract. A–C, C. cotopaxensis (Cerón et al. 3804, holotype US), showing porrect trichomes with spurred bases, and intermixed smaller stellate trichomes; C, Trichomes on stem with enlarged tips; D, E, C. dorrii (Harling & Andersson 12585, US), showing stellate trichomes; F, C. elbertiana (Little 8688, holotype US), inner involucral bract showing narrowed base. Vernonia elbertiana Cuatrec., Bot. Jahrb. Syst. 77:68. 1956. (Colombia. Putumayo: Alta cuenca del rı́o Putumayo, SE of Gigante, 10,000 ft, 20 Sep 1944, Little 8688, US). Distribution.—Napo, Tungurahua, Zamora (Colombia). Altitudinal range.—1800–2550 m. The species was noted by Cuatrecasas (1956) for the ‘‘auriculate’’ inner involucral bracts (Fig. 5F), but no other specimens besides the type show as extreme a devel- opment of this character. In this treatment we have extended the concept of this species to include specimens that match the number of florets in the heads, venation pattern and pubescence, but have smaller auricles on the inner involucral bracts. Critoniopsis floribunda (Kunth in H.B.K.) H. Rob. Figs. 6C–F, 7A, B Phytologia 46:440. 1980. 10 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 6. Critoniopsis. SEM micrographs of venation and trichomes on abaxial leaf surface. A, B, C. elbertiana (Little 8688, holotype US): abaxial leaf surface showing irregularly branched trichomes; C–E, C. floribunda var. sevillana (Camp 4344, US). C, Abaxial surface showing strongly prominulous tertiary and quaternary veins; D, E, Surface showing scarcely branched vermiform trichomes and intermixed stellate trichomes; F, C. floribunda var. floribunda (Dalessandro 657, US). Surface showing scarcely branched vermiform trichomes and intermixed stellate trichomes. Vernonia floribunda Kunth in H.B.K., Nov. Gen. Sp., ed. fol. 4:30. 1818. (Peru? 1802, Humboldt & Bonpland s.n. P-HBK) (IDC 6209:91:3:5). Vernonia affina Kunth in H.B.K., Nov. Gen. Sp., ed. fol. 4:30. 1818. (Peru? 1802, Humboldt & Bompland s.n. PHBK) (IDC 6209:91:3:7). Distribution.—Napo, Loja, Zamora. Altitudinal range.—2250–2900 m. The characteristic venation pattern of the species can be seen in photographs of the type specimen in the Humboldt and Bonpland collections in Paris. The abaxial leaf venation shows a high order of relief to the fourth order of veins that seems unique to the species. Critoniopsis floribunda var. sevillana (Cuatrec.) Haro-C. & H. Rob., comb. nov. Figs. 6C–E, 7A, B Vernonia sevillana Cuatrec., Bot. Jahrb. Syst. 77:78. 1956. (Ecuador. Azuay: VOLUME 121, NUMBER 1 11 Fig. 7. SEM micrographs of Critoniopsis trichomes. A, B, C. floribunda var. sevillana (Camp 4344, US), showing stellate trichomes; C, D, C. harlingii (Harling & Andersson 14408, holotype GB), showing simple trichomes; E, F, C. huairacajana (Prieto P-76, US), showing weakly branching vermiform trichomes. Eastern Cordillera, 1–8 km N of Sevilla de Oro, 8000–9000 ft, Jul–Aug 1945, Camp E-4507, NY) Critoniopsis sevillana (Cuatrec.) H. Rob., Phytologia 46:441. 1980. Distribution.—Azuay. Altitudinal range.—2400–2750 m. Vernonia sevillana was described by Cuatrecasas (1956), distinguished primarily by the number of florets in the heads. The number of florets in the type series certainly is greater than in other material of Critoniopsis floribunda, but the full range has no evident discontinuity on which a species could be reliably based. At the same time, the undersurfaces of the leaves have the same extreme relief of third and fourth order of veins seen in detailed photographs of the type specimen of C. floribunda. The variety is known only from the type and a paratype, both collected by Camp near Sevilla de Oro in 1945. Critoniopsis harlingii (H. Rob.) H. Rob. Fig. 7C, D Proc. Biol. Soc. Wash. 106(3):614. 1993. 12 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Vernonia harlingii H. Rob., Phytologia 44:66. 1979. (Ecuador. El Oro: Road Pasajo-Santa Isabel-Girón, valley of Rı́o Jubones, ca. 1600 m, 7 May 1974, Harling & Andersson 14408, GB). Distribution.—El Oro. Altitudinal range.—,1600 m. The species is known only from the type collection. It is very distinct from the other opposite leaved species by the blades being sessile, having 20 florets in the head and broad, whitish, dissected margins on the involucral bracts. The hairs of the leaves are simple, unlike those of other Ecuadorian species in Critoniopsis. The completely simple hairs of the leaves place the species outside of the most recent more restricted concept of the genus Critoniopsis (Robinson in Kadereit & Jeffrey 2007). Actual placement requires further study. Critoniopsis huairacajana (Hieron.) H. Rob. Fig. 7E, F Phytologia 46:440. 1980. Vernonia huairacajana Hieron., Bot. Jahrb. Syst. 19:43. 1894. (Ecuador. Azuay: Cuenca, Páramo Huaira-Caja, 2600–3000 m, 1880, Lehmann 4692, B destroyed, F, US). Distribution.—Azuay, Cañar. Altitudinal range.—2500–3500 m. The narrowly oblong leaves with roughened upper surfaces are distinctive. Critoniopsis jaramilloi Pruski & H. Rob. Fig. 8A, B Phytologia 78(5):338. 1995. (Ecuador: Azuay. Carretera Cuenca-San Joaquı́n-Angas, S de Cuenca entre Bayán y Heda, 30 Jul 1983, Jaramillo & Winnerskjold 5389, US). Distribution.—Azuay. Altitudinal range.—2000–2500 m. The species is known only from the type. The many rows of involucral bracts and the bracteoles on the peduncle are characteristic of the species. Critoniopsis occidentalis (Cuatrec.) H. Rob. Fig. 8C, D Phytologia 46:440. 1980. Vernonia occidentalis Cuatrec., Bot. Jahrb. Syst. 77:73. 1956. (Colombia. El Valle: Hoya de rı́o Digua, Quebrada del rı́o San Juan arriba Queremal: Las Colonias, 1950–2000 m, 10 Mar 1947, Cuatrecasas 23914, F). Distribution.—Carchi, Cotopaxi, Pichincha, Zamora (Colombia). Altitudinal range.—1100–2300 m. The species was described from southwestern Colombia and has been commonly collected in northern Ecuador. The species is a tree with distinctive broad, acuminate-tipped leaves and sordid pappus bristles. Critoniopsis palaciosii H. Rob. Fig. 8E Proc. Biol. Soc. Wash. 106(3):617–618. 1993. (Ecuador. Imbabura: Cantón Cotacachi, carretera Catacachi-Apuela, sitio Tabla Chuap, a 1 km de Hacienda La Providencia, 00u259N, 78u259W, 3100 m, 4 Apr 1990, Palacios & Iguago 4867, US). Distribution.—Imbabura, Cotopaxi. Altitudinal range.—2400–3100 m. The species is notable for the large, ovate, opposite leaves and the villous or woolly pubescence of porrect, basally spurred trichomes on the under surface of the leaves. Critoniopsis persetosa X. Haro-C. & H. Rob., sp. nov. Figs. 2, 8F, 9A, B Type locality.—Ecuador. Loja: Loja: Las Palmas, Cerro El Tambo, just south VOLUME 121, NUMBER 1 13 Fig. 8. SEM micrographs of Critoniopsis. A–E, Trichomes on abaxial leaf surfaces. A, B, C. jaramilloi (Jaramillo & Winnerskjold 5389, holotype US), showing irregularly branched trichomes; C, D, C. occidentalis (Webster & Castro 30246, US), showing irregularly branched stellate trichomes; E, C. palaciosii, showing vermiform trichomes with spurred bases; F, C. persetosa (Jørgensen et al. 92074, holotype US), inner involucral bract showing dense pubescence. of Cerro Villonaco. Roadside and secondary forest. 04u049S, 79u149W, 2570– 3020 m, tree 6 m, inflorescence white, 23 Jul 1990, Jørgensen, Ulloa, Gavilanes, Mena & Suarez L. 92074 (holotype US; isotype AAU, QCA) (Fig. 2). A C. dorrii in foliis plerumque oppositis et in bracteis interioribus involucri extus dense pilosis valde affinis sed in laminis foliorum base subtruncatis et in nervis secundariis basilaribus laminarum congestis et in bracteis involucri brevioribus distincta. Trees to 8 m tall. Leaves opposite to subopposite or alternate; petioles 1.5– 2.5 cm long; blades oblong-ovate, 8– 11 cm long, 4.5–8.0 cm wide, bases rounded to subtruncate, abruptly narrowed at petiole, margins entire, apex obtuse to subacute, upper surface green, glabrous on lamina and veins, undersurface gray with dense tomentellum of small, narrow, irregularly stellate scales, secondary veins 8–10 on each side, closely set, 3–7 mm apart and spreading at 70– 80u angles in basal 40% of blade, mostly 14 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 9. SEM micrographs of Critoniopsis trichomes. A, C. persetosa (holotype), trichomes on outer surface of inner involucral bract showing spurred bases; C, D, C. pycnantha (Ollgaard 58006, US), showing irregularly branching trichomes on abaxial leaf surface; E, F, C. sodiroi (Moran & Vaca et al. 38, US), showing goblet-shaped trichomes on abaxial leaf surface. 8–12 mm apart and spreading at 50–60u angles in distal 60%; tertiary and quaternary veinlets weakly to scarcely prominulous. Inflorescence a dense, terminal, corymbiform to subpyramidal panicle; branches spreading at 60–80u angles, gray with tomentellum of small hair-like scales. Heads ca. 9 mm high; involucre broadly campanulate, 3.5–4.0 mm high, 4.0– 4.5 mm wide, only as long as bodies of mature achenes; bracts ca. 25 in ca. 4 series, not spreading, broadly ovate to oblong, 1.0–3.0 mm long, 1.0–1.2 mm wide, apices obtuse, lower bracts mostly glabrous, inner bracts rather deciduous, rather glabrous at tips but with dense mass of hairs on middle of outside surface. Florets 8–11 in head; corollas brown with age, whitish at anthesis (?) ca. 5.5 mm long, glabrous, basal tube ca. 3.5 mm long, throat ca. 0.7 mm long, lobes ca. 2 mm long, strongly coiled at maturity; anther thecae ca. 1.7 mm long; apical appendage ca. 0.4 mm long, 0.2 mm wide, glabrous; style base with node; style branches with round-tipped sweeping hairs. Achenes 3.8–4.0 mm long, glabrous, with idioblasts on surface; VOLUME 121, NUMBER 1 15 pappus whitish, of ca. 40 capillary bristles slightly broadened distally, ca. 4 mm long, with outer series of narrow squamae ca. 0.7 mm long. Paratype.—Ecuador. Loja: Loja Canton. Base del Villonaco (Pan de Azúcar); Bosque Seco Premontano, 03u599S, 79u179W, 3000 m, 7 Sep 1990; Cerón & Ocampo 11911 (MO, QCNE, US). Distribution.—Loja. Altitudinal range.—2550–3000 m. The hairs on the inner involucral bracts (Figs. 8F, 9A), a feature shared to a lesser extent with C. dorrii, are unusual in being concentrated on the part of the bracts overlain by the outer bracts. Hairs on inner involucral bracts in most Asteraceae are found only on the more exposed apical parts. The leaf blades differ from those of the related C. dorrii by their abrupt bases with crowded lower secondary veins. A further difference from C. dorrii is the shorter involucral bracts. The more northern species of Azuay and northernmost Loja has longer, more acute inner involucral bracts, distinctly longer than the bodies of the mature achenes. The result is that the involucres of C. dorrii look distinctly longer than wide, while those of the new species are scarcely longer than wide. spaced basal secondary veins of the leaves. The species is known from a number of recent collections. Ecuador: Loja: Parque Nacional Podocarpus, S of Loja, 04u059S, 79u109W, 2800–2950 m, 23 Feb 1985, Øllgaard, Laegaard, Thomsen, Korning & Illum 58006 (AAU, QCNE, US); Parque Nacional Podocarpus, E of Nudo de Cajanuma, just N of ‘‘Centro de Informacion’’, 04u059S, 79u109W, 2900 m, 28 Oct 1988, Madsen 75524 (AAU, QCNE, US). Morona-Santiago: Along road GualaceoEl Limon, 2500 m, 15 Aug 1989, van der Werff & Gudiño 11104 (MO, US). Zamora-Chinchipe: Estación Cientifica San Francisco, 30 km E of Loja on road to Zamora, Quebrada El Milagro watershed, 03u589350S, 79u049140W, 2100 m, 20 Apr 2000, Neill et al. 12611 (MO, QCNE, US). Peru: Amazonas: Bongará, Tresleras, trocha Pomacochas-Leimebamba, 2200–2300 m, 18 Aug 1963, Ferreyra 15238 (US, USM). Piura: Huancabamba, Canchaque, entre ‘‘Chorro Blanco’’y ‘‘War War’’, 2000–2500 m, 18 Jan 1989, Dı́az, Pennington & Reynel 3184 (MO, US). Critoniopsis pycnantha (Benth.) H. Rob. Fig. 9C, D Phytologia 46:441. 1980. Vernonia pycnantha Benth., Pl. Hartw. 134. 1844. (Ecuador: Loja. Mountains of Paccha, May–June 1842?, Hartweg 754, K). Tephrothamnus pycnanthus (Benth.) Sch. Bip., Jahresber. Pollichia 20/21:433. 1863. Phytologia 69:105. 1990. Piptocarpha sodiroi Hieron. ex Sodiro, Bot. Jahrb. Syst. 29:2. 1900. (Ecuador, in silva andien & suband, Sodiro 1/7, B destroyed). Vernonia pichinchensis Cuatrec., Bot. Jahrb. Syst. 77:76. 1956 (Ecuador: Pichincha. Pululagua, 2800 ft, 6 Jun 1939, Asplund 6723, US). Critoniopsis pichinchensis (Cuatrec.) H. Rob., Phytologia 46:440. 1980. Distribution.—Morona-Santiago, Loja, Zamora-Chinchipe, Peru. Altitudinal range.—1500–3000 m. One of the distinctive features of the species, seen in photographs and scans of the type, is the usually more closely Distribution.—Bolivar, Chimborazo, Cotopaxi, Imbabura, Pichincha. Altitudinal range.—1600–3650 m. The consistently opposite leaves are very distinctive, and the goblet formed trichomes of the leaf undersurface resem- Critoniopsis sodiroi (Hieron) H. Rob. Fig. 9E, F 16 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 10. SEM micrographs of Critoniopsis trichomes on abaxial leaf surfaces. A, C. suaveolens (Zak & Jaramillo 2721, US), showing vermiform trichomes branching from base; B, C. tungurahuae (Benoist 4195, holotype P), showing goblet-shaped trichomes; C, D, C. yambonensis (Benoist 4497, holotype P), showing gobletshaped trichomes; E, F, C. zamorensis (Homeier 589, type MO), showing porrect trichomes with spurred bases. ble those of the Colombian type species of the genus, Critoniopsis lindenii Sch. Bip. Critoniopsis suaveolens (Kunth in H.B.K.) H. Rob. Fig. 10A Phytologia 46:441. 1980. Vernonia suaveolens Kunth in H.B.K., Nov. Gen. Sp., ed. fol. 4:30. 1818. (Ecuador [not Colombia as originally stated]: Azuay?. Ayavaca ad fluvium Cutaco, 1801, Humboldt & Bonpland s.n., P-HBK) (IDC 6209:91:3:5). Distribution.—Azuay, Bolivar, Chimborazo. Altitudinal range.—1700–2800 m. Similar to C. huairacajana in the general aspect and pubescence, but the elliptic leaf blades with acuminate apices distinguish it clearly. Recent collections include the following: Ecuador. Azuay: Cuenca Cantón, carretera Cuenca-Molleturo-Naranjal-Bajo Molleturo, 02u429180S, 79u279080W, 2000 m, 3 Aug 1996, Palacios & Cerón 13828 (MO, QCNE, US). Bolivar: Carretera VOLUME 121, NUMBER 1 Guaranda-San Pablo de Atenas-Chillanes, sector Sicoto, 01u509S, 79u059W, 2200–2450 m, 28 Aug 1987, Zak & Jaramillo 2533, 2538 (MO, US); carretera Chillanes-Tiquibuso, en el sector de San José de Guayabal, 01u559S, 79u059W, 2100 m, 3 Sep 1987, Zak & Jaramillo 2721 (MO, US); carretera ChillanesTambillo-Trgoloma, entre Bola de Oro y Panecillo, 01u559S, 79u059W, 2100– 2200 m, 5 Sep 1987, Zak & Jaramillo 2740 (MO, US). Bolivar-Chimborazo: Carretera Pallatanga-Yunguilla-Llimbe, a orillas de Rı́o Chimbo, 1720 m, 7 Sep 1987, Zak & Jaramillo 2802 (MO, US). Critoniopsis tungurahuae (Benoist) H. Rob. Fig. 10B Phytologia 46:441. 1980. Vernonia tungurahuae Benoist (Ecuador: Tungurahua. Baños, 3 Apr 1931, Benoist 4195, P). Distribution.—Tungurahua. Altitudinal range.—2500–3200 m. Only one more recent collection of the species has been seen in this study: Ecuador. Tungurahua: Cantón Baños, Rı́o Vascun Valley, northern slopes of Volcán Tungurahua, 01u269210S, 78u259580W, 2500–3200 m, 27 Apr 2003, Clark, Jost & Mailloux 7722 (US), a specimen that is almost a topotype. The distribution seems very restricted, but the species name has been applied tentatively to many additional specimens from adjacent areas during recent years. Many of the specimens are now recognized as being a variant of Critoniopsis elbertiana, but one very immature collection from low elevation in Lorocachi, Pastaza to which the name was applied, Jaramillo, Coello & Freire 30962 (AAU, QCNE, US), has been redetermined as Piptocarpha vasquezii H. Rob., previously known only from Peru. A primary characteristic of the species is the densely pubescent, flat appearing leaf undersurface with weakly prominu- 17 lous tertiary veins. At higher magnification, the goblet-shaped trichomes are distinctive. Critoniopsis yamboyensis (Benoist) H. Rob. Fig. 10C, D Proc. Biol. Soc. Wash. 106:626. 1993. Vernonia yamboyensis Benoist, Bull. Soc. Bot. France 83: 804. 1936 [1937] (Ecuador: Pichincha. Yamboya, ca. 2500 m, 3 Jul 1931, Benoist 4497, P). Distribution.—Pichincha. Altitudinal range.—2000–2500. The leaves are distinctively deltoidovate and thinly herbaceous. The presence of setulae on the achenes is also distinctive. Critoniopsis zamorensis X. Haro-C. & H. Rob., sp. nov. Figs. 3, 10E, F Type locality.—Ecuador. Zamora Chinchipe: Area of the Estación Cientifı́ca San Francisco (03u589S, 79u049W), road LojaZamora, ca. 30 km from Loja, montane tropical forest, 2050 m, tree. Homeier 589 (holotype MO) (Fig. 3). A C. cotopaxensis in pilis longis porrecti-stellatis simila sed in foliis longioribus base cuneatis et in floribus ca. 20 in capitulo differt. Tree; branches terete, densely velutinous with long brownish hairs. Leaves alternate, with velutinous petioles 2–3 cm long; blades narrowly obovate, mostly 15–22 cm long, 5.5–10.0 cm wide at widest part, base narrowly acute to cuneate, margins entire to remotely denticulate, apex short-acuminate; secondary veins ca. 10 on each side of primary vein, ascending at ca. 45u angles at base; upper surface rather glaucous-green, coriaceous, glabrous, except hirsute on primary and secondary veins; lower surface darker green, densely pilose with brownish hairs, hairs spurred at bases; main veins prominent below, veinlets of third and fourth order prominulous with shallow areoles. 18 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Inflorescence a pyramidal terminal panicle; branches alternate, brownish velutinous, spreading at 50–65u angles; branchlets often forked, becoming seriate-cymes of 5–8 sessile heads. Heads ca. 7 mm high; involucral bracts ca. 25, in ca. 5 series, short and triangular at base, densely puberulous outside, median bracts mostly persistent, widely spreading, oblong, ca. 4 mm long, ca. 2 mm wide, short-acute to obtuse apically, with numerous blackish hairs distally and near scarious margins, innermost bracts deciduous, lanceolate, ca. 5 mm long, narrowly acute, with many blackish hairs distally and marginally. Florets usually 19 in a head. Corollas from fragments in frass withered and brownish, white at anthesis (?) with lobes ca, 1.5 mm long, narrowly lanceolate, with numerous glandular dots distally; anther appendages oblong-ovate, ca. 0.4 mm long, 0.2 mm wide at base, glabrous. Achenes ca. 2 mm long, 4angled, glabrous; pappus with ca. 25 whitish capillary bristles to 3.5 mm long, slightly broadened in distal half, with series of numerous outer squamellae 0.5– 0.7 mm long. Distribution.—Zamora. Altitudinal range.—2050 m. The ovate to oblong leaves and the villous or woolly undersurface of the leaves with stellate or porrect basally spurred trichomes most closely resemble C. cotopaxensis. The number of florets in the head of around 19 in the species differs from the eight to nine florets found in C. cotopaxensis. The base of the leaf is also more acute in C.zamorensis. Acknowledgments We thank Marjorie Knowles for her assistance in many different aspects of this research, and Scott Whittaker for his assistance in the management of the SEM, and for the preparation of the SEM samples. The herbaria at Chicago (F), Kew (K), New York (NY), and Paris (P) are thanked for loans of material and scans of specimens. We thank the Smithsonian Latino Initiatives Fund for funding this project, and Mary Sangrey for directing the Research Training Program of the Smithsonian Institution under which this research was undertaken. We also thank Catherine F. Harris of The Office of Research and Training for making available a Short-Term Visitor Grant to allow completion of the project. Literature Cited Bentham, G. 1873. Ordo 88: Compositae. Pp. 163–533 in Genera Plantarum 2(1). Reeve & Co, London. Cuatrecasas, J. 1956. Neue Vernonia-Arten und Synopsis der andien Arten der Sektion Critoniopsis.—Botanische Jahrbücher für Systematik 77:52–84. Jørgensen, P. M. and S. León-Yánez (eds.). 1999. Catalogue of The Vascular Plants of Ecuador. Missouri Botanical Garden Press, St. Louis, 1181 pp. Robinson, H. 1980. Re-establishment of the genus Critoniopsis (Vernonieae: Asteraceae).—Phytologia 46:437–442. ———. 1993. A review of the genus Critoniopsis in Central and South America (Vernonieae: Asteraceae).—Proceedings of the Biological Society of Washington 106:606–627. ———. 2007[2006]. Tribe Vernonieae Cass. (1819). Pp. 165–192 in J. W. Kadereit and C. Jeffrey, eds., Families and Genera of Vascular Plants, Vol. VIII. Flowering Plants - Eudicots Asterales, 740 pp 131 illus. Part of series by Kubitzki, K., ed. Kubitzki’s Authoritative Encyclopedia of Vascular Plants. SpringerVerlag. Associate Editor: Carol Hotton