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The Genus Geraniuml. (Geraniaceae) in North Amemca

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The Genus Geraniuml. (Geraniaceae) in North Amemca THE GENUS GERANIUML. (GERANIACEAE) IN NORTH AMEMCA. II. PERENNIAL SPECIES by CARLOS AEDO Real Jardín Botánico, CSIC. Plaza de Murillo, 2. E-28014 Madrid ([email protected]) Resumen AEDO, C. (2001). El género Geranium L. (Geraniaceae) en Norteamérica. II. Especies perennes. Anales Jará. Bot. Madrid 59(1): 3-65 (en inglés). Se revisan las especies perennes de Geranium de Norteamérica al N de México. Se aceptan 19 especies de las cuales 17 pertenecen al subg. Geranium y dos al subg. Robertium. Geranium erianthum, G. maculatum, G. oreganum, G. sylvaticum, G. caespitosum, G. californicum, G. lentum, G. wislizenii, G. richardsonii, G. viscosissimum son las especies autóctonas, y G. pratense, G. sanguineum, G. potentilloides, G. sibiricum, G. thunbergii, G. retrorsum, G. solanderi, G. pyrenaicum y G. palmatum, las introducidas tanto desde Europa como desde Asia u Oceanía. Geranium eremophilum, G.j"remonta, G. parryi, G. margínale y G. cowenii, a veces aceptados en la literatura reciente, son considerados como sinónimos de G. caespitosum. Del mismo modo, se sinonimiza G. strigosum y G. attenuilobum a G. viscosissimum y G. concinnum a G. californicum. Se revisa la nomenclatura de todas las especies y se designan diecisiete lectótipos. Se incluyen descripciones completas, una clave, mapas de distribución e ilustraciones. Palabras clave: Geranium, Geraniaceae, taxonomía, tipificaciones, Norteamérica. Abstract AEDO, C. (2001). The genus Geranium L. (Geraniaceae) in North America. II. Perennial species. Anales Jard. Bot. Madrid 59(1): 3-65. Perennial species of Geranium from North America north of México are revised. Nineteen species are accepted, of which seventeen belong in subg. Geranium and two belong in subg. Robertium. Geranium erianthum, G. maculatum, G. oreganum, G. sylvaticum, G. caespito- sum, G. californicum, G. lentum, G. wislizenii, G. richardsonii, G. viscosissimum are native, whereas G. pratense, G. sanguineum, G. potentilloides, G. sibiricum, G. thunbergii, G. retrorsum, G. solanderi, G. pyrenaicum and G. palmatum were introduced from Europe, Asia, and Oceania. Geranium eremophilum, G. fremontü, G. parryi, G. marginóle and G. cowenii, sometimes accepted in currant literature, are considerad to be synonyms of G. caespitosum. Similarly, G. strigosum and G. attenuilobum are synonymized to G. visco- sissimum, and G. concinnum to G. californicum. Nomenclatura for all species is reviewed, and seventeen lectotypes are designated. Descriptions, key, distribution maps, and illustra- tions are included. Key words: Geranium, Geraniaceae, taxonomy, typification, North America. ANALES JARDÍN BOTÁNICO DE MADRID, 59(1) 2001 INTRODUCTION MATERIALS AND METHODS The study of Geranium from North Ameri- This revisión is based on more than 2000 ca has been divided in two parts for operative herbarium specimens from the following reasons. The first part, including annual (and herbaria: ARIZ, BM, CAN, CM, COLÓ, biennial) species was published in AEDO DAO, F, GH, ILL, JEPS, K, LE, MA, (2000). The second part revising perennial MEXU, MICH, MIN, MO, MSC, NEB, species is presented here. NMC, NY, OKL, P, PH, SD, TEX, UC, General information about North Ameri- UNM, US, V, VT, W, and WTU. Further- can Geranium was given in the introduction more, microfiches, photographs, and other of AEDO (2000), and is not repeated. Addi- material have been examined from BM and tionally, the study by JONES & JONES (1943) LINN. Curators from ALTA, ACAD. should be remarked. These authors revised BRY, MOAR, UAT, and UBC, kindly an- perennial species from the United States and swered our petition, but they did not find Canada. They considered eighteen species any of the requested specimens in their from which only one was introduced herbaria. (G. pratense). Their treatment is more ana- The ten native species have been rather lytic, since they split G. caespitosum in six thoroughly studied in some aspects such as species, G. viscosissimum in three species, morphology, synonymy, and distribution. In and G. californicum in two species. those native species, a search for micromor- Geranium comprises in the área of North phological characters has been carried out us- America flora 30 species (both annuals and ing scanning electrón microscopy (SEM). perennials), of which 23 belong in subg. Samples were glued to aluminium stubs, coat- Geranium and 7 belong in subg. Robertium. ed with 40-50 nm gold, and examined with a All native species (13) belong in subg. Gera- JEOL-TSM T330A scanning electrón micro- nium. Among the 17 introduced species, 10 scopeat 15 kV. belong in subg. Geranium and 7 belong in Short glandular hairs (<40 jim long), usu- subg. Robertium. From these introduced ally oonstituted by two cells (although they species 13 have arisen from the Old World sometimes have a bicellular foot, Fig. lf), (Europe and neighbouring áreas), 3 from are present in all species here studied but Oceania, and 1 from East Asia. they are not considered in descriptions Some remarks should be also made about since they are only evident at high magnifi- the distribution of native perennial species. cation. Two of them also occur in another conti- For introduced species only synonyms de- nent: G. erianthum in Asia, G. sylvaticum in scribed from North America are indicated, Europe. Among the remaining, two are while a comprehensive synonymy is provided widespread in North America, one is re- for natives ones. stricted to the Eastern half of the continent, The área covered in this paper includes and eight are restricted to the opposite half. North America north of México plus Green- Thus, the Rocky Mountains and western ár- land, since this study has been carried out for eas seem to be the richest in Geranium the Flora of North America project, which in- species. The number of endemic species is cludes the above mentioned área. For eight eight. Geranium caespitosum, G. caroli- native Geranium, maps cover the total range nianum, and G. wislizenii have their Southern of the each species. However, for G. erian- limit in México. Finally, six species do not thum and G. sylvaticum maps only include occur in Canada: G. caespitosum, G. cali- American localities. For the introduced fornicum, G. lentum, G. oreganum, G. te- species maps are restricted to its North Ame- xanum, and G. wislizenii. rican área. C. AEDO: THE GENUS GERANIUM L. IN NORTH AMERICA. II Fig. I.-SEM photomicrographK showing indumentiim of some perennial Geranium native from North America: a, pcdicel of G. erianthum [Calder 5411 (DAO)]; b, pedicel of G. viscosissimum {Peck 18320 (ORE)]; c, apex of a glandular hair on the sepal of G. maculatum [Ahles 82611 <VT)]; d, apex of a glandular haír on the pedicel of G. vis- cosissimum [Peck 18320 (ORE)]; e, pedicel of G. maculatum [Ahles 82611 (VT)J; f, small glandular hair on the pedicel of G. caespitosum [Spellenberg 6324 (NMC)J. ANALES JARDÍN BOTÁNICO DE MADRID, 59(1) 2001 KEY TO THE SPECIES 17. Nectarious dorsally láñate; petáis usually white 16. G. richardsonii l.Cymules 1-flowered 2 17. Nectarious with a tuft of hairs at the top, dor- 1. Cymules 2-flowered 4 sally glabrous; petáis white orpink 18 2. Petáis more than 15 nun long 18. Ratio fruit length / stigmatic remains length = 6. G. sanguineum 6-8.2; basal leaves 7.1-16 cm wide 2. Petáis 5-10 mm long 3 17. G. viscosissimum 3. Leaves with middle segment obtríangular, 3 18. Ratio fruit length / stigmatic remains length = (-5)-lobed at the apex 7. G. potentilloides 3.8-5.5; basal leaves 3-7.5 cm wide 19 3. Leaves with middle segment rhombic, 7-9- 19. Fruit 22-31(-33) mm long; rostrum with nar- lobed in the di s tal half 8. G. sibiricum rowed apex 2-4 mm long; petáis usually purpusii; 4. Mericarps without basal callus 5 pedicels short (1.8-12 cm)... 12. G. caespitosum 4. Mericarps with basal callus 7 19. Fruit 32-42 mm long; rostrum with narrowed 5. Petáis 7-11 mm long 18. G. pyrenaicum apex 5-6 mm long; petáis usually white; pedicels 5. Petáis 19-30 mm long 6 long (2.2-24 cm) 13. G. californicum 6. Mericarps reticulate 19. G. palmatum 6. Mericarps smooth G. x magnifícum TAXONOMY 7. Petáis glabrous on the adaxial surface, ciliate at the base 8 I. Geranium subg. Geranium 7. Petáis hairy on 1/3-3/4 of their adaxial surface Fruit of "seed-ejection-type," with seeds 15 actívely discharged by the explosive recoiling 8. Petáis 5-12 mm long; fruit 13-23 mm long ... 9 8. Petáis 12-21 mm long; fruit 27-52 mm long of the awn, which remains together with the 11 mericarp attached to the columella; mericarps 9. Calyx with glandular hairs ... 9. G. thunbergii with basal callus or prong. 9. Calyx without glandular hairs 10 1. Geranium erianthum DC, Prodr. 1: 641 10. Pedicels with retrorse, appressed, eglandular hairs; sepal muero 0.7-1 mm long, without bris- (1824) G. pratense var. erianthum (DC.) B. tles 10. G. retrorsum Boivin in Naturaliste Cañad. 93:1060 (1967) 10. Pedicels with patent or reflexed (never ap- Ind. loe: "in KamtschatM et Amer. bor. et pressed), eglandular hairs; sepal muero 0.4-0.5 occidentali. Nelson. (v. s.)" mm long ending in 1-3 bristles 0.3-0.4 mm long Typus: USA. Alaska, Newenham, Nelson 11. G. solanderi s.n. (lectotype, here designated, G-DC mi- 11. Staminal filaments with eglandular hairs 1-3 cro!) [58°39'N, 162°10'W] mm long 1. G. erianthum = G. erianthum var. elatum Maxim., Prim. 11. Staminal filaments with eglandular hairs up to Fl. Amur.: 71 (1859) 0.7 mm long 12 G. elatum (Maxim.) R. Knuth in Engl., 12. Pedicels with eglandular hairs, without glandu- Pflanzenr. IV.129 (Heft 53): 113 (1912) lar hairs 2. G. maculatum 12. Pedicels densely glandular-hairy 13 Ind.
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