Symplocos citrea Lex.

Kelly, Lawrence M., Almeda, Frank & Fritsch, Peter W., 2016, A taxonomic revision of Mexican and Central American Symplocos (Symplocaceae), Phytotaxa 264 (1), pp. 448-450 : 448-450

7. Symplocos citrea Lex. in La Llave & Lexarza (1824: 22). Neotype (designated by Díaz-Barriga & Cházaro B. 1993):— MEXICO. Michoacán: Mpio. Morelia, Ichaqueo, en bosque de encino humedo con elementos del bosque mesófilo de montaña, 2500 m, [19°34′22″N, 101°08′26″W], 23 September 1992, H. D í az-Barriga 7260 (neotype IEB-187899-n.v., online image!, isoneotypes CAS!, ENCB, IEB-115952-n.v., online image!, MEXU!, MICH-n.v., online image!, TEX-n.v., online image!, XAL)

= Symplocos prionophylla Hemsley (1881: 302) . Type:— MEXICO. Oaxaca: without precise locality, 1842, A. B. Ghiesbreght s.n. (holotype K, lost). Neotype (designated here): MEXICO. Guerrero: Atoyac , carretera entre El Paraiso y Filo de Caballos, 16–18 km al N de El Paraiso, 1200 m, [17°24′30″N, 100°11′36″W], 4 November 1979, S. Koch et al. 79325 (neotype CAS!, isoneotype NY!).

= Symplocos pringlei Robinson (1891: 168) . Type:— MEXICO. Michoacán: hills of Patzcuaro , [19°30′36″N, 101°36′W], 1890, C. G. Pringle 3345 (holotype GH!, isotypes A!, AC-n.v., online image!, BKL-n.v., online image!, BR-836920-n.v., online image!, BR-836906-n.v., online image!, CAS!, F!, GOET!, HBG-n.v., online image!, JE!, K!, LL-n.v., online image!, M- n.v., online image!, MEXU!, MO-2!, MIN-n.v., online image!, MSC-n.v., online image!, MU-n.v., online image!, NY!, PH-n.v., online image!, PUL-n.v., online image!, S!, US!, W-n.v., digital image!).

= Symplocos novogaliciana L.M.González in González-Villareal (2002: 21). Type:— MEXICO. Jalisco: San Sebastian del Oeste, camino de la Virgencita a Real Alto , 1940 m, 20 April 1999, J. J. Reynoso D. et al. 4032 (holotype IBUG-0162958- n.v., online image!, isotypes CAS!, ENCB, F-2!, IBUG-0162957-n.v., online image!, IEB, MEXU!, MICH-n.v., online image!, NY!, TEX-n.v., online image!, US!, WIS-n.v., online image!).

Trees 4–15 m tall; juvenile branchlets and vegetative buds sericeous to pilose, trichomes 0.75–1.5 mm long, appressed to antrorsely spreading, reddish brown. Petioles 0.8–1.3 cm long; leaf blades bicolorous, elliptic or oblong to lanceolate or obovate, (3–)6–13(–16) × (2.0–) 2.5–6.5 cm, coriaceous, abaxially appressed-pilose, densely spreading-pilose along midvein, adaxially glabrous, secondary veins not adaxially impressed, base acute to obtuse or rounded, margins serrulate-denticulate, with minute, narrowly conical, black deciduous glands (often persistent toward leaf apex), apex acute to short-acuminate. Inflorescences racemes 1–2.5(–5.5) cm long, 7–10- flowered; peduncle 0–5(–20) mm long; rachis 2–7 mm long, sericeous to pilose, trichomes 0.75–1 mm long; bracts deciduous, oblong to ovate, 2–4 × 2–3 mm, sericeous, ciliate to densely sericeous; bracteoles deciduous, 2–4, ovate, 2–4 × 2–3 mm, sericeous, ciliate to densely sericeous; pedicels 0–1.5 mm long. Hypanthium sericeous. Calyx lobes 5, broadly ovate to triangular, 2–3 × 2–3 mm, densely sericeous, margins ciliate. Corolla pink, 5–7- lobed, 8–12 mm long; tube 3–5 mm long; lobes adnate to filament tube for 5–7 mm, oblanceolate or rarely broadly elliptic, puberulent to sericeous distally. Stamens 4-seriate; filament tube 7–8 mm long; distinct portions of filaments 1–4 × ca. 0.5 mm. Disk densely pilose; style 8–10 mm long, sparsely pilose basally; stigma irregularly lobed. Fruits green maturing to dark bluish purple, narrowly cylindric to slightly obovoid (rarely fusiform), 1.3–1.6(–2) × (0.4–) 0.6–0.7 cm, strigillose, apex acute with calyx lobes forming a conical beak; disk conical, apex partly visible, surpassed by calyx; endocarp 3-locular, perimeter rounded.

Vernacular names —Boncillo (Soto & Roman 1429 [CAS, MO]), chocoyolillo (Lozano 516 [MEXU]), cucharillo (Cházaro B. & Zamudio 4724 [MEXU]), cucharo (González 48 [MEXU]), garrapato ( Magallanes & Parada 4454 [MEXU]), jaboncillo blanco (Rees s.n. [CAS, MO]), mamuyo (Hinton 7615 [GH, NY, US]), palo blanco (Madrigal 3091 [MEXU]), tchcari-urapiti (Madrigal 3091 [MEXU]), urápiti ucu (Madrigal 3123 [MEXU]).

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Illustrations — Díaz-Barriga (1993: 3); Díaz-Barriga & Cházaro (1993: 44). Photographic image — Figure 2f . Phenology —Flowering July through May; fruiting throughout the year. Distribution and habitat — Mexico (Guanajuato, Nayarit, Jalisco, Colima, Michoacán, México, Distrito

Federal, Morelos, Guerrero, and Oaxaca), common on the Pacific slope of the Eje Volcánico Transversal and in the

Sierra Madre del Sur, oak-pine forest, oak forest, broad leaved evergreen forest, cloud forest at (1100–)1500–2650

m elev. Figure 10 .

Conservation status —Although Symplocos citrea is endemic to Mexico, its distribution extends from the states of Nayarit southeastward to Oaxaca where it grows throughout a wide elevational gradient. It is one of the most commonly encountered species of Symplocos in Mexico and Central America, as evidenced by the over 230 collections that we have examined. Based on these data, we assign a classification of Least Concern (LC) to this species.

Discussion — Symplocos citrea is distinguished by the following combination of characters: leaves with the abaxial midvein densely pilose and the margins serrulate, flowers in dense, short racemes, corolla lobes puberulent to sericeous near the apex, ovaries consistently 3-locular, and fruits narrowly cylindric. This species is the most commonly collected Symplocos in western Mexico and is not likely to be confused with any other congeners that grow within its range ( S. hintonii , S. longipes , and S. sousae Almeda [1982b: 255] ), all of which are relatively rare and morphologically distinctive.

Symplocos novogaliciana , described from the Pacific slope of Nayarit and Jalisco, was distinguished from S. citrea by characters of its inflorescences, stamens, and sepals (González-Villarreal 2002). The differences cited included the pedunculate inflorescences of S. novogaliciana (with peduncles up to 20 mm), versus the subsessile fasciculate inflorescences of S. citrea ; sepals 3–5 × 1–2 mm, with triangular-attenuate lobes, acute apices, and

A TAXONOMIC REVISION OF SYMPLOCOS

Phytotaxa 264 (1) © 2016 Magnolia Press • 31 margins with vesicular glands between trichomes in S. novogaliciana , versus sepals 4–6 × 2.5–4 mm, with ovate obtuse lobes, rounded to acute apices, and margins without glands; and a smaller number of stamens in S. novogaliciana (45–50 versus 50–75; González-Villarreal 2002). Most of the distinctions used to recognize S. novogaliciana are unreliable. The shape and the size of the sepals vary throughout the range of S. citrea . Individuals with narrow lobes and acute apices are common, and many individuals have occasional sparse vesicular glands along the sepal margins. In addition, stamen number is highly variable throughout the range of S. citrea . The long peduncles of S. novogaliciana are strikingly different from most individuals of S. citrea , but even this is a conspicuous extreme on a continuum of variation from subsessile to pedunculate inflorescences in S. citrea (inflorescences that are generally condensed racemes, and not at all fasciculate). We thus treat S. novogaliciana as a synonym of S. citrea .

The only known original material of Symplocos prionophylla was a Ghiesbreght specimen at K that was photographed prior to being sent on loan to MEXU in 2000. This specimen was lost in transit. Based on this K photograph of the specimen and on the protologue, we are confident that S. prionophylla is a synonym of the earlier name S. citrea . We have designated a specimen from Guerrero as the neotype of S. prionophylla because there are few specimens and duplicates of S. citrea from Oaxaca.

Additional specimens examined — MEXICO. Colima: Volcán del Nevado , [19°32′N, 103°42′W], Bárcena 433 ( MEXU!) . Distrito Federal: Segundo Dinámo de Contreras , 2600 m, [19°17′02″N, 99°16′26″W], 7 December 1952, Gold 244 ( MEXU!) ; Autopista Mexico-Cuernavaca, Km 56 1/2, 2250 m, [10°06′45″N, 99°11′01″W], 7 December 1952, Gold 250 (MEXU!, MO!) ; Contreras , [19°16′48″N, 99°16′34″W], 22 October 1950, Gold s.n. ( MEXU!) ; Dinámo de Contreras , 2600 m, [19°16′48″N, 99°16′33″W], 29 October 1950, Matuda et al. 18692 (CAS!, MEXU!, NY!) ; Cañada de Contreras, cerca del Primer Dinámo , 2550 m, [19°17′17″N, 99°15′52″W], 16 April 1970, Rzedowski 27209 (CAS!, F!, NY!) . Guanajuato: Jerécuaro , 2 km al SE de Chilarillo, 2200 m, [20°06′26″N, 100°27′02″W], 13 September 1988, Rzedowski 47770 (MEXU!, MO!) . Guerrero: crest of ridge near Huapango on rd to San Vicente S of Omiltemi, 2290 m, [17°32′53″N, 99°41′43″W], 18 October 1984, Breedlove 61874 (CAS!, MO!) ; Taxco de Alarcón, Parque El Huizteco, 5 km al N de Taxco, en El Sombrero , 2480 m, 18°35′43″N, 99°36′54″W, 2 March 1998, Calónico S. 8080 ( MEXU!) ; Chilapa de Álvarez, Estación de microondas Pozo Largo , carretera Chilapa a Tlapa , 1935 m, 17°30′N, 99°03′W, 27 December 1993, Calzada 19013 ( MEXU!) ; Chilpancingo, 3 km adelante de Xocomanantlán, camino de Chilpancingo–Omiltemi , 2000 m, [17°33′21″N, 99°39′13″W], 14 October 1984, Casstelo 367 ( MEXU!) ; Taxco de Alarcón, Taxco , 8.2 km al NW, 2400 m, 18°35′16″N, 99°40′41″W, 28 May 1998, Cruz D. 2330 (MEXU-2!) ; Taxco de Alarcón, Taxco , 9 km al NW, 2290 m, [18°36′31″N, 99°40′05″W], 4 July 1998, Cruz D. 2511 ( MEXU!) ; along dirt rd between Taxco to Tetipac, 5.3 mi NW jct. rd to Ixcateopan and hwy through Taxco and 1.8 mi from reservoir above Casahuates , 6700 ft, [18°34′16″N, 99°37′20″W], 5 December 1983, Daniel & Lott 3323 ( CAS!) ; La Primavera , 5 km al SE de Las Palancas, 1900 m, [17°38′34″N, 100°07′55″W], 14 February 1974, González M. et al. 6766 (CAS!, MEXU!) ; Chilpancingo, Mojonera Plan de Potrerillos o 40° NE de Omiltemi, camino a Xocomotlán , 2400 m, [17°33′23″N, 99°38′33″W], 1 November 1984, Lozano 516 ( MEXU!) ; Taxco , [18°33′23″N, 99°36′18″W], 26 December 1943, Miranda 3064 ( MEXU!) ; Taxco de Alarcón, Arriba de Taxco, camino a Tenería , 2000 m, [18°33′37″N, 99°36′37″W], December 1943, Miranda 3079 ( MEXU!) ; Chilapa de Álvarez, Km 72–73 de la carretera Chilapa–Tilpa , 1900 m, [17°32′59″N, 98°52′14″W], 5 March 1994, Panero & Calzada 3911 (LL!, MEXU!) ; Taxco de Alarcón, El Aguacate , 11.6 km al W de Taxco, 2440 m, 18°34′30″N, 99°43′15″W, 1 May 1998, Ponce Vargas 40 (MEXU-2!) ; 2 km al E de Omiltemi, sobre el camino a Chilpancingo , 2250 m, [17°33′12″N, 99°40′21″W], 2 September 1962, Rzedowski 16030 ( CAS!) ; Taxco de Alarcón, 0.5 km al E de Cruz Alta, Puerto de Las Pitayas , rumbo a Puerto Oscuro , 2580 m, 25 May 1986, Valencia A. 24 ( MEXU!) . Jalisco: Tequila, N side of Volcán Tequila , 14.2 km S of Tequila on the rd to the microwave station, 2150 m, [20°47′54″N, 103°44′37″W], 5 October 1985, Bartholomew et al. 2685 (CAS!, GH!) ; Mpio. Tequila, N side of Volcán Tequila , 17.4 km S of Tequila on the rd to microwave station, 2440 m, [20°47′54″N, 103°44′37″W], 5 October 1985, Bartholomew et al. 2704 (CAS!, NY!) ; Tequila, N side of Volcán Tequila , 17.4 km S of Tequila on the rd to the microwave station, 2440 m, 5 October 1985, Bartholomew et al. 2710 (CAS!, GH!, NY!) ; Autlán de Navarro, along logging rd from El Chante to El Guisar in the Sierra de Manantlán , 2160 m, [20°17′N, 103°24′W], 17 August 1980, Breedlove & Almeda 45627 (CAS!, F!) ; Mpio. Autlán de Navarro, along logging rd from El Chante to El Guisar in the Sierra de Manantlán , 2160 m, [20°17′N, 103°24′W], 17 August 1980, Breedlove & Almeda 45628 (C!) ; Autlán de Navarro, along logging rd from El Chante to El Guisar in the Sierra de Manantlán , 2160 m, 17 August 1980, Breedlove & Almeda

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45672 (CAS!, GH!); Mpio. Autlán de Navarro, near El Guisar along logging rd from El Chante in the Sierra de Manantlán, 2620 m, 17 August 1980, Breedlove & Almeda 45703 (CAS!); 19 km SE of Mexican Hwy 110 S of Tecalitlán along rd to Jilotlán de los Dolores, 2130 m, [19°26′11″N, 103°11′23″W], 16 September 1986, Breedlove & Anderson 64274 (CAS!); Autlán de Navarro, Sierra de Manantlán, arriba del aserradero, ladera N, 9100 ft, 23 November 1968, Boutin & Brandt 2534 (CAS!, MEXU!); Tequila, Volcán de Tequila, 8 km S de Tequila por el camino que sube a las antenas, 2614 m, 20°48′30″N, 103°50′37″W, 5 September 2001, Carillo R. et al. 2614 (MEXU-2!); Tequila, Cerro de Tequila, brecha a las microondas, 2300 m, [20°47′54″N, 103°44′37″W], 1 September 1991, Cházaro B. et al. 6725 (MEXU!, MO!); Cerro El Almeal y las cercanías, 4.5 km por el camino al SE de la Estación Biológica Las Joyas, 2050 m, 22 December 1984, Cochrane et al. 10624 (MEXU!); Autlán de Navarro, R. B. Sierra de Manantlán, upper W edge of Cerro Grande, [19°29′27″N, 103°57′22″W], 7 June 1991, Cochrane et al. 12552 (F!, MO!); Cuautitlán, 52 km al NW de Colima, 3–4 km al NW de Telcruz, 1600 m, 19°25′38″N, 104°08′07″W, [19°29′53″N, 104°08′40″W], 10 February 1988, Cuevas & Guzmán 2163 (MEXU!); Cuautitlán, 17–18 km al NE de Cuautitlán, La Cumbre, 2100 m, 19°33′10″N, 104°14′20″W, 29 April 1988, Cuevas & Guzmán 2911 (MEXU!); 16–17 km al NE de Cuautitlán, 0.2 km al SW de La Cumbre, 1700 m, [19°32′45″N, 104°14′35″W], 22 July 1988, Cuevas & Guzmán 3167 (MEXU!); Cerro de Tequila, al E de Tequila, 2500 m, [20°47′39″N, 103°51′15″W], 15 September 1967, Díaz L. 413 (MEXU!); Tequila, Cerro de Tequila, 2300 m, [20°47′54″N, 103°44′37″W], 28 August 1972, Díaz L. 3471 (MO!); Sierra de Nayarit (Huichol Territory), Diquet s.n. (NY!); Tolimán, 4 km al NW del Terrero, camino a las microondas, 2200 m, [19°27′56″N, 103°57′34″W], 7 June 1991, Flores F. et al. 2634 (MEXU!, MO!); Tolimán, Rancho El Terrero, 1.5 km adelante del Aserradero, camino a las microondas, 2000 m, [19°27′42″N, 103°57′W], 14 June 1991, Flores F. et al. 2668 (MEXU!, NY!); Tequila, 12 km al SE de la estación de tren Tequila, camino a la estación de microondas, Cerro Tequila, 2200 m, [20°47′32″N, 103°50′33″W], 7 November 1993, Flores F. et al. 3101 (MEXU-8!); Barranca El Tejocote, 20 km al S de San Juan de la Montaña , 1735 m, [20°31′19″N, 103°08′20″W], 7 April 1988, García M. et al. 3879 (MEXU!); Sierra de Manantlán Biosphere Reserve, Las Joyas Section, 36 km S of Ahuaca TRANSECT #9-10, 1950 m, [19°24′36″N, 104°06′W], 25 March 1991, Gentry & Jardel 73554 (CAS!, MO!); Ayutla, Predio Santa Mónica, proximidad al campamento Las Iglesias, orilla del camino, 2040 m, [19°59′N, 104°28′W], 21 October 1979, Guízar N. 701 (MEXU!); Tequila, Cerro Tequila, 2650 m, [20°47′54″N, 103°44′37″W], 23 December 1967, González Villarreal et al. 1071 (MEXU-2!); Tonila, El Terrero, 2200 m, [19°27′42″N, 103°57′W], 2 April 1989, González Villarreal et al. 3658 (MO!); without precise locality, Hinton 3711 (F!); Tonila, brecha que va al Playón, Volcán de Fuego, 2350 m, 12 July 1992, Huerta et al. 232 (MEXU!); 2–4 km E of Cerro La Cumbre, 17.5 km of El Chante, top Sierra of Manantlán plateau, along lumber from N end of S. de Manantlán Central to arroyo de Neverias at W end of S. Manantlán W, 2350 m, 19°33′30″N, 104°11′13″W, 6 January 1980, Iltis et al. 2355 (CAS!, MEXU!); Autlán de Navarro, R. B. Sierra de Manantlán, 75 m to the N of rd fork to Las Joyas, 2100 m, 9 March 1987, Iltis et al. 29379 (GH!); on SW facing slope below the plateau top of Cerro Grande, on rd from Terrero to Mexicalitos (Los Picachos), 2300 m, 17°27′45″N, 103°56′W, 13 March 1987, Iltis et al. 29493 (F!, MEXU!, MO!, S!); Sierra de Manantlán, SW facing slope below the plateau top of Cerro Grande, 2300 m, [19°26′32″N, 103°57′21″W], 13 March 1987, Iltis et al. 29493a (CAS!, F!, MO!); Tecalitlán, 20 km al S de Llanitos por la brecha a Plan de Lego, y ca. 3 km al E por la cañada, 2150 m, [19°20′34″N, 103°15′45″W], 14 November 1989, Koch & Fryxell 89135 (NY!); Cuautitlán, 16–17 km NE de Cuautitlán, 2 km SW de La Cumbre, 1900 m, 19°33′07″N, 104°14′30″W, 15 February 1989, López & Guzmán 77 (MEXU!); Mazamitla, Fraccionamiento Natura, Mazamitla, 2000 m, 1 July 1996, Machuca N. 7727 (NY!); Autlán de Navarro, Las Joyas, Manantlán, Magallanes S. & Parada B. 4454 (MEXU!); Tecatitlán, Barranca del puerto de La Resbalosa, 2000 m, [19°30′40″N, 103°16′19″W], 6 July 1974, Mancera O. 361 (MEXU!); Venustiano Carranza, Loma de la Ailera, predio Hacienda del Jazmín, 2500 m, 10 July 1974, Mancera O. 419 (MEXU!); NW slopes of Nevada de Colima, in pine-fir zone above Jazmín, abandoned lumber rd 1 km above El Isote, 2500 m, [19°37′43″N, 103°38′05″W], 29 March 1949, McVaugh 10100 (GH!, MEXU!, NY!); 3–5 mi NW of San Miguel de la Sierra (ca. 40 km, airline, W of Ayutla), [20°07′28″N, 104°37′15″W], 2000 m, 3 November 1962, McVaugh 22055 (NY!); San Sebastián, segundo arroyo, 1500 m, [20°49′33″N, 104°07′07″W], 22 January 1927, Mexia 1547 (A!, CAS!, DS!, F!, MO!, NY!, US!); San Sebastian, arroyo del Triangulo, 1425 m, [20°49′33″N, 104°07′07″W], 20 March 1927, Mexia 1908 (A!, CAS!, DS!, F!, GH!, MO!, NY!); Ciudad Guzmán, Brecha a un lado de Los García , hacia la parte O del Nevado de Colima, 2155 m, [19°38′21″N, 103°40′09″W], 28 September 1993, Negrete et al. 49 (NY!); Tequila, Volcán Tequila, Km 9 de la carretera a la estación de microondas en la cima de la montaña , 2600 m, [20°47′54″N,

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Phytotaxa 264 (1) © 2016 Magnolia Press • 33 103°44′37″W], 8 October 1993, Panero et al. 3644 (CAS!, MEXU!, NY!); Tamazula, cerca de Agua Hedionda, 45 km al E de El Aserradero, sobre el camino a Manuel Diéguez, 1900 m, [19°52′43″N, 104°22′54″W], 26 May 1973, Rzedowski & McVaugh 1088 (NY!); Tolimán, 3 km al NW de El Terrero, 12–13 km NE de Minatitlán, 2300 m, 19°27′42″N, 103°57′48″W, 22 November 1993, Santana M. & Benz 6592 (MEXU!); La Manzanilla, 2 km al W de La Manzanilla, camino a Concepción Buenos Aires, 2100 m, [19°59′24″N, 103°10′22″W], 8 November 1987, Soltero 701 (MEXU-2!); camino de La Cumbre al Rincón de Manantlán, Cuautitlán, 2020 m, 3 April 1986, Vázquez 3909 (NY!); Tequila, Cerro de Tequila, 2500 m, [20°47′32″N, 103°50′33″W], 14 September 1969, Villareal de Puga 2650 (MEXU!); Tequila, Volcán Tequila, along rd to microwave station, 8500 ft, 20°47′N, 103°50′W, 24 October 1970, Webster & Breckon 15890 (GH!, MEXU-2!); Autlán de Navarro, R. B. Sierra de Manantlán, just E of Cerro La Cumbre on upper end of rd down to Rincón de Manantlán, 2.4 km by main rd E of Cerro El Almeal, 15.5 km (air) SE of Ahuacapán, 2100 m, 19°33′28″N, 104°14′14″W, 19 March 1989, Wetter et al. 1096 (CAS!, F!, GH!); R. B. Sierra de Manantlán, 7.3 km N of El Terrero on the rd to La Laguna, 4.5 km (air) NNW of El Terrero, 2372 m, [19°28′51″N, 103°58′21″W], 23 March 1989, Wetter et al. 2052 (F!); 26.5 km SE of jct. to Mex 110, on the way to Jilotlán de Dolores, 2040 m, [19°26′18″N, 103°11′03″W], 29 October 2000, Yahara et al. 2200 (MEXU!). Estado de México: Mpio. Valle de Bravo, 9.8 km N of Temascaltepec on rd to Valle de Bravo, 2100 m, [19°04′56″N, 100°04′31″W], 12 October 1985, Bartholomew et al. 2961B (CAS!); Temascaltepec, 16 km NE of Temascaltepec, 2290 m, [19°06′44″N, 100°09′58″W], 25 November 1983, Breedlove & Almeda 60478 (CAS!, MEXU!, NY!); Valle de Bravo, Temascaltepec, [19°11′34″N, 100°07′50″W], 21 October 1982, García R. 377 (MEXU!); Ocuilan de Arteaga, carretera Jococingo–Malinalco, desviación a Santa Cruz Tezontepec–Chalma, meseta basáltica del Holotepec, Tenango del Valle, 2310 m, 18°59′N, 99°26′W, 18 December 1989, González & Miranda 43 (MEXU!); Tejupilco, 3–4 km al E de Nanchititla, 2000 m, [18°52′24″N, 100°25′47″W], 19 November 1993, González M. et al. 6464 (CAS!, MEXU!); Tejupilco, 8–9 km al S de Nanchititla, en el camino a Bejucos, [18°49′59″N, 100°25′36″W], 19 November 1993, González M. et al. 6455 (MEXU!); Valle de Bravo–Avándaro, 1900 m, [19°10′44″N, 100°07′10″W], 12 November 1977, Hernández M. 3075 (MEXU!); Temascaltepec, Las Mesas, 2000 m, [19°02′60″N, 99°59′07″W], 2 November 1932, Hinton 2342 (F!, US!); Nanchititla, Dist. Temascalpetec, [18°52′N, 100°28′W], 17 March 1933, Hinton 3616 (NY!, US!); Temascaltepec, Hinton 3711 (F!); Temascaltepec, Rincón, 1980 m, [18°54′N, 100°07′12″W], 15 November 1933, Hinton 5092 (F!, MEXU!, S!); Tejupilco, Rincón, 1960 m, 12 November 1933, Hinton 5093 (A!, F!, MO!, S!); Dist. Temascaltepec, 20 March 1935, Hinton 7540 (US!); Temascaltepec, Nanchititla, [18°52′N, 100°28′W], 13 April 1935, Hinton 7615 (GH!, NY!, US!); 18 km W of Temascaltepec, 49 km E of Toluca at Km 49 along Toluca–Temascaltepec Hwy (#134), 2364 m, 19°06.719′N, 99°57.333′W, (19°06′43″N, 99°57′19″W), 22 January 2003, Kelly et al. 1375 (CAS!); Ocuilan de Arteaga, SE de Las Lagunas de Zempoala, Km 18 de la carretera Ocuilan a Cuernavaca, Barranca de Ocuilan, 2280 m, [18°59′15″N, 99°19′20″W], 1 November 1985, Lorence & Tejero 4892 (CAS!, MEXU!); Temascaltepec, Temascaltepec, [19°02′N, 100°03′W], 26 October 1952, Matuda et al. 26574 (MEXU-3!); Real de Abajo, Sultepec, 1800 m, 21 September 1953, Matuda et al. 29308 (MEXU-2!); Valle de Bravo, 1800 m, [19°11′34″N, 100°07′50″W], 23 November 1952, Matuda et al. 27309 (CAS!, MEXU-3!); Valle de Bravo, 1800 m, 21 November 1952, Matuda et al. 27817 (MEXU!); Santa Bárbara –Santo Tomás de los Plátanos, 1100 m, [19°10′37″N, 100°16′20″W], 11 October 1953, Matuda et al. 30385 (CAS!, MEXU!); Cerro de Mamatla, Zacualpan, 2000 m, [18°43′N, 99°46′39″W], 3 May 1954, Matuda et al. 30567 (MEXU!); Cerro de Ahuacatitlán, Almoloya de Alquisiras, 1700 m, [18°51′57″N, 99°53′38″W], 29 March 1954, Matuda et al. 30609 (MEXU!); Nanchititla, 1900 m, [18°52′N, 100°28′W], 26 May 1954, Matuda et al. 30867 (MEXU!); Cerro de Pinal, Otzoloapan, 2300 m, [19°07′34″N, 100°15′05″W], 18 October 1954, Matuda et al. 31893 (MEXU-2!); along Hwy 153 between Temascaltepec and Toluca, 53 km SW of Toluca, at turnoff to El Polvorin, 2000 m, [19°03′N, 100°01′48″W], 26 April 1987, Miller & Meyers 2621 (CAS!, MEXU!, MO!); Valle de Bravo, near Río Molino above reservoir, Valle de Bravo, 1 November 1949, Moore Jr. & Cetto 5484 (GH!); Valle de Bravo, ca. 6 km al S de carretera Avandaro–Toluca sobre la carretera a Temascaltepec, 2300 m, 5 October 1997, Panero et al. 7345 (NY!); Temascaltepec, Temascaltepec, 4 November 1956, Paray 2258 (MEXU!); Salto de Agua, [19°08′N, 98°46′W], November 1905, Purpus 1775 (F!, GH-2!, MO-2!, NY-2!, US!); Tejupilco, 1 km al W de Nanchititla [sic], 1800 m, [18°51′18″N, 100°27′29″W], 9 April 1966, Rzedowski 22099 (F!, MEXU!, MO!); Amecameca, 4 km al E de Santa Isabel Chalma, 2650 m, [19°09′02″N, 98°43′15″W], 25 February 1979, Rzedowski 36076 (CAS!, MEXU!, MO!); Desierto de Tenancingo, [18°55′23″N, 99°33′40″W], 1915, Salazar s.n. (MEXU!); 14.5 km al E de Ocuilan, sobre la carretera a Cuernavaca, 2250 m, [18°59′20″N, 99°20′05″W], 30 July 1983, Toledo M. 21 (MEXU!). Michoacán:

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Morelia , Cerro Azul, 2300 m, [19°25′40″N, 100°40′40″W], 6 September 1912 , Arsène s.n. ( JE!); Morelia , vicinity of Morelia, Cerro Azul, 2100 m, 4 November 1909 , [19°25′40″N, 100°40′40″W], Arsène 2842 (GH!, MEXU!, MO-2!, NY!); Morelia , vicinity of Morelia, Campanario, 2100 m, [19°23′10″N, 101°09′52″W], December 1910 , Arsène 5808 (A!, MO!, US!); along Mexico Hwy #15, 18 mi E of Morelia , 7200 ft, [19°39′14″N, 100°55′57″W], 2 November 1970 , Breedlove 18716 ( CAS!); 32 km SE of Puerto Las Cruces, N of Coalcoman along rd to Dos Aguas, Sierra de Coalcoman, 2195 m, [18°49′13″N, 103°03′42″W], 17 September 1986 , Breedlove & Anderson 64336 ( CAS!); 16 km N of Ejido Varaloso along rd to Dos Aguas , Sierra de Coalcoman, 2375 m, [18°48′27″N, 102°57′10.5″W], 17 September 1986 , Breedlove & Anderson 64412 ( CAS!); Morelia , 1–1.5 km al S de Atécuaro, 2260 m, [19°34′22″N, 101°11′21.5″W], 19 February 1992 , Carranza 3907 (CAS!, MEXU!); Los Reyes, Cerro Patambán, 3 km al E de Cirío, 2540 m, [19°45′13″N, 102°23′14″W], 27 April 1999 , Carranza & Torres 5733 ( MEXU!); Villa Madero , Brecha Villa Madero–Nocupetaro, aprox. 15 km después de Villa Madero, 2480 m, [19°20′15″N, 101°22′01″W], 4 May 1987 , Cházaro B. & Zamudio 4724 ( MEXU!); Santa Clara del Cobre , Rincón Agua Verde, [19°25′04″N, 101°45′03″W], 27 July 1994 , Cruz E. & Pérez Calix 3 ( MEXU!); Quiroga , Cerro La Acumara, 6 km Santa Fé de la Laguna, 2100 m, [19°41′34″N, 101°34′22″W], 29 May 1985 , Díaz B. 1058 ( MEXU!); 2 km SW de Los Tanques , 2300 m, [19°27′54″N, 101°36′37″W], 23 September 1986 , Díaz B. 3208 ( MEXU!); Tingambato , Ladera W del Cerro Cumburinda, 2000 m, [19°30′57″N, 101°53′14″W], 14 October 1988 , Díaz B. & Bello 5179 ( MEXU!); Pátzcuaro , entre los Cerros La Cantera y Los Lobos, 2200 m, [19°51′20″N, 102°22′33″W], 29 March 1989 , Díaz B. 5792 ( MEXU!); Santa Clara del Cobre , camino al Cerro de La Cantera, 2300 m, [19°18′28″N, 101°40′40″W], 23 October 1989 , Escobedo 1877 ( MEXU!); Pátzcuaro , 6 km al S de Pátzcuaro, carretera a Opopeo, 2340 m, [19°27′47″N, 101°36′42″W], 10 November 1985 , Espinosa Garduño 2014 ( MEXU!); Morelia , Cerro Peña de San Pedro, SW de San Miguel del Monte, 2400 m, [19°36′45″N, 101°09′11″W], 8 December 1990 , García L. 3557 ( MEXU!); Paracho , cerro al N de Ahuiran, 2300 m, [19°40′16″N, 102°04′22″W], 13 December 1990 , García & Pérez 3583 ( MEXU!); Morelia , Agua Zarca, cerca de San Miguel del Monte, 2200 m, 22 October 1991 , García & Pérez 4029 ( MEXU!); Uruapan , Malpais de Capacuaro, 2350 m, [19°32′55″N, 102°03′05″W], 13 November 1991 , García & Pérez 4118 ( MEXU!); Uruapan , 10 km de la desv. Anagahuan–Paricutín, 2250 m, [19°33′02″N, 102°13′05″W], October 1988 , González M. et al. 17060 (F!, MEXU- 2!); Uruapan , 10 km de la desviación Anagahuan–Paricutín, 2250 m, 13 July 1988 , González M. et al. 17068 (MEXU-2!); Sierra Toricillas , Dist. Coalcoman, [20°05′09″N, 102°00′46″W], 14 October 1938 , Hinton 12379 (NY!, US!); Zitacuaro – Los Orcados, Dist. Zitacuaro, 2400 m, [19°22′12″N, 100°20′30″W], 11 October 1938 , Hinton 13349 (NY!, US!); Coalcoman , S. Torricillas, 2200 m, [20°05′09″N, 102°00′46″W], 28 July 1939 , Hinton 15028 (F!, GH!, MO!, NY!); Mpio. Tancítaro , Tancítaro, 2100 m, [19°20′14″N, 102°21′50″W], 11 October 1940 , Hinton 15511 (F!, MO!, NY!); Barroloso , Coalcoman, 2400 m, 22 October 1939 , Hinton 15367 (GH!, US!); Rincón de Ahorcado , al S de Cerro Cacique, 2400 m, [19°22′17″N, 100°18′38″W], 21 October 1978 , Ibarra C. 225 ( MEXU!); Ladera S del Cerro Cacique, 2410 m, [19°22′15″N, 100°18′45″W], 28 April 1979 , Ibarra C. 463 ( MEXU!); entre Copétiro y Apo., 1860 m, [19°26′54″N, 102°26′14″W], 1 September 1980 , Ibarra C. 1162 ( MEXU!); Ciudad Hidalgo , 20 km al O de Cd. Hidalgo, sobre carretera a Morelia, 2300 m, [19°40′18″N, 100°43′54″W], 7 November 1977 , Koch & Fryxell 77383 ( NY!); Charopan , 5 km al S de Ocumicho, 2290 m, [19°46′05″N, 102°12′51″W], 25 May 1986 , Labat 822 ( MEXU!); Uruapan , La Alberca, Tiamba, 1960 m, [19°29′25″N, 102°01′59″W], 9 November 1978 , Madrigal S. 3091 ( MEXU!); San Juan Parangaricutiro, Agua Chiquita, 2300 m, [19°23′35″N, 102°13′05″W], 11 November 1978 , Madrigal S. 3100 ( MEXU!); Tlalpujahua , Cerro de Pamatácuaro , Pamatácuaro, Los Reyes, 2520 m, [19°42′22″N, 102°21′32″W], 16 November 1978 , Madrigal S. 3123 ( MEXU!); Cd. Hidalgo , en La Venta , a 17 km al W de Cd. Hidalgo, carretera Morelia, por Mil Cumbres, 2450 m, [19°39′23″N, 100°43′13″W], 11 October 1983 , Martínez S. et al. 4675 (CAS!, F!, MEXU!); Mpio. Patzcuaro , 2 km al N de la ex-Hacienda de Charahuen, 2115 m, 19°31′59″N, 101°42′45″W, 5 February 2004 , Molina & Zamudio 449 ( CAS!); Uruapan , 3.3 mi from Uruapan Hwy on rd to Paricutín, 2250 m, [19°30′32″N, 102°09′18″W], 13 September 1961 , Moore Jr. & Bunting 8816 ( MEXU!); Panahuacaro , borde del camino, 2450 m, 12 August 1981 , Motte 315a ( MEXU!); Santa Clara del Cobre , Cerro San Miguel, 2500 m, [19°19′N, 102°23′W], 28 October 1988 , Pérez Calix 354 ( MEXU!); Nahuatzen , 7 km W de Arantepacua, 2450 m, [19°34′24″N, 102°00′45″W], 19 June 1988 , Ramos s.n. ( MEXU!); Cañada Tsintzungua , Capácuaro, 2300 m, [19°32′55″N, 102°03′05″W], 17 December 1971 , Rees s.n. (CAS!, MEXU!, MO!); 20 km al W de Ciudad Hidalgo, sobre la carretera a Morelia , 2250 m, [19°40′18″N, 100°43′54″W], 21 July 1964 , Rzedowski & de la Sota 18395 ( MEXU!); San Miguel del Monte , 15 km al SE de Morelia, 2200 m, [19°39′22″N, 101°07′15″W], 17 September 1967 ,

A TAXONOMIC REVISION OF SYMPLOCOS

Phytotaxa 264 (1) © 2016 Magnolia Press • 35 Rzedowski 25164 (DS!); 12 km al E de Uruapan, sobre la carretera a Pátzcuaro, 1750 m, [19°27′59″N, 101°56′45″W], 23 January 1977, Rzedowski 34595 ( MEXU!); Morelia , 2 km SW de Atécuaro, 2300 m, [19°33′55″N, 101°11′30″W] , 18 May 1986, Rzedowski 39704 ( MEXU!); Morelia , 4 km al S de San Miguel del Monte, 2400 m, [19°35′06″N, 101°08′04″W] , 16 September 1987, Rzedowski 44909 ( MEXU!); Tlalpujahua, cerca de Pichardo , 5 km al SW de San Francisco Los Reyes, 2650 m, [19°44′18″N, 100°16′20"] , 20 December 1987, Rzedowski 46218 ( MEXU!); Morelia, El Salitrillo, cerca de San Miguel del Monte , 2200 m , 26 November 1985, Santos M. 1067 ( MEXU!); Morelia , 3 km al SE de Atécuaro, 2400 m, [19°34′10″N, 101°10′31″W] , 16 November 1986, Santos M. 1898 ( MEXU!); 2 km al S de Villa Escalante, carretera a La Huacana , [19°23′01″N, 101°39′W] , 27 May 1979, Soto N. & Román de S. 1429 (CAS!, MEXU!, MO!); aprox. 12 km al NE de Coalcoman, camino a Aguililla , 1750 m, [18°49′33″N, 103°05′46″W] , 20 October 1979, Soto N. & Silva R. 1921 (CAS!, MEXU!, MO!); 2 km S de Villa Escalante, Santa Clara del Cobre; carretera Pátzcuaro–Ario de Rosales , [19°23′01″N, 101°39′W] , 23 October 1981, Soto N. 3411 ( MEXU!); 10 km al SE de Villa Madero, carretera a Nocupétaro , 2500 m, [19°21′10″N, 101°21′21″W] , 30 September 1982, Soto N. 4740 ( MEXU!); Ocampo , en La Laguna Verde, 2500 m, [19°37′N, 100°24′W] , 11 August 1984, Soto N. et al. 6539 (MEXU!, MO!, NY!); Mpio. Hidalgo, carretera Cd. Hidalgo – Mil Cumbres , 2 km al O de La Venta, 2200 m, [19°39′55″N, 100°43′14″W] , 8 March 1985, Soto N. & Aureoles C. 7360 (CAS!, MEXU!, NY!); desviación a Laguna Verde, carretera San Felipe de Los Alzati–Angangueo , 10 km al NE de San Felipe, 2200 m, [19°34′10″N, 100°20′34″W] , 7 July 1985, Soto N. et al. 9057 (CAS!, MEXU!, NY-2!); Pátzcuaro, 6.5 km S de Pátzcuaro, carretera a Santa Clara del Cobre , 2300 m, [19°27′16″N, 101°36′40″W] , 13 November 1999, Zamudio R. 11234 ( MEXU!). Morelos: Tepoztlán, La Pera, Coajomulco , 1600 m, [19°01′12″N, 99°08′14″W] , 6 September 1985, Castillo P. s.n. ( MEXU!); Tepoztlán, Mirador, Coajomulco , 2360 m, [19°01′17″N, 99°08′34″W] , 4 November 1985, Cervantes T. s.n. ( MEXU!); Cuernavaca, Colonia del Bosque , al N de la Barranca El Tecolote, 1800 m , 29 October 1987, Estrada L. 1866 ( MEXU!); Cuernavaca, Sierra de Morelos, 1925 m, [18°58′37″N, 99°16′19″W] , 30 October 1969, Hinton et al. 17438bis ( MEXU!); Tepoztlán, 5.5 km al E de Coajomulco, 500 m al N de la carretera, Sierra de Chichinautzin , 2450 m, [19°01′18″N, 99°09′34″W] , 8 January 1984, Ishiki I. 793 ( MEXU!); Tepoztlán, carretera (cuota) México – Cuernavaca , 20 km al NE de Cuernavaca, 2160 m, [19°00′48″N, 99°09′39″W] , 15 April 1976, Koch 7644 ( MO!); Valle de Tepeite , [18°59′32″N, 99°17′01″W] , August 1932, Lyonnet 976 (C!, CAS!, MEXU!, MO-3!, US!); Huitzilac , [19°01′42″N, 99°16′04″W] , December 1930, Lyonnet 1074 ( MEXU!); hacía Santo Domingo , [19°00′38″N, 99°03′43″W] , December 1942, Miranda 3857 (MEXU-2!); Barranca de Atzompan , cerca de Huitzilac, 2460 m , 17 October 1964, Palacios s.n. ( DS!); lava fields above Cuernavaca , 8000 ft, [19°00′16″N, 99°18′04″W] , 19 November 1902, Pringle 11013 (CAS!, F!, GH!, MO!, NY!, US!); mtn. side above Cuernavaca , 8000 ft, [19°01′N, 99°16′53″W] , 13 October 1900, Pringle 9125 (F!, GH!, MEXU!, MO!); Mpio. Tepoztlan, 5 km al ESE de Cuajomulco, autopista Mexico – Cuernavaca , 2400 m, [19°01′18″N, 99°09′50″W] , 17 April 1966, Rzedowski 22162 ( DS!); Tepoztlán, camino a Cuernavaca , 2400 m, [18°58′23″N, 99°09′35″W] , 29 January 1977, Schwabe 77520 (CAS!, MEXU!); Huitzilac , 2500 m, [19°01′42″N, 99°16′04″W] , 13 October 1975, Sousa S. & Delgado S. 5118 ( MEXU!); Cuernavaca, Colonia Bosque , oriente , 8 November 1969, Vázquez 2332 ( MEXU!). Nayarit: Tepic, W side of Cerro San Juan, 9.2 km S of rd from Tepic to Santa Cruz , 1520 m, [21°28′56″N, 104°59′50″W] , 4 October 1985, Bartholomew et al. 2667 (CAS!, GH!, MEXU!); N slope of Cerro San Juan along Hwy 54 to Jalcocotán , 1110 m, [21°31′06″N, 104°58′46″W] , 4 December 1968, Boutin & Brandt 2650 ( CAS!); Tepic, W slope of Volcán San Juan ascended by small rd at Km 6 from Tepic to Jalcocotlán , 1380 m, [21°28′10″N, 105°00′19″W] , 23 September 1979, Breedlove 44345 (CAS!, MEXU!, MO!); Tepic, W slope of Volcán San Juan along small rd at Km 6 on rd from Tepic to Jalcocotán , 900 m [sic] , 8 August 1980, Breedlove 45199 (CAS!, MEXU!, MO!); Tepic, 5 km de la desviación al rancho La Noria–El Cuartereño, Cerro San Juan , 1500 m, 21°29′N , 104°59′, 4 April 1994, Calzada 19187 (MEXU-2!, MO!); Tepic, 10 km al E de la desviación para el poblado El Cuareteño, Cerro San Juan , 1400 m, 21°28′15″N, 105°00′18″W , 6 April 1994, Calzada 19218 (MEXU!, MO!); 5 km SW de la carretera El Izote – V. Carranza, camino a El Cuartereño, Cerro San Juan, [21°30′15″N, 104°58′40″W], Flores F. et al. 4451 (MEXU!, MO!); El Terrero, 4.7 km del Terrero , por la brecha hacia Los Picachos , 2420 m, [20°57′N, 104°39′18″W] , 13 December 1996, González G. 48 ( MEXU!); Tepic, Cerro San Juan, camino a El Cuarenteño , 8 km W de Tepic, 1330 m, [21°28′55″N, 104°58′43″W] , 4 April 1992, González & Lorea 1337 ( MEXU!); Tepic, 12–17 km NW de Tepic, sobre la terracería a El Cuarenteño, 300 m adelante del El Izote , camino Tepic–Miramar , 21°30′N, 104°55′W , [21°30′19″N, 104°58′29″W] , 9 September 1985, Téllez V. 9145 (MEXU!, NY!); Tepic, Km 5–10, terracería a El Cuarenteño , que inicia 300–400 km [sic] al W de El Izote, [21°29′40″N,

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104°59′32″W], 6 October 1987, Téllez V. 10868 (MEXU!, MO!); Tepic , 13.5 km al SW de El Izote, brecha a El Cuarenteño, 1430 m , 21°29′N, 104°58′W, [21°27′N, 104°54′W], 5 December 1989, Tenorio L. et al. 15694 (MEXU!, MO!); Tepic , 2 km al E de La Hacienda de la Noria, brecha a Tepic, 2200 m , [21°28′55″N, 104°58′43″W], 23 March 1989, Tenorio L. et al. 15694 (MEXU!, MO!); Tepic , 13.5 km al SW de El Izote, brecha al Cuarenteño, 1430 m , 21°29′N, 104°58′W, 5 December 1989, Tenorio L. et al. 16939 ( MO!). Oaxaca : near Rancho La Cienaga , 11 February 1945, Alexander 730 ( NY!); La Soledad (near Mitla ), 9 February 1966, Ernst 2580 ( MEXU!); without precise locality, Karwinski s.n. (photo ex B at US!); 9.6 km al SE del Cerro de Vidrio carretera Oaxaca– Puerto Escondido, 1850 m , [16°12′53″N, 97°08′10″W], 1 August 1984, Torres C. & C. Martínez 5821 (F! MEXU!). State unknown: without precise locality, Sessé & Mociño 3607 (F!), Sessé & Mociño 3610 (F!), Sessé & Mociño 3611 (F!) .

Figures

FIGURE 2. Flowers and fruits of some species of Mexican and Central American Symplocos. a. S. serrulata. b. S. speciosa. c. S. striata. d. S. tribracteolata. e. S. austromexicana. f. S. citrea. g. S. pycnantha. h. S. coccinea. i. S. serrulata. j. S. hartwegii.

FIGURE 10. Geographic distribution of Symplocos breedlovei, S. citrea, and S. coccinea.