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Flora Mesoamericana, Volume 2 (2), Rosaceae, page 1 of 61

First published on the Flora Mesoamericana Website, 19 Dec. 2011.

99. ROSACEAE (in part) By F.R. Barrie.

Herbs, shrubs or trees, often armed with thorns, spines or prickles. Stipules usually present, conspicuous. Leaves alternate, persistent or deciduous, simple or compound. Inflorescence a spike, raceme or cyme, or flowers solitary. Flowers perfect, rarely unisexual, pedicellate or sessile, the perianth perigynous, the axis sometimes enlarged to form a flat to conic or urceolate receptacle or a hypanthium bearing the bracteoles, calyx, corolla and stamens along the margin; bracteoles 4-5, sepals 4-5; petals 5 or sometimes absent; stamens 2 to many, often 10 or 20; carpels 1 to many, free or partially to wholly united, often connate with the receptacle; ovules 1-several per carpel; styles as many as the carpels, free or connate basally. Fruit a follicle, achene, drupe, hip or pome; seeds usually lacking endosperm; cotyledons often fleshy and convex. Approx. 100 genera, 3000 species, distributed world-wide but most strongly represented in the North temperate regions. Approx. 20 genera and 75 species in Mesoamerica. Bibliography: Kalkman, C. Fam.Gen. Vasc. Pl. VI: 343-386. 2007.

1. Leaves variously compound. 2. Plants armed with prickles. 3. Fruit an aggregate of fleshy druplets borne on an elarged receptacle. 3. Fruit of achenes borne inside a fleshy receptacle (hip). 2. Plants unarmed (fruits may have spines, prickles or awns). 4. Fruit of 1-several achenes within a persistent hypanthium bearing spines or barbs. 5. Leaves compound, the leaflets all of similar size. 1. Acaena 19. Rubus 18. Rosa

5. Leaves compound, the larger leaflets separated by one or more, significantly smaller, leaflet. 4. Fruit various, unarmed. 2. Agrimonia

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 2 of 61 6. Petals absent; the achenes enclosed within the persistent hypanthium. 10. Lachemilla 6. Petals present; the achenes not enclosed in the hypanthium. 7. Receptacle not greatly enlarged in fruit; obscured by the achenes. 8. Style persistent and elongate on mature achenes. 8. Style deciduous, not persistent on mature achenes. 7. Receptacle fleshy and enlarged in fruit; not obscured by the achenes. 9. Flowers solitary; petals yellow. 9. Flowers 3-10 per inflorescence, petals white. 1. Leaves simple. 10. Fruit dry, consisting of an aggregate of follicles or achenes, or of several to many fleshy druplets. 11. Fruit of fleshy druplets. 11. Fruit dry, of 1 or more follicles or achenes. 12. Fruit an aggregate of follicles; leaves not white-tomentose on underside. 20. Spiraea 12. Fruit of achenes; leaves white tomentose on underside. 10. Fruit fleshy and usually juicy; a drupe or pome. 13. Ovary superior, fruit a drupe. 13. Ovary inferior, fruit a pome. 14. Shrubs or trees, usually armed with thorns or the branches spine-tipped; carpels of the fruit bony at maturity. 15. Leaves less than 2 cm. 15. Leaves longer than 2 cm. 16. Leaves elliptic to obovate; flowers 3-8 per inflorescence. 3. Crataegus 8. Hesperomeles 15. Prunus 9. Holodiscus 19. Rubus 14. Potentilla (P. indica) 6. Fragaria 7. Geum 14. Potentilla

16. Leaves narrowly elliptic to oblong or oblanceolate; flowers 10-30 per inflorescence. 16. Pyracantha 14. Trees, usually unarmed; carpels of the fruit leathery or papery at maturity. 17. Inflorescence a compound corymb or panicle. 18. Mature leaves rusty-tomentose beneath; carpels of the ovary wholly connate. 5. Eriobotrya

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 3 of 61 18. Mature leaves not tomentose beneath; carpels free at the apex. 17. Inflorescence simple, not compound, or flowers solitary. 19. Carpels of the fruit containing 4 or more seeds. 19. Carpels of the fruit 1-2 seeded. 20. Ovary and fruit incompletely 6-10 celled. 20. Ovary and fruit 1-5 celled. 21. Styles connate at the base, fruit globose or subglobose, usually lacking stone cells. 12. Malus 21. Styles free at the base; fruit pyriform, containing numerous stone cells. 17. Pyrus 11. Malacomeles 4. Cydonia 13. Photinia

1. Acaena Mutis ex L. By F.R. Barrie.

Perennial herbs or subshrubs. Leaves alternate, imparipinnate; stipules adnate to the petioles, the tips free. Inflorescence a terminal or axillary spike or raceme. Flowers bracteate; hypanthium unarmed or with several to many spines; bracteoles absent; sepals 3 or 4; petals absent; stamens 2-4; carpels 1, rarely 2-4; ovules 1 per carpel, style terminal, stigma fimbriate. Fruit an achene enclosed in the persistent hypanthium, often armed with several to many glochidiate spines. Seed ovoid, testa thin. Approx.. 100 spp. Mesoamerica, South America, Australia, New Zealand.

1. Perennial herbs; leaves basal; inflorescence a compact, cylindrical head. 1. A. cylindristachya 1. Suffrutescent herbs or subshrubs; leaves cauline; inflorescence an elongate raceme, the flowers borne singly along its length. 2. A. elongata

1. Acaena cylindristachya A. Ruiz & Pavon, Fl. Peruv.1: 68 (1798). HT: Peru. Ruiz & Pavon s.n. (MA, photo F?). Illustr.: Fl. Peruv. t. 104, f. a (1798). (use as type?) Acaena cylindristachya var. nitidissima Bitter; A. cylindristachya subvar. pusilla Bitter; A. macrorhiza Hook. f.

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 4 of 61 Perennial herbs. Taproot stout; caudex simple, covered by the remains of old leaf bases. Leaves basal, the blade 10-15 2-5 cm; narrowly elliptic; leaflets 15-31, 10-25 5-8 mm, elliptic, subequal, sessile upper surface green, sparsely pubescent; lower surface silvery white, densely sericeous; margins serrate; apex acute; petioles sericeous. Inflorescence to c. 30 cm, peduncle erect, leafless or with 1-4 bractlike leaves on the lower half, terminating in a compact cylindrical head; 2-8 remote, lateral heads sometimes present; primary head with 50-150 flowers, secondary heads with 10-25 flowers. Flowers sessile; hypanthium c. 1 mm, sericeous; sepals 3, c. 1 mm, obovate; stamens 2; carpels 1. Fruit c. 3 mm, ellipsoid, puberulent, bearing 3-5 unequal, apical awns, to 5 m. Paramos. CR (Ramrez 446, F). 3000-3300 m. (Mesoamerica, Colombia, Venezuela.)

2. Acaena elongata L., Mant. Pl. 200 (1771). Neotype (designated by Barrie, 1993): Peru, Mutis no. 774, Acaena (MA). Illustr.: Ann. Missouri Bot. Gard. 37: 163, t. 69 (1950). Acaena agrimonioides Kunth, A. elongata subvar. compacta Bitter, A elongata var. robusta Bitter, A. lappacea Ruiz. & Pav. Suffrutescent herbs or more commonly a subshrub or shrub, to 1 m. Branches glabrate or with a few, scattered hairs, the bark red. Leaves cauline, the blade 2.5-4 .1.53 cm; elliptic to obovate; leaflets 7-11, 8-12 3-7 mm; elliptic; sessile; upper surface glabrate or with tufted hairs in the axes; lower surface sparsely sericeous; base cuneate; margin crenate-serrate; apex acute. Inflorescence an elongate raceme, 5-30 cm; flowers borne singly on short, lateral branches. Flowers pedicellate, the pedicels 1-4 mm; hypanthium 2-3 mm, reddish, pilose and uniformly covered with numerous, glochidiate spines, 2-3 mm; sepals 3 or 4, ovate, c. 1 mm., pilose. Fruit 5-8 3-4 mm, ellipsoid, reddish, uniformly covered with numerous, glochidiate spines. Open meadows or hillsides, thickets in wet forests. Ch (Breedlove 25819, MO); G (Williams et al. 41484, F); CR (Burger & Liesner 7437, F); P (White 62, MO). 1800-3700 m. (Mexico [Hidalgo, Distrito Federal, Mexico, Veracruz, Michoacan, Guererro, Oaxaca] Mesoamerica, Colombia, Venezuela, Ecuador, Peru, Bolivia.)

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 5 of 61 2. Agrimonia L. By F.R. Barrie.

Perennial herbs with short rhizomes. Roots fibrous, with or without slender, fusiform taproots. Stems 1 to several. Stipules prominent, leafy, more or less ovate in outline. Leaves alternate, cauline, imparipinnate, the lateral pairs of larger (major) leaflets increasing in size distally, separated by 1-several pairs of significantly smaller (minor) leaflets, the terminal leaflet the largest of the major leaflets. Inflorescence terminal or rarely axillary, racemose. Flowers short-pedicellate, bracteate; hypanthium stipitate, more or less ribbed, crowned by 2-5 ranks of hooked bristles; sepals 5, 1-3 mm, ovate, persistent in fruit; petals 5, yellow; stamens 5-15; carpels usually 2; style terminal. Fruit an achene, included in the persistent, bristly hypanthium. 18 spp., 1 sp. in Mesoamerica. Eurasia, North America, South America, South Africa. Bibliography: Kline, G.J. & Srensen, P.D., Brittonia 60: 11-33 (2008).

1. Agrimonia gryposepala Wallr., Beitr. Bot. 1: 49-50, pl. 1, f. 8 (1842). Lectotype (designated by Kline & Sorensen, 1990): United States, Poeppig s.n. (W). Illustr.: Britton & Brown, Ill. Fl. N. U.S., ed. 2. 2: 266, t. 2267 (1913). Agrimonia macrocarpa (Focke) Rydb., A. parviflora var. macrocarpa Focke. Herbs 30-150 cm; roots fibrous. Stems sparsely pubescent. Stipules 1.0-3.5 0.52.5 cm, ovate, incised, the apex acuminate. Leaves obovate in outline; abaxial surface glandular pubescent, hirsute along the veins; major leaflets 3-13, the terminal leaflet 3-10 1.5-6 cm, the lateral major leaflets progressively smaller towards the obovate to elliptic or rhombic, the base acute, the margins serrate, the apex acute to obtuse; minor leaflets in 1-3 pairs between the major leaflets; petioles xx-xx, glandular pubescent. Inflorescence rachis bearing short-stalked and sessile glandular hairs and scattered spreading hairs, 1-2 mm; flowers as many as 80, disposed singly along the rachis. Flowers pedicellate, the pedicels 1-2 mm; hypanthium obconic, glandular pubescent; bristles in 4-5 ranks, sepals elliptic, petals 2.5-4 1.4-3.0 mm, ovate to obovate; carpels 2. Fruit an achene enclosed within the hypanthium, this 3-5 mm, obconic to campanulate, ribbed, surmounted by rows of hooked bristles; reflexed at maturity. Deciduous forests and conifer forests. Ch

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 6 of 61 (Mndez 7917, MO); G (Molina & Molina 25017, MO). 1800-4500 m. (in Mesoamerica) (Canada, United States, Mexico [Coahuila], Mesoamerica.)

3. Crataegus L. By F.R. Barrie.

Shrubs or small trees. Trunks and larger branches commonly armed with stout, simple or branching thorns. Smaller branches bearing leaves on short, lateral shoots, the shoots typically subtended by straight, unbranched thorns. Leaves alternate, simple, stipulate, the stipules caducous or persistent, petiolate, the blades of leaves on mature branches consistent in form, those on juvenile shoots often inconsistently variable and different in form than the mature leaves; blades simple, dentate or lobate. Inflorescence terminal, corymbose. Flowers white or pink, hypanthium cupuliform or campanulate; sepals 5, petals 5, inserted on the margin of the disk, stamens (5)10-20(25), in 1 to 3 ranks; ovary inferior; carpels 1-5; styles 1-5, free to base; ovules usually 1 per carpel; fruit a pome, yellow or red; seeds 1-5. 250-500 spp., 2 spp. in Mesoamerica. Temperate regions of Eurasia and North America. Crataegus is a genus that is extremely complicated taxonomically. Approximately 1000 names have been published based on North American types, but the actual species count is probably 10% to 25% of that number. The natural southern range limit for Crataegus is the montane regions of Chiapas and western Guatemala. One species, Crataegus mexicana, is cultivated in other regions of Central and South America at higher elevations. Species of Crataegus commonly have two leaf forms, a juvenile form, found on seedlings young long shoots, and a mature form, found on the perennial short shoots (Phipps 1997). The latter generally are morphologically consistent, while the former may be quite variable and often unlike the mature leaves. In the descriptions below, the leaves described are from mature shoots. Bibliography: Eggleston, W.W., Bull. Torrey Bot. Club. 36: 501-514 (1909). Phipps, J.B., Sida, Botanical Miscellany 15: 1-94 (1997).

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 7 of 61 1. Leaf margins serrate or doubly serrate on upper 1/2; flowers 3-8, stamens 10; fruit 8-10 mm in diameter, red. 1. C. lindenii.

1. Leaf margin subentire or serrate on upper 1/2; flowers 3-5; stamens 20; fruit 15 mm in diameter, golden yellow. 2. C. mexicana.

1. Crataegus lindenii Stapf, Bull. Misc. Inform. Kew 1914: 326 (1914). Holotype: Mexico, Chiapas, Linden 708, K! Illustr.: not seen. Medium-sized trees. Branches armed; twigs stout; bark white or gray; thorns 2-6 cm, straight, unbranched. Leaves with blades 4-6 2.5-3.5 cm, chartaceous, green, (vestiture?), base narrowly cuneate to cuneate; margin entire along lower 1/3 to 1/2, serrate or doubly serrate along upper 2/3 to 1/2; apex obtuse to acute; petioles 10-25 mm. Inflorescence tomentose; flowers 3-8. Flowers pedicellate, the pedicels c. 9 mm, tomentose; hypanthium campanulate, tomentose; sepals c. 5 mm, narrowly triangular, the margins entire or with 1-3 glands; petals 7-8 mm in diameter, orbicular to obovate, clawed; stamens 10, c 5 mm; carpels 4-5; styles 4-5, c. 5 mm. Fruit 8-10 mm, globose, red. Deciduous forests. Ch (Mendez 8423, MO). 2100 m. (Endemic.)

2. Crataegus mexicana DC. Prodr. 2: 629 (1825). Nom. cons. prop. (designated by McVaugh, 2000): Illustr. No. 0879, Torner Collection of Biological Illustrations (Hunt Institute, Pittsburgh). Illustr.: Crataegus nelsonii Eggl., C. stipulosa (Kunth) Steud., nom. rejic. prop. Shrubs or more commonly a tree to 10 m. Branches armed; twigs stout, new growth red, ultimately gray or white; thorns to 8 cm, straight. Leaves with blades 2.5-8.0 1-4 cm; elliptic to obovate or oblanceolate; chartaceous; upper surface green, sparsely pilose or glabrate; lower surface paler, sparsely to densely pilose or tomentose; base cuneate; margins subentire or serrate along upper 1/2 to 3/4; apex obtuse or acute; petioles 4-5 mm, white-tomentose. Inflorescence corymbose, flowers 3-5 or solitary. Flowers pedicellate, the pedicels 8-10 mm in flower, to 25 mm in fruit, white tomentose; hypanthium c. 5 4 mm, campanulate, white tomentose; sepals 5-7 1-2 mm, triangular, tomentose without, glabrous within; petals 7-9 6-9 mm, orbicular, clawed, white; stamens 20, c. 5 mm; carpels 5; styles 5, c. 5 mm. Fruit globose, 15 15 mm, golden at

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 8 of 61 maturity. Dry or moist forests, pine-oak forests and secondary thickets. Also widely cultivated. Ch (Carlson 1990, MICH); G (Steyermark 35587, F); ES (Rodriguez DR01216, MO); CR (Echeverria 433, F). 1800-2800 m. (Mexico [Distrito Federal, Mxico, Michoacan, Morelos, Puebla, Oaxaca], Mesoamerica.) Crataegus mexicana is the source of manzanilla, a popular fruit drink base and dessert fruit in many parts of Latin America. Consequently, it has been widely cultivated from Mexico at least as far as Peru, at elevations above 1800 m. The species native range is calculated to be from central Mexico to Chiapas or western Guatemala. There has been considerable confusion regarding the authorship and type of Crataegus mexicana, both in print and on herbarium labels. Authorship is commonly ascribed to Mocio & Sess or to Sess & Mocio ex DC. However, Sess & Mocio supplied neither the name nor the validating description, thus the name is properly ascribed to Candolle alone (McVaugh 2000). The putative holotype cited by Phipps (1997: 21), Sess et al. 2083 (MA), is not type material, as it was not seen by Candolle. He based his description solely on the Sess & Mocio illustration, which is annotated in Candolles hand (McVaugh, 2000). Many herbarium collections of this species have been identified using the illegitimate name C. pubescens Steud. (1841), a later homonym of the mediterranean species, C. pubescens (C. Presl) C. Presl (1826). This usage is, in fact, doubly incorrect, as the type of Steudels name and the type of C. mexicana belong to different taxa (Phipps 1992). Phipps renamed C. pubescens, a species of central and northeast Mexico, C. gracilior.

4. Cydonia Miller By F.R. Barrie.

Deciduous, unarmed shrubs or small trees; branchlets and leaf buds pubescent. Leaves simple, entire; stipules persistent, prominent. Flowers terminal, solitary, bisexual; bracteoles absent; sepals 5, entire, reflexed; petals 5, obovate, white or pink; stamens 20; styles 5, free, pubescent basally; ovary inferior; carpels 5, ovules numerous. Fruit a pome. 1 species, native to East Asia. Widely cultivated elsewhere.

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 9 of 61

1. Cydonia oblonga Mill. Gard. Dict., ed. 8, Cydonia No. 1 (1768). Type BM? Illustr.: Fl. China Illustr. 9: 81, t. 81 (5-7) (2004). N.v.: Membrillo, quince. Shrubs or trees to c. 8 m; branchlets tomentose. Stipules c. 5 mm, oblanceolate or obovate, glabrate or glandular-ciliate. Leaves 5-10 3-6 cm, ovate to oblong; upper surface green, glabrous; lower surface densely gray-tomentose; base rounded to subcordate; margins entire; apex acute to rounded. Flowers 4-5 cm in diam; pedicels 2-5 mm, tomentose; hypanthium c. 1 cm, cupuliform, tomentose externally and internally; sepals c. 1 cm, ellptic, the apex acute; petals c. 2.5 1.5 cm, obovate. Fruit pyriform, yellow, villosolous, fragrant. Cultivated; locally naturalized; Quercus forests, Ch (Ton 330, DS); G (--, F), 2200-2600 m. (United States, Mesoamerica, Colombia, Ecuador, Bolivia.)

5. Eriobotrya Lindl. By F.R. Barrie.

Trees or shrubs. Leaves persistent, alternate, simple, short-petiolate to subsessile; veins prominent; margins coarsely toothed. Inflorescence a terminal, compound panicle, peduncles, pedicels and bracts densely tomentose or lanate. Flowers 5-merous; stamens 20; ovary inferior, carpels 2-5, styles connate at base. Fruit a fleshy pome; seeds 1-2, large. 10 spp., native to east Asia. 1 spp. cultivated and naturalized in Mesoamerica.

1. Eriobotrya japonica (Thunb.) Lindl., Trans. Linn. Soc. London 13: 102 (1821). Mespilus japonica Thunb., Nova Acta Regiae Soc. Sci. Upsal. 3: 199, 208 (1780). Holotype: Japan, Thunberg s.n., Herb. Thunberg No. 11908 (UPS). [Verify status] Illustr.: Flora of China Illustrations 9: 61, t. 61(1-5) (2004). N.v.: Nspero, nspero de japn. Tree to 5 or 10 m. Leaves alternate, simple, the blade oblanceolate, 15-25 5-7 cm; oblanceolate or obovate; coriaceous; upper surface green, glabrate; lower surface gray, sparsely to densely pubescent, the veins prominent; base narrowly cuneate, margins coarsely serrate; apex acuminate; petiole 5-10 mm. Inflorescence 10-15 cm, the lateral

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 10 of 61 branches c. 5 cm, branches, bracts, hypanthium, and calyx lanate, the hairs white or dull yellow; flowers typically 50-100. Hypanthium campanulate, c. 5 mm in diameter at anthesis, calyx lobes 5, 2-3 mm triangular, the apex acute; petals white, c. 10 5 mm, ovate to obovate, clawed; stamens 4-5 mm; styles 3-4 mm. Fruit 3-4 cm in diameter, pyriform or ellipsoid, yellow; seeds 1-15. cm. Cultivated and locally naturalized. Ch (Ton 6340, F); G (Steyermark 4658, F); ES (Martnez JBL02218, MO); H (Meja 165, MO); N (Moreno 3438, MO); CR (Gentry 952, MO). 300-4100 m. (Mesoamerica.) [Missing Meso country vouchers, also in USA, Canada, Carribean, South America, Africa, Japan, India Madagascar.]

6. Fragaria L. By F.R. Barrie.

Perennial herbs, rhizomatous and often stoloniferous. Leaves basal, trifoliate, stipules adnate to the petioles. Inflorescences terminal on leafless stems, cymose. Flowers bisexual or unisexual and plants dioecious or polygamodioecious; hypanthium obconic to saucer-shaped; bracteoles 5; sepals 5; petals 5; stamens 20 or more; receptacle conical or globose; carpels many, free; style lateral or subbasal. Fruit formed of numerous achenes on or embedded in the enlarged, fleshy receptacle. Approx. 10 spp., temperate Eurasia and North America, one sp. (F. chiloensis (L.) Miller) in South America. Fresa. Bibliography: Staudt, G. Univ. California Publ. Bot. 81: 1-162 (1999).

1. Leaves coriaceous or subcoriaceous; flowers and fruits borne below or just above the leaves. 1. F. ananassa

1. Leaves chartaceous; flowers and fruits borne above the leaves. 2. F. vesca

1. Fragaria ananassa Duchesne ex Rozier, Cours compl. agric. 5: 52, t. 5, fig. 1 (1785). F. chiloensis (L.) Mill. var. ananassa Weston, Bot. Univ. 2: 329 (1771). Lectotype (designated by Staudt, 1962): Collector ??? Herb. Linn. No. 654.19 (LINN). Illustr: Fragaria chiloensis (L.) Miller, sensu Fl. Guatemala. Fresa.

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 11 of 61 Herbs with short, thick rootstocks, stoloniferous. Stipules 1-2 cm, adnate basally, the apex free, acuminate, simple or 1-2-lobed. Leaves thick and coriaceous or subcoriaceous, petioles 5-20 cm, pilose; leaflets 3-6 3-6 cm; ovate to rhombic or obovate; upper surface dark green, nitid, glabrate; lower surface sparsely to densely pilose, sericeous along the veins, the veins prominent; petiolules 2-7 mm. Inflorescence 10-20 cm, erect or porrect, borne below or just above the leaves; flowers 3-10. Flowers perfect or rarely female; pedicels 3-10 cm, pilose, bracteoles and sepals subequal or the bracteoles slightly narrower, 6-10 2-4 mm, lanceolate or elliptic, uniformly pilose or glabrate distally, the margins entire, or 1-3-toothed, the apex acute to acuminate; petals 10-15 mm, broadly ovate. Fruit 2-6 cm in diameter, globose or conical, red to yellow or white; achenes embedded in the receptacle. Cultivated, perhaps escaping locally. Ch (); G (); H ( Molina 34003, MO); N (Rueda et al. 17416, MO); CR (); P (Sez 57,MO) 8002900 m. (Mesoamerica.) The commercially-grown strawberry is widely cultivated and possibly escapes occasionally in Mesoamerica. It is a hybrid between F. chiloensis (L.) Miller and F. virginiana Miller (Staudt, 1999).

2. Fragaria vesca L. Sp. Pl. 495 (1753). Lectotype (designated by Staudt, 1962): Collector ???? Herb. Linn. No. 654.2, (LINN). Illustr.: Holmgren, Illlustr. Companion to Gleason & Cronquist's Manual: 220, t. 1 (1998). F. mexicana Schltdl.

2a. Fragaria vesca subsp. bracteata (Heller) Staudt, Canad. J. Bot. 872 (1962). Fragaria bracteata Heller, Bull. Torrey Bot. Club 25: 194 (1898). Lectotype (designated by Staudt, 1962): United States, Heller & Heller 3615 (lower plant, GH.). Illustr.: Holmgren, Intermountain Flora, 3A: 119 (1997). Herbs with short rootstocks, stoloniferous and rooting at the nodes, the stolons slender, green, pubescent. Stipules 1.0-1.5 cm, adnate to the petiole, the apex free, apiculate. Leaves chartaceous, petioles 4-14 cm, pilose; leaflets subequal, 1.5-4.5 1-3.5 cm, obovate or rhombic; green, the upper surface glabrate or sparsely tomentose; the lower surface sparsely to densely tomentose, persistently sericeous along the veins; base

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 12 of 61 cuneate or the lateral leaflets oblique, margin coarsely dentate-serrate, apex rounded or broadly acute, petiolules 2-5 mm. Inflorescence axillary, racemose, c. 20 cm; flowers 510, borne above the leaves. Flowers perfect or rarely female; pedicels 1-3 cm, sericeous or pilose; bracteoles lanceolate to oblanceolate, 5-7 1-2 mm, green, the apex acute or acuminate; sepals ovate, 5.5-8 1.5-3 mm, green, the apex acuminate; petals 6-10 5-9 mm, broadly elliptic or obovate, white, the apex obtuse or rounded; stamens numerous receptacle conical, expanded in fruit. Fruits 1.1-1.5 cm in diameter, red or whitish; bracteoles and calyx reflexed, spreading or clasping the fruit or spreading; achenes superficial. Forest openings, montane slopes and meadows; also cultivated. Ch (Davidse 9451, MO); G (Croat 41019, MO). 1400-2400 m. (Canada, United States, Mexico [Chihuahua, Coahuila, Nuevo Leon, Durango, Zacatecas, San Luis Potosi, Hidalgo, Mxico, D.F., Puebla, Veracruz, Michoacn, Oaxaca] Mesoamerica.) Staudt (1999) recognized four subspecies and many varieties and formae under Fragaria vesca. The nominative subspecies is native to Eurasia, although it has been widely cultivated and naturalized in other parts of the world. Subspecies americana is native to the eastern and central United States and Canada, while subsp. californica is restricted to coastal areas in California and Baja California. The description and distribution above refer to subsp. bracteata, the subspecies that occurs in Mesoamerica, Mexico and the western United States and Canada. Subsp. bracteata is distinguished by flowers that are 1.5 to two times as large as those found in other subspecies, and by the calyx of mature fruit spreading or clasping, rather than reflexed. According to Standley and Steyermark (1946), F. vesca has been so widely cultivated in Guatemala, and has so readily escaped and naturalized, that it is unclear whether or not it is native to the region.

7. Geum L. By F.R. Barrie.

Perennial, rhizomatous herbs; roots fibrous. Leaves predominately basal, simple, pinnatifid or imparipinnately compound, the terminal leaflet larger than the lateral leaflets; cauline leaves few, reduced; stipules adnate to the petiole. Flowers few to

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 13 of 61 several, borne singly on elongate pedicels; hypanthium saucer-shaped, bracteoles 5; sepals 5; petals 5, yellow, white or red; stamens numerous; receptacle conical or cylindrical; carpels numerous, free; ovules 1 per carpel; styles filiform, elongate and persistent in fruit. Fruit an achene, aggregated into globose or cylindrical heads, the persistent style forming an elongate, hooked beak. 60 spp.; mostly Eurasia and North America, a few spp. in the montane regions of South America and southern Africa. 1 sp. in Mesoamerica.

1. Geum canadense Jacq. Hort. Bot. Vindob. 2: 82, pl. 175 (????). Type: the plate? Illustr: Holmgren, Illustr. Companion to Gleason & Cronquist's Manual, p. 227, t. 2 (1998). Stems 1-several, to c. 50 cm. Stems, leaves and pedicels pubescent. Basal leaves pinnately compound; lowermost cauline leave ternately compound, the uppermost simple, reduced, sessile. Flowers borne on pedicels 5-10 cm; bracteoles 1-2 mm, ligulate, green; sepals 5-6 1.5-3 mm, triangular, green; petals ligulate to obovate, 5-8 2-4 mm, ligulate to obovate, white, the apex rounded; stamens 2-3 mm; style 1-2 mm in flower, stigma c. 1 mm, bearing a few hairs. Fruit an aggregate of achenes in a globose head, the achenes 3-4 mm, ovate, sparsely sericeous, the persistent style 4-7 mm, looped or hooked apically. Pinus-Quercus forests. Ch (Breedlove 51928, F). 2000-2500 m. (Canada, United States, Mesoamerica.) In Mesoamerica, G. canadense is known only from Chiapas, where it is associated with other temperate, montane genera that are at or near the southern extreme of their range, such as Crataegus, Quercus and Pinus. Geum canadense is widely distributed in the eastern forests of Canada and the United States as far south and west as Texas, but is otherwise unknown from Mexico.

8. Hesperomeles Lindl. By F.R. Barrie.

Shrubs or small trees, often profusely branched, the branchlets unarmed or spine-tipped. Leaves simple, alternate, 1-5 cm long; coriaceous; margins crenate, dentate or serrate;

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 14 of 61 stipules small, caducous. Inflorescence terminal, cymose or flowers solitary. Flowers 5merous, pink or white; hypanthium campanulate, turbinate or urceolate; sepals persistent; petals white or cream-colored; stamens c. 20; carpels 5, free in flower; ovules 1 per carpel. Fruit a pome, c. 1 cm in diam; red, purple or black; the carpels partially fused by a stony endocarp, the free ends exserted from the open apex of the pome. Approx. 20 spp. Mesoamerica and South America, at elevations above 2000m. One species in Mesoamerica. Bibliography: Romoleroux, K. Fl. Ecuador 56: 143-151 (1996).

1. Hesperomeles obtusifolia (Pers.) Lindl., Bot. Reg, sub. fol. 1956 (1837). Crataegus obtusifolia Pers., Syn Pl. 2: 37 (1806). Lectotype (designated by Romoleroux, 1996): Peru, Dombey s.n. (P). Illustr.: Romoleroux, Fl. Ecuador 56: 145, t. 46, C-H. (1996). Eriobotrya obtusifolia (Pers.) DC, Hesperomeles chiriquensis Woodson, H. heterophylla Hook., H. obovata (Pittier) Standl., Osteomeles obovata Pittier. Shrubs to 5 or 6 m. Branchlets unarmed or commonly spine-tipped, initially puberulent, ultimately glabrescent; bark gray or black. Leaves glabrous, the blades 5-35 5-20 mm; obovate or oblanceolate to elliptic; initially sericeous, ultimately glabrate; upper surface lustrous, smooth or commonly rugose-reticulate; lower surface paler, less textured; base cuneate, decurrent into the petiole; margin crenate-dentate, the teeth often gland-tipped; apex broadly acute to rounded or obtuse; petiole 3-5 mm, conduplicate. Inflorescence cymose, pubescent, c. 3 cm in diameter at anthesis; flowers 2-20; bracts linear, 3.5-5 mm. Flowers pedicellate, the pedicels 1-10 mm; bracteoles 2, 2-5 c. 0.5 mm; hypanthium 3-5 mm, campanulate, pubescent; sepals 2-5 2-3 mm, triangular, pubescent basally, the apex commonly glabrous, the apex acute; petals 3-6 2-4 mm, elliptic to obovate, white or pale pink, the margin erose, the apex rounded; stamens c. 5 mm; disk tomentose, carpels 5, styles 3-5 mm. Fruits 5-12 5-10 mm, globose oblong, the calyx and styles persistent. Open hillsides and paramos. CR (Allen 5405, F); P (Woodson et al. 1078, MO). 1600-3400 m. (Mesoamerica, Colombia, Venezuela, Ecuador, Peru, Bolivia.)

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 15 of 61 9. Holodiscus (K. Koch) Maxim. Spiraea unranked Holodiscus K. Koch By G. Davidse.

Shrubs, not armed. Leaves simple, alternate, dentate; venation pinnate; stipules absent. Inflorescences racemes or panicles, bracteate. Flowers bisexual; hypanthium saucershaped or hemispheric; nectarines connate into a shallowly lobed, annular ring; sepals 5, valvate; petals 5, white or pink; stamens 15-20, seriate, 2-3 opposite the petals, 1 opposite the sepals, the filaments free, the outer series with the base dilated and, more or less connate, the anthers orbicular; carpels 5, free alternating with the calyx-lobes, the styles terminal, connate at the base, tomentose, the ovules 2. Fruit achene, short-stipitate, pilose, membranous; seeds 1, pendulous, oblong, lacking an aril, without endosperm, the testa? 5 spp. Canada to Colombia. I have adapted the treatment of Lis (1990). Literature: Ley, A.F. Bull. Torrey Bot. Club 70: 275-288 (1943). Lis, R.A. A taxonomic Revision of Holodiscus Maxim.(Spiraeoideae: Rosaceae) Based Upon Numerical Analyses of Leaf Morphometric Variation. Ph.D. thesis, University of California, Berkeley (1990). Silva, P.C. Taxon 40: 638-642 (1991).

1. Short-shoot leaf blades elliptic, with large primary teeth and frequent secondary teeth; tertiary veins mostly not perpendicular to the primary vein, ascending and irregular. 1. H. fissus 1. Short-shoot leaf blades obovate with very small primary teeth, rarely with any secondary teeth; tertiary veins perpendicular to the primary vein, with direct course and well developed. 2. Leaf blades pubescent or glabrous above; tertiary veins barely impressed above. 2a. H. argenteus var. argenteus 2. Leaf blades glabrous above; tertiary veins deeply impressed above. 2b. H. argenteus var. alpestris

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 16 of 61 1. Holodiscus fissus (Lindl.) C.K. Schneid., Ill. Handb. Laubhlzk 1: 495 (1905). Spiraea fissa Lindl., Edwards's Bot. Reg. 26: Misc. 73 (1840). Holotype: Mexico, state unknown, Hartweg 1840 (K). Illustr.:?. Holodiscus argenteus (L. f.) Maxim. var. bifrons Focke, H. loeseneri Dammer, Schizonotus argenteus (L. f.) Kuntze var. fissus (Lindl.) Kuntze, S. discolor (Pursh) Raf. var. fissus (Lindl.) Rehder, Sericotheca fissa (Lindl.) Rydb. Evergreen shrubs or trees 2-8 m; trunk up to 25 cm in diameter; branches angled, dark red to gray and brown, exfoliating; young stems deeply ribbed tan to reddish brown with short dense pubescence, occasionally villous; internodes 3-7 cm. Long-shoot leaves with the petiole distinct; blades 5-12 1.5-5 cm, lanceolate to ovate, the base cuneate, the margins serrate with many primary teeth 7-12 on each side and with secondary toothing, each tooth ending in a sharp mucro 1-2 mm, the apex acute to acuminate. Short-shoot leaves in fascicles of 5-8, subtended by long shoot leaves; petioles distinct to obscured by the decurrent blade; blades lanceolate to ovate-elliptic, the base cuneate, the apex acute, the margin serrate with 5-9 primary teeth on each side, secondary teeth rare; leaf blades of both long and short shoots glabrous adaxially or rarely sparsely pubescent on immature leaves, densely short pubescent abaxially and often villous especially along the veins. Panicles 9-20 5-15 cm, spreading, usually dense; pedicles 2-3 cm, the bracts 1-3 1 mm, lanceolate; sepals 1-2 mm, triangular, acute, sparsely short-pubescent; petals 1.5-2.5 mm, oval to orbicular, sparsely short pubescent across the back with fine hairs, denser along the veins. Achenes 1-2 mm, puberulent to glabrous on the faces with 0-10(15) glands, white hirsute on the margin, the upper edge slightly curved, the lower edge convex; styles 1-2 mm, often with hirsute hairs on the lower half. Pinus-Quercus forests, margins of cloud forests. CH (Breedlove 34675, MO); G. (Croat 41093, MO). 1700-3000 m. (Mexico, Guatemala.)

2. Holodiscus argenteus (L. f.) Maxim., Trudy Imp. S.-Peterburgsk. Bot. Sada 6: 254 (1879). Spiraea argentea L.f., Suppl. Pl. 261 (1781). Holotype: Colombia, Mutis s.n. (LINN?). Illsutr.:?. Schizonotus argenteus (L.f.) Kuntze, S. argenteus var. mutisianus Kuntze, Sericotheca argentea (L.f.) Rydb.

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 17 of 61 Spreading shrubs 1-5 m, rarely trees 4-10 m; branches arching, the older bark reddish brown to gray, exfoliating; young stems angled to ribbed, brown to gray with short dense pubescence or often villous; internodes 0.5-2.5 cm. Long-shoot leaves with the petioles 0.25-1.0 cm; blades 1.5-4.5 1-2 cm, the base narrowly cuneate, the margins serrate in the upper 1/4 to 1/2 with 4-5(-8) primary teeth on each side, the secondary teeth absent or rarely a few. Short-shoot leaves with the petioles 0.1-1.0 cm or obscured by the extremely decurrent blade; blades 0.5-3.0 0.4-1.5 cm, lanceolate to obovate, the base narrowly cuneate, decurrent on petiole, the margins serrate in the upper 1/4, the lower 3/4 entire, with 3-4 primary teeth on each side and secondary teeth absent, the apex acute to obtuse. Panicles diffuse, usually narrow, occasionally spreading, pubescent to villous; rachis 3-15 2-7 cm; bracts 2-5 mm, narrow or often leaf-like; pedicels 1-5 mm with 3 bracts, one larger than the other two. Flowers with the sepals 1.5-2 mm, broadly triangular-ovate, acute, densely pubescent to densely villous; petals 2-3 mm, oval to orbicular, with a thick row of short silky hairs along the back median vein; stamens as long as the sepals, the filaments 1.5-2 mm. Achenes 2-3 mm, pubescent to densely hirsute along the margin, puberulent to densely villous on either face, with glandular hairs, the upper surface slightly curved, the lower surface extremely convex; styles 1-1.5 mm. 2700-3850 m. Ch, G. H, N, P. (Mexico, Mesoamerica, Colombia.) Although most specimens can be readily assigned to the three varieties recognized by Lis (1990), there are some intermediates throughout the range of the species that are difficult to place. The typical variety is restricted to Colombia.

2a. Holodiscus argenteus (L. f.) Maxim. var. alpestris (Kuntze) F.A. Ley, Bull. Torrey Bot. Club 70: 288 (1943). Schizonotus argenteus (L.f.) Kuntze var. alpestris Kuntze, Revis. Gen. Pl. 1: 226 (1891). Lectotype (designated by Lis, 1990): Costa Rica, Kuntze 2358 (NY). Leaf blades narrowly obtrullate; tertiary veins deeply impressed into the adaxial surface; upper surface glabrous (glabrescent) and often very shiny; lower surface densely velutinous, bearing canescent hairs in the intercostals region with longer silky hairs along the primary and secondary veins. Pramos, margins of cloud forests, elfin forests, cinder

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 18 of 61 slopes of volcanos. CR (Davidse et al. 23132, MO). 2700-3700 m. (Mesoamerica, Colombia.)

2b. Holodiscus argenteus (L. f.) Maxim. var. velutinus (Rydb.) F.A. Ley, Bull. Torrey Bot. Club 70: 286 (1943). Sericotheca velutina Rydb. in Britton et al., N. Amer. Fl. 22: 265 (1908). Holotype: Mexico, Oaxaca, Smith 821 (US). Holodiscus argenteus (L. f.) Maxim. var. matudae F.A. Ley, H. velutinus (Rydb.) Standl. Leaf blades obtrullate to rhombic; tertiary veins not deeply impressed into the adaxial surface; upper surface glabrous to shortly pubescent; lower surface densely tomentose with sericeous hairs along the veins. Alpine grasslands, Abies forests, Pinus forests. Ch (Breedlove 29401, MO); G (Vliz et al. 7807); H (Lis, 199: 174); P (White 58, MO). 2800-3850 m. (Mexico [Oaxaca], Mesoamerica.) The only Panamanian collection seen was determined as var. alpestris by Lis but, despite the enormous distributional gap, seems closer morphologically to var. velutinus.

10. Lachemilla (Focke) Rydb. Alchemilla Sect. Lachemilla Focke By F.R. Barrie.

Perennial herbs or subshrubs. Branches creeping, caespitose, stoloniferous, decumbent or erect. Leaves simple 3-11-lobed, pinnate or reduced and forming verticillate, lobed sheathes. Inflorescences terminal or occasionally axillary cymes, lax or glomerate. Flowers bracteate, pedicellate or subsessile, hypanthium turbinate, campanulate or urceolate; bracteoles 4 or 5, slightly larger or smaller than, or subequal to, the sepals, or absent; sepals 4 or 5, spreading or erect; petals absent; stamens 2-4; carpels 1-6, ovules 1 per carpel; style basal. Fruit an achene, 1-6 per flower, which remain enclosed in the persistent hypanthium. Approx. 80 spp. found in montane regions of North and South America, from California to Argentina.

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 19 of 61 Bibliography: Perry, L.M. Contr. Gray Herb. 84: 1-57 (1929); Romoleroux, K. (1996). Lachemilla. Fl. Ecuador 56: 89-133 (1996).Rothmaler, W. Repert. Spec. Nov. Regni. Veg. 42: ??? (1937).

1. Plants suffrutescent herbs or shrubs, to 1 m; leaves clustered at branch tips. 7. L. polylepis. 1. Plants herbaceous or suffrutescent basally, to c. 50 cm; leaves basal and/or cauline, not clustered at branch tips. 2. Leaves reduced, uniting with the stipules to form a tube, sheathing at the base, with an apical whorl of 4-9 similar lobes. 2. Leaves not reduced, the lower leaves, at least, petiolate. 3. Basal leaves pinnately compound. 4. Leaves narrowly elliptic; leaflets 15-21, simple, cleft or bifid. 4. Leaves deltoid or ovate, leaflets 3-5, lobed or dissected. 5. Stems and leaves villose; flowers and fruits subsessile. 2. L. erodiifolia 6. L. pinnata 12. L. verticillata

5. Stems and leaves usually glabrous; flowers and fruits on pedicels 2-5 (-12) mm long. 4. L. mandoniana 3. Basal leaves simple or palmately divided. 6. Basal leaves shallowly 5-11-lobed or 5-7-cleft. 7. Leaves shallowly 5-11-lobed, the lobes broadly triangular. 7. Leaves 5-7-lobed, the lobes elliptic to obovate. 6. Basal leaves ternatisect or 3-5-cleft. 8. Inflorescence more or less open; flowers on pedicels 2-10 mm. 9. Leaves up to 4 cm wide, 3- lobate but appearing 5-lobate, the lateral leaflets deeply bifid. 9. Leaves less than 1.5 cm wide, distinctly 3-lobate. 8. L. procumbens 13. L. vulcanica 5. L. pectinata 11. L. venusta

8. Inflorescence aggregated or glomerate, flowers subsessile or on pedicels less than 2 mm. 10. Stems, stipules and the lower side of the leaves sericeous. 10. L. standleyi

10. Stems, stipules and leaves glabrous to pilose or sparingly pilose. 11. Bracteoles smaller than the sepals, fruits 2-2.5 mm, pilose. 3. L. fulvescens

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 20 of 61 11. Bracteoles and sepals subequal, fruits c. 1.5 mm, glabrous or pilose. 12. Hypanthium and fruits glabrous or sparingly pilose;sepals c. 0.5 mm, ovate. 1. L. aphanoides 12. Hypanthium and fruits pilose or sparingly pilose; sepals 0.5-1 mm, triangular. 9. L. sibbaldiifolia.

1. Lachemilla aphanoides (Mutis ex L.f.) Rothm., Repert. Spec. Nov. Regni. Veg. 42: 170 (1937). Alchemilla aphanoides Mutis ex. L. f. Suppl. 129 (1781). Lectotype (designated by Romoleroux, 1996), Peru, Herb. Linn. 166.4 (LINN). Illustr. (not seen) Alchemilla aphanoides var. subalpestris (Rose) L.M. Perry, A. hirsuta var. alpestris Schltdl & Cham., A. subalpestris Rose, A. tripartita Ruiz & Pav., Lachemilla subalpestris (Rose) Rydb., L. tripartita (Ruiz & Pav.) Rydb. Erect, prostrate or decumbent herbs; stems 10 to 75 cm, often numerous and profusely branched, green, glabrous or sparsely to densely pilose. Stipules sheathing, adnate to the petioles, the apices 2-4 cleft or lobed. Leaves ternatisect, the blades 1.0-1.4 0.7-1.2 cm; lobes subequal, deeply cleft or incised, obovate to oblanceolate; surfaces sparsely pilose; base cuneate; petioles of lower leaves c. 1.5 cm; upper leaves subsessile or sessile. Inflorescence an aggregate of terminal and axillary cymes; flowers borne in remote, terminal clusters of 3-10. Flowers sessile or pedicellate, the pedicels 0-2 mm; hypanthium urceolate, c. 1 mm, glabrous or sparingly pubescent; bracteoles 4, similar to the sepals; sepals 4, ovate, c. 0.5 mm, glabrous; stamens 2, c. 1 mm; styles 1-3, c. 1 mm. Fruits c. 1.5 mm, glabrous or sparingly pubescent. Achenes 1-3, c. 1 mm. Montane meadows, fields and rocky slopes Pinus-Quercus forests, paramos. (Ch (Breedlove 6869, MICH); G (Vliz et al. 8839, MO); CR (Sanchez & Estrada 777, MO); P (Stern et al. 1973, MICH). 1800-4000 m. (Mexico [Baja California Sur, Chihuahua, Nuevo Leon, Mxico, Michoacn, Oaxaca] Mesoamerica, Colombia, Venezuela, Ecuador, Peru, Bolivia.)

2. Lachemilla erodiifolia (Wedd.) Rothm., Repert. Spec. Nov. Regni Veg. 42: 172 (1937). Alchemilla erodiifolia Wedd., Chlor. Andina 2: 247 (1861). Lectotype

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 21 of 61 (designated by Romoleroux, 1996):??????? Illustr.: Fl.Ecuador, 56: 129, t. 42 A-C (1996). Decumbent, stoloniferous herbs. Stems rooting at nodes, reddish-green, villose. Basal stipules 5-10 mm, adnate to the petioles basally, villose, the apex free, leafy, simple and acute or bifid; cauline stipules 5-7 mm, leafy, villose, the apex 5-10 -lobed. Basal leaves pinnate, the blades 0.5-2.0 0.5-1.5 cm, ovate or deltoid; upper surface green, sparsely villose or glabrescent; lower surface densely villose; leaflets 5-9, 3-7 2-5 mm, pinnatifid, the lobes 2-5, ovate or elliptic; petioles 5-20 mm, villose. Cauline leaves similar but reduced; blades 0.5-1.0 0.5-1.0 cm, deltoid, leaflets 3-5, petioles 3-5 mm. Inflorescence terminal or axillary, few-flowered or flowers solitary. Flowers subsessile, the petioles less than 1 mm; hypanthium urceolate, 1-1.5 mm, villose externally, pilose within; bracteoles and sepals subequal, c. 1 mm, villous externally, glabrous within; yellow; carpels c. 4. Fruit 2.5-3 mm; achenes 2-4. Paramos. CR (Alfaro 1725, MO). 3400-3500 m. (Mesoamerica, Ecuador, Peru.)

3. Lachemilla fulvescens (L.M. Perry) Rothm., Repert. Spec. Nov. Regni Veg. 42: 171 (1937). Alchemilla venusta var. fulvescens L.M. Perry, Contr. Grey Herb. 84: 19 (1929). Isotype: Colombia, Pennell 10385 (US!). Illustr.: Fl. Ecuador 56: 113, t. 38 K-N (1996). Alchemilla fulvescens (L.M. Perry) Rothm. Decumbent herbs. Stems moderately branched, reddish-brown, pilose. Basal stipules 5-10 mm, papery, brown, simple or lobed; cauline stipules leafy, green, the apex bifid, the lobes 2-4 mm, elliptic, the apex acute. Basal leaves in rosettes; blades 1.0-1.5 cm in diameter, orbicular or widely ovate; palmately 3-5 -lobed, the lobes obovate, the upper surface sparsely pilose or glabrescent, the lower surface pilose, the margins with 28, irregular rounded teeth; petioles 0.3-2.5 cm, pilose. Cauline leaves 5-10 4-8 mm, obovate, trilobed, the lobes ovate; petioles 3-8 mm. Inflorescence axillary or terminal; flowers 1-15, in apical clusters. Flowers subsessile; hypanthium c. 1 mm, campanulate, glabrate or pilose exernally, bracteoles smaller than the sepals, c. 0.5 mm, ovate; sepals c. 1 mm, ovate, pilose or glabrate externally, glabrous within. Fruit 2-2.5 mm; achenes

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 22 of 61 usually 2. Paramos. CR (Quesada et al. 2004, INB); P (Gmez et al. 22444, MO). 30003400 m. (Mesoamerica, Colombia, Venezuela, Ecuador.) Collections of Lachemilla fulvescens seen from Panama and Costa Rica differ from South American populations in having glabrate flowers and fruits and somewhat less vestiture overall.

4. Lachemilla mandoniana (Wedd.) Rothm., Repert. Spec. Nov. Regni Veg. 42: 171 (1937). Alchemilla mandoniana Wedd., Chlor. Andina 2: 246 (1861). Holotype, Bolivia, Mandon 666 (P, photo MO!). Illustr.: Fl. Ecuador, No. 56: 129, t. 42 D-I. (1996). Alchemilla paludicola Rothm., Lachemilla paludicola (Rothm.) Rothm. Delicate, decumbent herbs, rooting at the nodes; plants uniformly and sparingly pilose or nearly glabrous, the hairs restricted to a few on the tips of the leaf lobes. Stems red. Stipules 2-10 mm, papery, brown basally, the apex greener, the apex free, acuminate. Basal leaves pinnatifid, the blades c. 1 1 cm, deltoid; the lobes 3-5, these cleft into 2-5 irregular, segments, the apices rounded; petioles 10-30 mm. Cauline leaves similar but smaller. Inflorescence a terminal, solitary flower cluster of 2-3 flowers. Flowers pedicellate, the pedicel 2-5 (-12) mm; hypanthium urceolate, green, yellow or red; bracteoles absent or greatly reduced, c. 1/4 the size of the sepals; sepals 0.4-1 mm, ovate, the apex bluntly acute. Fruit c. 2 mm, red; achenes 1 or 2. Quercus forests, paramos. CR (Gamboa 1525, MO); P (Weston 10176, MO). 3000-3400 m. (Mesoamerica, Colombia, Ecuador, Peru, Bolvia.)

5. Lachemilla pectinata (Kunth) Rothm., Repert. Spec. Nov. Regni. Veg. 42: 171 (1937). Alchemilla pectinata Kunth, in Humboldt, Bonpland & Kunth, Nov. Gen. Sp. 6: 226 (1824). Holotype: Colombia, Humboldt & Bonpland s.n. (P). Illustr.: Romoleroux, Fl. Ecuador, No. 56: 11326, t. 38 G-J. Alchemilla pectinata var. mexicana Perry. Caespitose, stoloniferous herbs, taproot slender. Basal stipules 5-15 mm; sheathing, chartaceous, brown, sparsely sericeous, adnate to the petiole basally, free apically, the apex with 3-5 short, acute lobes. Basal leaves simple, the blade 1-3.5 1.5-

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 23 of 61 5.5 cm; widely ovate or reniform; chartaceous; 5-11-lobed, the lobes broadly triangular, cut 1/4 to 1/3 into the blade; upper surface green, glabrescent; lower surface green, sericeous along the veins; base cordate; margins pectinate-serrate; apex of the lobes broadly acute to rounded; petioles 20-90 mm, sericeous or glabrescent. Cauline leaves similar to the basal leaves though significantly smaller, or absent. Inflorescence 3-7 cm; racemose; peduncles, pedicels, hypanthium, bracteoles and sepals uniformly pilose; bracts connate. Flowers subsessile, the pedicels less than 1 mm; hypanthium 1-2 mm, campanulate; bracteoles and sepals 1-2 mm, ovate-elliptic; carpels 2-4. Fruits, c. 3 mm, perisistently pilose. Montane slopes and alpine and subalpine meadows. Ch (Breedlove & Raven 8142, F); G. (Steyermark 35500, F); CR (Burger & Leisner 6506, F); P (Davidson 546, MO). 3000-3600 m. (Mesoamerica, Colombia, Ecuador, Peru, Bolivia.) Rydberg (1908) and McVaugh (1950) treated Lachemilla pectinata as a synonym of L. orbiculata (Ruiz & Pavon) Rydb., which here, following Perry (1929) and Romoleroux (1996), is considered a separate species, endemic to the northern Andes. The two differ most prominently in the floral bracts, which are sheathing in L. orbiculata, the carpel number, 1-2 in L. orbiculata vs. 4-6 in L. pectinata, and the scaly leaves on the stolons in L. orbiculata.

6. Lachemilla pinnata (Ruiz & Pav.) Rothm., Repert. Spec. Nov. Regni. Veg. 42: 171 (1937). Alchemilla pinnata Ruiz & Pav., Fl. Peruv. 1: 69 (179). Lectotype (designated by. Romoleroux, 1996): Peru, Ruiz & Pavon s.n. (M). Illustr. Romoleroux, Fl. Ecuador, No. 56: 126, t. 41 (????). Aphanes pinnata (Ruiz & Pav.) Pers., Zygalchemilla pinnata (Ruiz & Pav.) Rydb. Caespitose herb. Branches decumbent, rooting at the nodes; villous or glabrescent. Stipules of basal leaves 10-15 mm, adnate to the petiole chartaceous, 1-2 lobed, the lobes acute. Basal leaves pinnately compound, the blades 10-20 3-5 mm; narrowly elliptic; leaflets 15-21, 2-4 1-2 mm; simple or more commonly cleft or bifid; upper surface glabrous or sparsely villous; lower surface glabrous to villous; margins entire; apex acute; petioles 10-20 mm; villous. Stipules of distal leaves 4-7 mm, connate and adnate to the petiole at base, free at apex; 5-6-lobed, green and leaflike. Distal leaves pinnately compound, 3-5 c. 3 mm; leaflets 3-5; petioles 1-3 mm. Inflorescence of

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 24 of 61 solitary flowers or compact racemes of 3-5 flowers. Flowers petiolate, the petioles 1-2 mm; hypanthium 1-2 mm, urceolate to campanulate; green or reddish, villous on the outer surface, glabrous within; bracteoles and sepals c. 1 0.5 mm, erect, ovate, carpels 2. Open, alpine meadows Ch (Breedlove 26709, F); G (Veliz 95.4739, MO). 3100-3900 m. (Mexico [Mexico], Mesoamerica, Colombia, Ecuador, Peru, Bolivia, Chile, Argentina.)

7. Lachemilla polylepis (Wedd.) Rothm., Repert. Spec. Nov. Regni. Veg. 42: 169 (1937). Alchemilla polylepis Wedd., Chlor. And. 2: 246 (1857). TYPE: Illustr.: Weddell, Chloris Andina 2: t. 75A. (1857). Suffrutescent herbs or suffruticose shrubs; branches reddish-brown. Stipules sheathing, persistent on stems, the apices acute to acuminate. Leaves clustered at branch tips, 3-5-fid, the blades 1.0-1.5 0.5-0.8 cm, coriaceous, upper and lower surfaces silvery-sericeous or glabrate above; lobes ovate elliptic, the base cuneate, the margins entire, revolute, the apices acute, tufted-ciliate; petioles broad, less than 1 mm, sheathed by stipules; Inflorescence terminal, flowers 10-20, clustered. Flowers subsessile, the hypanthium, calyx, bracteoles and sepals sericeous; hypanthium c. 2 mm, sericeous; bracteoles and sepals subequal, c. 1 0.5 mm, ovate, yellow, sericeous on outer surface, glabrous within; carpels 1. Paramos. CR (Burger & Leisner 7447, F). 3300-3900 m. (Mesoamerica, Colombia, Venezuela.)

8. Lachemilla procumbens (Rose) Rydb. N. Amer. Fl. 22: 382 (1908). Alchemilla procumbens Rose, Contr. U.S. Nat. Herb. 10: 96 (1906). Holotype: Mexico, Hidalgo, Rose & Painter 9202 (US). Illustr. Rose, Contr. U.S. Nat. Herb. 10(3): t. 27 (1906). Lachemilla costaricensis Dammer, L. laxa Dammer. Procumbent herbs from a slender taproot. Branches 1-many, sparingly pilose or villous. Stipules sheathing, green, leafy, 2-5-lobed. Leaves all of similar size and shape; blades widely ovate, 1.0-2.0 1.2- 4.0 cm, ternatisect but superficially 5-lobed, the central lobe obovate, 10-20 4-10 mm, the lateral lobes deeply bifid, the segments obovate, 5-15 2-8 mm; chartaceous; upper surface green, sparingly pilose or glabrate;

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 25 of 61 lower surface paler, pilose or villous along the veins; leaflets coarsely dentate-serrate on upper 1/2; apex bluntly acute to obtuse; petioles 5-10 mm, pilose or sericeous. Inflorescence laxly racemose; flowers 5-10. Flowers pedicellate, the pedicels 2-10 mm; hypanthium c. 2 mm, campanulate, the outside sericeous, inside pilose; bracteoles 4-5, larger than the sepals, 1-1.5 mm, ovate, glabrescent, the apex acuminate; sepals 4-5, 0.7-1 mm, triangular to ovate, glabrescent, the apex acuminate. Achenes 3-8. Disturbed sites and meadows in Pinus, Quercus and Abies forests, paramos. Ch (Martinez et al. 19500, MO); G (Harmon 3582, MO); ES (Gonzles 175, MO); CR (Morales & Dauphin 4802, MO). 2200-3600 m. (Mexico [Chihuahua, Durango, Nuevo Leon, Tamaulipas, Hidalgo, D.F., Mxico, Tlaxcala, Veracruz, Jalisco, Michoacn Guererro, Oaxaca], Mesoamerica.)

9. Lachemilla sibbaldiifolia (Kunth) Rydb., N. Amer. Fl. 22: 384 (1908). Alchemilla sibbaldiifolia Kunth, in Humboldt, Bonpland & Kunth, Nov. gen. sp. (quarto ed.) 6: 225, t. 261 (1824). Type: Mexico, Mexico, Humboldt. & Bonpland s.n. (P). Illustr. Rzedowski & Rzedowski, Fl. del Bajo 135: 24 (2005). Alchemilla pringlei (Rydb.) Fedde, A. sibbaldiifoia var. tonduzii (Dammer) L.M. Perry, Lachemilla pringlei Rydb., L. tonduzii Dammer. Decumbent or prostrate herb, from a slender taproot, often profusely branched. Stems reddish or green, pilose. Stipules leafy, green, sheathing at the base, the apex divided into 2-4 lobes, appressed-pilose. Leaves ternatisect, the blade 1-3 1- 2 cm, triangular to widely ovate in outline, lobes obovate, or the lateral lobes of the lowermost leaves bifid; upper and lower surfaces green, sparingly pilose or glabrate, or sericeous along the veins below; apex rounded, 6-10-toothed; petiole 3-15 mm, pilose. Inflorescence terminal or axillary, flowers 4-10, in compact clusters borne on short, lateral branches, 0-10 mm. Flowers subsessile, pedicels 0- 2 mm; hypanthium c. 1 mm, urceolate, pilose; bracteoles subequal to or slightly smaller than the sepals triangular, glabrous; sepals 0.5-1 mm, triangular, glabrous. Fruits c. 1.5 mm, pilose; achenes 1-4. Pinus-Quercus forests, cloud forests. Ch (Breedlove 25861, MO); G (Skutch 197, MICH); ES (Croat 42445, MO). 1900-3300 m. (Mexico [Sinaloa, Chihauhau, Durango, San Luis Potos, Guanajuato, Guertaro, Hidalgo, Mxico, Districto Federal, Morelos, Puebla, Tlaxcala, Veracruz, Michoacn, Guerrero, Oaxaca], Mesoamerica.)

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 26 of 61

10. Lachemilla standleyi (L.M. Perry) Rothm., Repert. Spec. Nov. Regni Veg. 42: 171 (1937). Alchemilla standleyi L.M. Perry, Contr. Gray Herb. 84: 29 (1929). Holotype: Costa Rica, Standley and Valerio 43893 (US!). Illustr: Alchemilla kupperi Suess. Decumbent, profusely branched herbs, often forming dense mats. Stems sericeous. Stipules leafy, 3-4-lobed, sericeous on both surfaces. Leaves ternatisect, 5-10 5-10 mm, obovate, upper surface green, sericeous, lower surface white, densely sericeous; leaflets 3-5 2-3 mm, obovate, the upper 1/2 4-7-toothed; petioles 2-5 (-10) mm, densely sericeous. Inflorescence a terminal or axillary, congested cyme, flowers 110. Flowers subsessile or pedicellate, the pedicels less than 1 mm; hypanthium 1-2 mm, campanulate, densely sericeous externally, pubescent within; bracteoles and sepals subequal, 1-2 mm, densely sericeous externally, glabrous within; styles 4-10. Fruits 3-4 mm, sericeous; achenes 4-10. Paramos, wet depressions in alpine slopes. CR (Tonduz 11810, MICH); P (D'Arcy & Hammel 12398, MO). 2700-3500 m. (Endemic.)

11. Lachemilla venusta (Schltdl. & Cham.) Rydb., N. Amer. Fl. 22: 382 (1908). Alchemilla venusta Schltdl. & Cham., Linnaea 5: 572 (1830). Isotype: Mexico, Veracruz, Schiede 592 (US!). Illustr.: Ann. Missouri Bot. Gard. 37: 161, t. 68 (1950), as Alchemilla pascuorum. Alchemilla guatemalensis Rothm., A. pascuorum Standl., Lachemilla guatemalensis (Rothm.) Rothm., L. pascuorum (Standl.) Rothm. Prostrate, stoloniferous herbs from a slender taproot. Stems slender, green, sparsely pilose or sericeous. Stipules foliaceous, oblong, partially connate and sheathing. Leaves simple, 1.0-4.0 1.0-3.5 cm, reniform or widely ovate, 5-7 -lobed, the lobes elliptic-obovate, cut 2/3-3/4 into the blade, margin entire within the sinuses, pectinateserrate distally, the apex rounded or bluntly acute; upper surface green, sparsely pilose; lower surface gray, glaucous, sericeous; base obtuse to subcordate; petioles 1-7 cm, pilose, the hairs spreading or appressed. Inflorescence a dense to somewhat lax cyme, flowers 5-10, clustered at the tips of the inflorescence branches. Flowers pedicellate, the pedicels 2-10 mm, hypanthium 1-2 mm, campanulate, sericeous; bracteoles and sepals c.

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 27 of 61 1 mm, sparsely sericeous or glabrate. Fruit 3-4 mm; achenes 2-8. Moist or dry open slopes or fields. Ch (-,F); G (-, F); CR (-, F); P(Woodson 892, MO). 1300-4200 m. (Mexico (Puebla, Veracruz, Oaxaca), Mesoamerica.) Populations of Lachemilla venusta from Costa Rica and Panama, identified as L. pascuorum, tend to have appressed vestiture and more rounded leaf lobes than is found in specimens from Mexico, where the lobes are more bluntly acute and the stems have spreading hairs. Populations from Guatemala tend to be intermediate between the extremes, which a mixture of appressed and spreading hairs.

12. Lachemilla verticillata (Fielding & Gardner) Rothm., Repert. Spec. Nov. Regni Veg. 42: 170 (1937). Alchemilla verticillata Fielding & Gardner, Sert. Pl. t. 65 (1844). Holotype: Venezuela, Linden 442 (K isotype?; holotype, CGE?). Alchemilla ocreata Donn. Sm., Lachemilla ocreata (Donn.Sm.) Rydb. Caespitose herbs. Stems few to many from a branching caudex, erect, reclining or decumbent; reddish-brown, pilose. Leaves reduced, uniting with the stipules to form a tube, sheathing at the base, with an apical whorl of 4-9 lobes, 4-9 1-2 mm, narrowly ellipitic or lanceolate, carnose, the lower surface densely pilose, the margins revolute, the apex acute. Inflorescence terminal, few-flowered. Flowers subsessile; hypanthium 1-2 mm, campanulate, pubescent externally and internally; bracteoles and sepals subequal, c. 1 mm, ovate. Fruits c. 2.5 mm, pubescent; achenes 4-8. Paramos. CR (Almeda & Wilbur 3423, MO); P (Weston 10163, MO). 3200-3900 m. (Mesoamerica, Colombia, Venezuela, Ecuador, Peru.)

13. Lachemilla vulcanica (Schltdl. & Cham.) Rydb., N.Amer. Fl. 22: 382 (1908). Alchemilla vulcanica Schltdl. & Cham., Linnaea 5: 573 (1830). Holotype: Mexico, Veracruz, Schiede & Deppe 591 (HAL). Illustr. Decumbent, suffrutescent herbs, often profusely branched. Stems sericeous, reddish or green. Stipules leafy, sheathing at the base, the apex two-lobed. Leaves ternatisect, the blades 1.0-1.5 1.0-1.5 cm, broadly ovate; upper surface green and glabrescent; lower surface paler, sericeous along the veins; lobes obovate, base cuneate, the lateral margins entire, the apex 3-7-toothed; petioles 2-5 mm, sericeous. Inflorescence

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 28 of 61 terminal, cymose, flowers 3-10. Flowers pedicellate, the pedicels 2-5 mm; hypanthium 1.5-2 mm, campanulate, sericeous externally, pilose within; bracteoles and sepals 1-2 mm, subequal or the bracteoles slightly smaller and/or narrower, ovate, the apex acute, green or reddish, sericeous externally, glabrous within. Fruits 3-4 mm, sericeous basally, the calyx sparsely sericeous to glabrate; achenes 1-4. Open, conifer forests, alpine meadows and steep slopes. Ch (Breedlove 29364, MO); G (Vliz 97.5946, MO); ES (Villacorta et al. 2416, MO). 2700-4100 m. (Mexico [Coahuila, Durango, Quertaro, Jalisco, Michoacn, Mxico, Distrito Federal, Puebla, Tlaxcala, Veracruz, Oaxaca], Meosamerica, Colombia, Venezuela, Ecuador, Peru, Bolivia.)

11. Malacomeles (Decne.) Engl. Cotoneaster sect. Malacomeles Decne. By F.R. Barrie.

Shrubs, armed or unarmed. Stipules present. Leaves simple; coriaceous; lower surface glabrous to white-tomentose; margin entire or denticulate. Inflorescence a few-flowered raceme. Flowers pedicellate; braceolate; sepals 5, petals 5, stamens 20, ovary inferior, carpels 5, styles 5, free. Fruit a fleshy, berry-like pome. 3 spp. USA (Texas), Mexico, Mesoamerica. 1 sp. in Mesoamerica. Bibliography: Jones, G.N. Madroo 8: 33-39 (1945).

1. Malacomeles denticulata (Kunth) G.N. Jones, Madroo 8: 36 (1945). Cotoneaster denticulatus Kunth, Nov. Gen. Sp. (ed. qto.) 6: 214, pl. 556 (1824). Holotype: Mexico, Hidalgo, Humboldt & Bonpland s.n. (P). Illustr. Rzedowski & Rzedowski, Fl. del Bajo 135: 33 (2005). Amelanchier denticulata (Kunth) K. Koch; A. nervosa (Decne.) Standl.; Cotonaeaster nervosa Decne.; Malacomeles nervosa (Decne.) G.N. Jones Shrubs 1-3 m, unarmed. Stems tomentose, the hairs white, ultimately glabrate; bark reddish to black. Stipules c. 1 mm, triangular, caducous. Leaves simple, the blades 2-4 1.2- 2.5 cm; coriaceous; upper surface green, glossy or matte, glabrous, the midvein sulcate; lower surface tomentose, the hairs white or sordid, arachnoid hairs, the midvein

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 29 of 61 and lateral veins prominent; base cuneate; margin entire or remotely and minutely denticulate; apex obtuse, the tip apiculate; pedicels 3-10 mm, tomentose. Inflorescence terminal, racemose, flowers 3-4, often with solitary flowers in upper 1 or 2 leaf axils; peduncle white-tomentose. Flowers pedicellate, the pedicels 5-10 mm; hypanthium 3-5 mm in flower, tomentose; sepals c. 2.5 mm, triangular, tomentose on outer surface, glabrous within; petals 5-6 mm in diameter, suborbicular, white, glabrous;. Fruit 8-10 mm, globose to ellipsoid, sparsely pubescent, red. Montane deciduous forests, PinusQuercus forests. Ch (MacDougall H114, F); G (Standley 82544, F); H (Molina & Molina 22692, F); CR (Morales 5893, F). 1300-2300 m. (United States [Texas], Mexico [Nuevo Len, Coahuila, Chihuahua, San Luis Potos, Hidalgo, Quertaro, Guanajuato, Distrito Federal, Veracruz, Oaxaca], Mesoamerica.)

12. Malus Mill. By F.R. Barrie.

Deciduous trees or shrubs, the leaves and inflorescences sometimes borne on lateral short shoots; unarmed or the branchlets sometimes armed with spines. Leaves deciduous or persistent, simple, alternate, stipulate, petiolate, the margins serrate or lobed. Inflorescence umbelliform racemes. Flowers petiolate; bracteate; hypanthium cupuliform; bracteoles absent; sepals 5, petals 5, stamens 15-50; ovary inferior, carpels 3-5; styles 25,connate at the base. Fruit an oblate to globose pome, the calyx persistent or deciduous. Approx. 55 spp., Eurasia, United States, Canada; widely culitivated elsewhere. 1 cultivated species in Mesoamerica.

1. Malus pumila Mill., Gard. Dict., ed. 8, Malus No. 3 (1868). Pyrus malus L., Sp.Pl. 479 (1753). Lectotype (designated by Ghora & Panigrahi, 1995): Herb Linn. No. 647.3 (LINN). Illustr.: N.v.:Manzana, apple. Trees to c. 15 m; unarmed. Young shoots green, tomentose, ultimately reddishbrown, glabrate. Stipules caducous, c. 5 mm, linear. Leave emerging with or just following anthesis, the blades 3.0-7.5 3.0-4.5 cm; ovate to elliptic or obovate; chartaceous; upper surface dull to bright green, glabrate or tomentose along the midvein;

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 30 of 61 lower surface paler, sparsely to densely tomentose; base rounded to truncate; margins serrate or crenate; apex rounded to acute or acuminate; petioles 2-4 cm, tomentose. Flowers fragrant, pedicels 1.0-2.5 cm, tomentose; hypanthium c. 5 mm, cupuliform, densely tomentose; sepals 3-7 mm, narrowly triangular, tomentose, the apex acuminate; petals c. 3 2 cm, obovate, white or pink; stamens 5-8 mm; styles 6-10 mm. Fruit usually 5-10 cm or more, oblate or globose, red, yellow or green, glabrous. Cultivated, possibly locally naturalized. Ch (Ton 2242, CAS) G (--, F); ES (Rodriguez et al., 1217, MO); CR ( --, ). (Mesoamerica.)

13. Photinia By F.R. Barrie.

Trees or shrubs, to c. 20 m. Young growth densely tomentose, ultimately glabrous. Leaves simple, alternate, pedicellate; stipules triangular or ovate, small, persistent or caducous; blades ovate, lanceolate, elliptical, obovate or oblanceolate; coriaceous; base cuneate or narrowly cuneate; margins entire or weakly to strongly toothed; apex acute to acuminate. Inflorescence terminal, paniculate, densely tomentose, flowers 20-100 or more. Flowers pedicellate, hypanthium and calyx tomentose, sepals 5, persistent in fruit; petals 5, white, orbicular to elliptic, weakly clawed; stamens 20, 2-4 mm; carpels 2-3; stigmas 2-3. Fruit a pome, seed 1 per carpel. 40-45 spp; eastern Asia, Mexico (6 spp.), Mesoamerica (3 spp.) Bibliography: Phipps, J.B. Heteromeles and Photinia (Rosaceae subfam. Maloideae) of Mexico and Central America. Canad. J. Bot. 70: 2138-2162 (1992).

1. Leaves coriaceous, puberulent when young, ultimately glabrate; lowermost inflorescence bracts 5-30 mm long; fruit globose, 1.5-2.0 cm in diameter; calyx lobes erect. 2. P. mexicana

1. Leaves coriaceous or chartaceous, inflorescense bracts 1-5 mm long, vestiture persistent on petioles and along the midvein below; fruit ellipsoid, up to 1 cm in diameter; calyx lobes appressed.

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 31 of 61 2. Leaves coriaceous, the margins entire and weakly to strongly deflexed, vestiture of the calyx lobes typically whiter than the more reddish or brown hypanthium. 1. P. matudae 2. Leaves coriaceous or chartaceous, the margins entire to serrulate or serrate, not deflexed; hypanthium and calyx vestiture uniformly tan or reddish-brown. (P. microcarpa). 3. Leaves 5-7 cm, vestiture dark reddish brown; inflorescence 4-7 cm in diameter, flowers 20-40. 3b. P. microcarpa subsp. elliptica

3. Leaves 7-12 cm, vestiture tan or reddish brown, inflorescence 7-10 cm in diameter, flowers 30-80. 4. Vestiture dark reddish brown, inflorescence 7-8 cm in diameter, flowers 30-50. 3c. P. microcarpa subsp. hintonii 4. Vestiture tan to brown, but not dark, inflorescence 8-10 cm in diam,. flowers 40-80. 3a. P. microcarpa subsp. microcarpa

1. Photinia matudae Lundell, Contr. Univ. Michigan Herb. 4: 7 (1940). Holotype: Mexico, Chiapas Matuda 2937 (MICH!). Illustr.: Canad. J. Bot. 70:2152, t. 9 (1992). Trees to 25 m. Branches initially rufous-tomentose, ultimately gray, glabrate. Leaf blades 6-12 2-4 cm, narrowly ovate to elliptic or weakly obovate; coriaceous; midvein sulcate above, tomentose on both surfaces, the blade otherwise glabrate, the secondary veins partially or wholly inscribed in the above; base cuneate; margin entire and typically deflexed; apex acute to acuminate; petioles 5-10 mm, rufous-tomentose, the vestiture similar to the branches. Inflorescence a more or less flat-topped, compound panicle, 5-10 cm in diameter, the pedicels, bracts, hypanthium and calyx white or rufous-tomentose, the hairs on the calyx often whiter, contrasting with the reddish hypanthium; flowers 30100; bracts 1-2 mm, caducous, glabrate. Flower pedicellate, the pedicels 3-5 mm at anthesis; hypanthium 3-5 mm, campanulate; sepals triangular, 1-2 mm, the apex acute; petals 3-4 mm in diameter, white; carpels 3. Fruit 8-10 4-5 mm, ellipsoid, yellow or red, glabrous or persistently tomentose apically; sepals persistent, appressed. Montane rain forests; cloud forests. Ch (Breedlove 41765, F); G (Steyermark 49068, F). 25003000 m. (Endemic.)

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 32 of 61

2. Photinia mexicana (Baill.) Hemsl., Biol. Cent.-Amer., Bot. 1(5): 380 (1880). Chamaemeles mexicana Baill., Adansonia 9: 148 (1869). Isotype: Mexico, Veracruz, Galeotti 1660 (K!). Illustr.: Canad. J. Bot. 70: 2151, t. 8 (1992). Tree 5-15 m. Branches initially rufous-tomentose, ultimately gray, glabrate. Leaf blades 10-18 3.5-7 cm, oblanceolate or obovate, puberulent, ultimately glabrate; midvein flat or carinate above, the secondary veins flat; base narrowly cuneate; margins coarsely crenate-serrate; apex obtuse, acute or abruptly acuminate; petioles 5-20 mm, glabrate. Inflorescence paniculate, 7-10 cm in diameter, the pedicels, hypanthium and calyx densely rufous tomentose; bracts 5-30 1-5 mm, sparsely pubescent or glabrate at anthesis. Flowers pedicllate, the pedicels 1-5 mm at anthesis; hypanthium 5-6 mm in diameter, campanulate; sepals 2-3 mm, triangular, erect; petals 8-10 6-8 mm, the margins entire; carpels 3, white-tomentose apically. Fruit 1-2 cm in diameter, oblate to globose, rufous-tomentose; sepals persistent, erect. Pinus-Quercus forests Ch (Breedlove 9689, F). 1300-1600 m. (Mexico [Quertaro, Veracruz, Michoacn, Oaxaca], Mesoamerica.)

3. Photinia microcarpa Standl., Publ. Carnegie Inst. Wash. 461(4): 57 (1935). Holotype: Belize, Schipp 1291 (F!). Illustr.: Cand. J. Bot. 70: 2153, t. 1 (1992). Trees to c. 15 m. Branches tan- or rufous-tomentose initially, ultimately glabrate, the bark reddish-brown or gray. Leaves 5-15 2-4 cm, elliptic to obovate, chartaceous to coriaceous; the midvein sulcate above, the secondary veins usually flat; both surfaces initially tomentose, ultimately glabrate with persistent hairs along the midvein below; base cuneate; margins entire to serrulate or serrate; apex acute, acuminate or obtuse; petioles 2-15 mm. Inflorescence a flat-topped panicle, 4-8 cm in diameter, the pedicels, hypanthium and calyx densely white- or rufous-tomentose; flowers 20-80; bracts linear, 3-5 mm. Flowers pedicellate, the pedicels 2-5 mm; hypanthium conical, c. 4 mm; sepals triangular, 1-2 mm, the apex acute; petals 3-4 mm, white; carpels 3, white-tomentose apically. Fruit 8-12 5-7 mm, ellipsoid or obovoid, glabrate, red; the calyx persistent, white-tomentose. Wet forests. Vouchers ??? 500-3000 m. (Mesoamerica.)

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 33 of 61 Phipps (1992) recognized three subspecies that are most easily separated by leaf size and the number of flowers in an inflorescence. Although the typical form of each may be readily identified, they do intergrade and intermediate specimens can be difficult to place to subspecies.

3a. Photinia microcarpa subsp. microcarpa. Leaves 7-12 2-4 cm; margins entire to serrate; petioles 2-10 mm. Inflorescence 7-8 cm in diameter, flowers 30-50. Vegtation missing. B (Monro 937, MO); H (Meja 245, MO). 500-2500 m. (Endemic.)

3b. Photinia microcarpa subsp. elliptica J.B. Phipps, Canad. J. Bot. 70: 2154 (1992). Holotype: Panama, Lao 602 (MO). Illustr.: Cand. J. Bot. 70: 2157, t. 12 (1992) Leaves 5-7 2-2.5 cm; margins entire, cartilaginous; petioles 5-6 mm. Inflorescence 4-7 cm in diameter, flowers 20-40. Vegetation missing. B (Monro 1234, MO); G (Lundell & Contreras 70317, MO); N (Molina 20486, MO); P (Galdames 1955, F). 500-1400 m. (Endemic.)

3c. Photinia microcarpa subsp. hintonii J.B. Phipps, Canad. J. Bot. 70: 2154 (1992). Holotype: Mxico: Hinton 338 (GH). Illustr. Canad. J. Bot. 70: 2155, t. 11 (1992). Photinia microcarpa var brevis J.B. Phipps., P. microcarpa var. hintonii J.B. Phipps. Leaves 7-13 2-4 cm; margins entire to remotely serrate; petioles 5-15 mm. Inflorescence 8-10 cm in diameter, flowers 40-80. Vegetation missing. Ch (Breedlove 41284, MO); G (Steyermark 43087, F); H (Hazlett 1547, MO); N (Stevens 4519, MO). 1200-3000 m. (Mexico [Mexico, Michoacan, Oaxaca], Mesoamerica.)

14. Potentilla L. By F.R. Barrie.

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 34 of 61 Annual, biennial or perennial herbs, not armed, from a stout caudex or rhizome, rarely stoloniferous. Leaves basal or with 1 - several alternate, cauline leaves, these often reduced; blades pinnately or palmately compound, chartaceous, alternate, stipules present, leafy. Inflorescences cymes or panicles, or flowers solitary. Flowers bisexual; hypanthium salverform, campanulate or cupulate, bractoles 5, adnate to the hypanthium and alternating with the sepals, sepals 5; petals 5, white, yellow or red; stamens about 20; carpels many, free, the style subterminal. Fruit a head of achenes on the conical or hemispheric receptacle, often enclosed by the persistent hypanthium; achenes without a persistent style; seed 1. Bibliography: Dobles C. & J. Paule. Mol. Phylogenetics and Evol. 56: 156-175 (2010); Eriksson, T., et al., Int. J. Plant Sci. 164: 197-211 (2003); Rydberg, P.A. Potentilla. N. Amer. Fl. 22(4):293-352 (1908).

1. Stems trailing, stoloniferous and rooting at the nodes. 1. Stems erect, stolons absent. 2. Leaves palmately compound; leaflets 5-9. 3. Leaf undersurface pubescent, but not densely lanate. 3. Leaf undersurface densely lanate, the hairs white or grey. 2. Leaves ternately or pinnately compound, leaflets 3-9.

3. P. indica

1. P. goldmanii 4. P. staminea

4. Leaflets 3-9; flowers 10 or fewer; petals white, longer than sepals. 2. P. heterosepala 4. Leaflets 3, flowers 10 to 50 or more, petals yellow, shorter than sepals. 3. P. norvegica

1. Potentilla goldmanii Painter ex Rydb., N. Amer. Fl. 22(4): 314 (1908). Holotype: Mexico, Oaxaca, Nelson & Goldman 691 (US). Illustr.: Herbs with a stout caudex, typically shrouded by old leaf bases. Leaves predominately basal, the blade palmately compound, orbicular, 1.5-3.0 cm in diameter; leaflets 5, the 3 central leaflets subequal, 10-15 5-7 mm, the lateral leaflets 8-10 3-5 mm; surfaces green, pubescent, more densely so along the midvein and margins; base cuneate; margins crenate or serrate; apex acute; petioles 2-4 cm, green, pubescent.

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 35 of 61 Cauline leaves simple, 10-20 mm, linear, entire or 1-3 cleft. Inflorescence 1 to 4 or more, 2-20 cm; pubescent; flowers fewer than 10. Flowers pedicellate, the pedicels 2-5 cm, gracile, sericeous or pubescent; bracteoles c. 5 mm, oblong, the apex bluntly obtuse; calyx lobes 5-7 2-3 mm; ovate; green or flushed purple, sericeous or pubescent, the apex acute; petals 8-10 6-8 mm, obovate, white, glabrous, the apex entire or retuse; receptacle flat or convex, pilose; stamens 20-60, 3-4 mm; styles c. 2 mm. Achenes c. 3 mm, glabrous. Quercus and Juniperus forests. G (Steyermark 50232, F). 2900-3700 m. (Mesoamerica.)

2. Potentilla heterosepala Fritsch, Bot. Jahrb. Syst. 11: 314 (1890). Lectotype (1st stage?): Standley & Steyermark, Fieldiana 24 (4): 460 (????). Scherzer s.n. HERB? ST: Veracruz, Wawra 913, 936 [herb.?]. Illustr. Potentilla donnell-smithii Focke, P. heterosepala var. guatemalensis Fritsch, P. heterosepala var. mexicana Fritsch. Herbs with a simple or branching caudex, shrouded by old leaf bases. Stems green, pilose, Basal leaves 2-5 2-4 cm, deltoid; ternate or 5-9 pinnate, green or flushed red, the terminal and lateral leaflets subequal, 7-20 3-15 mm, ovate to obovate, subsessile; pilose, more densely so along the veins on the lower surface; the base cuneate, the margins deeply crenate; apex rounded or broadly acute; petioles 2-17 cm, slender, pilose. Cauline leaves similar to the basal leaves but reduced, apetiolate distally. Inflorescence cymose; flowers 4-8, remote. Flowers pedicellate, the pedicels 1-3 cm; tomentose; bracteoles 3-5 mm, ovate to elliptic, tri-lobed, the apex acute; sepals 4-5 mm, ovate, the margins entire, the apex acute to acuminate; petals yellow, 5-10 mm, obovate or obcordate, glabrous, the margin entire, the apex retuse; receptacle convex, pubescent; stamens 10-20, 2-3 mm. Achenes c. 50, 1-2 mm, glabrous. Pinus-Quercus and Abies forests, alpine slopes. Ch (Breedlove 29366, MICH); G (Matuda 2374, MICH); CR (Alfaro 1733, MO). 2200-4100 m. (Mxico [Puebla, Guererro, Oaxaca], Mesoamerica.)

3. Potentilla indica (Andrews) Th. Wolfe, Syn. Mitteleur. Fl. 6(1): 661 (1904). Fragaria indica Andrews, Bot. Repos. 7, pl. 479 (1807). Holotype: Illustration drawn from living material in England, Andrews, Bot. Repos. t. 479 (????).

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 36 of 61 Duchesnea indica (Andrews) Focke. Perennial herbs, stems trailing and stoloniferous, rooting at the nodes, green or reddish, sparsely to moderately pilose or sericeous. Leaves ternate, the leaflets subequal or the central leaftlet slightly larger, 1.5-5.5 1.4-3.5 cm, widely ovate to rhombic or elliptic; green, pilose; base cuneate; margins crenate or dentate, the teeth rounded; apex broadly acute or rounded; petioles 5-15 cm, pilose. Flowers axillary, solitary; pedicels 210 cm, green, pilose; hypanthium plane, pilose; bracteoles 3-10 2-7 mm, obovate, trilobed, green, pilose; sepals 7-9 3-4 mm, ovate, green, pilose, the margin entire, the apex acuminate; petals c. 10 3 mm, yellow, the apex rounded or truncate; stamens numerous; receptacle hemispheric, ovaries numerous. Achenes c. 1 mm, ovate, brown, superficial on the greatly expanded receptacle. Vegetation missing. Ch (Ruz 573, MO); G (Standley 58565, F). 1500-2200 m. (Eurasia, Canada, United States, Mxico [Nuevo Leon, Hidalgo, Distrito Federal, Mxico, Puebla, Veracruz], Mesoamerica, Peru, Bolivia, Chile, Argentina, West Indies.) Native to East Asia, Potentilla indica has become a weed of agricultural and disturbed sites in tropical and subtropical regions world-wide. The name most commonly applied to it has been Duchesnea indica. However, molecular analysis nests the species well within the Potentilla clade (Eriksson et al., 2003). The fruit greatly resembles a strawberry, but is inedible.

4. Potentilla norvegica L., Sp. Pl. 1: 499 (1753). Lectotype (designated by Jonsell & Jarvis, 2002): Herb. Linn. no. 655.41 (LINN). Illustr.: Reed, C. Common weeds of the United States 219, t. 107 (1971). P. monspeliensis L. Biennial or perennial herbs; stems erect, 0.5-1.0 m, green or reddish, hirsute, the hairs 1-3 mm. Leaves basal and cauline, ternate, the leaflets subequal or the central leaflet slightly larger, 2-8 1-3 cm, narrowly elliptic to obovate or oblanceolate, surfaces green, pubescent or hirsute, more densely so along the midvein and margins; base cuneate; margins crenate or serrate; apex acute; petioles of basal leaves 5-20 cm; petioles of cauline leaves 1-10 cm, subsessile apically. Inflorescence a multi-branched terminal cyme; flowers 10-50 or more; bracts leafy, lobed. Flowers pedicellate, the pedicels 1-3

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 37 of 61 cm; hypanthium cupulate, green, hirsute; bracteoles 3-7 2-3 mm, ovate, the apex acute or acuminate; sepals 5-8 3-4 mm, ovate, the apex acute or acuminate; petals 2.5-5 22.5 mm, widely ovate or oblong, yellow, the apex trucate or retuse; stamens usually 20; receptacle hemispheric, ovaries numerous. Achenes 0.8-1.3 mm, tan. Disturbed grounds. P (Johnston 1242, MO). 0-500 m. (Canada, United States, Mesoamerica, Eurasia.) A weed widespread in temperate regions, the single known Mesoamerican collection of Potentilla norvegica was collected near a "goat barn" on San Jose Island, Panama, in 1946. Collections from elsewhere in Mesoamerica have not been seen.

5. Potentilla staminea Rydb., Mem. Dept. Bot. Columbia Coll. 2: 67 (1898). Syntypes, So. Mexico, Ghiesbreght 131 & 681. Illustr.: Herbs to c. 50 cm, from a simple or branched caudex, shrouded in old leaf bases. Stems densely white sericeous. Basal leaves palmately compound, the blades 3-10 cm in diameter; leaflets 5-7, 1-5 0.5-1.5 cm, oblong or obovate, the upper surface green, pilose, the lower surface lanate, white or gray, pilose-sericeous along the veins; base cuneate; margins coarsely serrate, apex rounded; petioles 5-11 cm, densely white pilose or tomentose. Inflorescence racemose, flowers 5-18, remote. Flowers petiolate, the petioles 1-6 cm, gracile, white-tomentose; bractoles 4-5 1.5-2 mm, oblong or ovate, pilose, the apex acute; sepals 5-6 2-3 mm, ovate, pilose, the apex acute or acuminate; petals 10-15 10-15 mm, yellow, widely obovate, glabrous, the base cuneate, the margin irregular, the apex rounded; stamens 40-50, 2-3 mm; receptacle convex, pilose; styles c. 3 mm. Achenes c. 2 mm. Alpine meadows Ch (Breedlove & Strother 46359, MO); G (Smith 201, F). 2500-3700 m. (Mexico [Distrito Federal, Hidalgo, Oaxaca], Mesoamerica.)

15. Prunus

To be added. 16. Pyracantha M. Roem. By F.R. Barrie.

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 38 of 61 Shrubs or small trees. Unarmed or more commonly armed with thorns or thorny branchlets. Leaves alternate or fasciculate, often borne on short lateral branches, simple, coriaceous, the margin entire, crenulate or crenate; stipules minute, caducous. Inflorescences axillary or terminating lateral shoots, corymbose, bracts minute, caducous. Flowers pedicellate, hypanthium cupulate or campanulate; bracteoles absent; sepals 5; petals 5, white, shortly clawed; stamens 15-20; carpels 5, adnate to the hypanthium basally, free apically, white-pubescent or tomentose, ovules 2 per locule; styles 5, free. Fruit a pome, globose, red or orange, the sepals persistent. Pyrenes 5. Approx. 10 spp., native to Eurasia, widely planted as ornamentals and occasionally naturalizing locally. Pyracantha spp. are shrubs or small trees native to Eurasia grown for their attractive, glossy green leaves, showy inflorescences of small, white flowers and bright red or orange fruits. In Mesoamerica, they appear to naturalize infrequently and spread rarely. Bibliography: Gu Cuizhi & S.A. Spongberg, in Flora of China Editorial Committee, Flora of China, 9: 108-111 (2003). Nesom, G.L. Phytoneuron 2010-2: 1-6 (2010).

1. Leaves glabrous, the leaf margin crenulate.

2. P. crenulata

1. Leaves tomentose or sparsely pubescent on lower surface, the leaf margin entire or minutely serrulate. 2. Leaves densely tomentose on lower surface, the blade 4-8 mm wide. 1. P. angustifolia 2. Leaves sparsely pubescent on lower surface, the blade 7-12 mm wide. 3. P. koidzumii

1. Pyracantha angustifolia (Franch.) C.K. Schneid., Ill. Handb. Laubholzk. 1(5): 761 (1906). Cotoneaster angustifolius Franch., Pl. Delavay 221 (1890). Three syntypes listed in Tropicos. Check ref. Illustr.: not found Shrubs or small trees to 4 m. Branchlets densely yellowish-tomentose when young, ultimately glabrate, the bark purplish-black. Leaf blades 1.5-5.0 0.4-0.8 cm; narrowly oblong to oblanceolate; initially densely grey-tomentose on both surfaces, the

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 39 of 61 upper surface ultimately glabrescent; base cuneate; margin entire; apex obuse, apiculate or emarginate; petioles 1-3 mm or blades sessile. Inflorescence 2-4 cm in diameter in flower, flowers c. 12; peduncles tomentose. Flowers pedicellate, the pedicels 1-2 mm, the pedicels, hypanthium and calyx densely grey tomentose; hypanthium campanulate; sepals c. 1 1 mm, triangular, the apex acute; petals orbicular, c. 4 mm in diameter; stamens 20, 1.5-2 mm; styles 1-2 mm. Fruit 5-6 mm in diameter, oblate, red-orange. Cultivated, naturalized locally. Ch (Breedlove & McClintock 23628, MO). 2200 m. (Mesoamerica, China, cultivated elsewhere.)

2. Pyracantha crenulata (D. Don) Roem., Fam. Nat. Syn. Monogr. 3: 220 (1937). Mespilus crenulata D.Don, Prodr. Fl. Nepal. 238 (1825). Type:?? China, Henry 10625. Illustr.: not found. Shrubs or small trees to 5 m. Branchlets pubescent when young, the hairs red, ultimately glabrate, the bark dark brown. Leaf blades 2-7 0.8-1.8 cm; oblong to oblanceolate; glabrous; base cuneate; margins crenulate; apex acute or obtuse; petioles 36 mm, glabrous. Inflorescence 3-5 cm in diameter in flower; flowers 10-30; peduncles pubescent basally or glabrescent. Flowers pedicellate, the pedicels 4-10 mm, glabrous; hypanthium campanulate, glabrous; sepals triangular, 1.0-1.2 1.0-1.2 mm; petals 4-5 3-4 mm, broadly elliptic; stamens 20, 2-3 mm; styles 1.5-2.5 mm. Fruits 3-8 mm in diameter, oblate or globose, orange-yellow, orange-red or red. Cultivated, naturalized locally. G (Steyermark 46759, F) 1600-1700 m. (Mesoamerica, China, cultivated elsewhere.)

3. Pyracantha koidzumii (Hayata) Rehder, J. Arnold Arbor. 1: 261 (1920). Cotoneaster koidzumii Hayata (koizumii), J. Coll. Sci. Imp. Tokyo 30: 101 (1911). Holotype: Taiwan, ????? Illustr.: not found. Shrubs to c. 4 m. Branchlets pubescent when young, ultimately glabrate, the bark dark grey. Leaf blades 3.0-7.0 0.7-2.5 cm; narrowly elliptic to narrowly obovate; upper surface glabrous; lower surface densely pubescent when young, ultimately with scattered hairs; base cuneate; margin entire or occasionally with minute, remote teeth; apex emarginate or truncate; petiole 2-5 mm, pubescent. Inflorescence 3-4 cm in diameter in

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 40 of 61 flower; flowers 10-30; peduncle sparsely pubescent. Flowers pedicellate, the pedicels 511 mm, the pedicel, hypanthium and calyx sparsely pubescent, hypanthium campanulate, sepals triangular, 1.0-1.2 1.0-1.2 mm; petals 3.5-4.0 3.0-4.0 mm, suborbicular or broadly elliptic; stamens 20, 2-3 mm; styles 2-3 mm. Fruit 4-5 mm in diameter, oblate, red-orange. Cultivated, naturalized locally. G (Molina 15320, F); CR (Rodriguez & Vargas 386, MO). 1200-1500 m. (Mesoamerica, China, Taiwan, cultivated elsewhere.)

17. Pyrus L. By F.R. Barrie.

Shrubs or trees, unarmed or with thorns. Leaves deciduous; simple, alternate, chartaceous; margins entire, denticulate or dentate; stipules persistent[?]. Inflorescence racemose. Flowers appearing with or before the leaves; petiolate; hypanthium cupuliform; bracteoles absent; sepals 5; petals 5; stamens 20-30; carpels 2-5, connate basally; styles free. Fruit a pyriform pome with a cartilaginous core, the flesh with numerous stone cells; seeds 1-2 per carpel. Approx. 75 spp., Eurasia, Canada and United States. Widely cultivated elsewhere. 1 cultivated species in Mesoamerica.

1. Pyrus communis L., Sp.Pl. 470 (1753). Lectotype (designated by Amaral Franco & Rocha Afonso, 1965): Herb. Linn. No. 647.1 (LINN). Illustr.: N.v.: Pera; pear. Small to medium trees; branches unarmed or with thorns; branchlets glabrous or sparsely pubescent. Stipules caducous, 5-10 mm, linear, pubescent. Leaves 2-8 2-5 cm, ovate to widely ovate or elliptic; chartaceous or subcoriaceous; surfaces pubescent or tomentose when young, ultimately glabrate; base rounded or truncate; margins crenateserrulate; apex acute to acuminate; petioles 1.5-5 cm, tomentose or glabrate. Inflorescence of 5-15 flowers; pedicels 1.5-4 cm, glabrous to sparsely puberulent; hypanthium 2-4 mm, glabrous to sparsely tomentose externally, densely tomentose on inner surface; sepals triangular 2-4 mm, the inner surface tomentose; petals 10-15 mm, obovate, white; stamens 20-30. Fruit 5-12 3-9 cm, pyriform, the calyx persistent. Cultivated. Ch (Breedlove & McClintock 34014, MO); G (---, F). 1500-3000 m. (Mesoamerica, Eurasia, cultivated elsewhere.)

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 41 of 61 The common pear is planted for its fruit and as an ornamental street tree in the montane regions of Chiapas and western Guatemala. It is unclear whether or not it naturalizes in the region.

18. Rosa L. By F.R. Barrie.

Shrubs erect, straggling or prostrate, or woody lianas; unarmed or more commonly armed with straight or recurved prickles. Leaves persistent or deciduous, ternate or imparipinnate, leaflets 3-9, stipules present, partially adnate to the petioles, the distal end free. Inflorescences corymbs or flowers solitary; bracteoles absent; sepals 5, often pinnately lobed; petals 5 (10 or more in many cultivars), white or pink (or red or yellow); stamens numerous; hypanthium globose, campanulate or urceolate, carpels superior, numerous, free; style attached at apex; ovule 1. Fruit a pometum (hip), consisting of the fleshy, persistent hypanthium enclosing the numerous, bony achenes. Approx. 200 spp., Eurasia, Canada, United States, Mexico. No species of Rosa is native to Mesoamerica, but members of the genus are often planted as ornamentals. These are cultivars of hybrid origin, bearing large, brightly colored flowers with numerous petals, far exceeding the wild type number of 5. A cultivar of Rosa banksiae R. Br., a white-petaled native of China, has been collected in Chiapas (Santz 746, MO). Bibliography; Gu Cuizhi & K.P. Robertson, 2003. Rosa, in Flora of China Editorial Committee, Flora of China, vol. 9: 339-381; Lewis, W.H. 2012(?) Rosa. Flora of North America, vol 9:

1. Leaflets 7-9, 2-3 1.0-1.5 cm, inflorescence of 10-30 flowers. 2. R. multiflora 1. Leaflets 3-7, 2.0-5.0 1.5-3.5 cm; inflorescence of 1-5 flowers. 2. Flowers usually 4 or 5, rarely solitary, with little or no fragrance. Erect shrubs. 1. R. chinensis 2. Flowers solitary, rarely 2 or 3, fragrant; semiscandent shrubs. 3. R. odorata

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 42 of 61 1. Rosa chinensis Jacq., Obs. Bot. 3: 7 (1768). Type: not found? Illustr.: Flora of China Illustrations, vol. 9, t. 148, nos. 4-7 (????). Rosa montezumae Bertol. Shrubs to c. 2 m; branchlets green to purple-brown, sparesely puberulent or glabrescent; prickles 5-7 mm, sparse to abundant, recurved, flat. Leaves ternate or pinnate, leaflets 3-7; terminal leaflet 3-5 2.0-3.5 cm, elliptic, green, puberulent or glabrate; base cuneate or rounded; margin serrate; apex acuminate; lateral leaflets similar but smaller, 2.5-4 1.5-3 cm; petioles 2-5 cm, unarmed or with scattered prickles, 2-3 mm, stipules adnate to the petiole, the margin glandular-ciliate, the apex auriculate. Inflorescence terminal, flowers 4 or 5 or rarely flowers solitary. Flowers 5-9 cm in diameter, white, yellow, pink, red or purple, faintly fragrant or none; hypanthium ovoid or pyriform, green, glabrous; sepals 2-9 cm, lanceolate, tomentose adaxially, the apex attenuate; petals numerous, 2-4 1.5-4 cm, obovate; stamens few to numerous, some or all replaced by petals; styles free, about half as long as the stamens. Fruit 1-2 cm in diameter, pyriform or ovoid, red, glabrous. Cultivated; naturalized locally. B (Balick et al. 2333, MO); G (Vliz 94.3777, MO); H (Molina & Molina 31357, MO); N (Hernndez 30, MO). 0-1400 m. (Mesoamerica, China; widely cultivated elsewhere.) Commonly planted as an ornamental. Standley & Steyermark (Fl. Guatemala, p. 471) report that Rosa chinensis has naturalized there, particularly in the Pacific foothills.

2. Rosa multiflora Thunb., in Murray, Syst. Veg. ed. 14: 474 (1784). Type: Japan. Thunberg s.n., Herb. Thunberg 12168 (UPS). Illustr.: Rosa cathayensis (Rehder & E.H. Wilson) L.H. Bailey, R. multiflora var. cathayensis Rehder & E.H. Wilson. Erect or semiscandent shrub, to 3 m or more. Stems terete, green or reddish, armed with scattered, retrorsely curved prickles, glabrous. Leaves pinnately compound, leaflets 7-9, subequal, 2-3 1-1.5 cm, elliptic or obovate, chartaceous, puberulent or glabrescent; base cuneate; margin serrate; apex acute; petiole 2-5 cm, pubescent or glandular- pubescent; stipules adnate to the petioles, the margin fimbriate, often glandular-ciliate, the apex auriculate. Inflorescence terminal, flowers 10-30. Flowers 3-5 cm in diameter, white or pink; pedicels 5-20 mm, pubescent, hypanthium campanulate,

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 43 of 61 glabrous or pubescent, sepals 5-8 mm, elliptic or ovate, tomentose abaxially, the margins entire or infrequently 2-3-lobed, the apex aciculate; petals 5, or 10 or more in cultivars, 612 mm, obovate, stamens numerous, or some replaced by petals in cultivars; styles united to form a column, about as long as the stamens. Fruit c. 1 cm, ellipsoid, red. Cultivated; naturalized locally. Ch (Santz 902, MO); G (Standley 65753, F); H (Standley & Willliams 4598, F); N (Araquistain 44, MO); CR (Kahn et al. 589, MO). 0-1600 m. (Mesoamerica, Japan and Korea; widely cultivated elsewhere.) In sites where it becomes established, Rosa multiflora can be quite aggressive. It often forms dense hedgerows along roadsides.

3. Rosa odorata (Andrews) Sweet, Hort. Suburb. Lond. 119 (1818). Rosa indica var. odorata Andrews. Roses 2: t. 77 (1810). Illustr.: Flora of China Illustrations 9: fig. 148, 1-3 (????). Semiscandent shrub. Branches repent, glabrous, prickles 5-7 mm, scattered, recurved. Leaves ternate or pinnate, leaflets 3-7; terminal leaflet 3-5 2.0-3.5 cm, elliptic, green, puberulent or glabrate; base cuneate or rounded; margin serrate; apex acuminate; lateral leaflets similar but smaller, 2.5-4 1.5-3 cm; petioles 2-5 cm, unarmed or with scattered prickles, 2-3 mm, stipules adnate to the petiole, the margin glandularciliate, the apex auriculate. Flowers 3-10 cm in diameter white, pink, yellow or orange, solitary, or rarely in fascicles of 2 or 3, fragrant; hypanthium oblate, glabrous; sepals 2-5 cm, lanceolate, villose abaxially, exernally glabrous, the apex attenuate. Petals numerous, 2-3 cm, obovate; styles free, about as long as the stamens. Fruit 1-2 cm in diameter, oblate, red. Cultivated; naturalized locally. H (Sanchez 188, MO). ???? m. (Mesoamerica.)

19. Rubus L. By F.R. Barrie.

Erect or semiscandent shrubs or trailing or scandent vines. Stems, leaves and inflorescences unarmed or more commonly armed with slender to stout, straight or recurved prickles, and vested with simple, white to reddish hairs and also often with

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 44 of 61 reddish, glandular hairs. Stems green, red or red-purple. Leaves 3- or 5- foliolate, pinnate or simple; margin singly or doubly serrate; stipules persistent. Inflorescences terminal or axillary, racemose or paniculiform, or flowers solitary. Flowers pedicellate; hypanthium flat; bracteoles absent; sepals 5, green, persistent in fruit; petals 5, white, pink or purple; stamens numerous; carpels 10 to 100 or more, free, the style terminal. Fruits an aggregate of drupelets, these falling singly or as a unit, either attached to or free of the receptacle; yellow, orange, red or purple-black at maturity. Bibliography: Rydberg, P.A. N. Amer. Fl. 22(5): 428-480 (1913). Focke, W.O., Biblioteca Bot. 72: 1-120 (1910); 121-220 (1911); Biblioteca Bot. 73: 1-274 (1914).

1. Plants unarmed; leaves simple. 1. Plants armed with prickles; leaves compound. 2. Leaves pinnately compound.

21. R. trilobus

3. Underside of leaflets densely velutinous, white; corolla pink to red-purple; mature fruits deep purple. 14. R. niveus

3. Underside of leaflets sparsely pilose, green, corolla white; mature fruits red or orange. 17. R. rosifolius 2. Leaves digitally 3- or 5- foliolate. 4. Stems with glandular hairs. 5. Leaves simply serrate; inflorescence racemose; fruits 1.5-2.5 cm, cylindrical or oblong. 13. R. miser 5. Leaves doubly serrate; inflorescence paniculate; fruits 1-1.5 cm, subglobose. 6. Leaflets caudate, sepals 10-30 mm. 6. Leaflets acuminate; sepals 4-10 mm. 7. Stems with glandular hairs 2-5 mm long, green or yellow. 7. Stems with glandular hairs 1-2 mm long, reddish. 4. Stems without glandular hairs. 8. Leaves uniformly ternate. 9. Leaflets broadly elliptic to broadly obovate, the apex acute rounded. 4. R. ellipticus 9. Leaflets narrower, ovate to elliptic or broadly ovate, the apex acuminate. 1. R. adenotrichos 3. R. costaricanus 10. R. leptosepalos

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 45 of 61 10. Inflorescence of 1 to 3 flowers; fruit ovoid, the sepals clasping. 16. R. pringlei 10. Inflorescence of 5-30 flowers, fruity pyramidal or ellipsoid; sepals reflexed or spreading. 11. Upper surface of leaflets puberulent. 11. Upper surface of leaflets glabrous. 5. R. eriocarpus 7. R. glaucus

8. Leaves mostly 5- foliolate, 3-foliolate leaves commonly occurring near the inflorescences. 12. Stems and petioles densely hispid, the hairs 2-3 mm, red. 12. Stems and petioles pilose, the hairs white, or glabrate. 13. Stipules amplexicaul, reniform. 13. Stipules not amplexicaul, linear to lanceolate. 14. Leaves simply serrate, coriaceous. 15. Leaves glossy green on upper surface. 15. Leaves not glossy, drying brown. 14. Leaves doubly serrate, chartaceous. 16. Sepals 3-6 mm. 17. Inflorescences sparingly glandular-hirsute; petals 10-15 mm. 17. Inflorescences lacking glandular hairs; petals 5-6 mm. 16. Sepals 5-12 mm. 18. Inflorescences compact, cylindrical, pedicels less than 1 cm long; fruits villous or with a few apical hairs. 8. R. hadrocarpus 2. R. coriifolius 23. R. vulcanicola 6. R. fagifolius 15. R. ostumensis 12. R. malacocarpus 22. R. urticifolius

18. Inflorescences more open, pedicels 1-3 cm long; fruits pubescent or glabrous. 19. Inflorescences bearing 1 to 8 flowers; fruits 5-10 mm in diameter; drupelets fewer than 30. 9. R. humistratus

19. Inflorescences bearing (5)-10 to c. 100 flowers; fruits 10-25 mm; drupelets more than 30. 20. Sepal apices as long or longer than the blade, caudate or foliose. 20. R. shankii 20. Sepal apices shorter than the blade, cuspidate to caudate-acuminate.

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 46 of 61 21. Stems and inflorescences puberulent or glabrate; inflorescence racemose, flowers (5) 10-20. 12. R. macrogongylus

21. Stems and inflorescences pilose; inflorescence paniculate or corymbose; fls 10 to c. 100. 22. Inflorescences armed with stout prickles; drupelets pubescent apically. 18. R. sapidus. 22. Inflorescences unarmed or armed with a few, weak prickles; drupelets glabrous. 19. R. schiedeanus.

1. Rubus adenodrichos Schltdl., Linnaea 13: 267 (1839). Isotype: Mexico, Schiede s.n. (NY) (web image!). Illustr.: Ann. Missouri Bot. Gard. 37: 157, t. 66 (sub R. praecipuus). Rubus praecipuus L.H.Bailey, R. roseorum A. Berger. Arching shrub; stems 1-3 m, armed with recurved prickles, densely hirsute, the hairs 1-5 mm, erect, red and glandular, also densely short-pilose, the hairs white. Leaves digitally 3- or 5-foliolate, the leaflets 4-15 1.5-7 cm, elliptic to oblong or ovate, chartaceous, the upper green, sparsely pilose, the lower surface pale green, densely pilose, especially along the veins; base cuneate; margin doubly serrate; apex acuminate; petioles 2-9 cm, petiolules 1.0-4.5 cm, glandular hispid and pilose, armed with recurved prickles; stipules 3-5 mm, linear-lanceolate, pilose. Inflorescence terminal, paniculate, peduncles and petioles glandular hispid and pilose, armed with recurved prickles or more commonly unarmed; flowers typically 30-100, sometimes fewer; bracts 5-8 mm, linear, pilose on both surfaces; pedicels c. 1 cm. Flowers with sepals 5-10 mm, ovate, densely pilose on both surfaces, often glandular hispid on outer surface, apex acute to acuminate; petals 6-12 3-8 mm, elliptic, pink or white. Fruits ellipsoid, 1-1.5 cm, the sepals reflexed; drupelets 20-70, black at maturity, glabrous. Moist or wet thickets, PinusQuercus forests, disturbed areas. Ch (Ton 2173, F); G (Steyermark 43416, F); ES (Fasset 28547, F); H (Molina 13758, F); N (Williams et al. 27692, F); CR (Burger et al. 12022, MO); P (Woodson et al. 869, MO). 1000-2500 m. (Mexico [Morelos, Puebla, Veracruz, Michoacn, Guerrero, Oaxaca], Mesoamerica, Colombia, Ecuador, Venezuela.)

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 47 of 61 2. Rubus coriifolius Liebm., Vidensk. Meddel. Naturhisk. Foren. Kjbenhavn 1852: 157 (1853). Syntype: Mexico, Leibmann s.n. (GH!). Illustr.: Gentes Herb. 6: 357, t. 181 (1944). Suberect, arching or semiscandent shrub; stems sparsely armed with recurved prickles, eglandular, moderately to densely pilose, the hairs white. Leaves digitally 3- or 5- foliolate, the leaflets 5-12 2.5-7.5 cm; ovate to elliptic; chartaceous; upper surface dark green, sparsely pubescent; lower surface pale olive, pubescent, the veins prominent; base subcordate to cuneate; margin doubly serrate; apex acuminate; petioles 3-9 cm, petiolules 0.5-4.0 cm, armed, pilose; stipules 3-7 mm, linear-lanceolate, pilose. Inflorescences terminal, paniculate; flowers 10 to c. 100; peduncles and petioles pilose, bracts 5-7 mm, lanceolate; puberulent on both surfaces; pedicels 5-30 mm. Flowers with sepals 4-6 mm, oblong-elliptic, pilose on both surfaces, the apex acute to acuminate; petals 10-15 5-8 mm, white or pink, elliptic to obovate, the apex rounded. Fruits c. 1 cm in diameter, subglobose, the sepals reflexed, the drupelets 8-40, black at maturity, glabrous. Pinus-Quercus forests, cloud forests. Ch (Moore & Brossard 6346, F); G (Williams et al. 22245, F); H (Molina 18552, F); N (Rueda 11743, MO). 500-2500 m. (Mxico [Tamaulipas, Hidalgo, Mxico, Veracruz, Michoacn, Guerrero, Oaxaca], Mesoamerica.)

3. Rubus costaricanus Liebm., Vidensk. Meddel. Naturhisk. Foren. Kjbenhavn 1852: 159 (1853). Holotype: Costa Rica, Oersted s.n. (C; photo F!). Illustr.: Gentes Herb. 6: 357, t. 181 (1944). Rubus hondurensis L.H. Bailey, R. irasuensis Liebm., R. laxus Rydb., R. pittieri Rydb., R. tantus L.H. Bailey. Arching shrub or semiscandent vine; stems sparingly armed with recurved prickles, moderately to densely glandular setose, the hairs 1-2 mm, and densely pilose, the hairs white. Leaves digitally 3- or 5-foliolate, the leaflets 4-11 1.5-6 cm; ovate or oblong to widely ovate or widely oblong; chartaceous; upper surface green, pilose; lower surface olive, densely pilose; base subcordate or rounded; margin doubly serrate; apex acuminate; petioles 6-12 cm, petiolules 0.5-2.0 cm, armed, glandular setose and pilose; stipules 3-7 mm, linear-lanceolate, pilose. Inflorescence terminal, a simple or compound

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 48 of 61 panicle; flowers 20-100 or more; peduncles and pedicels unarmed, densely pilose and glandular setose, bracts 5-13 mm, linear-lanceolate, pilose; pedicels 5-20 mm, Flowers with sepals 4-10 mm, elliptic or ovate, densely tomentose on both surfaces, the apex cuspidate; petals 6-10 mm, elliptic, white or pink. Fruit c. 1 cm in diameter, subglobose to obloid, the sepals reflexed; drupelets c. 50, black at maturity, glabrous or pubescent apically. Pinus-Quercus forests, montane forest margins. Ch (Ton 798, F); G (Standley 82506, F); ES (Molina & Montalvo 21508, F); H (Molina 11275, F); CR (Morales 6232, F); P (Davidson 72, MO ). 1200-2700 m. (Endemic.) Rubus costaricanus may be distinguished by the sparingly armed stems with glandular hairs that are 1-3 mm long, and the subglobose fruits with relatively few drupelets. As the number of synonyms suggests, this species is somewhat variable across its range. Specimens with oblong leaves and apically pubescent fruits have gone by the name R. irasuensis, and Honduran plants with robust, large and spreading, compound inflorescences have been segregated as R. hondurensis. However, a comparison of photographs of the types of R. costaricanus and R. irasuensis indicates that the two represent the same taxon, and plants with large, compound inflorescences occur throughout the range of the species.

4. Rubus ellipticus Sm. in Rees, Cycl. 30.: Rubus no. 16 (1815). Type: Herb Smith? [verify] Illustr.: not seen. Subscandent shrub or vine; stems armed with recurved prickles, reddish, hirsute, the hairs 2-3 mm, red, eglandular, and short-pilose, the hairs white. Leaves ternate, the terminal leaflet 4-9 3-6 cm, the lateral leaflets 2.5-6.0 2-4 cm; broadly elliptic to widely obovate; chartaceous; upper surface green, sparsely pubescent; lower surface pale green or olive, tomentulose; base cuneate; margin serrulate; apex rounded; petiole 2-6 cm, armed, hirsute and short-pilose; petiolule of terminal leaflet 1-5 cm, that of lateral leaflets 2-5 mm; stipules linear, 5-7 mm, pilose. Inflorescences terminal and axillary, paniculiform; flowers 5-30; bracts 3-4 mm, linear, peduncles and pedicels pilose. Flowers with sepals 4-7 2-5 mm, ovate to elliptic, pilose on both surfaces with a few erect red hairs basally; margin entire; apex cuspidate; petals 5-10 mm, obovate, white. Fruits 1-2 cm in diameter, subglobose, the sepals ascending; druplets 50-150, yellow or orange at

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 49 of 61 maturity, glabrous. Cultivated; escaped and naturalized locally. CR (Williams et al. 28563, F). 1600-3200 m. (Mesoamerica, India; cultivated elsewhere.)

5. Rubus eriocarpus Leibm., Vidensk. Meddel. Naturhisk. Foren. Kjbenhavn 1852: 159 (1853). Syntype: Leibmann, s.n. (F!). Illustr.: not seen. Erect to semiscandent shrub; stems armed with retrorse prickles, glabrous and often glaucous. Leaves ternate, the terminal leaflet 4-11 2-7.5 cm, the lateral leaflets 4-8.5 2-5.5 cm, elliptic or obovate to broadly elliptic or obovate; chartaceous; upper surface green, sparsely to moderately puberulent; lower surface velutinous, white; base truncate or subcordate; margin doubly serrate; apex acuminate; petioles 4.5-9.0 cm, glabrous, armed; petiolule of terminal leaflet 1-3 cm, those of lateral leaflets 2-3 mm; stipules 3-5 mm, linear, glabrate. Inflorescences terminal, paniculate; flowers usually fewer than 20; peduncles and pedicels unarmed or armed, the prickles sparse, slender, tomentulose; bracts 1-2 mm; pedicels 1-2 cm. Flowers with sepals 5-6 mm, lanceolate, tomentulose on both surfaces, the apex caudate-acuminate; petals 5-7 mm, obovate, white. Fruit 1-2 cm, pyramidal or ellipsoid, the sepals reflexed; drupelets 50-80, black at maturity, tomentose. Wet thickets in Pinus-Quercus forests, cloud forests. Ch (Ton 3989, F); G (Standley 70024, F); ES (Tucker 1068, F); H (Molina 25724, F); CR (Burger & Liesner 6300, F); P (Davidson 881, F). 1500-2500 m. (Mexico [Distrito Federal, Mexico, Veracruz, Guerrero, Oaxaca], Mesoamerica.) Treated as conspecific with Rubus glaucus in the Flora de Nicaragua (Pankhurst, 2001), Rubus eriocarpus is distinguished from R. glaucus in the Flora of Guatemala (Standley & Steyermark, 1946) predominately on differences in the upper leaf surface, here being more or less puberulent, and glabrous or with only a few hairs in R. glaucus. A comparison of the types suggests that the two species are distinct, separated not only by the leaf character but also by floral characters. The petioles on the syntype of R. eriocarpus are less than 2 cm long and virtually without armament, while the petioles on the holotype or R. glaucus are about 5 cm long and armed with stout, retrorse prickles. An inspection of a larger suite of specimens suggests that the leaf and floral characters are usually found together, although the pedicels on some plants with glabrous leaves are

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 50 of 61 bereft of prickles. Here, two species are recognized, but a more detailed investigation of the relationship between them would be welcome.

6. Rubus fagifolius Schltd. & Cham., Linnaea 5: 571 (1830). Syntype: Mexico, Schiede 589 (HAL) (web image!). Illustr.: Arching shrubs or more commonly scandent vines; stems up to 18 or 20 m, armed with compressed, recurved prickles, tomentose, eglandular. Leaves digitally 3- or 5foliolate, the leaflets 3-10 2-6 cm, elliptic to oblong; coriaceous; the upper surface green, glossy, glabrous; lower surface tan, velutinous; base cuneate or rounded; margin remotely serrate to finely and doubly serrulate; apex caudate-acuminate; petiole 3-10 cm, armed, pilose; petiolules 2-30 mm, armed, pilose; stipules 3-7 mm, linear-lanceolate, tomentose. Inflorescences terminal and axillary, compound panicles, flowers usually several hundred, peduncles and pedicels pilose, prickles few or absent; bracts 3-5 mm, lanceolate; pedicels 5-15 mm. Flowers with sepals 5-7 mm, ovate, velutinous on both surfaces, the apex acute or acuminate in fruit; petals 10-12 mm, elliptic to obovate, white or pink, the apex rounded. Fruit c. 1 cm, globose, the sepals reflexed; drupelets 5-20, red at maturity, sparsely pubescent, ultimately glabrate. Moist or wet forests. Ch (Ton 1878, F); B (Schipp 8704, F); G (Steyermark 37153, F); ES (Molina et al. 16944, F). 200-2700 m. (Mexico [Veracruz] Mesoamerica.)

7. Rubus glaucus Benth., Pl. Hartw. 173 (1845). Holotype: Ecuador, Hartweg 973 (K) (web image!) Illustr.: Romoleroux, Fl. Ecuador, no. 56. 30: t. 11 (1996). Erect, arching or semiscandent shrubs; stems armed with retrorse prickles, glabrous, often glaucous. Leaves ternate, the terminal leaflet (2-)6-12 (1.5-) -6 cm, the lateral leaflets (1.5-)5-8 (1-)2-4 cm; lanceolate to ovate or broadly ovate; chartaceous; upper surface dark green, glabrate to sparsely pilose; lower surface densely white-velutinous, the veins prominent, red; base cuneate, rounded or subcordate; margin serrulate or doubly serrate; apex acuminate; petioles armed, (1-)5-8 cm; petiolules of terminal leaflets 1-2 cm, petiolules of lateral leaflets 2-3 mm; stipules 2-5 mm, linear, pubescent. Inflorescences terminal and axillary; flowers solitary or 5-30; peduncles and pedicels armed, the prickles slender or more commonly stout, recurved, pilose; bracts 2-3 mm,

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 51 of 61 linear; pedicels 1-5 cm. Flowers with sepals 5-10 mm, lanceolate to elliptic, tomentose on both surfaces, the apex caudate-acuminate or caudate; petals 5-8 mm, obovate, white. Fruits 1-2 cm, pyramidal or ellipsoid, the sepals reflexed; drupelets 40-80, black at maturity, tomentose. Wet forests, cloud forest and paramos. G (Williams et al. 41969, F); ES (Carlson 467, F); H (Molina 12730, F); CR (Burger 7934, F); P (Davidson 448, F). 1800-3000 m. (Mesoamerica, Colombia, Ecuador, Peru, Bolivia.) See comments under Rubus eriocarpus.

8. Rubus hadrocarpus Standl. & Steyerm., Fieldiana, Bot. 24(4): 477 (1946). Holotype: Guatemala, Standley 86270 (F!). Illustr.: Rubus hadrocarpus fo. adenophorus Standl. & Steyerm. Semiscandent shrub, 1-2 m; stems armed with retrorse prickles, finely pubescent, eglandular. Leaves digitally 3- or 5- foliolate; leaflets 5-12 3- 6 cm; oblong to broadly elliptic; chartaceous; upper surface green, sparsely pilose; lower surface slightly pale green; pilose along the veins; base rounded; margin serrate, apex abruptly acuminate to caudate-acuminate; petioles 2-10 cm, armed, pubescent; petiolules 5-30 mm, armed, pubescent; stipules 5-9 mm, linear, pubescent. Inflorescences terminal and axillary, paniculate, somewhat compact and cylindrical; flowers 5-20; peduncles and pedicels armed, pubescent; bracts 3-8 mm, linear; pedicels 2-10 mm. Flowers with sepals 5-11 mm, ovate, tomentose on both surfaces, the apex acuminate to caudate; petals 7-8 mm, elliptic, white. Fruits 1.5-2.5 cm; cylindrical, the sepals reflexed; drupelets 50-75, black at maturity, villous or with a few apical hairs. Wet thickets and montane forests. Ch (Breedlove 1608, F); G (Steyermark 46979, F). 2100-3000 m. (Endemic.) Standley and Steyermark (Fieldiana, Bot. 24(4): 478.) described Rubus hadrocarpus fo. adenophorus from a single specimen (Holotype: Steyermark 49859, F!). It differs from the typical form in having erect, red, glandular hairs along the stem. No similar collection has been found. It seems likely that this plant was the result of hybridization with a glandular-haired species, which was not sustained. The typical form was collected from the same population (Steyermark 46960, F).

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 52 of 61 9. Rubus humistratus Steud., Nomencl. Bot, ed 2, 2:478 (1841). R. humifusus Schltdl., Linnaea 13: 270 (1839), non Wiehe & Nees (1821). Syntype: Mexico, Schiede, s.n. (HAL) (web image!). Illustr.: not seen. Shrub or semiscandent or trailing vine; stems red, glabrate, armed with recurved prickles, eglandular, new growth green, pilose. Leaves digitally 3- or 5- foliolate; leaflets 1.5-4 0.8-2.5 cm; ovate to elliptic; chartaceous; upper surfaces sparsely pilose; lower surface pilose along the veins; base rounded; margin doubly serrate; apex acuminate; petioles 2-3 cm, armed, pilose; stipules 5-10 mm, linear, pilose. Inflorescences terminating lateral branches; flowers 1-8; peduncles and pedicels pilose, armed; bracts 510 mm, linear to oblanceolate; pedicels 10-20 mm. Flowers with sepals 5-10 mm, ovate, pilose on both surfaces, the apex cuspidate to caudate; petals 7-10 mm, elliptic to obovate, white. Fruits 5-10 mm in diameter, subconical or subglobose, the sepals reflexed; druplets 15-30, black at maturity, pilose or glabrate. Vegetation missing Ch (Carlson 1634, F); G (Martinez & Ramrez 20802, MEXU). 700-2500 m. (Mexico [Nuevo Leon, Coahuila, Tamaulipas, San Luis Potosi, Guanajuato, Quertaro, Hidalgo, Mexico, Distrito Federal, Puebla, Veracruz, Jalico, Michoacan, Guererro, Oaxaca], Mesoamerica.)

10. Rubus leptosepalus Donn. Sm. Bot. Gaz. 57: 421 (1841). Holotype: Guatemala, Turckheim 2452 (US!). Illustr.: not seen. Arching shrub; stems armed with retrorse prickles, glandular-hirsute, the hairs 1-3 mm, red, also pilose, the hairs white. Leaves digitally 3- or 5- foliolate, the leaflets 6-15 2-6 cm; ovate to elliptic; chartaceous; upper surface dark green, sparsely pilose; lower surface pale green or olive, pilose; base cuneate or obtuse; margin doubly serrate; apex acuminate to caudate-acuminate; petioles 2-6 cm, armed, glandular-hirsute and pilose; stipules 5-9 mm, linear-lanceolate. Inflorescence a terminal panicle, flowers to c. 100; peduncles and pedicels glandular-hirsute; bracts linear, 10-20 mm; pedicels 1-2 cm. Flowers with sepals 1-3 cm, narrowly lanceolate, conspicuously nerved, pilose on both surfaces, the apex caudate; petals 1-1.5 cm elliptic or obovate, pink or white. Fruits c. 1 cm in diameter, subglobose to subconical, sepals reflexed; drupelets 25-40, black at

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 53 of 61 maturity, glabrous. Wet thickets and pastures. Ch (Raven & Breedlove 19811, F); G (Standley 69513, F); H (Molina 6086, F). 1200-1700 m. (Endemic.)

11. Rubus macrogongylus Focke, Feddes Rep. 9: 236 (1911). Type: not seen. Illustr.: not seen. Arching shrub to 3. m; stems sparsely armed, reddish or gray, pilose, eglandular. Leaves 3- or 5- foliolate, the leaflets 4-7 1-4 cm; ovate to elliptic; chartaceous; upper surface green, sparsely pilose or glabrate; lower surface olive, pilose; base subcordate; margin serrate; apex acuminate; petioles 3-5 cm armed, pubescent; stipules 5-8 mm, linear-lanceolate, pilose. Inflorescence terminal, racemose; flowers 5-20; peduncles and pedicels unarmed, pilose; bracts c. 5 mm, linear; pedicels 1-2 cm. Flowers with sepals 510 mm, ovate, pilose on both surfaces, the apex caudate-acuminate; petals c. 1 cm, obovate, white. Fruits 1-2 cm, oblong, sepals reflexed; drupes 50-80, black at maturity, glabrous. Moist thickets, mixed forests. Ch (Breedlove 8885, F); G (Steyermark 31078, F); N (Moreno 14336, MO). 1300-2400 m. (Endemic.)

12. Rubus malacocarpus Standl. & L.O. Williams, Ceiba 1: 239 (1951). Holotype: Costa Rica, Williams 16116 (F!) Illustr.: not seen. Semi-scandent shrub, 2-3 m; stems densely armed with retrorse prickles, pilose, eglandular. Leaves ternate, the leaflets 5-12 2-5 cm; ovate to narrowly elliptic; subcoriaceous; upper surface green, sparsely pilose; lower surface paler, pilose; base cuneate or obtuse; margin doubly serrate; apex acuminate; petioles 3-5 cm, armed, pilose; stipules amplexicaul, prominent, 5-10 mm, reniform. Inflorescences terminal and axillary, racemose; flowers 1-5; bracts amplexicaul, 5-10 mm, reniform; pedicels 3-9 cm, densely armed, tomentose. Flowers with sepals 1.2-2 cm, ovate, densely tomentose on both surfaces, the apex caudate-acuminate; petals c. 1.5 cm, ovate, pink to red-purple. Fruits 3-4 cm, ovoid to pyramidal, the sepals reflexed; druplets c. 100, black at maturity, densely pilose. Quercus forests, forest margins and disturbed sites. CR (Jimenez M. 3224, F) 2300-3000 m. (Endemic.)

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 54 of 61 13. Rubus miser Liebm., Vidensk. Meddel. Naturhisk. Foren. Kjbenhavn 1852: 156 (1853). Holotype: Costa Rica, Oersted s.n. (C, n.v., photo F!) Illustr: not seen. Rubus breviglandulifer L.H. Bailey. Arching shrub or small vine; stems densely glandular-hirsute, the hairs c. 1 mm or less, very uniform in length, red, also densely tomentose, the hairs white. Leaves digitally 3- or 5- foliolate, the leaflets 5-13 1.5-6 cm; lanceolate or narrowly elliptic to ovate or elliptic; chartaceous; upper surface dark green, pilose; lower surface pale green or olive, pilose; base rounded or cuneate; margin serrate; apex acuminate; petioles 2-8 cm, armed, glandular-hispid and tomentose; petiolules 0.5 -5 cm; stipules 5-10 mm, linear-lanceolate, tometose. Inflorescences terminal, racemose; flowers usually 20 or fewer; peduncles and pedicels unarmed, glandular-hirsute and pilose; bracts 5-10 mm, linear-lanceolate; pedicels 1-3 cm. Flowers with sepals 5-10 mm, ovate, densely tomentose on both surfaces, the apex caudate-acuminate; petals 5-10 mm, ellipitical, white. Fruits 1.5-2.5 cm, cylindrical or oblong, the sepals reflexed; drupelets 70-150, black at maturity, glabrous. Pinus-Quercus forests, wet and dry thickets. G (Standley 84046, F); ES (Molina 12603, F); H (Molina 1487, F); CR (Lent 3928, F). 1400-2300 m. (Endemic.) Rubus miser is characterized by the densely glandular-hirsute stems. The hairs are erect, dark red and both shorter, less than 1mm, and much more uniform than the glandular vestiture found on the stems of other Mesoamerican species.

14. Rubus niveus Thunb., Rubo 9, t. 3: ???? (????).Type: Indonesia, Thunberg s.n. (UPS) (not seen). Illustr.: Romoleroux, Fl. Ecuador, no. 56, 28, t. 10 (1996). Rubus albescens Roxb. Arching or scandent shrubs to c. 2 m; stems red to purple, armed with erect to slightly recurved prickles, glabrous, sometimes glaucous. Leaves imparipinnate, elliptic; chartaceous; leaflets 5-7, 3-6 2-5 cm, ovate or elliptic; upper surface green, sparsely pilose; lower surface white, densely velutinous; base cuneate or obtuse; margin doubly serrate; apex acute to acuminate; petioles 3-5 cm, sparsely armed, stipules 6-15 mm, lanceolate, glabrescent. Inflorescences terminal and lateral, compactly racemose; flowers 20-50; peduncles and pedicels moderately armed, pilose; bracts c. 5 mm, lanceolate; pedicels 5-10 mm. Flowers with sepals 3-6 mm, ovate, pilose on both surfaces, the apex

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 55 of 61 cuspidate; petals 3-5 mm, obovate or suborbicular, pink to purple. Fruits 8-15 10-15 mm, the sepals spreading or clasping; druplets 50-80, purple-black at maturity, tomentose to glabrate. Cultivated; escaped locally. G (Williams et al. 42127, F); ES (Molina & Montalvo 21650, F); H (Molina 7871, F); N (Stevens & Segovia 29796, MO); CR (Morales 1142, MO); P (Castillo 39, F). 1300-1900 m. (Mesoamerica, Eurasia, cultivated elsewhere.)

15. Rubus ostumensis Ant. Molina, Ceiba 18: 97 (1984). Holotype: Honduras, Molina 20335, F! Illustr: not seen. Trailing or semiscandent vine. Stem unarmed or armed with scattered, recurved prickles; puberulent; with or without a few erect glandular hairs. Leaves digitally 3- or 5foliolate, the leaflets 5.5-9.5 3-5 cm; oblong to elliptic; coriaceous; upper surface drying brown, sparsely pilose; lower surface pilose, especially along the veins; base rounded to obtuse; margin serrate, the teeth well spaced; apex acuminate; petioles 5-8 cm, unarmed or with a few prickles, pilose; petiolules 5-35 mm; stipules 4-6 mm, triangular. Inflorescences terminal and axillary, compound panicles; flowers 70 - 100 or more; peduncles and pedicels pilose, bracts 1.5-5 mm, linear or lanceolate; pedicels 3-10 mm. Flowers with sepals 4-5 mm, ovate or elliptic, tomentose on both surfaces, the apex cuspidate; petals 5-10 mm, oblong or obovate, white. Fruits 5-10 mm in diameter subglobose or ovoid, sepals reflexed; druplets c. 30 or fewer, black at maturity, glabrous. Cloud forest N (Williams et al. 27795, F) 1200-1500 m. (Endemic)

16. Rubus pringlei Rydb., N. Amer. Fl. 22: 443 (1913). R. occidentalis L. var. grandiflora Focke, Bot. Gaz. 16: 3 (1891). Syntype: Guatemala, Donnell Smith 2168 (US!). Illustr.: R. occidentalis var. mexicanus Focke. Erect or subscandent shrubs, to c. 2 m; stems red, armed with slender, erect or recurved prickles, initially pubescent, ultimately glabrate. Leaves ternate, the leaflets 310 1.5-6 cm; lanceolate to ovate; chartaceous; upper surface dark green, pilose; lower surface white or tan, densely velutinous; base cuneate or rounded; margin doubly serrate; apex acute or acuminate; petioles 2-6 cm; armed with small prickles, pilose; stipules 5

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 56 of 61 10 mm, linear-lanceolate, pilose. Inflorescences terminating lateral branches; flowers 1-3; bracts 3-5 mm linear, pilose; pedicels 1-3 cm, armed with small prickles, pilose. Flowers with sepals 6-7 mm, ovate, pilose on both surfaces, the apex caudate-acuminate; petals 67 mm, elliptic, white. Fruits ca 2 1.5 cm; ovoid; the sepals erect, clasping; druplets 4050, dark purple at maturity, tomentose. Moist montane thickets. Ch (Goldman 943, MO ); G (Steyermark 47352, F). 2400-3000 m. (Mexico [Mexico, Puebla, Oaxaca], Mesoamerica.)

17. Rubus rosifolius Sm., Pl. Icon. ined. 3: t. 60 (1791). Type: Mauritius, Commerson s.n., Herb. Smith No. 902.63, LINN., Illustr.: Erect or arching shrub to c. 2 m. Stems sparingly armed, eglandular, sparingly pilose, the hairs white, or glabrate. Leaves imparipinnately compound, the leaflets 5-15, 4-9 23 cm; lanceolate to narrowly elliptic; chartaceous; green, sparsely pilose along the veins on both surfaces; base cuneate; margin doubly serrate; apex acuminate; petioles 3-4 cm, sparingly armed, pilose; stipules 5-12 mm, acicular, sparingly pilose. Inflorescence terminal; flowers 1-3; peduncles and pedicels pilose, unarmed or with a few, delicate prickles; bracts 3-5 mm, linear, pilose; petioles 1-3 cm. Flowers with sepals 10-20 mm, ovate or lanceolate, sparingly pilose, the apex caudate-acuminate; petals 1-2 cm, widely elliptic or obovate, white. Fruits 1.5-2.0 cm, cylindrical, sepals spreading; drupelets c. 100 or more, red or orange at maturity, glabrous. Cultivated, naturalized locally. G (Steyermark 36938, F); ES (Molina et al. 16815, F); H ( ); N (Molina 38624, F); CR (Burger et al. 10768, F); P (Croat 66696, MO). 800-3800 m. (Mesoamerica, South and East Asia, cultivated elsewhere.)

18. Rubus sapidus Schltdl. Linnaea 13: 269 (1839). Syntype: Mexico, Schiede, s.n. (HAL, web image!). Illustr.: Erect, arching or semiscandent shrub; stems green, turning purple, armed with numerous, stout prickles, pilose. Leaves digitally 3- or 5- foliolate, the leaflets 5-14 39 cm; broadly ovate to broadly elliptic; chartaceous; upper surface dull green, sparsely pilose or glabrate; lower surface olive, pilose; base obtuse to subcordate; margin serrate; apex acuminate; petioles 3-8 cm; armed with stout prickles, pilose; petiolules 5-20 mm;

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 57 of 61 stipules 5-12 mm, linear to lanceolate, pilose. Inflorescences terminal, corymbiform or paniculiform; flowers 10 to c. 100; peduncles and pedicels armed, pilose; bracts 5-8 mm, linear, pilose; pedicels 1-3 cm. Flowers with sepals 5-10 mm, elliptic, tomentose on both surfaces, the apex cuspidate or caudate; petals 1-1.3 cm, elliptic to obovate, white. Fruit 1-1.8 cm, subglobose to cylindrical, the sepals reflexed; drupelets c. 30- 50, black at maturity, pubescent apically. Moist or dry thickets,open fields, Pinus-Quercus forests. Ch (Breedlove 8406, F); G (Williams et al. 41800, F); H (Molina 26131, F). 1500-2000 m. (Mexico [Veracruz], Mesoamerica.)

19. Rubus schiedeanus Steud., Nomecl. Bot, ed. 2, 2: 4781(841). Rubus dumetorum Schltdl., Linnaea 13: 267 (1839), non Wiehe ex. Boenn (1824). Holotype: Mexico, Schiede s.n. (HAL, web image!) Illustr: Rubus superbus Focke. Suberect or arching shrub, often a large vine, trailing over shrubs or trees; stems purplish, armed with recurved prickles, eglandular, puberulent, ultimately glabrate. Leaves digitally 3- or 5- foliolate, the leaflets 6-13 2.5-8 cm; ovate to elliptic or obovate; chartaceous; upper surface green, glossy, pubescent along the veins; lower surface dull green, pilose along the veins; base cuneate or subcordate; margin serrate; apex acute to acuminate; petioles 4-12 cm, reddish, armed with recurved prickles, pilose; petiolules 0.5-4.0 cm; stipules 4-8 mm, lilnear to lanceolate, puberulent. Inflorescence terminal and axillary, paniculiform, simple and few-flowered or compound and pyramidal; flowers 10-75; bracts c. 5 mm, ovate; pedicels 1-3 cm. Flowers with sepals 58 mm, ovate to elliptic, pilose on both surfaces, the apex acute to acuminate; petals c. 1 cm, obovate, white. Fruits 1-1.5 cm, ellipsoid, the sepals reflexed; drupelets 30-80, black at maturity, glabrous. Damp or wet montane thickets, open fields. Ch (Breedlove7816, F); G (Standley 68062, F); H (Molina 22307, F); N (Rueda et al. 8443, MO); CR (Brenes 678, F). 1000-2500 m. (Mexico [Veracruz], Mesoamerica.) This species was treated in the Flora of Guatemala under the name Rubus alpinus Macfad., which was not validly published (Stafleu & Cowan 1981: 219).

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 58 of 61 20. Rubus shankii Standl. & Steyerm., Ceiba 1: 143 (1950). Holotype: Honduras, Shank 71 (F!). Illustr.: not seen. Arching or subscandent shrubs; stems armed with coarse, recurved prickles, eglandular, pilose. Leaves ternately compound; leaflets 3-12 2-9 cm; elliptic to widely elliptic or widely ovate; chartaceous; upper surface dark green, sparsely pilose; lower surface pale green, tomentose; base obtuse; margin doubly serrate; apex acuminate; petioles 3-9 cm, armed with recurved prickles, pilose; stipules 5-15 mm, linear to lanceolate, pilose. Inflorescensces terminal and axillary; paniculate; flowers 10-30; peduncles and pedicels armed, pilose; bracts 5-15 mm, linear, pilose; pedicels 1-2 cm. Flowers with sepals 7-12 mm, pilose on both surfaces, apex caudate-acuminate or flattened and foliose; petals c. 10mm, elliptic of obovate, white. Fruits 1.5-2.5 cm, cylindrical, the sepals reflexed; drupelets 50-75, black at maturity, glabrous. Open montane forests and cloud forest. H (Williams & Molina 15654, F) 1700-2000 m. (Endemic.)

21. Rubus trilobus Seringe in de Candolle, Prodr. 2: 566 (1825). Holotype: Mexico, Sesse & Mocio s.n., t. 290 (G). Illustr: not seen. Slender suberect or subscandent shrub; stems unarmed, eglandular or rare with reddish glandular hairs on new growth, pilose the hairs white, soon glabrate. Leaves simple, the blades 3-12 3-12 cm; broadly triangular, trilobed; chartaceous; both surfaces green, pilose along the veins; base cordate; margin doubly serrate; apex acute or acuminate; petioles 3-5 cm, pilose; stipules 5-10 mm, lanceolate or narrowly elliptic. Flowers solitary; pedicels 2-4 cm, pilose, eglandular or rarely with reddish glandular hairs; sepals 1.0-1.5 cm, ovate to broadly ovate, pilose on both surfaces, the apices caudate-acuminate with prominent, leafy appendages, 1-2 cm, the margin lobed or serrate; petals 2-3 cm, broadly elliptic, white. Fruit 1.5-2.0 cm in diameter, subglobose, the sepals clasping; drupelets 25-40, black at maturity, glabrous. Montane forests. Ch (Breedlove 12387, F); G (Smith 578, F). 2500-3900 m. (Mexico [Puebla, Veracruz, Oaxaca], Mesoamerica.)

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 59 of 61 22. Rubus urticifolius Poir. Encyl. 6: 246 (1804). Holotype: Peru, Dombey s.n. (P, web image!) Illustr.: Ann. Missouri Bot. Gard. 37: 158, t. 67 (????). (sub. R. trichomallus). Rubus trichomallus Schltdl. Arching shrub to 3 m or more; stems sparingly armed with coarse, retrorse prickles, hirsute, the hairs c. 2 mm, red, eglandular, and pilose, the hairs white. Leaves 3- or 5foliolate, the leaflets 4-16 2-6 cm; elliptic to ovate; chartaceous; upper surface green, pilose; lower surface gray or white, densely tomentose; base rounded to obtuse; margin doubly serrate; apex acute to acuminate; petioles 3-9 cm, hirsute, densely armed; stipules 4-10 mm, linear-lanceolate. Inflorescence terminal, paniculate, pyramidal; flowers 20-40 or more; peduncles and pedicels sparingly armed, hirsute; bracts 5-8 mm, lanceolate, pilose; pedicels c. 1 cm. Fruits subglobose, 5-10 mm in diameter; the sepals reflexed; drupelets 50-75; purple-black at maturity, glabrous. Montane forest margins and clearings, disturbed sites. Ch (Breedlove 7572, F); G (Standley 68046, F); ES (Molina & Montalvo 21828, F); H (Molina 24091, F); N (Grant 832, F); CR (Brenes 5241, F); P (Vayama & Torres 26, F). 1000-2100 m. (Mexico [Oaxaca], Mesoamerica, Colombia, Eucador, Peru, Brazil, Paraguay.)

23. Rubus vulcanicola (Donn. Sm.) Rydb., N. Amer. Fl. 22(5): 455 (1913). R. guianensis var. vulcanicola Donn. Sm., Bot. Gaz. 23: 243 (1897). Syntype: Costa Rica, Pittier 806 (CR). Illustr.: not seen. Rubus panamanus L.H. Bailey; R. poliophyllus Focke non Kuntze, R. smithii Rydb. non Backer. Shrub or semiscandent vine. Stems armed with retrorse prickles, eglandular, puberulent to pilose, the hairs white. Leaves 3- or 5- foliolate, the leaflets 4-10 1.5-4 cm; chartaceous; upper surface dark green, pilose to glabrate; lower surface pale green, pilose, densely pilose along the veins; base cuneate to rounded or subcordate; margin serrate; apex acuminate; petioles 2.5-9 cm, armed, puberulent to pilose; petiolules 5-35 mm; stipules 3-8 mm, acicular, puberulent. Inflorescences terminal and axillary, paniculate; flowers 30-60 or more; peduncles and pedicels puberulent to pilose; bracts 38 mm, linear, puberulent; pedicels 1-2 cm. Flowers with sepals 3-4 mm, ovate, pilose on

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 60 of 61 both surfaces, the apex cuspidate; petals 5-6 mm, broadly elliptic to broadly obovate, white, pink or reddish purple. Fruits 5-10 mm, subglobose, sepals reflexed; druplets c. 50, purple-black at maturity, glabrous. Forest margins, cloud forest and disturbed sites. CR (Burger 7918, F); P (Davidson 72, F). 1000-3200 m. (Endemic.)

20. Spiraea L. By G. Davidse.

Shrubs, not armed. Leaves simple, alternate, denticulate or dentate; venation usually pinnate; stipules absent. Inflorescences corymbs or panicles, bracteate. Flowers bisexual; hypanthium hemispheric to campanulate; nectaries numerous, separate; sepals 5, valvate; petals 5, white; stamens 15-60, the filaments free except at the base; carpels 3-8, free, the styles free, the ovules 2 to several. Fruits follicles, dehiscent ventrally and sometimes tardily dorsally near the apex; seeds several, pendulous, linear to oblong, without an aril, the endosperm thin or absent, the testa membranous. Approx. 100+ spp. North temperate zone; 1 sp. Cultivated in Mesoamerica.

1. Spiraea cantoniensis Lour., Fl. Cochinch. 1: 322 (1790). Holotype: China, Loureiro s.n. (P?). Illustr.: Shrubs 1-1.5 m, glabrous; branches slender. Leaves petiolate; blades 2-6.3 0.62.1 cm, rhombic-oblong or rhombic-lanceolate, deep green above, pale bluish green beneath, the venation impressed above, raised below, the tertiary veins prominent, the base cuneate, the margins incised-serrate, the apex obtuse or subacute petioles 2-11 mm. Corymbs 1-3 cm, umbel-like, dense. Flowers c. 1 cm wide, 5-merous in the wild type, or with numerous petals and few or no stamens or pistils in cultivars. Cultivated. Ch (Mndez T. 9453, MO); G (Standley y Steyermark, 1946: 483); H (Garcs 99, MO); N (Grijalva et al. 2999, MO); CR (Chavarra y Herrera 20, MO); P (D'Arcy et al. 12712, MO). (Mesoamerica, native of China and Japan; widely introduced elsewhere as an ornamental.)

Flora Mesoamericana, Volume 2 (2), Rosaceae, page 61 of 61 Standley and Steyermark, (Fl. Guatemala, p. 483, 1946), report that the form planted in Mesoamerica is S. cantoniensis var. lanceata Zabel, Strauch. Spiraen. 41 (1893), which was described from ornamental material grown in Germany. The variety differs chiefly from the wild form in having double flowers with numerous petals, relatively few or no stamens, and usually no pistils.