-.” PHYTOLOGIA #3 __ An international journal to expedite plant systematic, phytogeographical and ecological publication Vol. 70 March 1991 No. 3 CONTENTS 4 v G.L. NESOM, A new Bolivian species of A/oysia (Verbenaceae)...145 E.A. CHRISTENSON, The last Physingia Lindley (Orchidaceae)...147 D> A.H. LIOGIER, Novitates Antillanae. XVIucccsssscssssssesssssssssesessesee 149 B.L. TURNER, Recension of the Cronquistianthus group of Eupatorium (ASteraceae) ...........:..ccccssecceceecessscccecssesceeeceseeseessees 158 F.L. PEREZ, Prostrate growth in caulescent Andean rosettes of . Coespeletia (Asteraceae, Heliantheae)....................:ccceseeeteees 178 -G.L. NESOM, Taxonomic adjustments in Bouchea and Lippia (Verbenaceae) Of MEXICO ..........eccecceseescscseeescescescsecssesseenecenenseass 185 ed, Ee. LAFERRIERE, Range extensions in the Sierra Madre Occidental, Chihuahua, tS a een ate Sc eR tea 28 188 J.T. KARTESZ & K.N. GANDHI, Nomenclatural notes for the North AM@THGOP FIO Kay patra sonnetacn sea eters a Scccte slit nceiivcsiocnet aiselie 194 / M.A. HERSHKOVITZ, Taxonomic notes in Cistanthe, Calandrinia, and Talinunt{(POmMmulaCaCOae) <50..23.5.ciies cscs Moen vescovancvaseneagescodcccenes 209 BOOKS TEOCOIVO Ge rceretrie mach enlebaet ad ee crenaladtc dons osencdeds aleodoethtodauees 226 LIBRARY JUN 71 1931 Published by Michael J. Warno®EW YORK 185 Westridge Drive Huntsville, T WRACALSGARDEN PHYTOLOGIA is printed ona TA paper. PHYTOLOGIA (ISSN 00319430) is published monthly by Michael J. Warnock, 185 Westridge Drive, Huntsville, TX 77340-8916. Second Class postage at Huntsville, TX. Copyright ©1990 by PHYTOLOGIA. Domestic individual subscription (6 issues): $18.00. Domestic institutional subscription (6 issues): $20.00. Foreign and/or airmail postage extra. Single copy sales: Current issue and back issues volume 67 to present, $3.50; Back issues (previous to volume 67), $3.00 (add $.50 per copy postage and handling US [$1.00 per copy foreign]}). Back issue sales by volume: $17.00 per yolume 42-66 (not all available as complete volumes); $21.00 per volume 67-present; add $2.00 per volume postage US ($4.00 per volume foreign). POSTMASTER: darts changes to Phytologia, 185 Westridge Drive,Huntsville, TX 40-8916. Phytologia (March 1991) 70(3):145-146. A NEW BOLIVIAN SPECIES OF ALOYSIA (VERBENACEAE) Guy L. Nesom Department of Botany, University of Texas, Austin, Texas 78713 U.S.A. ABSTRACT A new species of Aloysia, A. arcuifolia, is described from Bolivia in South America. KEY WORDS: Aloysia, Verbenaceae, Bolivia, South America Aloysia arcuifolia Nesom, sp. nov. TYPE: BOLIVIA. Depto. Potosi, Vallea de Palqui, 3500 m, suelo limo-arenoso, pH 6.4, 7 Feb 1987, R. Ehrich 889 (HOLOTYPE: TEX!; Isotype: LPB). Species foliis aliquanto arcuatis 3-verticillatis, spicis brevibus, bracteolis calyces aequantibus, et calycibus brevibus dense glan- diferis minute strigosisque a congeneribus diversa. Subshrubs to 2.5 m tall; stems reddish brown, deeply grooved, sparsely strigulose with stiff hairs 0.2-0.3 mm long. Leaves mostly in whorls of 3, oppo- site near the apex, on a swollen saccate petiole 1 mm long, the blades narrowly oblong-lanceolate, entire, 2.5-4.5 mm wide, 20-30 mm long, rounded-acute at the apex, spreading arcuate to slightly deflexed, densely granular glandular beneath and sparsely strigose on the midvein, the upper surfaces lustrous, strigose-hirsute with hairs 0.2-0.3 mm long, arising from pustulate bases. In- florescence of numerous, ascending, axillary spikes, the rachis 7-17 mm long, with flowers mostly clustered toward the apex; bractlets ovate-attenuate, equal or slightly longer than the calyces, sparsely glandular compared to the calyces. Calyx 1.2-1.6 mm long, densely granular glandular and strigulose-hirtellous, the lobes deltate, purplish. Corollas light blue, the tube 2.5-3.0 mm long, densely and evenly puberulous to minutely strigillose, granular glandular at the throat, the lobes hairy abaxially and adaxially. Aloysia arcuifolia Nesom is distinctive within the genus in its high ele- vation habitat, somewhat arcuate leaves in whorls of three, its short spikes, very short calyces with densely glandular and minutely strigose vestiture, and bractlets equaling the length of the calyx. Most of the species of Aloysia 145 146 PHYT@LOGIA volume 70(3):145-146 March 1991 have densely pilose-hirsute calyces. The remaining taxa produce calyces with greatly reduced vestiture, varying from closely puberulent to short strigose. While these latter species may not form a natural group, they are easily dis- tinguished from those with stiffly spreading calycine vestiture. The only other species of Aloysia described from Bolivia, A. beck Mold. and A. boliviensis Mold., both produce hirsute calycine vestiture and differ from A. arcuzfolia in other features as well. Both occur at significantly lower elevations. Among other species in the genus with reduced vestiture, A. fiebrigii (Hayek) Mold. and A. triphylla (L’Her.) Britton both occur in Peru, although the latter is found in many other areas as well. Both of these species, however, have signif- icantly longer calyces than A. arcuifolia and qualitatively different vestiture. ACKNOWLEDGMENTS I thank Dr. Billie Turner and Dr. Linda Escobar for their review and com- ments on the manuscript. Phytologia (March 1991) 70(3):147-148. THE LAST PHYSINGA LINDLEY (ORCHIDACEAE) Eric A. Christenson 1646 Oak Street, Sarasota, Florida 34236 U.S.A. ABSTRACT In rejoining Physinga Lindley with Epidendrum L., one Brazilian species has been left without an applicable name in Epidendrum. Hence, the combination Epidendrum acreense (Brieger & Bicalho) E.A. Christenson is proposed. KEY WORDS: Brazil, Epidendrum, Physinga, nomenclature, Or- chidaceae The genus Epidendrum L. is one of the largest Neotropical orchid genera with more than 1,000 species recorded from the southern United States, the Caribbean, and Central and South America. Because of its size and diversity, many attempts have been made to divide Epidendrum into smaller units with varying success (Hagsater 1985). Some of these segregate genera are now well established, such as Barkeria Know. & Westc., Encyclia W.J. Hook., Jacquiniella Schltr., and Oerstedella Reichb. f. Others have been generally rejected by consensus, such as Auliza Salisb., Lanium Lindl., Nanodes Lindl., and Neolehmannia Kraenzl. One segregate genus, Physinga Lindley, was resurrected by Brieger & Bi- calho (1978) for a group of species having long flexuous inflorescences, se- quential solitary flowers, a prominent vesicle, and usually a long clinandrium. While defining a cohesive species group, the characters are not worthy of generic recognition (Hagsater 1985, 1987). In eliminating Physinga, authors “orphaned” one Brazilian species without an applicable name in Epidendrum. The combination is made here: Epidendrum acreense (Brieger & Bicalho) E.A. Christenson, comb. nov. BASIONYM: Physinga acreensis Brieger & Bicalho, Bradea 2(34):234. 1978. 147 148 PHY TOLOGIA volume 70(3):147-148 March 1991 LITERATURE CITED Brieger, F.G. & H.D. Bicalho. 1978. Restabelecimento do Genero Physinga Lindl. (Orchidaceae). Bradea 2(34):231-237. Hagsater, E. 1985. Towards an understanding of the genus Epidendrum. In: Proc. Eleventh World Orchid Conference (K. Tan, ed.). pp. 195-201. . 1987. Epidendra Nova et Criticae 2: Epidendrum macroclinium, a new species of the Physinga group, confused with Epidendrum physodes. Orquidea (Mex.) 10:330-336. Phytologia (March 1991) 70(3):149-157. NOVITATES ANTILLANAE. XVI. Alain H. Liogier Botanic Garden — Central Administration, University of Puerto Rico, P.O. Box 364984, San Juan, Puerto Rico 00936-4984 U.S.A. ABSTRACT New species and new combinations from the island of Hispaniola, reported here, as a result of studies in preparation of coming volumes of the Flora of Hispaniola. KEY WORDS: Asclepiadaceae, Solanaceae, Rubiaceae, Hispan- iola, taxonomy, floristics ASCLEPIADACEAE Following the latest opinion on the genus Metastelma vs. Cynanchum, the following combinations are necessary: Metastelma calcicola (Alain) Alain, comb. nov. BASIONYM: Cynanchum calcicolum Alain, Phytologia 64(5):345. 1988. Metastelma leptophyllum (Schltr.) Alain, comb. nov. BASIONYM: Aste- phanus leptophyllum Schltr. in Urban, Symb. Ant. 1:241. 1899. Cy- nanchum leptophyllum (Schltr.) Jiménez, Rhodora 62:238. 1960. Metastelma occidentale (Spreng.) Alain, comb. nov. BASIONYM: Seca- mone occidentalis Spreng., Neue Entd. 3:34. 1822. Tainionema occiden- tale (Spreng.) Schltr. in Urban, Symb. Ant. 1:241. 1899. Cynanchum occidentale (Spreng.) Jiménez, Rhodora 62:238. 1960. 149 150 PHYTOLOGIA volume 70(3):149-157 March 1991 SOLANACEAE Lycianthes speciosa (Dunal) Lourteig, Phytologia 62(6):442. 1987. This species reported by Plumier from Haiti: Port de Paix and Léogane, has been confused by different authors, and al- though Lourteig did not see the specimens reported by Plumier, it seems different from L. pauciflora (Vahl) Bitter. Howard (Fi. Less. Ant. 6:278. 1989.) cited Solanum speciosum Dunal in the synonymy of Lycianthes pauciflora (Vahl) Bitter; he also cites Solanum neglectum Dunal (= Lycianthes neglecta [Dunal] Lourteig) in the synonymy. Lourteig creates Lycianthes ne- glecta (Dunal) Lourteig (Phytologia 62(6]:442. 1987.). It seems that Lourteig thinks that the plants cited by Plumier from Martinique and two specimens from Dominica (Toepfer 896 and Plée s.n.) belong to a distinct species, named by her Ly- cianthes neglecta. She thus creates two species, basing the first one on drawings and two old specimens, and the second one on drawings only. As for our species, Lycianthes speciosa (Dunal) Lourteig, as we have no specimens at hand, | have decided to consider it as a dubious record. Furthermore, on page 443 of her paper on Lycianthes, she divides the genus in the Antilles into two sections: a) species with glabrous petals, and b) species with pubescence on the adaxial side of the petals; but there is no mention of Lycianthes speciosain either of the sections. Where does it belong? Howard (Fl. Less. Ant. 6:278. 1989.) cites Lycianthes pauciflora (Vahl) Bitter from the Lesser Antilles, with Solanum speciosum Dunal and S. neglectum Dunal in the synonymy. No mention is made of Lycianthes neglecta, cited by Lourteig from Dominica and Martinica. Solanum aquartia Dunal var. fuertesii (O.E. Schulz) Alain, comb. nov. BASIONYM: Solanum aculeatum O.E. Schulz var. fuerteszi O.E. Schulz wm Urban, Symb. Ant. 7:370. 1912. Solanum aquartia Dunal var. luxurians (O.E. Schulz) Alain, comb. nov. BASIONYM: Solanum aculeatum O.E. Schulz var. lururians O.B. Schulz in Urban, Symb. Ant. 7:370. 1912. Liogier: Novitates Antillanae XVI 151 RUBIACEAE Exostema myrtoides Alain, sp. nov. Frutex 3 m altus, glaberrimus; rami teretes striati lenticel- losi, vetustiores cinerascentes cortice fisso sed adhaerente; stipulae ovatae apice rotundatae, obtusae vel acutae, 1 mm longae; folia opposita usque 5 mm longae petiolata, petiolo supra canaliculato; lamina elliptico-oblonga 2.0-3.5 cm longa, 1.0-1.5 cm lata, apice ro- tundata, basi attenuata in petiolum protracta, nervo medio supra prominulo subtus prominente, lateralibus nulls vel utroque latere 2-3 arcuatis non anastomosantibus, margine anguste recurva, supra nitentia obscure viridia, subtus pallidiora glaberrima; inflorescen- tiae plerumque ad ramos terminales in corymbum usque 2.5 cm latum collectae, in juventute pulverulentae; bracteae filiformes 1.5 mm longae; pedicelli 1-2 mm longi; calycis tubus in fructo turbina- tus 1.5 mm longus, glaber, nitens, venis prominentibus auctus; lobi 4 lineares 1 mm longi; corolla et caetera non visa; semina elliptica apice ala 1 mm longa, lateraliter 0.5 mm alata. TYPE: DOMINICAN REPUBLIC. Prov. Barahona: In thick- ets, on slope, Polo-Los Arroyos, alt. 800-1000 m, 28 Feb 1969, A. Liogier 14842 (HOLOTYPUS: NY). PARATYPE: DOMINICAN REPUBLIC: between Boucan Calice and Hoyo de Peiempito, 13 Feb 1969, A. Liogier 13866 (NY), old fruits only. This species is in the same group as Ezostema elegans Krug & Urban, from which it is distinguished by being entirely glabrous, its leaves are obovate- elliptic, up to 3.5 cm long, instead of oval to oval elliptic and up to 9 cm long in E. elegans; this last species has domatiate nerve axils beneath the leaves. Ezostema selleanum Urban & Ekman has acute to acuminate leaves, and these are also domatiate in the nerve axils beneath; in both EF. elegans and E. selleanum, the branchlets are pubescent. The genus Stevensia Poiteau. The genus Stevensia was created by Poiteau in 1804, with a single species, S. buzifolia Poiteau (Ann. Mus. Hist. Nat. Paris 4:1835. 1804.). Urban (Ber. Deutsch. Bot. Ges. 15:264. 1897.) cites this genus and species. In the mean- time, Grisebach (Fl. Brit. W. Ind. 328. 1861.) described Rondeletia pozteat, a name applied in part to S. buzifolia, and erroneously cited by Grisebach as occurring in Cuba. Several authors included S. buzifolia in their publications: 152 PHY DPOLOG FA volume 70(3):149-157 March 1991 Gaertner f. (1807), Poiret in Lamarck, Encyclopédie (1806), A. Richard (1829), P. DeCandolle (1830). Urban again describes the genus in Symb. Ant. (1:420. 1899.). Standley (N. Amer. Fl. 32:42. 1918.) describes the genus with the only known species (S. buzifolia) as part of the tribe Rondeletieae. After the extensive collections by Ekman in Haiti (1917), and later (1924- 1931) both in Haiti and the Dominican Republic, Urban described five more species: Stevensia samanensis Urban (1924), S. ovatifolia Urban & Ekman (1926), S. hotteana Urban & Ekman (1931), 5S. ekmaniana Urban (1931), and S. ebracteata Urban & Ekman (1931). In my explorations in Hispaniola, I have collected and studied three species new to science, giving a total of nine species in this genus which is endemic to Hispaniola. Recent collections confirm my diagnoses and add new specimens to my own collections. Stevensia Poiteau is a member of the Rondeletieae and differs from Ron- deletia L. in having a calyx closed before anthesis and strongly rostrate, open- ing in two or three lobes and deciduous, usually densely sericeous within; the corolla presents 5-7 lobes, the flowers are solitary in the leaf axils; in Rondele- tia, the calyx is 4-5(-7) lobed, not rostrate and persistent, the corolla cymose, paniculate, thyrsoid, racemose or spiciform. While Rondeletza is widespread throughout tropical America, Stevensia is restricted to the island of Hispan- iola. Key to the species of Stevensia 1. Twigs glabrous; fruits glabrous; pedicels 5-12 mm long. ............... 2 2. Stipule tube truncate, 2-3 mm long; prophylla forming a truncate tube Lovmm: long 422023005. cokes nn a0 oe S. ebracteata 2’ Stipules connate at base, with triangular-acute to deltoid lobes. ..3 3. Prophylla leafy, linear-lanceolate, 7-10 mm long; stipules 5 mm long, including the connate base; leaves up to 8 cm long. S. hotteana 3’ Prophylla connate forming a tube, with minute, narrowly trian- gular lobes; stipules 2 mm long, with deltoid lobes; leaves up fo, V2 em lone, on. cc scsenss seenne een S. grandiflora 4. Leaves elliptic to oblong, up to 1.5 cm long, rounded to obtuse at apex; corolla 8 mm long; flowers sessile or nearly so. S. minutifolia 4’ Leaves acute to acuminate at apex, 2.5-10. cm long; corolla 1.0-2.5 CMV TONG ec ees cence reese sconce nntuat t= 5 Liogier: Novitates Antillanae X VI 153 5. Twigs usually not resinous; corolla 20-25 mm long; leaves brown ented tli: spl cOuisciern i-th kaintase thi Tei a2 ai S. ekmaniana 5' Twigs resinous; corolla 10-20 mm long; leaves whitish to grayish Rene it lad 2 cote eet Sys ic ep acetals: eet its clos diy BIAS otic SRS 2 6 6. Leaves 2.5-5.0 cm long; corolla 1.0-1.5 cm long. ........ 7 7. Stipule tube 1 mm long, the lobes triangular, 1 mm long; corolla 1 cm long; prophylla connate, tube 2 mm long, lobes 1.5 mm long; pedicels 2 mm long. ......... TP eee «oad stare ae arate Sane Rais eels S. buzifolia 7 Stipules 2.5-4.0 mm long, including the connate base; prophylla free; pedicels up to 0.5 mm long. .......... 8 8. Leaves ovate, 1.5-4.0 cm long, 1.0-2.2 cm broad; pro- phylla triangular, 1 mm long; twigs smooth, papillose. Sse cee ihe? ates oe reise he S. ovatifolia 8/ Leaves elliptic oblong to lanceolate, 2.5-5.0 cm long, 5-15 mm broad; prophylla oblong-elliptic, 6-8 mm long, acuminate; twigs aculeolate, retrorse strigose. ....... Boge Re Se tt vc cpcyac hs i i asaya Chara ots ea S. aculeolata 6’ Leaves lanceolate to linear lanceolate, 4-10 cm long; corolla 2 cm long or longer; prophylla lanceolate to linear lanceo- latex Summing Palisa ais sass. 0 5.saa te ne S. samanensis Stevensia aculeolata Alain, sp. nov. Frutex 1.5 m altus; rami hornotini valde resinam exsudantes, retrorse-strigosi aculeolati teretes; stipulae interpetiolares basi sub- connatae, e basi triangulari cuspidatae pulverulentae, 4 mm longae; folia petiolis usque 3 mm longis supra canaliculatis suffulta, lamina anguste elliptico-oblonga vel lanceolata, 2.5-5.0 cm longa, 5-15 mm lata, basi versus attenuata, basi ipsa rotundata, obtusa vel subcor- data, apice acuta vel breve acuminata, nervo medio supra impresso subtus prominente, lateralibus supra obsoletis, subtus utroque lat- ere 15-20, sub angulo 60-65° abeuntibus non anastomosantibus, lamina valde discoloria, supra glabra in sicco pallide brunnea, sub- tus pallide fulva dense tomentulosa, margine recurva, coriacea; flores in axillis solitarii sessiles; prophylla 2 oblongo-elliptica, 6-8 mm longa apice acuminata dorso brevissime puberula, cum stipulis aequilongis alternantia; calycis alabastra ovata in rostrum 3 mm longum contracta, posterius in lobos 2 anguste ovato-cuspidati 4-5 mm longi, ciliati, dorso strigulosi; corolla alba 12-14 mm longa, extus dense retrorso-sericea, lobi 6 anguste elliptici 3 mm longi, 1.5 mm lati; capsulae sphaeroideae 4 mm diam., extus parce et brevissime pilosa, calycis lobis coronata, demum nuda. 154 PHY TOL OGLiA volume 70(3):149-157 March 1991 TYPE: DOMINICAN REPUBLIC. On limestone rocks, at sea level, Samana Bay, Los Haitises, 19 Jan 1977, Alasn & Perfa Liogier 26829 (HOLOTYPUS: UPR; Isotypi: NY, SD). Paratypes: DO- MINICAN REPUBLIC. On limestone rocks, at sea level, Samana Bay, Los Haitises, 25 Apr 1985, Zanoni, et al. 84895 (NY, SD); On limestone rocks, at sea level, Samana Bay, Los Haitises, 24 Apr 1985, Zanoni, et al. 34202 (NY, SD); Boca del Infierno, Samana Bay, 24 Jun 1930, Ekman 15421 (S, NY). Stevensia aculeolata Alain is similar to S. buzifolia. However, the latter species has but slightly resinous branchlets, not aculeolate, short pilose; the stipular sheath is 1 mm long, and the lobes are triangular, 1 mm long; the pedicels are up to 2 mm long, the prophylla are connate at the base, the lobes 2-4, triangular, acute; the nerves on the leaf blade are 8-10 on each side. The new species is also near to S. samanensis Urban, which has larger leaves, 4-10 cm long; the corolla is longer (2 cm long or longer), the prophylla are lanceolate to linear lanceolate, 8 mm long. Stevensia buzifolia Poiteau, Ann. Mus. Hist. Nat. Paris 4:235. 1804. (Ron- deletia poiteai Griseb., Fl. Brit. W. Ind. 328. 1861. p.p.). - “Cuabilla,” RD. - This is by far the most common species in the genus, growing in the eastern half of the Dominican Republic and throughout Haiti, at altitudes from sea level to about 1000 m alt., in forests and thickets, mostly on metamorphic soil. Stevensia ekmaniana Urban, Ark. Bot. 24A(4):41. 1931. TYPE: HAITI. Massif du Nord, near Bayeux on limestone rocks at Morne Brigand, 800 m alt., Ekman 2849 (HOLOTYPE: S). Known only from the type collection. Stevensia grandiflora Alain, sp. nov. Frutex 2 m altus, rami hornotini teretes, in sicco striati, resinam exsudantes, glabri; stipulae infra petiolos breve connatae, late ovatae apice breve cuspidatae ciliatae, 3.0-3.5 mm longae; petioli 1.0- 1.3 mm longi, supra canaliculati glabri; foliorum lamina oblongo- elliptica 4.5-8.0 cm longa 1.5-3.0 cm lata, apice acuminata, basin versus attenuata in petiolum protracta, nervo medio supra im- presso lateralibus obsoletis, subtus nervo medio bene prominente, lateralibus utroque latere 10-12, brunneis ad marginem paullo ar- cuatis non anastomosantibus, supra obscure viridia (in sicco ob- scure brunnea), subtus albo-grisea, margine integra paullo recurva, Liogier: Novitates Antillanae X VI 155 glabra; flores solitarii in foliorum axillas, pedunculi ca. 1 cm longi glabri, prophylla sub calycem posita in cupulam 1.5 mm longam connata, lobi lineares 2 mm longi; alabastra anguste ellipsoidea 4 mm longa 2 mm lata in rostrum 4 mm longum protracta, in lobos 2 subaequales 3 nerves, glabri usque 1.5 mm longos aperiens; corolla alba 2.5 cm longa, extus dense retrorso-sericea, tubus cylindraceus ad apicem paullo ampliatus, in statu compresso usque 3 mm latus, lobi 6 oblongo-obovati usque 8 mm longi, 4 mm lati; stamina sub fauce corollae inserta, filamenta brevissima; capsulae globulosae 8 mm diametro, glabrae, nitidae, septicide dehiscentes, apice nudae. TYPE: DOMINICAN REPUBLIC. La Vega Prov.: La Man- aclita, on lateritic soil and serpentine, alt. 600 m, 25 Jun 1969, Alain H. Liogier 15816 (HOLOTYPUS: NY; Isotypi: G, J, P, US). Paratypes: DOMINICAN REPUBLIC. La Vega Prov.: La Man- aclita, on lateritic soil and serpentine, alt. 600 m, 28 May 1968, Alain H. Liogier 11461(G, J, NY, P, US). Bonao: banks of Hato Viejo River, Hato Viejo, 10 May 1969, J. Jiménez & A. Liogier 5678 (NY); Barrancon, Hato Viejo, alt. 250-300 m, 10 May 1969, Alain H. Liogier & J. Jiménez 15186 (G, J, NY, P, US); Loma Peguera, alt. 500 m, 17 Oct 1970, Alain H. Liogier 17595 (G, J, NY, P, US); Loma Peguera, alt. 450 m, 29 Nov 1972, Alain H. Liogier 18553 (G, J, NY, P, US); Falconbriage Dominicana, Loma Peguera, alt. 500 m, 23 Apr 1981, Zanoni, Mejia, & Ramirez 12950 (NT, SD). Piedra Blanca: Rio Mahoma, alt. 900-1100 m, 26 Aug 1982, Zanoni, et al. 22903 (NY, SD). Distrito Nacional: Loma Mariana Chica, alt. 300-800 m, 19 Jun 1984, Zanonz, et al. 30565 (N'Y ;SD): The specimens cited as paratypes present larger leaves than the type spec- imen: up to 12 cm long and 4 cm broad. This species has the largest flowers and leaves of any described member of Stevensia. Plants are entirely glabrous; stipules are short and deltoid, connate at the base, with narrowly triangular lobes. The specimens cited as types of S. grandiflora had previously been named S. ebracteata Urban & Ekman. How- ever, the latter species has stipules and prophylla connate to form a truncate tube, and smaller leaves (up to 8 cm long). Stevensia hotteana Urban & Ekman, Ark. Bot. 24A(4):40. 1931. TYPE: HAITI. Massif de la Hotte, near Jérémie at about 150 m alt., between Source Chaude and Source Cahouane, Ekman 10222. Another collection not seen by this author, but identified by T. Zanoni as this species is: HAITI. Morne Formon, in cloud forest, at 1530-1650 m alt., W. Judd 3653. 156 PHY? OLDOGTA volume 70(3):149-157 March 1991 Stevensia minutifolia Alain, sp. nov. Frutex 1.25 m altus; rami hornotini teretes, in sicco striati, ad apicem resinam exsudantes, pilis minutissimis pilosuli; stipulae basi breve intra petiolos connatae, triangulares, apice rotundatae, 1.5 mm longae, ciliatae; petioli 1-2 mm longi, supra sulcati, pilosull; lamina foliorum elliptica vel anguste elliptica, 1.0-1.5 cm longa, 0.3- 0.5 cm lata, apice rotundata, obtusa vel acuta, basi acuta, obtusa vel rotundata, nervo medio valde prominente, lateralibus utroque latere 5-6, ad marginem curvatis non anastomosantibus brunneis; lamina supra obscure brunnea, subtus albogrisea utroque latere glabra, margine paullo incrassata chartacea; flores in axillis folio- rum solitarii, pedicelli vix 1 mm longi; prophylla sub ovario posita in cupulam 1 mm longam connata, pars libera subulata 2 mm longa; calyx in alabastro ellipticus 3 mm longus in rostrum 2 mm longum contractus, papillosus, posterius in lobos 2 ellipticos cusp- idatos usque 3 mm longos intus dense sericeos divisus; corolla alba 8-9 mm longa, extrinsecus dense retrorso-sericea, tubus turbinatus 5 mm longus ad apicem 2 mm crassus, lobi 5 obovato-elliptici 2 mm longi, 1.5 mm lati, apice rotundati; stamina fauci corollae in- serta filamenta perbrevia, stylus ad apicem bifidus, lobis linearibus; capsulae sphaeroideae 4 mm diam., apicae nudae, minute pilosulae. TYPE: HAITI. In pine forest, 1650 m alt., from Furcy to Seguin, 17 Nov 1982, M. Mejia & J. Pimentel 24394 (HOLOTY- PUS: NY; Isotypus: SD). Paratypes: HAITI. Massif de la Selle, Morne Tranchant, on limestone on slope, 1750 m alt., 6 Jun 1924, Ekman 1779 (NY, S). Additional paratypes: DOMINICAN REPUBLIC. Prov. Ped- ernales: Hoyo de Pelempito, 1100 m alt., 26 Feb 1971, A.H. Lrogier 17936 (G, J, NY, P, US). Bahoruco Mountains: Aceitillar-Cayo, on limestone rocks mixed with bauxite, 9 Feb 1969, A.H. Lio- gier 19698 (NY); Aceitillar-Cayo, on limestone rocks mixed with bauxite, 9 Feb 1969, A.H. Liogier 19663 (NY); Aceitillar-Cayo, on limestone rocks mixed with bauxite, 24-25 Jul 1973, A. & P. Li- ogier 19653 (NY, SD); Aceitillar-Cayo, on limestone rocks mixed with bauxite, Feb 1971, F. Votava & A.H. Liogier 103 (NY). This species is readily recognized among the other species in Stevensia by its small leaves, rounded stipules, and small, nearly sessile flowers. The nearest species seems to be S. buzifolia. Stevensia ovatifolia Urban & Ekman, Ark. Bot. 20A(5):54. 1924. TYPE: HAITI. ad Montagnes Trou d’Eau, near Glore, on top of the mountain near 800 m alt, Ekman 1064. Liogier: Novitates Antillanae X VI 157 Additional specimens examined: HAITI. Morne a Cabrits, Ekman 5648; Montagnes Noires, near Croix des Bouquets, alt. 550 m, Zanonz, et al. 24011 (NY, SD): Stevensia samanensis Urban, Repert. Spec. Nov. Regni Veg. 20:351. 1924. TYPE: DOMINICAN REPUBLIC. Samana Peninsula, on Pilon de Azucar mountain, near Laguna, Abbott 2354. Paratypes: DOMINICAN REPUBLIC. Samana Peninsula, on Pilon de Azucar mountain, near Laguna, Ekman 14554; Los Haitises, Samana Bay, Zanoni, et al. 35958; Los Haitises, Samana Bay, Mejia & Pimentel 239538. Mejia & Pimentel 23953 has capsules globose, 4 mm diam., minutely pilose, grayish, the calyx lobes persistent, 3-4 mm long, acuminate, densely sericeous within, ciliate, resinous without; seed elliptic, 1.5 mm long, acuminate at both ends. A doubtful specimen collected by Ekman on Tortue Island, Haiti (no. 4189) seems different from Stevensia buzifolia. The specimen has young flowers on 3 mm long pedicels with linear prophylla 1.5 mm long, the bud is oblong, 3 mm long, 1.75 mm wide, with a 2.5 mm long rostrum; the leaves are elliptic, rounded to obtuse at apex, acute at base, 3.0-3.5 cm long, 2.0-2.7 cm broad, glabrous, dark brown above, light brown beneath, reticulate veined; the twigs are papillose. Probably a hybrid between S. ebracteata and S. ovatifolia Urban & Ekman. Phytologia (March 1991) 70(3):158-177. RECENSION OF THE CRONQUISTIANTHUS GROUP OF EUPATORIUM (ASTERACEAE) B. L. Turner Department of Botany, University of Texas, Austin, Texas 78713 U.S.A. ABSTRACT A taxonomic treatment of the largely Andean genus Cronquistianthus King & H. Robins. is rendered. Because it is retained in Eupatorium, s.l., the following new combinations and names are made: Eupato- rium celendinense (King & H. Robins.) B. Turner, comb. nov.; E. chachapoyense (King & H. Robins.) B. Turner, comb. nov.; E. bish- opii (King & H. Robins.) B. Turner, comb. nov.; E. determinatum (B.L. Robins.) B. Turner, comb. nov.; E. ferreyrii (King & H. Robins.) B. Turner, comb. nov.; E. huanucanum B. Turner, nom. nov.; E. ro- sei (King & H. Robins.) B. Turner, comb. nov. In all, twenty species are recognized. Brief descriptions, distribution maps, and a key to the species are provided, along with complete synonymies. KEY WORDS: Asteraceae, Eupatorieae, Eupatorium, Cronquis- tianthus King & Robinson (1987) recognized 25 Andean species of Colombia, Ecua- dor, and Peru in their segregate genus Cronquistianthus, most of these having previously resided in Eupatorium. The group was positioned in their subtribe Critoniinae between Ophryosporus Meyer (which contains about forty species) and Steyermarkia King & H. Robins. (with four species); it is readily distin- guished from the former by its imbricate involucres and appendaged anthers, and from the latter by its broad, strap shaped stylar appendages. In real- ity, the Cronguistianthus group would appear closest to Aristeguietia King & H. Robins. (containing about twenty species), the latter having very similar stylar appendages, but largely distinguished from the former by its acute in- volucral bracts (vs. obtuse or rounded) and rather symmetrical carpopodia (vs. + asymmetrical). Indeed, future workers, if inclined to accept at least some of the splintering of Eupatorium by King & Robinson, may want to combine these two segregate groups, both being essentially Andean taxa and the characters 158 Turner: Recension of Cronquistianthus 159 which distinguish between them somewhat arbitrary at best. Regardless, in attempting to provide names for the various plants sent to me for identifica- tion, I have felt the need to position the various new taxa proposed by King & Robinson for Cronguistianthus within Eupatorium, s.l., until such time as more definitive data bearing upon relationships within the latter complex is forthcoming (hopefully that obtained from DNA restriction site studies, etc.). CRONQUISTIANTHUS King & H. Robins. King & Robinson (1987) have provided an excellent description of this segregate from Eupatorium, s.l., and such is not duplicated here. They also provided an exceptional illustration of the generitype, Eupatorium niveum H.B.K. The group has a base chromosome number of z = 10, much as in Eupatorium, s.st. Distributional maps for al] of the species are presented here, these being arranged alphabetically in figures 1-4. KEY TO SPECIES OF CRONQUISTIANTHUS GROUP OF EUPATORIUM 1, Plants ‘of Eevadorior Wolombian yo. oosc0n ss ae geen ee eta d eee (2) 1 Plants of Perm soupumande so toe g od sas hoor ccs bese Aten eet (7) 2. Leaves clearly bicolored, the lower surfaces loosely to densely white tomentose ns sesso eee; be oe. BRbw Milani seems oe (3) 2. Leaves about equally green on both surfaces, if somewhat bicolored then the lower surface densely puberulent, not tomentose, clearly VEU GUlaLEWERGSe ye Soon c oe es is cae nenbat.cne 2 28 es ce eee (4) 3. Leaves with upper surfaces bullate; achenes glabrous or nearly so; northern Eedadoriand vadjacent (Colombia. 1230220 0.,5- a Se + te ceeds s E. niveum 3. Leaves with upper surfaces smooth; achenes clearly hispidulous; southern Bemadors, en cn ene aca Pe ermenes te cates ote ca eaten E. leucophyllum 4. Petioles mostly 4-12 mm long; involucral bracts thin, mostly 1.0-1.5 INT Wide os sce mtr tat oh ie ea niece nna Maral ae E. pseudoriganoides 4. Petioles mostly 1-4 mm long; involucral bracts thick, mostly 1.5-4.0 TAIT NG gies tH oS gs EUR ea eo EA cscs velit me ergs ul (5) 160 PHY TOLOGIA volume 70(3):158-177 March 1991 5. Heads with 10-12 florets; involucres 5-6 mm high. .............. E. rosei 5. Heads with 18-20 florets; involucres 6-8 mm high. .......... BE. bullatum 6. Leaves mostly 10-25 mm long., ......\...2.0.229. 50ers (7) 6. Leaves mostly 25-100 mm long. ... 2... 22% «cheer eee (12) 7. Blades linear to linear oblanceolate, 3-8 times as long as wide. ......... Sfcaier= Se ea ean Oe aap e nase win mys H i: uf =I . oa a ae ret "-=/ = ' . . . ’ \ Y ’ 5 . ‘ .. ‘ . 7) . / E } a ‘ ‘ ‘ ‘ PERU 4} EUPATORIUM Uyeer Ochamaedrifolium / 4 desmophyllum : be Adeterminatum i i @ferreyrii ak @glomeratum sa oon = <<" jones ‘ cr Wiescaes - : ‘ ' = .< Fo Se . Ss . =z 4 WAS = 55 _ / ‘ hls = _+! sees = Se! ' ™— Sets y ‘ Se Se ‘ —— BOLIVIA ; ne ’ == Necsaoe — cf. = .% Fig. 2. Distribution of Eupatorium Turner: Recension of Cronguistianthus 169 Eupatorium macbridei B.L. Robins., 1923). TYPE: PERU. Huanuco: Huacachi, a station near Muna, open rocky ridge, ca. 6500 ft, 20 May-1 Jun 1923, Macbride 3881 (HOLOTYPE: US; Isotype: F!). Shrub or shrublet to 1.5 m high. Leaves 3-4 cm long, 2-3 cm wide; peti- oles 1-4 mm long; blades triangular to subcordate; trinervate, densely long hispidulous beneath, the veins raised reticulate, the margins crenulate. Heads numerous in congested corymbose panicles. Involucres 4-5 mm high, the bracts 3-4 seriate, markedly deciduous, pilose, the apices broadly rounded. Florets ca. 10 per head. Corollas 4-5 mm long. Achenes ca. 1 mm long, sparsely hispidulous, the pappus of 30-40 bristles 4-5 mm long. ADDITIONAL SPECIMEN EXAMINED: PERU. Hudnuco: Acomayo, 2100 m, 14 Apr 1946, Woytkowks: 34231 (F, MO). This taxon might be mistaken for a large leaved form of Eupatorium chamae- drifolium, but the latter has fewer, larger heads with glabrous involucral bracts. It might also appear to be a small leaved form of E. callacatense, but the latter has leaves with smoother undersurfaces and appressed soft hairs. Its original] describers compared it to E. glomeratum DC., the latter having longer more sagittate blades with longer petioles (8-30 mm long vs. 1-4 mm long). Eupatorium lavandulaefolium DC., Prodr. 5:15. 1836. Crongqutstianthus lavandulaefolius (DC.) King & H. Robins., Phytologia 23:411. 1972. TYPE: PERU. without locality, without date, Dombey 51 [8147] (LEC- TOTYPE [selected here]: G-DC; Photolectotypes: F!, MO!, NY!, TEX!). Two specimens were examined by DC., the other a Haenke collection, also from Peru; they clearly belong to the same taxon. Eupatorium chotense Hieron., Bot. Jahrb. Syst. 36:466. 1905. Cron- quistianthus chotensis (Hieron.) King & H. Robins., Phytologia 23:410. 1972. TYPE: PERU. Cajamarca: between Chota & Cutervo, Jun 1979, Jelski 794 (HOLOTYPE: B, destroyed; Photo- holotypes: F!, MO!, NY!). Suffruticose herbs or subshrubs to 1 m high. Because of its bicolored lin- ear lanceolate to linear leaves, which are often deflexed, an easily recognized species. REPRESENTATIVE SPECIMENS: PERU. Ancash: Pachacoto to Catac, ca. 3400 m, 13 May 1976, Lopez M. 8333 (F, MO). Cajamarca: Llama- Huambos, ca. 2280 m, 22 May 1965, Lopez 5261 (F, MO); Las Quinuas to El] Mejon, ca. 3200 m, 14 Jun 1981, Sagastegui A. 10109 (F). Junin: ca. 18 km by air SSE of Tarma, ca. 4000 m, 27 Nov 1962, Jltts 132 (F). La Liber- tad: Huacas, ca. 2800 m, 15 Jun 1984, Sagastegui A. 11902 (F, MO). Loreto: Rio Blanco, ca. 4000 m, 8-19 May 1922, Macbride 794 (F). Piura: between 170 PHYTO YOOERA volume 70(3):158-177 March 1991 Canchaque and Huancabamba, 1900-2200 m, 17 Apr 1987, Diaz S. 2480 (MO, Nyy) A variable species, especially in leaf shape and vestiture. Relatively broad leafed forms have been called Eupatorium chotense Hieron. Eupatorium leucophyllum H.B.K., Nov. Gen. & Sp. 4:90. 1818. |ed. folio]. Cronquistianthus leucophyllus (H.B.K.) King & H. Robins., Phytologia 23:411. 1972. TYPE: ECUADOR. Loja: “Juxta pagos Caxanumae et Gonzanama, alt. 1080 hex,” Aug 1802, Humboldt & Bonpland s.n. (HOLOTYPE: P; Photoholotypes: F!, LL!, MO!). This relatively poorly collected species resembles Eupatorium niveum H.B.K. but is distinguished by its weakly bicolored leaves with more nearly ovate blades which taper upon the petioles, and yet other characters, as noted in the key. ADDITIONAL COLLECTIONS EXAMINED: ECUADOR. Loja: in moun- tains about Loja, without date, Hartweg s.n. (NY); Argelia, 2230 m, 4 May 1946, Espinosa 301 (NY); entrance to Parque Nacional Podocarpus, 2600 m, 30 Apr 1987, van der Werff 9205 (MO). The type and Hartweg collections cited above were said to be from Peru but presumably were collected in southernmost Ecuador (District of Loja). The taxon is likely also to be found in northernmost Peru. Eupatorium lopez-mirandae Cabrera, Bol. Soc. Argentina Bot. 10:21. 1962. Cronqutstianthus lopez-mirandae (Cabrera) King & H. Robins. Phytolo- gia 54:40. 1983. TYPE: PERU. La Libertad: Haciendo Cochabambo, ca. 2750 m, 26 Jun 1958, A. Lopez & A. Sagastegui 2782 (HOLOTYPE: LP). This taxon, what with its small ashy white foliage, is superficially similar to Eupatorium ferreyrit but is readily distinguished by its broader ovate leaves and smaller heads (ca. 10 vs. 15-20); in addition the pappus is quite short, ca. 1 mm long (vs. 3-5 mm long in E. ferreyriz). The original description is accompanied by an excellent illustration. SPECIMENS EXAMINED: PERU. Cajamarca: road to Langla, ca. 2600 m, 3 Jul 1985, Mostacero L. § (F); La Encanada, ca. 2600 m, 17 Aug 1973, Sagastegui A. 7775 (F); Pullucana (Arriba de Banos del Inca), ca. 2650 m, 17 Aug 1984, Sagastegui A. 12001 (F, MO, TEX). So far as known, both Eupatorium lopez-mirandae Cabrera and E. ferreyru are largely confined to the Dept. of Cajamarca, the former to areas south of Cajamarca, extending into the border regions of La Libertad (from whence the type); the latter is largely centered along the Rio Maranon, along the Celendin-Balsas road, above the Rio Maranon. Turner: Recension of Cronquistianthus 171 Eupatorium marrubifolium Hieron., Bot. Jahrb. Syst. 36:466. 1905. Cron- quistianthus marrubifolius (Hieron.) King & H. Robins., Phytologia 27: 396. 1974. TYPE: PERU. Cajamarca: between Chota and Cutervo, Jun 1979, Jelski 799 (HOLOTYPE: B, destroyed; Photoholotypes F'!, MO!, NY!, TEX!). Suffruticose herbs or shrublets 40-80 cm high. Leaves mostly 2.5-6.0 cm long, 1.3-3.0 cm wide; petioles 3-10 mm long; blades ovate to cordate, mod- erately to sparsely pubescent beneath with mostly soft appressed hairs (often densely so when young), the margins crenulate. Heads lavender to pale blue, numerous and mostly sessile in terminal corymbose panicles. Involucres 7-8 mm high, the bracts glabrous or nearly so, often purplish apically. Florets 10-16 per head. Corollas 4-5 mm long. Achenes ca. 2.5 mm long, pubescent with short hispid hairs mostly to 0.1 mm long, the pappus of ca. 40 bristles 4-5 mm long. REPRESENTATIVE SPECIMENS: PERU. Cajamarca: arriba de San Juan, ca. 2800 m, 17 May 1976, Sagastegui A. 8366 (F); Fila la Conga, ca. 2800 m, 6 Jul 1977, Sagastegui A. 8976 (F); Cerro Huacaris, ca. 2600 m, 23 May 1965, Vega 65(F). Huanuco: Llata, ca. 2600 m, 21 Aug 1922, Macbride 2236 (F). La Libertad: Usquil, 3000-3100 m, 9 Jun 1950, Ferreyra 7643 (F); Salpo-Chanchacap, ca. 7800 m, 24 May 1984, Sagastegui A. 11655 (F, MO, TEX); vicinity of Santiago de Chuco, 2800 m, 13 Jun 1984, Sagastegu: A. 11766 (F, MO, NY). Lima: Canta, 2600-2800 m, 20-26 Jun 1975, Pennell 14742 (F, MO, NY). This taxon is closely related to Eupatorium callacatense but is readily dis- tinguished from the latter by its mostly smaller thicker leaves with shorter petioles. Eupatorium niveum H.B.K., Nov. Gen. & Sp. 4:90. 1818 [ed. folio}. Cronquts- tianthus niveus (H.B.K.) King & H. Robins., Phytologia 23:411. 1972. TYPE: COLOMBIA. Cauca: near Popavan, 1801, Humboldt & Bon- pland s.n. (HOLOTYPE: P (microfiche!); Photoisotypes: F!, MO!). Eupatorium origanoides H.B.K., Nov. Gen. & Sp. 4:89. 1818. [ed. folio]. Cronguistianthus origanoides (H.B.K) King & H. Robins., Phytologia 23:409-412. TYPE: ECUADOR. without locality, 1802, Humboldt & Bonpland s.n. (PHOTOHOLOTYPE: P (microfiche!); Photoisotypes: MO!, NY!). Cronquistianthus trianae King & H. Robins., Phytologia 56:252. 1984. TYPE: COLOMBIA. Narino: Turquerres, 3200 m, Triana 1210 (HOLOTYPE: US!; Photoholotype: MO!). Eupatorium niveum H.B.K. var. setosocarpum Gilli, Feddes Repert. Spec. Nov. Regni. Veg. 94:308. 1983. TYPE: ECUADOR. Im- PHY TOL@CGTA volume 70(3):158-177 March 1991 . ‘ pina! ‘ame. . ne er ye WW" < m Po mM N os m ie QD si 2 : re z = Se is ? H == / ¢ SSS OS ‘ ‘ FE = i Py —a = s een fend #22 COLOMBIA ; es => 5 i -= dou f 22> BSR a x — oe iv 24, = ws ae a ’ 1 / EUPATORIUM @ huanucanum Xx! O lavandulaefolium 4 leucophyllum 4 lopez-mirandae @ marrubifolium _—-- ‘ ‘ S beet + ‘ = = yl ‘ SESS = - . . ———— = . a 1 idee * “SS =. 3 = / Aa ~~ sE a — =! eee = _—— aS = \ -, = Sa be! ‘ —S Satin \ SS = 1s = 1 = = =) = : S32 FY = BOLIVIA $ —— — en = > d _ = aS a —“‘—C — a f = ——— ‘ = — = nS Fig. 3. Distribution of Eupatorium Turner: Recension of Cronqutstianthus 17s babura: Berg bei Otavalo, 2600 m, 16 Jul 1975, A. Gilhk 426 (PHO- TOHOLOTYPE: MO!). Clambering shrubs 1-2 m high. Stems cottony pubescent to glabrescent. Leaves 3-8 cm long, 2-4 cm wide; petioles 1.0-2.5 cm long; blades broadly ovate, deltoid or cordate, bicolored, trinervate, the lower surfaces densely white to- mentose, the upper surfaces glabrous, rugose or subrugose, the margins evenly crenulate. Heads numerous-in congested corymbose panicles. Involucres 4-5 mm high, the bracts 9-12, 2-3 seriate, graduate. Florets 8-12 per head, the corollas glabrous except for the lobes, white, ca. 3 mm long. Achenes ca. 1.5 mm long, glabrous or sparsely hispid at apices, the pappus of 30-40 bristles 2-3 mm long. REPRESENTATIVE SPECIMENS: COLOMBIA. Narino: near Pun, high bank above Rio Pun, 2895 m, 15 Aug 1935, Mezia 7623 (LL). ECUADOR. Carchi: Las Penas, 3150 m, 22 Jan 1939, Asplund 7201 (NY). Imbabura: 3 km toward San José de Minas from the Otavalho-Selva Alegre road, 2900 m, 24 Jan 1984, Grimes 2535 (NY, TEX). Pichincha: E] Pongo, 2800 m, 25 Jul 1987, Zak 2260 (F, MO, TEX). To judge from collections (22 sheets from NY alone), this is a relatively common species in northern Ecuador and closely adjacent Colombia where it occurs at elevations from 2900-3000 m. King & Robinson (1972, p. 409) con- tended that “One species of Cronguistianthus, C. organoides (sic), has scat- tered hairs inside of the corolla, a feature not found in any other species of the genus, even the closely related C. niveus.” If one plots those specimens cited by King & Robinson which reportedly contain scattered hairs in the corolla (an observation which I can confirm, but these are usually widely disposed within the throat and do not occur in all corollas) these will mostly cluster in Colom- bia and closely adjacent Ecuador, whilst collections which lack such hairs are mostly found in the vicinity of Quito, Ecuador (Depto. Pichincha and closely adjacent Carchi). Subsequently, King & Robinson (1984) acknowledged that their original evaluation of C. origanoides (H.B.K.) King & H. Robins. was erroneous, and that the latter name is synonymous with C. niveus (H.B.K.) King & H. Robins. This decision was presumably made after examinations of type material for hairs in the corolla, although this was not stated. The holotype of Eupatorium origanoides H.B.K. is at P; the microfiche of this spec- imen is difficult to assess as regards leaf shape, much less corolla hairs, but a photoisotype at NY reveals the leaves to have a leaf shape suggestive of what King & Robinson might call Crongutstianthus trianae King & H. Robins., thus if they maintain the latter taxon it might ultimately be again replaced with C. origanoides. Nevertheless, they maintained that specimens with scattered hairs within the throat represented a good species and provided the name C. itrianae for these. Other than the throat hairs (which are variable and occa- sionally occur in plants from about Quito, Ecuador) there appears to be little 174 POR YOR OE OUGA A volume 70(3):158-177 March 1991 else to distinguish between their C. trianae and C. niveus, although in general, the leaves from the more northern populations of Eupatorium niveum H.B.K. (as considered here) have somewhat longer, less rugose, thinner blades with mostly rounded bases (vs. blades triangular to cordate) and hairy throats ap- pear to correlate somewhat with such leaves. Regardless, the plants concerned are so similar and the characters which mark them so variable that I have little hesitation in treating these as but members of a single variable species, although future workers may find other characters or attributes that might warrant varietal status. Eupatorium pseudoriganoides Hieron., Bot. Jahrb. Syst. 29:10. 1900. Cron- guistianthus pseudoriganoides (Hieron.) King & H. Robins., Phytologia 23:411. 1972. TYPE: ECUADOR. without locality, without date, Sodero 6/24 (HOLOTYPE: B, destroyed?; Photoholotypes: MO!, NY!). Intricately branched low clambering shrubs to 3 m high. Leaves mostly 2.5- 6.0 cm long, 1.0-2.5 cm wide; petioles 4-12 mm long; blades ovate to deltoid, 3(5) nervate, about equally green on both surfaces, sparsely to densely pilose puberulent beneath, the margins crenulate, the apices acute. Heads numerous, white to pale lavender, arranged in broad, mostly trichotomous assemblages, the ultimate peduncles 0-4 mm long. Involucres (4)5-6 mm high, the bracts 3- 4 seriate, straminous, puberulent, mostly 1.0-1.6 mm wide, the apices obtuse. Florets 10-15 per head. Corollas 3-4 mm long. Achenes 2.5-3.0 mm long, glabrous or nearly so, the pappus of 30-40 bristles (2)3-4 mm long. REPRESENTATIVE SPECIMENS: ECUADOR. Azuay: Ganadel, ca. 3100 m, 23 Sep 1955, Asplund 17731 (LL, NY); ridge between E] Pan and Guachapala, ca. 2800 m, 4 Sep 1945, Camp E-5235 (NY); ca. 15 km SW of Cuenca, ca. 3000 m, 29-30 Jun 1945, Camp E-3956 (MO, NY); Cuenca, 15 Sep 1920, Holway 998 (NY); ca. 17 km S of Cumbe, ca. 3000 m, 24 Jan 1976, King 6892 (F, MO, NY). Canar: between Biblian and Canar, ca. 3300 m, 18 Sep 1944, Camp E-445 (NY); near E] Tambo, ca. 3300 m, 5 Jul 1945, Camp E-3989 (F). Chimborazo: Paramo de Cacheaco, ca. 3200 m, 27 Jul 1959, Barclay 8300 (MO). A variable but readily distinguished species by its nonbicolored, distinctly petiolate leaves and relatively thin, narrow, involucral bracts. A collection from Azuay Province (Jaramillo 5447, F) is noteworthy for its relatively small heads (4-5 mm high) and short pappus (ca. 3 mm long), otherwise it is very similar to the species. Eupatorium rosei (King & H. Robins.) B. Turner, comb. nov. BASIONYM: Cronqutstianthus rose: King & H. Robins., Phytologia 51:182. 1982. TYPE: ECUADOR. Azuay: vicinity of Cuenca, 17-24 Sep 1918, J.N. Rose, A. Pachano & G. Rose 22919 (HOLOTYPE: US). Turner: Recension of Cronqutstianthus 175 This taxon is suspiciously close to Eupatortum chamaedrifolium and is said to differ from the latter in having smaller heads with fewer florets (ca. 12 vs. 18-20) and more strongly costate involucral bracts. Since E. roseum (King & H. Robins.) B. Turner occurs in the same area as EF. chamaedrifolium, the latter being quite variable, it is likely that these will ultimately prove synonymous. Eupatorium urubambense B.L. Robins., Proc. Amer. Acad. Arts 55:38. 1919. Cronquistzanthus urubambensis (B.L. Robins.) King & H. Robins., Phy- tologia 23:411. 1972. TYPE: PERU. Cusco: in the valley of Ymay [7}, without date, Urubamba, Pentland s.n. (HOLOTYPE: K; Photoholo- type: GH!; Fragment holotype: GH!). Eupatorium urubambense B.L. Robins. var. mantarense B.L. Robins., Contr. Gray Herb. 90:33. 1930. TYPE: PERU. Huancavelica: Mantaro Valley, near Mejorada, 2800-2900 m, 21 Mar 1926, W’eber- bauer 7614 (HOLOTYPE: F!; Photoholotype: NY!; Isotype: GH). Erect shrubs or subshrubs 0.5-3.5 m high. Leaves mostly 4-6 cm long, 1.5-2.5 cm wide; petioles 0.5-1.0 cm long; blades narrowly triangular, triner- vate from the base, densely puberulent beneath, the margins crenulodentate. Heads bluish, numerous in terminal cymules, the ultimate peduncles mostly puberulent, 1-5 mm long. Involucres 5-7 mm high, the bracts 3-4 seriate, the inner series glabrous, gradually narrowed and obtuse apically. Florets 10-15 per head. Corollas ca. 5 mm long. Achenes 3-4 mm long, hispidulous, the pappus of 30-40 bristles 4-5 mm long. REPRESENTATIVE SPECIMENS: PERU. Ayacucho: Ayacucho, 1951, Soukup 4010 (F). Cusco: 13 km N of Pisaq on road to Ollantaytambo, scree slope and adjacent cliffs overlooking Rio Urubamba, 2900 m, 27 Apr 1977, Gentry 19802 (F, MO); Chicon Canyon, 2800-3000 m, 7 May 1939, Vargas (Goodspeed) 11048 (F); Quebrada de Paucartambo, 3000 m, 20 Jun 1940, Vargas 1936 (F, MO). Eupatorium volkensu Hieron. in Urban, Bot. Jahrb. Syst. 40:370. 1908. Cronqguistianthus volkensi (Hieron.) King & H. Robins., Phytologia 23: 411. 1972. TYPE: PERU. without locality, without date, W’eber- bauer 4850 (HOLOTYPE: B, destroyed; Photoholotypes: F!, MO!, NY!, TEX): Scandent suffruticose perennial herb or shrublet to 1 m high. Leaves mostly 6-12 cm long, 0.8-2.0 cm wide; petioles 1-5 mm long; blades narrowly lanceolate to linear lanceolate, pinnately and reticulately veined beneath, moderately to densely puberulent, the margins crenulate to nearly entire. Heads blue or purplish, numerous in tight terminal cymules, the ultimate peduncles mostly 0-8 mm long. Involucres 7-8 mm high, the bracts 4-5 seriate, the inner series 176 PAY TOLOGLA volume 70(3):158-177 _ VENEZUELA” i" = a. a i < : a ‘ Sa ee COLOMBIA °: fai ‘ i pooner’ a b rae ee EUPATORIUM Oniveum @ pseudoriganoides Srosei Aurubambense Ovolkensii | pti aif ‘ a ‘ = ——ae’ a 3 -_ —-*=z ‘ ‘ 3 = Les ‘ \ — ee ‘ = = .s 5! => —— S ae ——— S—= —— ——_—s | SS see = > = Fig. 4. Distribution of Eupatorium March 1991 Turner: Recension of Cronquistianthus 177 glabrous or nearly so with obtuse or rounded apices. Florets ca. 12 per head. Corollas 5-6 mm long. Achenes 3-4 mm long, glabrous to sparsely hispidulous above, the pappus of 40-50 bristles, 5-6 mm long. REPRESENTATIVE SPECIMENS: PERU. Ayacucho: Huanta, 2800 m, 19-21 May 1929, Kilhp 23321 (NY). Cusco: about the ruins of Sacsahuaman, 3500 m, 6 May 1983, Beck 8353 (F); trail from Chincheros plaza to Antakillga, 3700 m, 13 Jan 1982, Davis, et al. 1414 (F); near San Jeremino, 3200 m, 14 Feb 1975, Mayolo 106 (F); Sacsahuaman, above Cusco, 3500-3600 m, 24 Apr 1925, Pennell 13542 (F); Quebrado de Juayoccari - Laguna de Yanaccocha, 2900-3860 m, 20 Dec 1988, Tupayachi 789 (MO). ACKNOWLEDGMENTS This study is based upon approximately 300 herbarium sheets from the following herbaria: F, GH, LL, MO, NY, TEX. I am grateful to Guy Nesom and Linda Escobar for reviewing the manuscript. LITERATURE CITED King, R. & H. Robinson. 1972. Studies in the Eupatorieae (Asteraceae). LXXV. A new genus, Cronguistianthus. Phytologia 23:409-412. . 1984. Studies in the Eupatorieae (Asteraceae). CCXIX. A new species of Cronquistianthus. Phytologia 56:252-254. . 1987. Cronquistianthus, in Monographs Syst. Bot. Missouri Bot. Gard 22:366-368. Phytologia (March 1991) 70(3):178-184. PROSTRATE GROWTH IN CAULESCENT ANDEAN ROSETTES OF COESPELETIA (ASTERACEAE, HELIANTHEAE) Francisco L. Pérez Department of Geography, University of Texas, Austin, Texas 78712 U.S.A. ABSTRACT This note reports on the prostrate growth of caulescent Andean rosettes ( Coespeletia tirmotensis Cuatr.) in the Venezuelan Andes. These plants normally grow erect and have a stem up to 3.5 m tall, but they can be tilted and toppled by frequent soil cryoturbation. Prostrate individuals tend to die as a result of toppling, but some can survive and continue growing in a decumbent position for a long time. KEY WORDS: Coespeletia trmotensis, Espeletiinae, Asteraceae, Heliantheae, stem morphology, paramo, Andes, Venezuela INTRODUCTION The north Andean paramos are commonly dominated by “giant” caulescent rosettes of Espeletia and related genera (subtribe Espeletiinae, Cuatrecasas 1976). In Venezuela, many high paramos are occupied by dense stands of Coespeletia timotensis Cuatr., which are found up to 4600 m (Pérez 1987a). Superficial soil movement due to recurrent frost activity can be a significant disturbance factor in the high paramo, both preventing the establishment of rosette seedlings and increasing the risk of adult mortality. Tall rosettes—up to 3.5 m-of C. timotensis seem to be easily destabilized by soil movement because their root systems are shallow, usually < 25-30 cm (Monasterio 1980; Pérez 1987b). Soil cryoturbation commonly results in progressive tilting of the larger individuals, which eventually become completely prostrate on the soil surface and then die (Goldstein, ef al. 1985). Some plants may escape this fate by germinating next to large rocks embedded in the ground, which can provide a stable anchorage (Pérez 1987b). Tilting of caulescent rosette plants also occurs in similar Afroalpine areas (Mahaney 1980; Beck, et al. 1983). 178 Pérez: Prostrate growth in Coespeletia 179 SURVIVAL OF TOPPLED ROSETTES In the Paramo de Piedras Blancas (Venezuela), I have often found large toppled rosettes lying dead on the ground; in most cases, dead stems are con- centrated in a few stands (cf. Monasterio 1980). These high mortality stands are more common on steep slopes and at higher elevations, where soil insta- bility and frost activity are presumably greater. However, prostrate rosettes are not always dead, and | have found several examples of large, decumbent, living plants. Figure 1 shows a recently toppled plant, about 180 cm long, lying on an 18° slope at 4470 m. Several dead stems were present in the vicinity, and the prostrate rosette itself was flanked by the remains of two dead plants. When visited, the slope was completely covered by striated soils. nubbins, and nonsorted stripes (Schubert 1975; Pérez 1987c); all are signs of recent needle ice activity, which is widespread in this paramo. The leaf crown of the decumbent rosette had phototropically tilted upwards about 35°; the plant showed no signs of senescence and seemed to be growing vigorously. Near this plant there were several more live fallen rosettes. One of them (Figure 2) showed a sharp, about 90° inflection along its stem, which grew horizontally for 130 cm and then vertically for 95 cm more (crown included). The roots of both rosettes were partially broken as a result of toppling, but clearly maintained enough of a connection with the soil to allow for continuous growth. In both cases, the plants were attached to the ground only at the stem base, and the trunks could be easily pulled and gently swung sideways; thus, no adventitious roots had been produced. This was expected, since Coespeletia timotensis appears to lack the capacity for vegetative reproduction (Monasterio 1980; Pérez 1987b). A nearby group of live, prostrate plants was also examined at 4450 m. The slope here is steeper, about 26°, and has repeatedly shown, for the past eleven years, widespread evidence of needle ice growth, miniature debris flows, and superficial solifluction, which produces small “mud grooves” (Hastenrath 1973, 1989, pers. comm.). Measurements of needle ice activity on this slope for nearly six years (Pérez 1987c) indicate that a shallow soil layer < 10 cm thick moves rapidly downhill, at rates of up to 15-18 cm/yvear. One decumbent rosette had a stem inflection of > 70°; the horizontal section near the ground was 140 cm long, the vertical one 105 cm (Figure 3). Fine debris, stones, and pieces of dead stems were piled upslope from the prostrate stem, where the ground stood up to 20 cm higher than on its downhill side. This attests again to the intensity of soil disturbance on this steep slope. The morphological adaptations discussed here are by no means unique to Coespeletia timotensis or to Piedras Blancas. Although no similar reports have been found in the literature, Cuatrecasas (1990, pers. comm.) has occasionally seen decumbent Espeletiinae rosette plants with their leaf crowns erect and in 180 PHYTOLOGIiA volume 70(3):178-184 March 1991 Figure 1 (above). Prostrate, recently toppled Coespeletia timotensis at 4470 m; the plant is 180 cm long. Note several dead rosettes in the background, and the leaning trunk of a large plant on the right side of the picture. Jan. 1990. Figure 2 (below). Rosette specimen with sharply inflected stem, on same general location as that of Figure 1. The measuring stick is 180 cm long. Noie two dead plant stems in the background, and the nubbin texture covering the soil surface. Jan. 1990. Pérez: Prostrate growth in Coespeletia 181 Figure 3. Decumbent Coespeletia timotensis at 4450 m. The horizontal section is 140 cm long, the vertical one 105 cm. Note the piling of mineral and organic debris on the upslope side. Dec. 1985. 182 PAY ROL: OG FA volume 70(3):178-184 March 1991 a vertical position after bending about 90° (c!. Figures 2, 3). He noticed this in a 1.2 + 0.3 m long (horizonta] + vertical sections) specimen of Espeletia ar- belaezui Cuatrecasas in the Paramo de Guanti-a (Boyaca, Colombia), and also in a 1.0+1.2 m Jong Ruzlopezia figuerrasi: (Cuatrecasas) Cuatrecasas (Cuatre- casas 1987) in the upper Andean forest of Siexra Santo Domingo (Venezuela). Cuatrecasas’ observations agree with mine in that prostrate individuals with sharp stem inflections are always much scarcer than toppled, living plants lying on the ground with an unmodified stem (1.¢.. Figure 1). This indicates that most rosettes probably do not survive topplirg, and that only a few continue growing afterwards. DISCUSSION AND CONCLUSIONS The tilting model of Goldstein, et al. (1955) proposes that adult rosettes become gradually tilted until they are eventuclly prostrate on the soil surface. The examples presented here indicate that this is not always true. The sharp stem inflections observed in some surviving prostrate plants suggest that top- pling was a rapid, discrete event. This may well have taken place following a period of progressive tilting, after which thresholds of root strength and/or soil stability were surpassed. The large, heavy rosettes fell to the ground suddenly, and the apical bud of those which survived became reoriented through nega- tive geotropism and phototropic response, producing bent trunks afterwards. Sharp stem deflections would not be expected if tilting had always been grad- ual; a progressive curving of the stem should probably result instead. Further field observations support this hypothesis. In September 1980, a 230 cm tall rosette only 3 m away from that shown in Figure 3 was leaning slightly (ca. 25°) from the vertical. When the slope was revisited 17 months later (Febru- ary 1982), this plant was found prostrate on the ground with its leaf crown tilted upwards ca. 45° (cf. Figure 1). Thus, the rosette toppled completely in a period of 1.5 years, at most. However. this plant did not survive the disturbance event; when the slope was revisited in 1990, the prostrate stem was dead, having become detached from the ground and rolled downslope a few meters. The observations reported here show that toppling is not necessarily fol- lowed immediately by plant death. Coespeletia timotensis grows at about 15 mm /year (Smith 1974), thus, the largest prostate individuals with bent stems continued growing for 50 to 70 years after toppling. Goldstein, ed al. (1985) ar- gue that death of fallen rosettes would occur Cue to root exposure and partial breakage, and the consequent drop in water and nutrient uptake. Also, the water storing pith tissue inside the stem woulc experience lower temperatures at the ground surface. Assuming not al] the roots are irreparably damaged, Pérez: Prostrate growth in Coespeletia 183 several effects could counterbalance the exposure of the stem pith to the lower temperatures at the soil surface. First, the stem would be insulated from freez- ing along its underside; second, gradual piling of debris on the uphill side of plants falling perpendicular to the slope would partially bury the stem, fur- ther contributing to insulating it (Figure 3) (cf. Pérez 1987c, photo 2). In addition, if several plants topple synchronously, the stems at the edges of the fallen group may help insulate the prostrate plants at the center (Figure 1), further minimizing their exposure to extreme cold at least until the dead stems decompose. It must be pointed out that the assumption that soil cryoturbation is the sole cause of rosette toppling is simplistic. Although frost induced creep and gelifluction are certainly significant geomorphic agents in this paramo, many other processes, including debris- and mudflows, surface runoff, throughflow, dry grain flows, rockfalls, and cattle disturbance can affect substrate stability (Pérez 1987b, 1987c) and thus cause plant tilting and toppling. Survival and continued growth of prostrate caulescent rosettes in the periglacial paramo occurs only occasionally, but merits attention as a significant morphological adaptation which partially counteracts the noxious effects of ground distur- bance on the tallest plants. ACKNOWLEDGMENTS The Research Institute (University of Texas, Austin) provided a travel grant. I am grateful to O. Vera and A. Lucchetti for their help in the field, I especially thank J. Cuatrecasas (Smithsonian Institution) for kindly sharing valuable field observations and for reading the manuscript, which also profited from reviews by B.L. Turner and I.L. Bergquist (both at the University of Texas). S. Hastenrath (University of Wisconsin, Madison) provided interesting comments on several paramo geomorphic features. LITERATURE CITED Beck, E., R. Scheibe, & M. Senser. 1983. The vegetation of the Shira Plateau and the western slopes of Kibo (Mt. Kilimanjaro, Tanzania). Phyto- coenologia 11:1-30. Cuatrecasas, J. 1976. A new subtribe in the Heliantheae (Compositae): Espeletiinae. Phytologia 35:43-47. . 1987. Clave diagnostica de las especies de Ruilopezia (Espeletiinae, Heliantheae, Compositae). Anales Jard. Bot. Madrid 44:401-419. 184 PHYTO Lb OvGt A volume 70(3):178-184 March 1991 Goldstein, G., F. Meinzer, & M. Monasterio. 1985. Physiological and me- chanical factors in relation to size-dependent mortality in an Andean giant rosette species. Oecol. Plant. 6:263-275. Hastenrath, S. 1973. Observations on periglacial morphology of Mts. Kenya and Kilimanjaro, East Africa. Z. Geomorph. 16:161-179. Mahaney, W.C. 1980. Las formaciones vegetales de ]os paramos de Venezuela. Pp. 93-158 In M. Monasterio. Estudios Ecoldgicos en los Padramos Andi- nos. Univ. de Los Andes. Mérida, Venezuela. Pérez, F.L. 1987a. Soil moisture and the upper altitudinal limit of giant paramo rosettes. J. Biogeogr. 14:173-186. . 1987b. Needle-ice activity and distribution of stem-rosette species in a Venezuelan paramo. Arct. Alp. Res. 19:135-153. . 1987c. Downslope stone transport by needle ice in a high Andean area (Venezuela). Rev. Geomorph. Dynam. 36:33-51. Schubert, C. 1975. Glaciation and periglacial morphology in the northwest- ern Venezuelan Andes. Eiszeitalter Ggw. 26:196-211. Smith, A.P. 1974. Population dynamics and life form of Espeletia in the Venezuelan Andes. Ph.D. Dissertation, Duke University, Durham. Phytologia (March 1991) 70(3):185-187. TAXONOMIC ADJUSTMENTS IN BOUCHEA AND LIPPIA (VERBENACEAE) OF MEXICO Guy L. Nesom Department of Botany, University of Texas, Austin, Texas 78713 U.S.A. ABSTRACT New nomenclatural combinations are proposed for the following taxa of Verbenaceae occurring in México: Bouchea longiflora (Mold.) Nesom and Lippia carterae (Mold.) Nesom. The distribution of B. longifiora and its closest relative, B. spathulata Torrey, are mapped. KEY WORDS: Verbenaceae, Bouchea, Lippia, Mexico Curation of LL,TEX collections of Mexican Verbenaceae has revealed that several taxa should be accorded a higher rank. The rationale with necessary nomenclatural combinations are provided below. Other comments on taxon- omy of the family, primarily on the occurrence of numerous synonyms, will be forthcoming as part of a “Checklist of Verbenaceae in México” (Nesom, ms.). 1. Bouchea longiflora (Mold.) Nesom, comb. et stat. nov. BASIONYM: Bouchea spathulata Torrey var. longiflora Mold., Phytologia 23:210. 1972. TYPE: MEXICO. Coahuila: Sierra de la Fragua, 1-3 Sep 1941, 1.M. Johnston 8724 (HOLOTYPE: TEX!). Bouchea longiflora (Mold.) Nesom clearly is closely related to B. spathulata Torrey, but the former differs in its much longer calyces (14-17 mm vs. 10- 12 mm) and corolla tubes (30-45 mm vs. 15-25 mm). Although each taxon displays some variability in these features, there is no specimen that could be interpreted as morphologically intermediate. The two species probably _ are reproductively isolated by the marked difference in corolla length, which suggests a corresponding difference in pollinators. They also appear to be completely separated in geographical distribution, although their ranges very closely approach each other at several points (Map 1). Bouchea longiflora is the only species in the genus with such a distinctly elongated corolla tube, and it is probable that it has arisen as an evolutionary derivative of B. spathulata, particularly in view of the strong similarity in 185 186 PHY TOLOGIA volume 70(3}:185-187 March 1991 TEX a B. longiflora]\ spathulata Map 1. Distribution of Bouchea longiflora and B. spathulata. All data points are from LL, TEX specimens, except for three localities in Texas for B. spathu- lata added from Moldenke (1940). Abbreviations are for Texas (TEX) and the Mexican states Coahuila (COA), Chihuahua (CHI), Durango (DUR), and Nuevo Leon (NUE). Nesom: Taxonomic adjustments in Bouchea and Lippia 187 other morphological features between these two species. Both species flower from about Apri] to November and both occur in essentially the same type of habitats of matorral or chaparral at the same elevations (ca. 1200-1900 meters), although B. spathulata, with more collections available, has been recorded at somewhat higher and lower sites. 2. Lippia carterae (Mold.) Nesom, comb. nov. BASIONYM: Lippia alba (Mill.) N.E. Britt. var. carterae Mold., Phytologia 7:430. 1961. TYPE: MEXICO. Baja California Sur: Sierra de la Giganta, dry rocky bed of Arroyo Santo Domingo, Rancho El] Horno (northeast of San Xavier), 17 Mar 1960, A. Carter & R. Ferris 3864 (HOLOTYPE: UC; Isotype: TEX!). Lippia alba (Mill.) N.E. Britt. is a widespread and somewhat variable species, but the plants regarded here as L. carterae (Mold.) Nesom are clearly outside the bounds of variation of any population of L. alba. Most promi- nently, while the pubescence of L. alba is strigose, composed of stiff, relatively thick based hairs, L. carterae produces a dense, close tomentum of fine, white, crinkled hairs on the stems, leaves, and floral bracts. The leaf surfaces have an understory of fine tomentum, with much longer, straight, stiffer, “alba-like” hairs intermixed on the upper surface and along the veins of the lower surface. Further, the corollas of L. carterae are distinctly yellow, as noted by Mold- enke in the original description, in contrast to the white to pink or purphsh corollas of L. alba. Lippia carterae is endemic to Baja California Sur although Wiggins (1980) treated it as L. alba. The latter is a common species from southern Texas through México and into Central America; it is known from the southwestern Mexican states of Sinaloa, Nayarit, Jalisco. and Colima, but over its whole range it apparently grows in much more humid habitats than does L. carterae. There are no records of the occurrence of L. alba in Baja California. ACKNOWLEDGMENTS I thank Dr. B.L. Turner and Dr. B.B. Simpson for their review and com- ments on the manuscript. LITERATURE CITED Moldenke, H.N. 1940. A monograph of the genus Bouchea. II. Feddes Repert. Spec. Nov. Regni Veg. 49:91-139. Wiggins, I.L. 1980. Flora of Baja California. Stanford Univ. Press, Stanford, California. Phytologia (March 1991) 70(3):188-193. RANGE EXTENSIONS IN THE SIERRA MADRE OCCIDENTAL, CHIHUAHUA, MEXICO Joseph E. Laferriere Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, Arizona 85721 U.S.A. ABSTRACT Opuntia durangensis Britt. & Rose, Smilar moranensis Mart. & Gal., Muhlenbergia durangensis Herrera vel aff., Erigeron delphinifolius Willd., and Browallia sp. are reported from Chihuahua for the first time. Range extensions within Chihuahua are reported for Jlez tolucana Hemsl. and Leibnitzia occimadrensis Nesom. All were collected in the vicinity of Nabogame, Municipio Temésachi, Chihuahua, México, in the Sierra Madre Occidental. RESUMEN Opuntia durangensis Britt. & Rose, Smilaz moranensis Mart. & Gal., Muhlenbergia durangensts Herrera vel aff., Erigeron delphintfolius Willd., y Browallia sp. se relatan de Chihuahua por primera vez. La dis- tribucién geografica de Jlez tolucana Hems]. y Leibnitzia occimadrensis Nesom se extienden hasta esta parte de Chihuahua. Todos se recolec- taron de la vicinidad de Nabogame, Municipio Temésachi, Chihuahua, México, en la Sierra Madre Occidental. KEY WORDS: México, Opuntia, Smilaz, Muhlenbergia, Browallia, Tlez, Leibnitzia 188 Laferriere: Range extensions in Chihuahua, México 189 INTRODUCTION Intensive collecting by the author in the Mountain Pima community of Nabogame in the Sierra Madre Occidental of Chihuahua, México, between Oc- tober 1986 and November 1988 has resulted in the collection of several species not previously reported from Chihuahua. These include Opuntia durangensis Britt. & Rose (Cactaceae), Smilaz moranensis Mart. & Gal. (Smilacaceae), Muhlenbergia durangensis Herrera vel aff. (Poaceae), Erigeron delphinifolius Willd. (Asteraceae), and Browallia sp. (Solanaceae). In addition, range exten- sions within Chihuahua are reported for Jler tolucana Hemsl. (Aquifoliaceae) and Leibnitzia occimadrensis Nesom (Asteraceae). Each is known from south- ern Chihuahua but not the northern or central parts of the state. Opuntia durangensis, new to Chihuahua Opuntia durangensis Britt. & Rose, Smiths. Misc. Coll. 50:518. 1908. Specimens examined: MEXICO. Chihuahua: Municipio Temésachi: Nabo- game, 28° 30’ N, 108° 30’ W, elev. 1800 m, fruit yellow, oak woodland, 11 Sep 1987, Laferriére 1119 (ARIZ, ASU, MEXU); 1 km S of Nabogame, field, fruit red, 20 Sep 1987, Laferriére 1198 (ARIZ, ASU, MEXU); Nabogame, oak wood- land, in flower, 17 Jun 1988, Laferriére 1438 (ARIZ, ASU, MEXU): Nabogame, canyon under Cupressus arizonica E. Greene, in flower, 18 Jun 1988, Laferriere 1444 (ARIZ, ASU, MEXU); Nabogame, oak woodland, fruit yellow, 28 Aug 1988, Laferriére 1846 (ARIZ, ASU, MEXU); Nabogame, canyon under Cupres- sus arizonica, fruit red, 18 Oct 1988, Laferriére 2107 (ARIZ, ASU, MEXU); 2 km S of Nabogame, field, fruit yellow, 18 Oct 1988, Laferriére 2114 (ARIZ, ASU, MEXU); same site, fruit red, Laferriére 2115 (ARIZ, ASU, MEXU). This species is known previously only from Durango (Britton & Rose 1963; Bravo-Hollis 1978; Backeberg 1982). This report constitutes a range extension not only for the species but also for the series Macdougalanae. Both red fruited and yellow fruited specimens are present in Nabogame. The original description of the species describes the fruits as “white or red” (Britton & Rose 1908). The local Pima names for the yellow and red forms are “duraznillo blanco” and “duraznillo colorado;” the use of the word “white” in Britton & Rose’s description may reflect this folk taxonomy. 190 PY 70 -L-0. GI A volume 70(3):188-193 March 1991] Smilaz moranensis, new to Chihuahua Smilaz moranensis Mart. & Gal., Bull. Acad. Brux. 9(2):389. 1842. Specimens examined: MEXICO. Chihuahua: Municipio Temésachi: South side of Rio Yepachi, 4 km S of Nabogame, under Acer grandidentatum Nutt. and Cupressus arizonica, 6 Dec 1987, Laferriére 1823 (ENCB, UC); same site, 3 Nov 1988, Laferrzére 2252 (ANSM, ARIZ, CHAPA, MEXU, SD, TEX). The site from which this species was collected is more mesic than much of the surrounding area. The plant is known from Sonora, Sinaloa, and Coahuila south to Oaxaca, but not previously from Chihuahua (Morton 1962). Browallia sp. nov. ined., new to Chihuahua Browallia sp. nov. ined. Specimens cited: MEXICO. Chihuahua: Municipio Temésachi: 1 km N of Nabogame, field, 29 Aug 1987, Laferriére 1003 (ARIZ, VDB); 3 km N of Nabogame, sunlit rocky area above waterfall, 27 Jul 1988, Laferriére 1562 (ARIZ, VDB); Nabogame, oak woodland, 8 Aug 1988, Laferriére 1629 (ARIZ, MEXU, VDB). This is the first report of this genus from northern México. It is previously known from southern México and Central America to Bolivia and the Antilles (D’Arcy 1973, 1978; Gentry & Standley 1974; Hunziker 1979). The specimens cited will soon be described as a new taxon (W. D’Arcy, R. Van Devender & P. Jenkins, personal communication). Muhlenbergia durangensis vel aff., new to Chihuahua Muhlenbergia durangensis Herrera, Phytologia 63:457. 1987. Specimens examined: MEXICO. Chihuahua: Municipio Temédsachi: 1 km NE of Nabogame, pine-oak forest, 25 Mar 1988, Laferriére 1416 (ANSM, ARIZ, MEXU, TEX). This species is known only from Durango (Herrera 1987). The Nabogame specimen differs from the type description in having smaller glumes (3-4 mm Laferriére: Range extensions in Chihuahua, México 191 long) and longer ligules (3.5-4.5 mm long). The Nabogame specimens are either Muhlenbergza durangensis or a new taxon. More collections would be necessary to determine which. Erigeron delphinifolius, new to Chihuahua Erigeron delphinifolius Willd., Enum. Hort. Berol. 2:873. 1809. Specimens examined: MEXICO. Chihuahua: Municipio Temésachi: 2 km N of Nabogame, oak woodland on steep slope, 6 Nov 1988, Laferréere 2267 (ARIZ, MEXU, TEX). This species is known from Durango to Michoacan and Puebla (Nesom 1989). This is the first published report from Chihuahua. Range Extension of Jlez tolucana Tlez tolucana Hemsl., Diagn. Pl. Nov. 5. 1878. Specimens examined: MEXICO. Chihuahua: Municipio Temésachi: 1 km E of Nabogame, along streambank under Cupressus arizonica, 25 Aug 1987, Laferriére 948 (ANSM, ARIZ, CHAPA, MEXU, TEX). Nabogame is the most northerly known location of this species. It is pre- viously reported from the Sierra Charuco 60 km to the south of Nabogame, as well as from southern México (Gentry 1942). Range Extension of Leibnitzia occimadrensis Leibnitzia occtmadrensis Nesom, Brittonia 35:130. 1983. Specimens examined: MEXICO. Chihuahua: Municipio Temésachi: South side of Rio Yepachi, 4 km S of Nabogame, under Acer grandidentatum Nutt. and Cupressus arizonica, 18 Oct 1988, Laferriére 2136 (ARIZ); 1 km E of Nabogame, under C. arizonica, 28 Oct 1988, Laferriére 2227 (MEXU, TEX). This is a range extension of 300 km for this species, previously known from Sinaloa and extreme southern Chihuahua (Nesom 1983). The site for this collection is the same as that for Smilaz moranensis. 192 PHYTOLOGIA volume 70(3):188-193 March 1991 ACKNOWLEDGMENTS Fieldwork was supported by an NSF doctoral dissertation grant to the author and Willard Van Asdall. I thank Charles Mason, Rebecca Van Deven- der, Dale McNeal, Phil Jenkins, David Brunner, Guy Nesom, Wilham D’Arcy, Charlotte Reeder, John Reeder, and Richard Felger for assistance in identify- ing specimens and review of the manuscript. REFERENCES Backeberg, C. 1982. Die Cactaceae, Handbuch der Kakteenkunde, Band I: Ernleitung und Beschreibung der Pereskioideae und Opuntioideae. Gus- tav Fischer Verlag, Stuttgart. Britton, N.L. & J.N. Rose. 1908. A preliminary treatment of the Op- untioideae of North America. Smithsonian Miscellaneous Collections 50(4):503-539. . 1963. The Cactaceae, second edition, Dover, New York. Bravo-Hollis, H. 1978. Las Cactdceas de Mézico. Universidad Autonoma de México, Mexico, D.F. D’Arcy, W.G. 1973. Flora of Panama, part IX: Solanaceae. Annals of the Missouri Botanical Garden 60:573-780. . 1978. A preliminary synopsis of Salpiglossis and other Cestreae (Solanaceae). Annals of the Missouri Botanical Garden 65:698-724. Gentry, H.S. 1942. Rio Mayo Plants. Carnegie Institute of Washington Publication 527, Washington. Gentry, J.L. & P.C. Standley. 1974. Flora of Guatemala: Solanaceae. Fiel- diana: Botany 24(part X, numbers 1+2):1-151. Herrera Arrieta, Y. 1987. Una nueva especie de Muhlenbergia (Gramineae) del Estado de Durango. Phytologia 63:457-459. Hunziker, A.J. 1979. South American Solanaceae: a synoptic survey. In: J.G. Hawkes, R.N. Lester, & A.D. Skelding (eds.), The Biology and Taz- onomy of the Solanaceae, pp. 49-85. Academic Press, New York. McVaugh, R. 1974. Flora Novo-Galiciana. Contributions of the University of Michigan Herbarium 12, Ann Arbor. Laferri€ére: Range extensions in Chihuahua, México 193 Morton, C.V. 1962. A reexamination of Mexican Smilaz (Smilacaceae). Brittonia 14:299-309. Nesom, G.L. 1983. Biology and taxonomy of American Lezbnitzia (Aster- aceae: Mutisieae). Brittonia 35:126-139. . 1989. Taxonomy of Erigeron sect. Polyactts (Compositae: Aster- eae). Phytologia 66:415-455. Phytologia (March 1991) 70(3):194-208. NOMENCLATURAL NOTES FOR THE NORTH AMERICAN FLORA - V John T. Kartesz & Kancheepuram N. Gandhi The North Carolina Botanical Garden, Dept. of Biology, Coker Hall, University of North Carolina, Chapel Hill, North Carolina 27599-3280 U.S.A. ABSTRACT The authorship of Actinostachys pennula (Sw.) Hook., Athyrium alpestre (Hoppe) Milde, and Equisetum x trachyodon (A. Braun) W.D.J. Koch and the taxonomy of Athyrium distentifollum Tausch ez Opiz and Athyrium americanum (Butters) Maxon are discussed. The name Equisetum x trachyodon is replaced by Equisetum x mackati (Newm.) Brichan, and the name Phyllitis scolopendrium (L.) Newm. var. emargi- nata Fern. ez Boivin is considered to be a nomen nudum. Ten new com- binations are proposed: Antrophyum intramarginale (Baker ez Jen- man) Kartesz & Gandhi; Asplenium scolopendrium L. var. amer- icanum (Fern.) Kartesz & Gandhi; Huperzia x bartleyi (Cusick) Kartesz & Gandhi; Huperzia x buttersii (Abbe) Kartesz & Gandhi; Huperzia x helleri (Herter) Kartesz & Gandhi; Huperzia mannii (Hillebr.) Kartesz & Gandhi; Huperzia occidentalis (Clute) Kartesz & Gandhi; Huperzia x serrata (Thunb. ez Murray) Trevisan var. dentata (Hillebr.) Kartesz & Gandhi; Lycopodiella cernua Pichi Sermolli var. curvata (Sw.) Kartesz & Gandhi; and Pecluma campto- phyllaria (Fee) M. Price ssp. abbreviata (Evans) Kartesz & Gandhi. KEY WORDS: Floristics, Nomenclature, Ferns, Pteridophyta, As- pleniaceae, Drvopteridaceae, Equisetaceae, Lycopodiaceae, Poly- podiaceae, Schizaeaceae, Vittariaceae, Actinostachys, Antrophyum, Asplenium, Athyrium, Equisetum, Huperzia, Lycopodiella, Lycopo- dium, Pecluma, Phyllitis, Polypodium, Polytaenium, and Vittaria. 194 Kartesz & Gandhi: Nomenclature of North American flora - V 195 Introduction Continuing with the “NOMENCLATURAL NOTES FOR THE NORTH AMERICAN FLORA” (Kartesz & Gandhi 1989, 1990a, 1990b, 1990c), a fifth note in the series is presented here in the hope of advancing our understanding of North American plant names. PTERIDOPHYTES Over the past few decades, our understanding of pteridophyte character- istics (including habit, habitat, external morphology, anatomy, chromosome number, spore and gametophyte structure, and hybridization) has increased dramatically. The application of modern techniques, such as genome anal- ysis, isozyme gel electrophoresis, and chloroplast DNA studies have further advanced our knowledge of fern evolution. The rapid accumulation of system- atic data has led to significant changes in pteridophyte classification, including splitting of large, heterogeneous genera and lumping of smaller, homogenous ones. For the North American flora, we have atterrpted to follow most of the revisions that we consider to be appropriate. In this connection, we provide the following taxonomical and nomenclatural notes. ASPLENIACEAE Asplenium and Phyllitis According to Lellinger (1985), the genus Phyllitzs Hill. comprises about eight species, mostly found in the tropics and subtropics (excluding South America). He remarked that with its double sori and unique lamina shape, Phyllitis can be differentiated from all the North American species of the genus Asplenium L. However, these two genera hybridize freely in Europe suggesting that they may be more closely related than some of the genera traditionally included within Asplentum. Rather than recognizing interbreeding genera, we concur with Proctor (1989, p. 217) that Asplentum is a large, homogenous genus that “is not readily split into discontinuous subgeneric groups, ....” Hence, we recognize Asplenium in the broad sense (including Phyllitis) for the North American flora and propose the following new combination for the American hart’s-tongue. 196 PHYTOLOGIA volume 70(3):194-208 March 1991 Asplenium scolopendrium L. var. americanum (Fern.) Kartesz & Gandhi, comb. nov. BASIONYM: Phyllitis scolopendrium (L.) Newm. var. amer- tcana Fern., Rhodora 37:220. 1935. TYPE: CANADA. Ontario: Grey Co., Inga Falls, 19 Jun 1934, Fernald, Thompson, & Wright s.n. (GH). Phyllitts scolopendrium (L.) Newm. var. emarginata Fern. ez Boivin, Nat. Canad. 93:268. 1966. nom. nud. DRYOPTERIDACEAE Athyrium alpestre and A. distentifolium Fuchs (1974) presented a detailed discussion on the correct name of the European alpine lady fern. Until 1956, this fern was known by the name Athyrium alpestre, which was based on Aspidium alpestre Hoppe (1805). The authorship of the name Athyrium alpestre was credited to several authors, in- cluding “Rylands ez Moore” and “(Hoppe) Rylands” (Fernald 1929). However, Fuchs (Pp. 181 and 189) indicated “(Hoppe) Milde” to be the author. In 1956, E. Janchen corrected the name Athyrium alpestre (Hoppe) Milde (1867); non Athyrium alpestre De Clairv. (1811) to Athyrium distentifolium (fide Fuchs; p. 182). Although Athyrium alpestre (Hoppe) Milde and Athyrium alpestre De Clairv. share the same specific epithet, these two names were based on different types. According to Fuchs (1974, pp. 181 and 187-188), Athyrium alpestre De Clairv. was based on a form of A. filiz-femina (L.) Roth, whereas the name Athyrzum alpestre (Hoppe) Milde was based on Aspidium alpestre Hoppe (1805), which is possibly a superfluous name (Fuchs 1974, pp. 186-187). Regarding De Clairville’s taxon, we quote from Fuchs (1974, pp. 187-188): “The taxonomic position of this Athyrium alpestre Clairv., taxon entirely independent from As- pidium alpestre Hoppe, is proven by a specimen in De Clairville’s own herbar- ium now at Z (cf. pl. IIb). This herbarium specimen is labeled in De Clairville’s handwriting: ‘Athyrium alpestre - Polypodium. Linn. hall. no. 1703 Les bois montagneux,’ and belongs clearly to Athyrium Filiz-fernina (L., sub Polypo- dio) Roth, as someone had recognized already earlier when annotating De Clairville’s specimen with the note ‘appartient a Aspl. filiz-foemina”’ Lellinger (1981, p. 91) rejected Fuchs’ analysis and concluded that Athyrium alpestre De Clairv. was indirectly based on Aspidium alpestre Hoppe. With this assertion, he reinstated the name “Athyrium alpestre (Hoppe) De Clairville.” The following is quoted from Lellinger: “According to R.M. Tryon, Jr. (pers. comm.), De Clairville’s work is an excursion flora in which the introduction states that it is not a technical work. De Clairville’s intent was not to name a Kartesz & Gandhi: Nomenclature of North American flora - V 197 new species, but to transfer Hoppe’s name. Therefore, it seems clear that the epithet alpestre (Hoppe) de Clairville should be reinstated.” With reference to Tryon’s comments on De Clairville’s excursion flora, we invoke ICBN Art. 29.4 (Greuter 1988). According to this article, new names that were proposed in nonscientific journals prior to 1 Jan 1953, must be considered to be effectively published. Hence, De Clairville’s excursion publi- cation, not being a technical work, is irrelevant. Moreover, we emphasize the fact that the name A. alpestre De Clairv. is associated with a type specimen, extant at Z (fide Fuchs 1974, p. 188). We conclude that Lellinger’s rejection of Fuchs’ analysis was unjustified, and thus we follow Fuchs’ treatment of this species. Athyrium distentifolium Tausch ez Opiz, Kratos 2(1):14, no. 41. 1820. TYPE: “Tausch ...‘no. 1838 Polypodium rhaeticum L. Brunberg zn Rresengeb.”’ (fide Fuchs 1974). Aspidium alpestre Hoppe, Neues Bot. Taschenb. Anfanger Wiss. Apo- thekerkunst. 216, no. 11. 1805, nom. superfl. Athyrium alpestre Milde, Fil. Eur. 53, no. 2. 1867, non De Clairv. 1811. Athyrium americanum In his study of North American ferns, Butters (1917, pp. 203-205) stated that the common arctic alpine fern of North America, previously assigned to the European species Athyrium alpestre (Hoppe) “Rylands er Moore” ( = A. distentifolium Tausch ez Opiz), must be considered as a distinct taxon. He characterized the American expression as follows: “ultimate segments of the fronds conspicuously narrower, and more widely separated from one another, and the sori ...(0.5-0.7 mm. in diameter ...), submarginal and protected by a teflexed tooth of the pinnule (false indusium). Careful study has failed to disclose any vestige of indusium.” In contrast, the sori in A. distentzfolium are 0.75-1.0 mm in diameter and have vestigial indusia. Butters noted the existence of intermediates between the American and European expressions and recognized the American expression to be a new variety: Athyrium alpestre (Hoppe) Rylands ez Moore var. americanum But- ters. However, Maxon (1918, pp. 120-121) stated that the invariable absence of indusia in the American expression was a substantiating character of some worth. He further argued that differences in gross structure, such as the strict, skeletonlike aspect of the American expression with its narrow, oblique, widely separated pinna segments (vs. leafy European plant with spreading segments 198 PUY TOL-0.G TA volume 70(3):194-208 March 1991 that are more broadly attached) and with its oblique, elongate deltoid pin- nae (vs. spreading, oblong-acuminate pinnae of the European plant) war- ranted specific recognition. Although Lellinger (1981) recognized this fern at subspecific rank (Athyrium alpestre [Hoppe] Milde ssp. americanum [But- ters] Lellinger), Dr. M.D. Windham (Univ. of Utah; pers. comm.) argues that the morphological features of the American alpine lady fern are sufficient for specific recognition. Our observation of herbarium material of the American element supports Windham’s treatment and we provide the following refer- ences. Athyrium americanum (Butters) Maxon, Amer. Fern J. 8:120. 1918. BA- SIONYM: Athyrium alpestre (Hoppe) Milde var. americanum Butters, Rhodora 19:204. 1917. Athyrium distentifoltum Tausch ez Opiz var. americanum (Butters) Crong., Univ. Wash. Publ. Biol. 17(1):63. 1969. Athyrium distentifolium Tausch er Opiz ssp. americanum (Butters) Hul- tén, Bot. Notiser 126:462. 1973. Athyrium alpestre (Hoppe) Milde ssp. americanum (Butters) Lellinger, Amer. Fern J. 71:91. 1981. LECTO- TYPE (vide Maxon 1918): CANADA. British Columbia: Rogers Pass, 23 Aug 1904, Heacock (in Shaw’s Selkirk Flora no. 554) (GH). EQUISETACEAE Equisetum x mackaw and E. x trachyodon The authorship of Eqguisetum x trachyodon has been generally assigned to A. Braun. Since this taxon is considered to be a hybrid, we researched the original literature to determine whether Braun proposed it as a species or as a hybrid. Braun (1838, p. 160) proposed “Equisetum brachyodon - subspecies nova E. hiemalis,” which was a nomen nudum. Subsequently, Braun (1839, p. 305) corrected the subspecific epithet to trachyodon (“Ueber ein neues Eq- uisetum [Equisetum trachyodon|”), and discussed its habit, habitat, and few diagnostic characters (pp. 305-306). Although his usage of the name “EL. tra- chyodon” suggests that he proposed a new species, he did not specifically state its rank on these two pages. However, on p. 308, he explicitly indicated that he treated “E. trachyodon” as a subspecies of EP. hiemale (“E. hiemale cujus subspecies: 1) E. ramosum...2) E. hiemale (genuinum) ...3) E. trachyodon; 4) E. variegatum”). It was an accepted practice in Braun’s time to indicate the subspecies rank as follows: “E. hiemale ssp. E. trachyodon” A. Braun. In present practice, the above citation must be altered, and cited as E. hyemale ssp. trachyodon A. Braun (ICBN, Art. 24.4). Even Hauke (1963) and Reed (1971) stated that Braun described his taxon at the subspecific rank, but these authors did not attempt to correct the nomenclature. Kartesz & Gandhi: Nomenclature of North American flora - V 199 It is evident from Braun’s classification that he proposed Equisetum trachy- odon at subspecific rank rather than at specific rank. With this established, we searched for the earliest reference (subsequent to Braun’s publication) treat- ing E. trachyodon at specific rank, and concluded that W.D.J. Koch (1845) was first to do so, and thus Koch inadvertently elevated Braun’s subspecies to species. Following the ICBN Art. 32.4, Ex. 5, the authorship of this taxon at the species rank is: E. trachyodon (A. Braun) W.D.J. Koch. Koch cited Equisetum mackavi as a synonym of E. trachyodon, as did Ben- tham (1892, p. 554), Broun (1938), Hauke (1963), and Reed (1971). Although Koch, Bentham, and Broun attributed the name FE. mackau to Newman, he used the epithet mackaz at varietal rank, and it was Brichan, who ultimately elevated Newman’s variety to specific rank. We concur with the above authors in considering E. mackazi and E. trachyodon to be conspecific, and conclude that, at specific rank, E. mackai clearly has priority over E. trachyodon. We provide the following references: Equisetum x mackazi (Newm.) Brichan, Phytologist 1:369. 1843 (Nov 1842). BASIONYM: Equisetum hyemale L. var. mackau Newm., Phytologist 1:305. 1843 (Sep 1842). TYPE: NORTHERN IRELAND. Belfast, Colin Glen, Aug 1833, Mackay & Whitla (7K). Equisetum hyemale L. ssp. trachyodon A. Braun, Flora 20(22):308. 1839. Equisetum trachyodon (A. Braun) W.D.J. Koch, Syn. Fi. Germ. Helv., ed. 2. 3:967. 1845. TYPE: Ad Rhenum Prope Carl- sruhe, Aug 1837, Braun (B) (fide Reed 1971). LYCOPODIACEAE Huperzia, Lycopodiella, Lycopodium In modern treatments of the genus Lycopodium L. sensu lato, several seg- regates, such as Diphasium C. Presl, Diphasiastrum Holub, Huperzia Bernh., Lateristachys Holub, Lepidotis P. Beauv., Lycopodzastrum Holub, Lycopodiella Holub, Palhinhaea Vascon. & Franco, Phlegmariurus (Herter) Holub, Pseu- dodiphasium Holub, Pseudolycopodiella Holub, and Pseudolycopodium Holub are recognized at generic ranks. Based on morphological, anatomical, phyto- chemical (Towers & Maass 1965; Pedersen & Ollgaard 1982), and chromosomal number analysis, Ollgaard (1987) recognized three genera in the Lycopodium complex: Huperzia, Lycopodiella, and Lycopodium. He (1989) also provided an index of the family Lycopodiaceae. Based on our observation, we concur with Ollgaard’s analysis and recognize three genera for North America. Seven new combinations are thus proposed in Huperzia and Lycopodiella. 200 PHY EOLOGT A volume 70(3):194-208 March 1991 KEY TO THE GENERA 1. Stems simple or isodichotomously branched, the main stems of determinate growth; strobili present or absent; sporophylls sessile, paleate, decidu- ous or persistent; sporangia axillary; spore surface foveolate-fossulate; plants epiphytic and often pendent, or terrestrial, confined to tropics or Widespread ’s |i aides conor 4) saat she gob eae Huperzia 1’ Stems anisodichotomously branched, the main stems of indeterminate growth, creeping, with erect, lateral branches of determinate growth; strobili present; sporophylls peltate or nearly so, deciduous; sporangia axillary or on sporophyll stalks; spore surface rugulate or reticulate; plants terrestrial, widespread. -..........:5.-0000c0se so eet meen 2 2. Strobilus solitary, erect and subsessile-pedunculate or deflexed and sessile; spore surface rugulate; leaves di- or monomorphic. ....... siputhinh tas doa, njalohe fs, some sepsge 3 ey isin’ seeks kts are ee Lycopodiella 2’ Strobili usually in 1 or 2 pairs, erect and sessile or pedunculate, or deflexed and pedunculate; spore surface reticulate; leaves monomor- PACH oh a Me AES sh ess oe Or Lycopodium Huperzia Huperzia Bernhardi, J. Bot. (Schrader) 1800(2):126. 1801. Plananthus P. Beauv. ez Mirbel in Lam. & Mirbel, Hist. Nat. Veg. 3:476. 1802, nom. superfil. Urostachys Herter, Beih. Bot. Centralbl. 39(2):249. 1922, nom. superfl. All with Lycopodium selago L. as lectotype (vide Rothmaker, Feddes Repert. Spec. Nov. Regni Veg. 54:59. 1944). Phlegmariurus Holub, Preslia 36:21. 1964. TYPE: Lycopodium phleg- maria L. Sporangia with sinuate, lignified side cell walls, the dehiscence isovalvate at maturity; gametophytes cylindrical, holosaprophytic, with paraphyses and long archegonia possessing persistent necks; main stem stele mostly radial; stem sclerenchyma nonlignified and peripheral; main root steles possessing crescent shaped xylem; mucilage canals absent; an ester of dihydrocaffeic acid often found; syringic acid not reported; chlorogenic acid, not the dominant Kartesz & Gandhi: Nomenclature of North American flora - V 201 phenol; n = 67, 68, 128, 130-140, 165-170 (fide Bruce 1976; Ollgaard 1987). Additional characters are provided in the key. Holub (1964) proposed the genus Phlegmariurus as a segregate from Hu- perzia. In this article, he argued that differences in habit, gametophyte, spore type, and basic chromosome number warranted generic distinction. Subse- quently, in his 1985 article, Holub stated that further studies on these two groups demonstrated that such differences were not consistent; hence, he aban- doned the genus Phlegmariurus and merged it with Huperzia. Based on our work, the following new combinations are proposed in the genus Huperziza. Huperzia x bartleyi (Cusick) Kartesz & Gandhi, comb. nov. (H. lucidula [Michx.] Trevisan x H. porophila {Lloyd & Underwood] Holub). BA- SIONYM: Lycopodium bartleyi Cusick, Amer. Fern J. 77:100. 1987. TYPE: U.S.A. Ohio: Hocking Co., 18 Mar 1987, Cusick 26204 (HOLO- DYPE:OS): Huperzia x buttersii (Abbe) Kartesz & Gandhi, comb. nov. (H. lucidula [Michx.] Trevisan x H. selago {L.] Mart. & Schrank). BASIONYM: Ly- copodium buttersii Abbe, Rhodora 55:91. 1953. TYPE: U.S.A. Min- nesota: Cook Co., 11 Jul 1938, Butters, Burns, & Hendrickson 111a (MIN). Huperzia x helleri (Herter) Kartesz & Gandhi, comb. nov. (H. serrata (Thunb. er Murray} Trevisan x H. somai [Hayata] Ching) BASIONYM: Lycopodium hellera Herter, Bot. Jahrb. Syst. 43 (Beiblat. no. 98, Heft 1 & 2):43. 1909. TYPE: U.S.A. Hawaii: Sandwich Islands, Oahu, Konahuanui, 1895, Heller s.n. (G, P). Huperzia mannii (Hillebr.) Kartesz & Gandhi, comb. nov. BASIONYM: Lycopodium phlegmaria L. var. mannii Hillebr., Fl. Hawazzan Isl. 645. 1888. Lycopodium mannii (Hillebr.) Skootsb., Acta Horti Gothob. 15:131- 132. 1942. TYPE: U.S.A. Hawaii: mountains above Maalaea Bay, Maui, Mann Enum. no. 656 (HOLOTYPE: CU). Huperzia occidentalis (Clute) Kartesz & Gandhi, comb. nov. BASIONYM: Lycopodium lucidulum Michx. forma occidentalis Clute, Fern Bull. 11:13. 1903. TYPE: U.S.A. Washington: Near the base of Mt. Rainier, 15 Aug 190%, Fleet, sin. (NY): Huperzia serrata (Thunb. ez Murray) Trevisan var. dentata (Hillebr.) Kartesz & Gandhi, comb. nov. BASIONYM: Lycopodium serratum Thunb. ez Murray var. dentatum Hillebr., Fl. Hawazan Isl. 642-643. 1888. TYPE: U.S.A. Hawaii: high mountains of Kauia, ? Hillebrand (7B). 202 Poa YT OL OuG tA volume 70(3):194-208 March 1991 Lycopodiella Lycopodiella Holub, Preslia 36:22. 1964. TYPE: Lycopodium inundatum L. Lepidotis auct. non Mirbel Palhinhaea Vascon. & Franco, Bol. Soc. Broter., I]. 41:24. 1967. TYPE: Lycopodium cernuum L. Lateristachys Holub, Folia Geobot. Phytotax. 18:440. 1983. TYPE: Lycopodium laterale R. Br. Pseudolycopodtella Holub, Folia Geobot. Phytotax. 18:441. 1983. TYPE: Lycopodium carolinianum L. Sporangia with straight, nonlignified side cell walls, the dehiscence mostly anisovalvate at maturity; gametophyte tuberous with multicellular lobes, hemis- aprophytic, with short archegonia possessing ephemeral] necks, lacking para- physes; main stem stele mostly radial; stem sclerenchyma lignified and subpe- ripheral or close to endodermis; main root steles possessing crescent shaped xylem; both basal and veinal mucilage canals often present; chlorogenic acid, the dominant phenol; dihvdrocaffeic acid and svringic acid not reported; n = 35, 68, 70, 78, 104, 108, 110, 136, 156, 165, 208 (fide Bruce 1976; Ollgaard 1987). Additional characters are provided in the key. Lycopodiella cernua (L.) Pichi Sermolli var. curvata (Sw.) Kartesz & Gandhi, comb. nov. BASIONYM: Lycopodium curvatum Sw., J. Bot. (Schrader) 1800(2):116. 1801. TYPE: ?JAMAICA. ? Swartz. Lycopodium Lycopodium L., Sp. Pl. 2:1100. 1753. LECTOTYPE (vide Britton in Britton & Brown, Ill. Fl. N. U.S., ed. 2. 1:43. 1913): Lycopodium clavatum L. Lepidotis P. Beauv. ez Mirbel in Lam. & Mirbel, Nat. Hist. Veg. 3:477. 1802. LECTOTYPE (vide Pichi Sermolli, Webbia 26:145. 1971.): Lycopodium clavatum L. Diphasiastrum Holub, Preslia 47:104. 1975. TYPE: Lycopodium com- planatum L. Kartesz & Gandhi: Nomenclature of North American flora - V 203 Diphasium Rothm., Feddes Repert. Spec. Nov. Regni Veg. 54:64. 1944. TYPE: Lycopodium jussiae? Poir. Lycopodiastrum Holub, Folia Geobot. Phytotax. 18:440. 1983. TYPE: Lycopodium casuarinoides Spring. Pseudodiphasium Holub, Folia Geobot. Phytotax. 18:440. 1983. TYPE: Lycopodium volubile G. Forster. Pseudolycopodium Holub, Folia Geobot. Phytotax. 18:441. 1983. TYPE: Lycopodium densum Labill., non Lam. Sporangia with sinuate, lignified side cell walls, the dehiscence isovalvate at maturity; gametophytes carrot or convoluted button shaped, holosaprophytic, with long archegonia possessing persistent neck, lacking paraphyses; main stem steles arranged in fixed parallel bands; stem sclerenchyma nonlignified, close to the endodermis; main root steles similar to the stem steles; basal mucilage canals generally present; veinal mucilage canals absent; dihydrocaffeic acid not reported; syringic acid reported; chlorogenic acid, not the dominant phenol; n = 23-34, 90 (fide Bruce 1976; Ollgaard 1987). Additional characters are provided in the key. POLYPODIACEAE Pecluma and Polypodium Lellinger (1981) considered the Polypodium pectinatum L. - plumula Humb. & Bonpl. ez Willd. group to be sharply distinct from other polypodies and re- marked that the group “deserved to be placed in a subgenus of its own:” Poly- podium subgenus Pectinatum Lellinger. Price (1983) elevated this subgenus to the generic rank as Pecluma and transferred 28 Polypodium species to the new genus, including Polypodium camptophyllarium Fee. Following Price’s treat- ment, we consider that Pecluma, with its short rhizomes, pectinate lamina, and linear Jamina segments (1-8 mm wide), can be segregated from the genus Polypodium, and thus transfer Polypodium camptophyllarium Fee var. abbre- viatum Evans to Pecluma. The var. abbreviata is found in Puerto Rico. Pecluma camptophyllaria (Fee) M. Price ssp. abbreviata (A.M. Evans) Kartesz & Gandhi, stat. & comb. nov. BASIONYM: Polypodium camp- tophyllarium Fee var. abbreviatum Evans, Ann. Missouri Bot. Gard. 55:256. 1969. TYPE: PERU. Herrera 872 (US). 204 PHYTOLOGIA volume 70(3):194-208 March 1991 SCHIZAEACEAE. Actinostachys pennula Christensen (1906) cited the authorship of the name Actinostachys pennula as Hooker (1842), whereas Proctor (1989) attributed the name to Hooker & Bauer. The name in question was published in Genera Filicum (also called JI- lustrations of the Ferns and Other Alhed Genera). On the title page of the pub- lication, we found the following information: “from the original coloured draw- ings of F. Bauer, with additions and descriptive letterpress by W.J. Hooker.” It is clear that Bauer should be cited as the senior author of this work. The dedication and preface were by Hooker. From the preface, we quote the fol- lowing: “Whatever may be merits of the present publication, it will be seen that they are entirely due to the distinguished Natural History Painter, whose name appears upon the plates ...” There are 120 plates in this publication, each plate with notes by Hooker. The copy found at the Botany Library (NCU) showed Bauer’s name for plates 1-40 and 50-51; the painter’s name (Bauer) is not seen for other plates. After analyzing the contents, we have concluded that if the new taxa were to be validated by Bauer’s illustration, then Bauer alone should be the author of such taxa. Conversely, if taxa were to be validated by Hooker’s notes, then Hooker should be the author. The taxon Actinostachys pennula was illustrated on plate 111-A, accompa- nied by Hooker’s notes. In this publication, A. pennula was not a new species, but rather, a new combination based on Schizea pennula Sw. Hooker cited the basionym and used the name A. pennula; thus, a new combination was properly made, which mei the requirements of ICBN Art. 33.2, for new com- binations proposed prior to Jan 1953. Since Bauer’s illustration is secondary here, we conclude that Hooker is the sole author for the new combination: Actinostachys pennula (Sw.) Hook., Gen. Fil. t. 111. 1842. VITTARIACEAE Antrophyum, Polytaenium, and Vittaria The genera Antrophyum Kaulf., Polytaenium Desv., and Vittaria J.E. Sm. belong to the family Vittariaceae. In both Antrophyum and Vittaria, para- physes are present and spores have either trilete or monolete apertures, but in Antrophyum the sori are usually multiple on the anastomosing veins, whereas in Vittaria, the sori are submarginal and form a single line along each margin. Kartesz & Gandhi: Nomenclature of North American flora - V 205 Polytaenium differs from Antrophyum in lacking paraphyses and in possess- ing a costa (usually extending almost to the apex of the lamina) and trilete apertured spores. In Antrophyum, a costa is absent or if present, then usu- ally partially developed. The separation of Polytaenium from Antrophyum at generic rank has been in dispute. Tryon (1964) and Proctor (1989) recognized its generic status; however, Tryon & Tryon (1982) remarked that Polytaenium was not a distinctive genus and treated Polyiaenium as a subgenus: Antro- phyum subgenus Polytaenium (Desv.) Bened. We concur with Tryon & Tryon in recognizing Antrophyum sens. lat. (including Polytaenium). Vittaria intramarginalis Baker ez Jenman possesses submarginal sori (as in other vittarias), but soral condition in Vittariaceae is of little importance in determining relationship relative to venation (fide Benedict 1907, p. 451, pro- tologue of Antrophyum lineatum |Sw.] Kaulf). In V2ttaria, two submarginal veins are formed by the interlocking of pinnate branches from the axial vein; occasionally, as in the narrow leaved vittarias, the pinnate veins are obscure and the venation consists of three primary longitudinal veins. In contrast, Antrophyum usually shows reticulated venation. In V. intramarginalis, there are few longitudinal veins that are connected by irregular cross veinlets, and such a venation pattern is comparable to the venation of narrow leaved expres- sions of Viltaria. However, V. intramarginalis differs from all other vittarias in lacking paraphyses. It is possible that both V. intramarginalis and other vittarias might have had a common ancestor. Due to the absence of paraphyses in Witdaria intramarginalis, Alston (1952) transferred it to Polytaenium and made the combination: Polylaenium intra- marginale (Baker er Jenman) Alston. Tryon & Tryon (1982, p. 359) treated V. intramarginalis as a synonym of A. lineatum. Initially, Proctor (1982) rec- ognized V. intramarginalis at varietal status, but in 1989 (p. 148) remarked that field work in Puerto Rico had convincingly demonstrated that V. antra- marginalis did not intergrade with A. hneatum, had its own allopatric distri- bution pattern involving different ecological parameters, and hence, deserved specific recognition. We concur with Proctor (1989) in providing specific rank to Baker’s plant. The transfer of V. intramarginalis to Antrophyum necessi- tates the following new combination. Antrophyum intramarginale (Baker ez Jenman) Kartesz & Gandhi, comb. nov. BASIONYM: Vittaria intramarginalis Baker ex Jenman, J. Bot. 15:266. 1877. Polytaenium intramarginale (Baker ez Jenman) Alston, Mutisia no. 7:9. 1952. Polytaentum lneatum (Sw.) J.E. Sm. var. an- tramarginale (Baker ez Jenman) Proctor, Amer. Fern J. 72:114. 1982. TYPE: JAMAICA. 1877, Jenman 58 (HOLOTYPE: K). 206 PBST O4,0:64 A volume 70(3):194-208 March 1991 ACKNOWLEDGMENTS The authors are grateful Dr. Michael D. Windham (University of Utah), Dr. Alan R. Smith (University of California, Berkeley), and Dr. Larry E. Brown (Houston Community College) for valuable help on the manuscript. We also thank the Librarian of the Biology-Forestry Library, Duke University for copies of Alston’s article. REFERENCES Alston, A.H.G. 1952. Pteridophyta of the Macarena Mountains, Colombia. Mutisia 1(7). Benedict, R.C. 1907. The genus Antrophyum - I. Synopsis of subgenera, and the American species. Bull. Torrey Bot. Club 34:445-458. Bentham, G. 1892. Handb. Brit. Fl., ed. 6, revised by J.D. Hooker. Lovell Reeve & Co., Limited, London. Braun, A. 1838. Botanische Notizen. Flora 21(Band I, no. 10):160. . 1839. Original - Abhandlungen. Flora 22(Band I, no. 20):305-320. Broun, M. 1938. Inder N. Amer. Ferns. Orleans, MA. (Published by the author). Bruce, J.G. 1976. Gametophytes and subgeneric concepts in Lycopodium. Amer. J. Bot. 63:919-924. Butters, F.K. 1917. Taxonomic and geographic studies in North American ferns. Rhodora 19:169-216. Christensen, C. 1906. Jndez Filic. Reprinted in 1973 by Otto Koeltz Anti- quariat, Koenigstein - Ts./B.R.D. Clairville, J.P. de. 1811. Man. Herbor. Suisse. Steiner Ziegler Publishers, Winterthur. Fernald, M.L. 1929. The author of Athyrium alpestre. Rhodora 31:165-166. Fuchs, H.P. 1974. The correct name of the alpine lady fern. Candollea 29:181-205. Kartesz & Gandhi: Nomenclature of North American flora - V 207 Greuter, W. (Chairman, Editorial Committee). 1988. International Code of Botanical Nomenclature. Regnum Veg. vol. 118. Bohn, Scheltema & Holkema, Utrecht. Hauke, R. 1963. A taxonomic monograph of the genus Equisetum subgenus Hippochaete. Nova Hedwigia Beihefte 8:1-123. Holub, J. 1964. Lycopodiella, novy rod radu Lycopodiales. Preslia 36:16-22. . 1983. Validation of generic names in Lycopodiaceae: with a de- scription of a new genus Pseudolycopodiella. Folia Geobot. Phytotax. 18:439-442. . 1985. Transfers of Lycopodium species to Huperzia: with a note on generic classification in Huperziaceae. Folia Geobot. Phytotax. 20:67-80. Hooker, W.J. 1842. Gen. Fil. Henry G. Bohn, London. Hoope, D.H. 1805. Verzeichniscder in Deutschland wild wachsenden Far- renkrauter. Neues Bot. Taschenb. Anfanger Wiss. Apothekerkunst. 1805: 199-226. Janchen, E. 1956. Cat. Fl. Austriae, 1:]. Teil: Pteridophyten und an- thophyten (edited by Austrian Academy of Sciences). Springer-Verlag, Wien. Kartesz, J.T. 1991. Syn. Checkl. Vasc. Fl. U.S., Canada, and Greenland, ed. 2 (in press). Timber Press, Portland, OR. & K.N. Gandhi. 1989. Nomenclatural notes for the North American flora - I. Phytologia 67:461-468. . 1990a. Nomenclatural notes for the North American flora - II. Phytologia 68:421-427. . 1990b. Nomenclatural notes for the North American flora - III. Phytologia 69:129-137. . 1990c. Nomenclatural notes for the North American flora - IV. Phytologia 69:301-312. Kartesz, J.T. & R. Kartesz. 1980. Syn. Checkl. Vasc. Fl. U.S., Canada, and Greenland. The University of North Carolina Press, Chapel Hill, NC. Koch, W.D.J. 1845. Syn. Fl. Germ. Helv., ed. 2. vol. 3. Frankfurt. Lellinger, D.B. 1981. Notes on North American ferns. Amer. Fern J. 71:90- 94. 208 PHY TOL O GLA volume 70(3):194-208 March 1991 . 1985. Field Man. Ferns. Smithsonian Institution Press, Washing- ton, DC. Maxon, W.R. 1918. Notes on American ferns - XII. Amer. Fern J. 8:114-121. Milde, C.A.J. 1867. Fl. Eur. Sumptibus A. Felix Publishers, Leipzig. Oligaard, B. 1987. A revised classification of the Lycopodiaceae s. lat. Critica Bot. 92:153-178. . 1989. Index of the Lycopodiaceae. Biol. Skr., vol. 34. Pedersen, J.A. & B. Oligaard. 1982. Phenolic acids in the genus Lycopodium. Biochem. Syst. Ecol. 10:3-9. Price, M.G. 1983. Pecluma, a new tropical American fern genus. Amer. Fern J. 73:109-116. Proctor, G.R. 1982. Notes on Jamaican ferns-[V. Amer. Fern J. 72:111-114. . 1989. Ferns of Puerto Rico and the Virgin Islands. Mem. New York Bot. Gard. vol. 53. Reed, C.F. 1971. Inder Equisetophyta, part I]. Reed Herbarium, Baltimore, MD. Towers, G.H.N. & W.S.G. Maass. 1965. Phenolic acids and lignins in the Lycopodiales. Phytochem. 4:57-66. Tryon, R.M. 1964. Taxonomic fern notes, IV. Some American vittarioid ferns, 1. The genus Polytaenium. Rhodora 66:110-114. & A.F. Tryon. 1982. Ferns and Allied Plants. Springer-Verlag New York Inc., New York. Phytologia (March 1991) 70(3):209-225. TAXONOMIC NOTES ON CISTANTHE, CALANDRINIA, AND TALINUM (PORTULACACEAE) Mark A. Hershkovitz Department of Botany, University of California, Davis, California 95616 USA ABSTRACT New combinations for South American species of Ctstanthe are pro- vided along with taxonomic notes. Also, a new section of Calandrinia and a new combination in Talinum are established. The nomenclatural changes reflect new morphological and cladistic data on Portulacaceae. KEY WORDS: Argentina, Calandrinia, Chile, Cistanthe, Mono- cosmia, Peru, Portulacaceae, Talinum, taxonomy In a previous paper (Hershkovitz 1990a), I established a sectional taxonomy and several combinations for Crstanthe Spach (Portulacaceae). These proposed changes reflected corroboration by leaf morphological evidence (Hershkovitz 1990b) of Carolin’s (1987) cladistic analysis of Portulacaceae, in which he concluded that Cistanthe should be segregated from Calandrinia Kunth in H.B.K. I also proposed (Hershkovitz 1990b) that Monocosmia Fenzl in Endl. & Fenz] is phylogenetically nested in Calandrinia and should be included therein. I provide below: (1) additional combinations for South American species of Cistanthe, along with miscellaneous notes and lists of materia] examined; (2) a sectional status for Monocosmia in Calandrinia; and (3) a new combination in Talinum Adans. for a species currently classified in Calandrinia. 1. Cistanthe NEW COMBINATIONS AND NOTES In my studies of leaf morphology in Portulacaceae (Hershkovitz 1990b), I noted that specimen identification and species delimitations of South American Cistanthe, heretofore classified in Calandrinia (cf. Hershkovitz 1990a, 1990b) and Philippiamra Kuntze (= Silvaea Philippi), were not adequately facilitated 209 210 Pu YAO, OG. TA volume 70(3):209-225 March 1991 by existing floristic treatments. At the same time, | concurred with Carolin (1987, in press) that Portulacaceae taxonomy was poorly served by the reten- tion of these species under their existing generic assignments. The first prob- lem I circumvented by sampling leaves representative of the range of gross morphological variation in Crstanthe as evident from herbarium specimens. The second problem is more difficult to resolve because there is simply no means short of a detailed monographic study to determine which of at least 60 combinations of Calandrinia and Philippiamra/Silvaea referable to Cistanthe will prove to correspond to distinct species. An example of the sort of taxonomic problems J encountered can be demon- strated by comparing accounts by Reiche (1898) and Johnston (1929) of the Cistanthe grandiflora (Lindley) Carolin ez Hershkovitz complex. According to Reiche (1898), Cistanthe grandiflora represents a single polymorphic species with a total of eleven synonyms variously referable to three more or less dis- tinct varieties corresponding to Calandrinza grandiflora Lindley, Calandrinia discolor Schrad., and Calandrinia speciosa Lehm. Johnston (1929, pp. 36, 37, 147), however, made severa] remarks indicating that he recognized at least some of the variants as distinct species. In addition, Johnston (1929) charac- terized two new species as members of the Cistanthe grandiflora complex. The difference in opinion cannot be attributed merely to a tendency by Johnston to split taxa. For example, Reiche (1898) recognized four species in Czstanthe sect. Philippramra, while Johnston (1929, p. 39) wrote that “ further studies will show [three of] these species ...to be phases of one variable species ....” In the present paper, I provide new combinations for South American species of Cistanthe examined in my leaf morphological studies, along with remarks on possible synonyms and lists of examined specimens. This paper brings the total number of recombined names for South American species to 17 - twelve for Cistanthe sect. Crstanthe, three for C. sect. Amarantozdes, and two for C. sect. Philippiamra (cf. also Hershkovitz 1990a). These figures will increase eventually — I suspect additional research may well establish that the actual numbers in each section are roughly twice these numbers. Unless other- wise noted, the species recombined here are presumed to include the synonyms listed elsewhere, and no judgment is implicit in my failure to mention other pertinent species of Calandrinza elsewhere recognized. There has never been a comprehensive key to the South American species of Cistanthe. The most extensive key, provided by Reiche (1898) for the Chilean species, emphasizes variable vegetative and sometimes nebulous traits (cf. Carolin 1987). Given the preliminary nature of the present work, however, 1 am unable to offer a useful alternative. Identifications, unfortunately, will continue to require the use of the keys in the cited floristic literature and simple description /specimen comparisons. For the present work, I consulted type descriptions and regional floristic literature provided by Reiche (1898) and Philippi (1860a, 1860b, 1894a, 1894b) Hershkovitz: Taxonomic notes in Portulacaceae Dili for Chile; Johnston (1928, 1929) for northern Chile and Peru; Macbride (1931, 1937) for Peru; Cullen (1953) for Argentina; and Peralta (1988) for the Ar- gentinian province of Mendoza. I also considered descriptions of seed and trichome morphology provided by Kelley (1973) for Cistanthe sections Cistan- the and Amarantoides, and my own observations of herbarium material and anatomical preparations thereof (Hershkovitz 1990b). I have seen photographs of supposed type material of many of the species (see below), but I generally cannot confirm their authenticity. A few years ago, Roger Carolin (SYD) provided me with an unpublished manuscript that included several of the combinations listed below and several others. Carolin proposed several combinations in Cistanthe that represent, I believe, synonyms of other species. In any case, I employ the “ez” notation to indicate Carolin’s authorship of the names contained in his manuscript. A. Cistanthe sect. Cistanthe Cistanthe sect. Cistanthe comprises all species of Calandrinia sect. Crstan- the (Reiche), Calandrinia sect. Rosulatae Reiche, Calandrinia sect. Arenarzae Reiche, and Calandrinia sect. Andinae Reiche (cf. Hershkovitz 1990b; Kelley 1973; Reiche 1897, 1898). Cistanthe arenaria (Diels) Carolin ez Hershkovitz, comb. nov. BASIONYM: Calandrinia arenaria Cham., Linnaea 6: 563. 1831. SPECIMENS EXAMINED: CHILE. Aconcagua: Bertero 1348 (photo- graph of type of Calandrinia chamissoi Barnéoud in Gay, US ex G-DEL); Morrison 16917 (CAS, NA). Atacama: Philippi s.n. (photograph of type of Calandrinia polyclados Philippi, US er G-DEL). Coquimbo: Joseph 4507 (US); Joseph 4511 (US); Simon 441 (UC); Simon 454 (UC); Wagenknecht 18444 (F - two sheets, UC); Worth & Morrison 16855 (CAS, NA); Zollner 6088 (L). Metropolitana: Joseph 2788 (US). Nuble: Joseph 3990 (US). Cistanthe arenaria is an annual with numerous branches spreading from the base and rhombic to linear leaves. The species is apparently closely re- lated to Cistanthe fenzlii (Barnéoud in Gay) Carolin ez Hershkovitz, which is perennial and has more erect branches (cf. Reiche, 1898). Calandrinia solisi Philippi appears to be somewhat intermediate between Cistanthe arenaria and Cistanthe fenzlii, so for now, I refer all intermediate plants to one species or the other. Cistanthe arenaria is widely distributed and highly variable (cf. Reiche, 1898). Some of the cited specimens (e.g., Wagenknecht 18444) correspond perfectly to Reiche’s (1898) key and description. The largest plants, however, 212 PHY LOO GA volume 70(3):209-225 March 1991 appear similar to supposed type material of Calandrinza polyclados Philippi, which Reiche regarded as a synonym of Cistanthe fenzlit. Joseph 4507 and 4511 key to and had been determined as Calandrinia glaucopurpurea Reiche. Reiche’s (1898) description of Calandrinia glaucopur- purea, however, also corresponds to that of Calandrinia chamissoi Barnéoud in Gay, which Reiche (1898) recognized as a variety of Cistanthe arenaria. Re- iche (1898) recognized two other varieties of Cistanthe arenaria, but I cannot determine whether these are discrete. Joseph 8990 more or less keys to and was determined as Calandrinza solisii, but the specimen has a slender, rather than thick (perennial?; cf. Reiche 1897) root. ] am not convinced that the key trait (sterile inflorescence bracts present vs. absent; Reiche 1898) and the difference in root morphology that distinguish Calandrinia soltsi from Cistanthe arenaria collectively warrant specific recog- nition of the former. I have elsewhere (Hershkovitz 1990b) cited Neger s.n. (M) and Zollner 10636 (NA) as Cistanthe arenarza. The former specimen I believe is Crstanthe fenzlu, although it also conceivably keys to Calandrinia solisu. The latter is a mixed collection with some arenaria-like plants (see indets. listed below). I also cited Zollner 9807 incorrectly as Cistanthe glaucopurpurea (see indets. listed below). Cistanthe cephalophora (I.M. Johnston) Carolin ez Hershkovitz, comb. nov. BASIONYM: Calandrinia cephalophora 1.M. Johnston, Contr. Gray Herb. 85:35. 1929. SPECIMENS EXAMINED: CHILE. Antofagasta: Pennell 13032 (US); Werdermann 855 (CAS, UC, US). This distinctive species has a capitate inflorescence and hairy seeds, but otherwise resembles Cistanthe longiscapa (Barnéoud in Gay) Carolin ez Her- shkovitz. Cistanthe coquimbensis (Barnéoud in Gay) Carolin ez Hershkovitz, comb. nov. BASIONYM: Calandrinia coqguimbensis Barnéoud in Gay, Fl. Chile [Hist. Fisica y Politica de Chile, Botanica] 2: 483. 1847 (“1846”). SPECIMENS EXAMINED: CHILE. Locality not determined: Gay s.n. (photograph of type, US ez P); Joseph 4973 (US). Coquimbo: Werdermann 881 (CAS, F, UC). This species seems to be related to the Cistanthe longiscapa alliance (Ca- landrinia sect. Rosulatae Reiche, cf. Reiche 1898; see also below). The cited specimens, both previously identified as C. coguimbensis, key to this species more or less by default rather than by critical traits - the other species in this section (Reiche’s nos. 22-26) are quite distinct from Cistanthe coquimben- sis. The cited specimens differ from Cistanthe longiscapa in: (1) having more Hershkovitz: Taxonomic notes in Portulacaceae 213 spreading or prostrate, rather than erect, flowering branches; (2) having longer, relatively narrower, and more distinctly petiolate leaves; and (3) having more foliage leaves scattered along the flowering branches rather than just near the base. In these respects, the specimens approach members of Calandrinia sect. Arenariae Reiche. The leaf venation pattern also appears distinct from that of Crstanthe longiscapa (cf. Hershkovitz 1990b). The examined material has ca. 15 stamens, rather than 6-8 as reported by Reiche (1898). Cistanthe cymosa (Philippi) Hershkovitz, comb. nov. BASIONYM: Ca- landrinia cymosa Philippi, Ann. Univ. Chile 85:192. 1893. SPECIMENS EXAMINED: CHILE. Antofagasta: von Bayern s.n., “11.x. 1898” (M); Werdermann 853(B, CAS, M, UC, US); Worth & Morrison 15816 (NA, UC). Atacama: Johnston 4787 (US). This species is related to the Cistanthe longiscapa alliance (cf. Reiche, 1898). Its distinctive feature is its nearly corymbiform inflorescence with nu- merous congested flowers. The flowers are yellow and the seeds black and shiny. The sepals in the cited specimens appear waxy to glutinous, a feature noted for Calandrinia chrysantha I.M. Johnston, which Johnston (1929) re- garded as most closely related to Cistanthe cymosa - they may prove to be the same species. Cistanthe fenzlii (Barnéoud in Gay) Carolin ez Hershkovitz, comb. nov. BASIONYM: Calandrinia fenzlii Barnéoud in Gay, Fl. Chile | Hist. Fisica y Politica de Chile, Botanica} 2:493. 1847 (“1846”). SPECIMENS EXAMINED: CHILE. Locality not determined: Gay s.n. (photograph of type, US ez P). Bio-Bio: Joseph 4015 (US); Neger s.n., “Nov 1896” (two sheets, probably duplicates, M); Philippi s.n. (US). In Reiche’s (1898) key, Cistanthe fenzli is set off from Cistanthe arenaria only by virtue of the more erect stems in the former. The description, however, also notes that Cistanthe fenzlii has a thick root, and, indeed, the cited spec- imens appear to be perennials. Judging from a Philippi-determined specimen (Philippi s.n.), Reiche (1898) was correct in regarding Calandrinia sanguinea Philippi as a synonym of Cistanthe fenzlit. Reiche (1898) also listed Calandrinia polyclados Philippi in synonymy with Cistanthe fenzlii, but the photograph of the type of the former (see Cistanthe arenaria above) is more highly branched, has broader leaves, and has less congested inflorescences than the photograph of the type of the latter and the specimens cited here. Cistanthe grandiflora (Lindley) Carolin ez Hershkovitz, Phytologia 68:269. 1990. 214 PRY TOdk0O GTA volume 70(3):209-225 March 1991 SPECIMENS EXAMINED: Cultivated: Peele 154 (NA). Without data: one sheet (B); one sheet, no. 906253... 39(L). Without collector and locality: 998 (L). CHILE. Without locality: Bernhard: 3 (B). Aconcagua: Behn s.n. (UC); Blood & Tremelling 427 (NA); Buchtien s.n. (US); Eyerdam & Beetle 24642 (UC); Hutchinson 176 (UC); Joseph 1764 (US); Morrison 16872 (NA, UC); Rose & Rose 19138 (US); Simon 446 (UC); Stebbins 8547 (UC); Stebbins 8592 (UC); Werdermann 39 (UC); West 3959 (F, UC - two sheets, US - two sheets); West 4574 (UC); Zollner 8364 (NA). Antofagasta: Worth & Morrison 16183(NA, UC). Atacama: Beetle 26143 (UC); Harding 22847 (US); Werdermann 405 (F, US); West 8868 (UC); Zollner 4079 (L). Bio-Bio: Hutchinson 234 (UC, US). Coquimbo: Simon 447 (UC); Simon 462 (UC); Simon 465 (UC); Wagenknecht 18562 (UC); Zollner 6327 (L); Zollner 10284 (NA). Metropolitana: Hastings 268 (UC); Morrison 16816 (UC); Philippi s.n. (US). O’Higgins: Aravena 33343 (UC). I alluded above to the taxonomic problems in the Cistanthe grandiflora complex. J] am unable to sort these specimens into finer, discrete specific en- tities. The Antofagasta and Atacama specimens, however, seem coarser and more succulent than the rest. Three of these (Beetle 26143, West 3868, Worth & Morrison 16133) were annotated as Calandrinia taltalensis 1.M. Johnston fide Johnston. I have elsewhere (Hershkovitz 1990b) cited West 3959 as Cis- tanthe glauca (Schrad.) ined., and Worth & Morrison 16138 as Cistanthe tal- talensis (1.M. Johnston) ined., but 1 chose not to recognize these combinations at present. Cistanthe lingulata (Ruiz Lopez & Pavon) Hershkovitz, comb. nov. BA- SIONYM: Talinum lngulatum Ruiz Lopez & Pavon, Syst. Veg. Fl. Pe- ruv. Chil. 115. 1798. SPECIMENS EXAMINED: PERU. Ancash: Ferreyra 18532 (US). La Libertad: Lopez Miranda 374 (US). Lima: Dombey (number illegible, possibly 170; P); Dombey (number illegible, possibly 716; P); Ferreyra 10486 (US); Ruiz Lopez & Pavon s.n. (photograph of type, US ez G-DEL); Weberbauer 7482 (US). According to Macbride (1937), Cistanthe lingulata differs from Calandrinia guivensis Macbride only in having a less exserted capsule. He noted, however, that capsule exsertion may reflect capsule maturity, so that the two species might be the same. Therefore, I have included all material keying to either species in Cistanthe lingulata. Of the cited specimens, the most extreme exser- tion of the capsule occurs in both of the Ferreyra specimens. Cistanthe lingulata also apparently intergrades with Cistanthe weberbaueri (Diels) Carolin ez Hershkovitz. According to Macbride (1937), Crstanthe we- berbaueri differs from C. lingulata in having an “umbellate-fasciculate” rather than “racemose or paniculate” inflorescence. Such a difference typifies younger Hershkovitz: Taxonomic notes in Portulacaceae 215 and more mature inflorescences, respectively, in Czstanthe. My determinations emphasize apparent distinctions in leaf morphology: The leaves of specimens referable to Cistanthe lingulata are generally narrower (<10 mm) than those of Cistanthe weberbaueri (>10 mm), and, while their venation patterns are sim- ilar, the vein orders of Cistanthe weberbauerz are more distinct and the vein reticulum appears denser even in comparably sized leaves (cf. Hershkovitz 1990b). Weberbauer 7432. however, was cited as Calandrinia weberbaueri by Macbride (1937) but has linear-filiform leaves less than 5 mm wide. The in- florescence is immature in this specimen, so ] cannot evaluate the other key characters. Cistanthe longiscapa (Barnéoud in Gay) Carolin ez Hershkovitz, comb. nov. BASIONYM: Calandrinia longiscapa Barnéoud in Gay, Fl. Chile (Hist. Fisica y Politica de Chile, Botanica) 2:492. 1847 (“1846”). SPECIMENS EXAMINED: CHILE. Without locality: Gay s.n. (photo- graph of type of Calandrinia longifolia [sic], US ex P). Antofagasta: Werder- mann 1046 (leg. Francke; UC, US). Atacama: Johnston 5034 (US); Morong 1267(NA, US); Philippi s.n. (US); Werdermann 445 (F, UC); Worth & Mor- rison 16164 (NA, UC); Worth & Morrison 16267(NA, UC); Zollner 4083 (L); Zollner 9291 (NA); Zollner 11917 (NA). Cistanthe longzscapa is distinguished from C. cephalophora and C. cymosa by its more open inflorescence. Vegetatively, these three species are similar in having relatively broad and tending towards orbicular leaves crowded near the base of the plant and typically a pair of subopposite leaves at the base of the inflorescence. I tentatively refer Calandrinia litoralis Philippi to Cistanthe longiscapa. From the examined material, I cannot identify the putative differences be- tween these species in bract morphology (cf. Reiche 1898). Pubescent seeds supposedly characterize Cistanthe longiscapa, while glabrous seeds character- ize Calandrinia litoralis (Kelley 1973; Reiche 1898). The specimens cited above have variously pusticulate-tomentose or glabrous seeds, but are other- wise very similar. According to Reiche (1898), Cistanthe longiscapa flowers in September-October, while Calandrinia litoralis flowers in December-January. Most of the cited specimens have September-October collection dates, while Johnston 5034 lists November 25, and Zollner 4083 lists January 12. The last two specimens are clearly in more mature stages than the September-October collections. The Atacama collections are mostly from the vicinity of Copiapo. The Antofagasta collection (Werdermann 1046) is unusual and may be misdeter- mined. This specimen includes a fragment of an inflorescence and a whole plant that appears to have been prostrate with a flowering scape barely 5 cm long. The bracts and sepals are not conspicuously etched with black markings. The absence of these etchings is unusual in Cistanthe sect. Cistanthe. 216 PH Y T.0O.LO\G-A volume 70(3):209-225 March 1991 Cistanthe macrocalyx (Hauman) Carolin ez Hershkovitz, comb. nov. BA- SIONYM: Calandrinia macrocalyr Hauman, Ann. Soc. Cient. Argentina 86:253. 1919. SPECIMEN EXAMINED: ARGENTINA. San Juan: Castellanos 15587 (US). Cullen (1953) and Peralta (1988) recognized this species as distinct from Cistanthe picta (Gillies er Arn. in Cheek) Carolin ez Hershkovitz in their treat- ments for Argentina and Mendoza, respectively. It is distinguished from the latter by its smaller, oblanceolate leaves and larger, relatively broader calyx. The specimen cited above agrees with the description in leaf morphology, but the flowers have largely been lost. Cistanthe paniculata (Ruiz Lopez & Pavon) Carolin ex Hershkovitz, comb. nov. BASIONYM: Calandrinia paniculata Ruiz Lopez & Pavon, Syst. Veg. Fl. Peruv. Chil. 114. 1798. SPECIMENS EXAMINED: PERU. Ancash: Dillon, et al. 4627 (F); Dul- lon, et al. 4633(F); Dillon & Santistebau 4706 (F); Ferreyra 6350 (US); Fer- reyra 11576(US); Ferreyra 12022 (US); Ferreyra 12611(US); Hutchinson 1292 (US, UC); Stork, et al. 9169 (UC); Weberbauer 1484 (B). Arequipa: Dillon, et al. $265 (F); Dillon, et al. $292 (F); Dillon & Dillon 3843 (F); Worth & Morrison 15761(NA, UC). La Libertad: Dillon, et al. 2718 (F); Dillon, et al. 4656 (F); Dillon, et al. 4693 (F). Lima: Soukup 2156 (US); Stork & Vargas 9363(UC). Tacna: Dillon, et al. 8884 (F); Dillon & Dillon 3662 (F); Ferreyra 12502 (US); Ferreyra 12507 (US); Ferreyra 12519 (US). I tentatively regard Cistanthe paniculata and Calandrinia ruizii Macbride as belonging to a single polymorphic species based on hints to this effect in the literature and my inability to readily sort the specimens. According to Macbride’s (1937) key and description, Cistanthe paniculata differs from Ca- landrinia ruizii in having a more exserted capsule, more densely pubescent seeds, and “obovate-oblong” rather than “broadly obovate or subrotund” leaves with acuminate rather than “merely apiculate” apices. Macbride (1937) listed Calandrinia adenosperma I.M. Johnston as a synonym of Cistanthe panicu- lata, but he had previously noted (Macbride 1931) that Calandrinia ruiz was also similar to Calandrinia adenosperma. Johnston’s (1928) description of Calandrinia adenosperma essentially bridges the gap between the descrip- tions of Cistanthe paniculata and Calandrinia ruizi: As in Calandrinia rutzit, the sepal and capsule lengths in Calandrinia adenosperma are equal, and the upper leaves are subamplexicaul and orbicular-ovate with obtuse apices. Cistanthe paniculata, Calandrinia ruizii, and Calandrinia adenosperma all reportedly have hairy seeds (Johnston 1928; Kelley 1973; Macbride 1937), but I have found glabrous seeds, apparently not correlated with any other trait in Hershkovitz: Taxonomic notes in Portulacaceae 217 several of the specimens. Label data from the cited collections also indicate that flower color (yellow vs. red) also varies independently of other traits (cf. Macbride 1937). Macbride (1931, 1937) and Johnston (1928) both believed Cistanthe pan- iculata to be especially closely related to Cistanthe grandiflora. Preliminary studies (Hershkovitz 1990b) indicate that the latter, also a polymorphic species complex, is distinctive in terms of leaf venation and perhaps also stomatal morphology and stomatal] density. In leaf venation, Czstanthe paniculata more closely resembles Cistanthe longiscapa than Crstanthe lamprosperma(I.M. John- ston), ined. (Hershkovitz 1990b). I have elsewhere (Hershkovitz 1990b) cited Ferreyra 10486 as Custanthe cf. ruizit (Macbride) ined. Cistanthe picta (Gillies er Arn. in Cheek) Carolin ez Hershkovitz, comb. nov. BASIONYM: Calandrinia picta Gillies ez Arn. 2n Cheek, Edin- burgh J. Nat. Geogr. Sci. 3:355. 1831. SPECIMENS EXAMINED: ARGENTINA. Mendoza: Gerth 107(L). San Juan: Castellanos 15588 (US). CHILE. Without locality: Gay s.n. (photograph of type of Calandrinia ob- longifolia Barnéoud in Gay, US ez P). Without province: Meyen s.n. “Cord. St. Fernando” (B). Aconcagua: Hutchinson 9& (UC, US); Morrison 17304 (NA, UC); West 5169 (UC); Zollner 3642 (L); Zollner 6548 (NA); Zollner 11082(NA). Antofagasta: Werdermann 1038 (US, UC). Atacama: Johnston 6218 (US). Coquimbo: Worth & Morrison 16584 (NA, UC). Metropolitana: Grandjot s.n., “XII. 32” (M); Grandjot s.n., “XII. 34” (M): Hastings 471 (UC); Joseph 2956 (US); Kuntze s.n., “I 92” (US); Morrison 16786 (NA, UC); Wer- dermann 648 (UC, CAS). O’Higgins: Gay s.n. (photograph, not type material fide Peralta [1988], US er G-DEL); Pennell 12279 (F). Cistanthe picta and related members of Reiche’s (1897, 1898) Calandrinia sect. Andinae are readily distinguishable from other Czstanthe by their: Thick, perennial roots; spreading habit; short stems bearing numerous, usually obo- vate to ovate, petiolate leaves; and congested cymules occurring immediately above the leafy stems and bearing relatively short pedicelled flowers and per- sistent, conspicuously black etched bracts. Also, the plants occur at higher elevation (mostly 2500 to 3500 m) than other species of Cistanthe. Despite the distinctiveness of this group, Reiche’s (1898) key separates Calandrimia sect. Andinae from other members of Cistanthe sect. Cistanthe only with re- spect to leaf shape, although leaf shape is highly variable throughout the genus (Hershkovitz 1990b). Cistanthe picta appears to represent a highly polymorphic entity from which notable variants have been segregated. Without seeing additional and type material, ] cannot make any judgment regarding the distinctiveness of 218 PH YTOLDO'CGLA volume 70(3):209-225 March 1991 most of the segregates recognized or listed in synonymy by Reiche (1898). Those recognized here are C. picta var. frigida (Barnéoud in Gay) Hershkovitz (see below) and Czstanthe macrocalyz (see above). I do note, however, that Johnston annotated his Atacama collection (J. 6218) as possibly representing a new species. It has conspicuous reniform bracts ca. 5 mm wide - roughly twice the norm. A surprisingly large number of specimens of Cistanthe picta have been annotated, sometimes emphatically, mostly anonymously, as Calandrinia are- naria Cham. The type and subsequent descriptions and illustrations of Calan- drinia arenaria Cham. (Chamisso 1831; Hooker & Arnott 1833; Lindley 1833) all clearly pertain to Cistanthe arenaria as delimited by Reiche (1898). Cistanthe picta (Gillies ez Arn. in Cheek) Carolin ez Hershkovitz var. frigida (Barnéoud in Gay) Hershkovitz, comb. nov. BASIONYM: Ca- landrinia frigida Barnéoud in Gay, Fl. Chil. '|Hist. Fisica y Politica de Chile, Botanica, 2:495. 1847 (“1846”). SPECIMENS EXAMINED: CHILE. Without province: Gay s.n., “1333 34” (P). Coquimbo: Gay s.n. (photograph of type of Calandrinia frigida, fide Peralta {1988}, US ez P): Morrison 16992 (NA, UC). Peralta (1988) listed this variety in synonymy with the species. This vari- ety occurs at higher altitudes than the rest of the species, which may account for its reduced morphology. Kelley (1973), however, reported glabrous, collic- ulate seeds, like those of Morrison 16992, in Cistanthe picta var. frigida. The Gay collection has no seeds. Leaves of this specimen also lack sinuous and ribbonlike veins that occur in Cistanthe picta and nearly all other species of Cistanthe (Hershkovitz 1990b). Cistanthe weberbaueri (Diels) Carolin ez Hershkovitz, comb. nov. BA- SIONYM: Calandrinia weberbauer: Diels, Bot. Jahrb. Syst. 37:399. 1906. SPECIMENS EXAMINED: PERU. Arequipa: Ferreyra 11723 (US); Fer- reyra 12006 (US). Lima: Weberbauer 5321 (F). As noted above, the distinction between Cistanthe weberbaueri and C. lin- gulata is tenuous. Weberbauer 5321 is unusual: It has the broadest leaves of any specimens I have examined in the lingulata/weberbaueri complex, and it has distinctly hairy seeds. The seed morphology of the other cited spec- imens accords with Kelley (1973), who reported pusticulate-tomentose seeds in Cistanthe lingulata, Cistanthe weberbaueri, and Calandrinia quivensis. We- berbauer 5321 was apparently miscited by Macbride (1937) as “5221.” Specimens not determined: The specimens listed below belong to species of Cistanthe sect. Cistanthe listed above, species not yet transferred from Ca- landrinia, or possibly to new species. The specimens are all from Chile. Hershkovitz: Taxonomic notes in Portulacaceae 219 Beetle 26145 (NA, UC), Region Atacama - Johnston annotated these sheets as Cistanthe cymosa, probably because of the congested inflorescence and yellow flowers. The habit, stems, leaves, and seeds are not as in Cistanthe cymosa, however. The plant looks more like a large, grotesque specimen of Cistanthe arenarza. Bertero 683 (P), Region O’Higgins - suggests Cistanthe arenaria or Calan- drinia solisz, and possibly a mixture of both forms. Joseph 2785 (US), Region Metropolitana - lax, spreading habit of Cistanthe arenarza but seeds and pedicels of Cistanthe grandiflora. Philippi s.n. (B), locality illegible - I have elsewhere (Hershkovitz 1990b) referred to this collection as Cistanthe glaucopurpurea (Reiche), ined. — it clearly is not that species, but I do not know what to call it. Rose & Rose 19318 (US), Region Coquimbo - suggests either Cistanthe fenzli or Cistanthe arenarza, but the hairy seeds indicate a relationship to the Cistanthe grandiflora complex. Simon 454 (UC), Region Coquimbo - suggests Cistanthe arenaria but much larger. Worth & Morrison 16184 (NA, UC), Region Atacama - I have elsewhere (Hershkovitz 1990b) cited this specimen as Cistanthe lamprosperma(I.M. Johnston), ined. - it is not that species. It is very similar to Cistanthe longiscapa and was determined by Johnston as Calandrinia litoralis. The leaf epidermis, however, is prominently papillate - a feature Johnston (1929) emphasized in his description of Calandrinia lamprosperma. As Johnston (1929) noted, however, the latter species is otherwise most similar to members of the Cistanthe grandiflora complex. Worth & Morrison 16568 (G, NA, US), Region Coquimbo - Johnston an- notated this specimen “nsp.?” In habit and leaves, it is identical to Cistanthe picta, but the few flowered inflorescence, long pedicels, and large flowers with unmarked sepals are unusual. Zollner 6885 (L, NA), Region Coquimbo - like Rose & Rose 19318 (see above). Zollner 9807 (NA), Region Coquimbo - like Philippi s.n. (B — see above). Zollner 10636 (NA), Region Coquimbo - mixed collection including one plant similar to Joseph 2785 (see above) and three plants suggesting Cistanthe arenaria but with hairy rather than merely pusticulate seeds. 220 PHY TP0g-8 CiA volume 70(3):209-225 March 1991] B. Cistanthe sect. Amarantoizdes Cistanthe sect. Amarantoides (Reiche) Carolin ez Hershkovitz, Phytologia 68:269. 1990. This section includes all members of Calandrinia sect. Amarantoideae. | had previously (Hershkovitz 1990a, 1990b) employed Reiche’s (1897) spelling for this section, but the suffix “-oides” is correct. Cistanthe calycina (Philippi) Carolin ez Hershkovitz, comb. nov. BASIONYM: Calandrinia calycina Philippi, Fl. Atacam. 21. 1860, and Viage al De- sierto de Atacama 196. 1860. SPECIMENS EXAMINED: CHILE. Antofagasta: Johnston 3590 (US); Johnston 5252 (US); Johnston 5258 (US); Johnston 5818 (US): Lourtezg 2580 (US); Werdermann 868 (CAS, UC); Werdermann 1033 (CAS; UC); Worth & Morrison 15825 (NA, UC). Atacama: Joseph 4976 (US); Werdermann 418 (CAS, UC, US). PERU. Arequipa: Vargas 7992 (US). Cistanthe calycina is distinguished from other species of Cistanthe sect. Amarantoides by its diminutive habit and linear to linear lanceolate leaves. Macbride (1937) did not list this species for Peru. Cullen (1953) listed it for Argentina, but the specimen cited may belong to Calandrinia minuscula Cullen (D. Ford, pers. comm.). I have not seen adequate material of the latter species, so I have not provided a new combination here. Cistanthe densiflora (Barnéoud in Gay) Hershkovitz, comb. nov. BA- SIONYM: Calandrinia densiflora Barnéoud in Gay, Fl. Chil. | Hist. Fisica y Politica de Chile, Botanica] 2:502. 1847 (“1846”). SPECIMENS EXAMINED: ARGENTINA. San Juan: Cabrera, et al. 29553 (US); Castellanos 15520 (US). CHILE. Antofagasta: Martin 540 (UC). Atacama: Philippi s.n. “1888” (B); Zollner 4282 (L); Zollner 5053 (L). Coquimbo: Gay s.n. (photograph of type, US ex P); Gay s.n. (photograph of type of Calandrinia fasciculata Philippi, fide Peralta [1988], US ez W). Peralta (1988) listed two species in synonymy with Cuistanthe denszflora that Reiche (1898) had recognized as distinct: Calandrinia fasciculata Philippi and Calandrinia barneoudi Philippi. Reiche (1898), however, mentioned Cis- tanthe densiflora as a problematic species pertaining to section Amarantozdes. The inflorescence morphology in the type specimen of Cistanthe densiflora is intermediate between the more capitate cymes of Calandrinia fasciculata Hershkovitz: Taxonomic notes in Portulacaceae 22M and the more open inflorescence with several spikelike cymules of Calandrinza spicata Philippi (also recognized by Reiche, 1898). If Calandrinza fasciculata is conspecific with Cistanthe denszflora, then Calandrinia spicata probably is also. The extremes, however, are quite distinct. Cistanthe salsoloides (Barnéoud in Gay) Carolin ez Hershkovitz. Phytologia 68:269. 1990. SPECIMENS EXAMINED: ARGENTINA. San Juan: Fabris & Mar- chionni 2279 (US). CHILE. Without data: one sheet (US no. 1038131). Antofagasta: Wer- dermann 1048 (leg. Francke; CAS, F, UC - two sheets, US - two sheets); Werdermann 1015 (US). Atacama: Biese 720 (CAS); Greninger 13 (CAS); Harding 22850 (US); Zollner 10633 (NA). Coquimbo: Gay 918 (photograph of type, US ez P). Tarapaca: Brandegee s.n. (fragment, UC). Two additional species of Cistanthe sect. Amarantozdes listed by Reiche (1898), Calandrinia acuminata Philippi and Calandrinia spicigera Philippi, pertain to Cistanthe salsoloides. Calandrinia acuminata supposedly differs in having more elongate basal leaves, but these leaves possibly represent ephemeral basal leaves that also occur in Crstanthe salsoloides, as evident from one of the US sheets of the widely distributed Werdermann 1048 collection. Based on Reiche’s (1898) description, Calandrinia spicigera is to Cistanthe salsoloides what Calandrinia spicata is to Cistanthe densiflora — the inflorescences in Ca- landrinia spicigera and Calandrinia spicata are similar (see above). I have not seen any material that corresponds to the description of Calandrinia spicigera, however. C. Cistanthe sect. Philippiamra (Kuntze) Hershkovitz Cistanthe sect. Philippiamra (Kuntze) Hershkovitz, Phytologia 68:269. 1990. Vegetatively, members of this section are extremely similar to Cistanthe (Amarantoides) salsoloides. I doubt that sterile material could be confidently identified as belonging to one or the other section. The geographic range of Cistanthe sect. Philippiamra is smaller than that of Cistanthe sect. Amaran- toides, though, and the former apparently does not occur in Argentina. Cistanthe amarantoides (Philippi) Carolin ez Hershkovitz, comb. nov. BASIONYM: Silvaea amarantoides Philippi, Fl. Atacam. 22. 1860, and Viage al Desierto de Atacama 196. 1860. 222 PHYTOLOGIA volume 70(3):209-225 March 1991 SPECIMENS EXAMINED: CHILE. Antofagasta: Philippi s.n. (number illegible, possibly 34; poor photograph of type, det. C. Munoz-P., UC ex SGO). Atacama: Werdermann 477 (CAS). Philippi (1860a, 1860b) and Reiche (1898) emphasized a distinction in growth form between Cistanthe amarantoides and Cistanthe celosioides, the former having simple and prostrate rather than ramified and erect branches, respectively. Johnston (1929), however, distinguished the species on the basis of bract morphology - pale, elliptic-oblong and relatively small in Crstanthe amarantoides vs. colored and broadly ovate in Cistanthe celosioides. This bract size and shape distinction is not emphasized in the type descriptions (cf. Philippi 1860a, 1860b). The only specimen IJ have seen that stands out as having the bract morphology described by Johnston (1929) is that listed above. I have elsewhere (Hershkovitz 1990b) listed Werdermann 862 (see below) under Cistanthe amarantozdes because of its branching habit, but Johnston (1929) cited this specimen as Cistanthe celosiozdes. Cistanthe celosioides (Philippi) Carolin ez Hershkovitz, Phytologia 68:269. 1990. SPECIMENS EXAMINED: CHILE. Antofagasta: Biese 613 (UC); Mor- rison 17092 (NA); Philippi s.n. (poor photograph of type, det. C. Munoz-P., UC ez SGO); Werdermann 862 (CAS, UC); Werdermann 998 (CAS); West 3859 (UC); Worth & Morrison 15820 (UC). Atacama: Beetle 26152 (NA); Beetle 26183 (NA; UC); Marticorena, et al. 40623 (F); Philippi s.n. (leg. San Roman; poor photograph of type of Sizlvaea capitata Philippi, det. C. Munoz- P.; UC ex SGO); Philippi s.n. (leg. F. Ph.; poor photograph of type of Silvaea fastigata Philippi, det. C. Munoz-P.; UC ex SGO); Werdermann 165 (Silvaea capitata Philippi fide Werdermann; CAS, UC); Werdermann 875 (CAS, UC). Tacna: Werdermann 731 (CAS, UC). Tarapaca: Zollner 7811 (NA); Zollner 9603 (NA). According to Johnston (1929), Cistanthe celosioides, Silvaea pachyphylla Philippi and Silvaea fastigaia Philippi, which Reiche (1898) recognized as dis- tinct, are phases of one variable species. ] cannot confidently sort specimens into the three species, so I have referred them all to Cistanthe celosioides. 2. New combination and status for Monocosmia Morphological evidence (Carolin 1987; Hershkovitz 1990b) indicates that the monotypic genus Monocosmiais closely related to Calandrinia sect. Calan- drinia (syn. C. sect. Compressae Reiche), and that both are probably derived from a common ancestor in Calandrinia sect. Acaules Reiche. On cladistic Hershkovitz: Taxonomic notes in Portulacaceae 223 principles, therefore, Afonocosmiza should be included in Calandrinia. In rec- ognizing Monocosmia as a distinct section of Calandrinia, I realize that Ca- landrinia sect. Acaules and/or Calandrinia might then be paraphyletic (cf. Hershkovitz 1990b). Additional investigation will be required to confirm this, however. Calandrinia Kunth in H.B.K. sect. Monocosmia (Fenzl in Endl. & Fenzl) Hershkovitz, comb. & stat. nov. BASIONYM: Monocosmia Fenzl in Endl. & Fenzl, Nov. stirp. dec. 84. 1839. TYPE: Talinum monandrum Ruiz Lopez & Pavén (= Calandrinia monandra [Ruiz Lopez & Pavon|} DC.). 3. New combination in Talisnum Talinum galapagosum (H. St. John) Hershkovitz, comb. nov. BASIONYM: Calandrinia galapagosa H. St. John, Amer. J. Bot. 24:95. 1937. Kelley & Swanson (1986) established Calandrinza sect. Pachypodae to ac- commodate two species, Calandrinia pachypoda Diels (with two subspecies) and Calandrinia galapagosa H. St. John. The capsule, seed, and pollen traits (cf. Kelley & Swanson 1986) that distinguish Calandrinza sect. Pachypodae from other sections of Calandrinia sensu lato (cf. Kelley 1973; Hershkovitz 1990b) characterize species of the genus Talinum. Calandrinia pachypoda seems to represent high altitude and/or xeric forms of the pantropical weed Talinum triangulare (Jacq.) Willd. Whether these forms deserve taxonomic recognition is unclear but should depend on the overall degree and pattern of variation in the latter species. Talinum galapagosum has small linear leaves and is thus most likely related to vegetatively similar species that are fairly well diversified in North America, although at least one of these species occurs in Argentina. While the precise taxonomic relationship of Talinum galapagosum remains uncertain, its geographic isolation from similar congeners warrants its continued recognition as a distinct species. ACKNOWLEDGMENTS I thank Walter Kelley, (especially) Donna Ford, Alan Smith, and someone else for assistance in working up this manuscript. 224 PHY TOLOGIA volume 70(3):209-225 March 1991 LITERATURE CITED Carolin, R.C. 1987. A review of the family Portulacaceae. Austral. J. Bot. 35:383-412. . in press. Portulacaceae, in K. Kubitzki (ed.), Families and genera of flowering plants. Chamisso, A.L. von. 1831. De plantas in Expeditione Romanzoffiana obser- vatis disserere pergitum. Linnaea 6:545-592. Cullen, D.A.S. de. 1953. Las especies Argentinas del género Calandrinia (Portulacaceae). Bol. Soc. Argent. Bot. 5:1-29. Hershkovitz, M.A. 1990a. Nomenclatural changes in Portulacaceae. Phy- tologia 68:267-270. 1990. . 1990b. Phylogenetic and morphological studies in Portulacaceae. Ph. D. dissertation, University of California, Davis. Hooker, W.J. & G.A.W. Arnott. 1833. Contributions towards a flora of South America and the islands of the Pacific [part 2]. Bot. Misc. 3:302-367. Johnston, I.M. 1928. Some undescribed American spermatophytes. Contr. Gray Herb. 85:1-164. . 1929. Papers on the flora of northern Chile. Contr. Gray Herb. 85:1-164. Kelley, W.A. 1973. Pollen morphology and relationships in the genus Ca- landrinia H.B.K. (Portulacaceae). M.S. thesis, California State Univ., Northridge. & J.R. Swanson. 1986. A new section and subspecies of Calandrinia (Portulacaceae). Phytologia 60:171-179. Lindley, J. 1833. Calandrinia arenaria. Bot. Reg. 19:1605. Macbride, J.F. 1931. Spermatophytes, mostly Peruvian-III. Publ. Field Mus. Nat. Hist., Bot. Ser. 11:1-35 [publication 288). . 1937. Portulacaceae. Pp. 562-573 in J.F. Macbride, Flora of Peru. Vol. 13, pt. 2 [Publ. Field Mus. Nat. Hist., Bot. Ser.], Field Museum of Natural History, Chicago. Peralta, I.E. 1988. Sinopsis de las especies de Calandrinia (Portulacaceae) de los Andes Mendocinos. Bol. Soc. Argent. Bot. 25:511-537. Hershkovitz: Taxonomic notes in Portulacaceae 225 Philippi, R.A. 1860a. Florula atacamensis. Halis saxonum [Halle], Sumptibus Eduardi Anton. - 1860b. Vrage al Deszerto de Atacama. Halle en Sajonia, Libraria de Eduardo Anton. [includes reprinting of Philippi (1860a)). . 1894a. Plantas nuevas chilenas (continuacién). Anales Univ. Chile 85:167-195. . 1894b. Plantas nuevas chilenas (continuacion). Anales Univ. Chile 85:299-324. Reiche, K. 1897. Zur systematik der Chilenischen Arten der Gattung Calan- drinia. Ber. Deutsch. Bot. Ges. 15:493-503. ———. 1898. Flora de Chile. Vol. 2. University of Chile, Santiago. Phytologia (March 1991) 70(3):226-228. BOOKS RECEIVED Current Research on the Tribe Delphineae Warming (Ranuncu- laceae). Collecteana Botanica, volume 19. Cesar Blanché & Angel M. Romo (eds.). Institut Botanic de Barcelona, Avgda. Muntanyans s/n, Montjuic, 08038 Barcelona, Spain. 1990. 160 pp. $20.00 (paper) ISSN 0010-0730. Eighteen authors have contributed ten papers on current re- search in the Delphineae (Aconitum, Consolida, and Delphinium. The scope of the papers range from taxonomy (summaries based on morphology, chemistry, and other features) and ecology (pollina- tion biology and distributions) to economic uses (stock poisoning and cultivation). This volume represents the first in a series of special volumes of Collecteana Botanica in which particular taxo- nomic groups will be featured. The regular volumes of contributed papers will also be continued. Elton’s Ecologists: A History of the Bureau of Animal Population. Peter Crowcroft. The University of Chicago Press, 5801 S. Ellis Ave., Chicago, IL 60637. 1991. xx. 177 pp. $14.95 (paper) ISBN 0-226-12148-8; $35.00 (cloth) ISBN 0-226-12146-1. Crowcroft’s book is a sentimental summary of the history of the Bureau of Animal Population at Oxford University. Many projects undertaken at the Bureau during its lifetime are described. The leading character in the book and in the past activities of the Bureau is Crowcroft’s mentor, Charles Elton. 226 Books received PTOR Gardening with Native Wild Flowers. Samuel B. Jones, Jr. & Leonard E. Foote. Timber Press, 9999 SW Wilshire, Portland, OR 97225. 1990. 195 pp. $32.95 + $3.00 shipping (hardcover) ISBN 0-88192-175-0. This book enumerates some of the features of wildflowers, and advantages of growing native plants over introduced species. The text continues with an outline and background information for de- velopment of a plan for a wildflower garden. Extensive information is included to summarize the physical and chemical requirements for growth and propagation of native plants in the garden. An en- tire chapter is devoted to producing meadowlike effects in garden situations. Six chapters list native species suitable for use in the garden. One chapter each is devoted to shade tolerant wildflowers, sun loving wildflowers, grasses and grasslike plants, ferns and al- hes, groundcovers, and wetland plants. One of the most important features of the book is a listing of commercial sources for seed and plants of wildflowers. Although the scope of the book is primarily directed at gardens and native plants from the eastern half of the United States, the listing of commercial] sources for plants includes many companies from the western part of the country. Hosta: The Flowering Foliage Plant. Diana Grenfell. Timber Press, 9999 SW Wilshire, Portland, OR 97225. 1991. 240 pp. $39.95 + $3.00 ship- ping (hardcover) ISBN 0-88192-188-2. This book is a compendium of current knowledge about the cultivation of Hosta. In addition to a summary of the history of cultivation of the genus, descriptions of over 150 cultivars are given along with cultivation notes and references to illustrations. Cultivation methods and descriptions of garden situations where Hosta does well are included. Many of the plants are shown in garden settings in a series of excellent colored plates. Many others are shown in drawings. 228 PAY TOLOGIA volume 70(3):226-228 March 1991 Plant Physiology, Biochemistry and Molecular Biology. David T. Dennis & David H. Turpin (eds.). Longman Scientific & Technical, Ltd., available in the United States from John Wiley & Sons, Inc., 1 Wiley Drive, Somerset, NJ 08875-1272. 1991. xi. 529 pp. $49.95 (hardcover) ISBN 0-470-21578-X. A total of 47 authors have contributed to this volume. The book is organized into nine major sections with each section in- cluding chapters on topics including metabolic control, descriptions of metabolic pathways, and prospects for “improvement” of plant metabolism. ard New York Botanical il 4920 | | | len Libra | | i 3 5185 00 | | | 288 information for Authors Articles from botanical systematics and ecology, including — biographical sketches, critical reviews and summaries of literature — will be considered for publication in PHYTOLOGIA. Manuscripts — may be submitted either on computer diskette, or as typescript. 3 Diskettes will be returned to authors after action has been taken on. the manuscript. 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