NUMBER 20 MUNN-ESTRADA: FLORA OF THE SIERRA MAZATECA OF OAXACA, MEXICO 25

CONTRIBUTION TO THE FLORISTIC KNOWLEDGE OF THE SIERRA MAZATECA OF OAXACA,MEXICO

Diana Xochitl Munn-Estrada Harvard Museums of Science & Culture, 26 Oxford St., Cambridge, Massachusetts 02138 Email: [email protected] Abstract: The Sierra Mazateca is located in the northern mountainous region of Oaxaca, Mexico, between the Valley of Tehuaca´n-Cuicatla´n and the Gulf Coastal Plains of Veracruz. It is part of the more extensive Sierra Madre de Oaxaca, a priority region for biological research and conservation efforts because of its high levels of biodiversity. A floristic study was conducted in the highlands of the Sierra Mazateca (at altitudes of ca. 1,000–2,750 m) between September 1999 and April 2002, with the objective of producing an inventory of the vascular plants found in this region. Cloud forests are the predominant vegetation type in the highland areas, but due to widespread changes in land use, these are found in different levels of succession. This contribution presents a general description of the sampled area and a checklist of the vascular flora collected during this study that includes 648 species distributed among 136 families and 389 genera. The five most species-rich angiosperm families found in the region are: Asteraceae, Orchidaceae, Rubiaceae, Melastomataceae, and Piperaceae, while the largest fern family is Polypodiaceae.

Resumen: La Sierra Mazateca se ubica en el noreste de Oaxaca, Mexico,´ entre el Valle de Tehuaca´n-Cuicatla´n y la Planicie Costera del Golfo de Mexico.´ La region´ forma parte de una ma´s extensa, la Sierra Madre de Oaxaca, que por su alta biodiversidad es considerada como prioritaria para la investigacion´ biologica´ y la conservacion.´ Se realizo´ un estudio en la Sierra Mazateca (a alturas de ca. 1,000–2,750 m) entre septiembre de 1999 y abril del 2002, que tuvo como objetivo compilar un inventario de las plantas vasculares de esta zona. El bosque mesofilo´ es el tipo de vegetacion´ predominante en esta a´rea, pero a causa de cambios extensos en el uso de suelo, esta vegetacion´ se encuentra en diferentes etapas de sucesion.´ Esta contribucion´ presenta una descripcion´ general del a´rea muestreada y un cata´logo de la flora vascular de la region´ colectada durante el presente estudio que incluye 648 especies, distribuidas en 136 familias y 389 generos.´ Las cinco familias de angiospermas ma´s abundantes en la region´ en terminos´ de su diversidad de especies son: Asteraceae, Orchidaceae, Rubiaceae, Melastomataceae y Piperaceae. La familia de helechos con la mayor diversidad es Polypodiaceae.

Keywords: Sierra Mazateca, Oaxaca, cloud forests

In Mexico, cloud forests (bosques where precipitation levels range from 1,000– mesofilos´ de montan˜a, bosques de neblina or 3,000 mm, and the average annual temper- bosques humedos´ de montan˜a) occupy 0.6– ature is 12–238C (53–738F). They typically 1% of the country’s territory and harbor have a constant cloud cover and are approximately 10% of its known plant characterized by high levels of humidity species (Rzedowski, 1991; Villasen˜or, (Torres-Colın,´ 2004; Villasen˜or, 2010; Gual- 2010). Human activities have reduced these Dıaz´ & Rendon-Correa,´ 2014). The states of cloud forests to mere fragments, making Chiapas, Oaxaca, and Veracruz have the them highly vulnerable and a priority for largest areas of cloud forests (Villasen˜or, conservation (Rzedowski, 1991, 1996; CON- 2010). The Sierra Madre de Oaxaca has an ABIO 2010; Ponce-Reyes et al., 2012; Gual- extensive area of cloud forests that has been Dıaz´ & Rendon-Correa,´ 2014). Cloud forests identified as a priority region for biological in Mexico are generally found in mountain- research and conservation actions (Arriaga ous regions, at elevations of 1,000–2,500 m, et al., 2000; Ponce-Reyes et al., 2012). While

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FIG. 1. A map of the study areas.

several floristic studies have been conducted The region encompasses an altitudinal range in this region, there are still areas that of 250–2,750 m (800–9,000 ft). Generally, its remain under collected. Such was the case of rugged topography is characterized by steep the Sierra Mazateca, located in northeastern slopes and deep ravines, although in some Oaxaca. The objective of the present study is areas there are small open valleys. The to contribute to the knowledge and docu- inhabitants of the Sierra Mazateca make a mentation of the vascular plant diversity distinction between two zones within the found in the highlands of the Sierra region, based on climatic differences: a tierra Mazateca, at altitudes of 1,000–2,750 m fr´ıa or Mazateca Alta (high-altitude areas), (3,250–9,000 ft). where the climate is generally temperate; and The Sierra Mazateca is located between a tierra caliente or Mazateca Baja (low- 188000–188220 N latitude, and 968300–978150 altitude areas), where it is generally much W longitude and occupies approximately warmer. The altitudinal limits of these areas 1,050 km2 (105,000 ha). Its northern limit is are subjective, but the transition zone the Petlapa River (Fig. 1), which separates it between lowland and highland areas gener- from the Sierra de Zongolica of Puebla- ally begins at about 1,000 m. The average Veracruz, and its southern limit is the annual temperature in the highlands is 16– canyon of the Santo Domingo River, which 228 C (62–728 F), and the average annual separates it from the southern Cuicatec precipitation is 1,498–4,942 mm, making it region and Sierra de la Chinantla (Fig. 1). one of the highest-precipitation regions in NUMBER 20 MUNN-ESTRADA: FLORA OF THE SIERRA MAZATECA OF OAXACA, MEXICO 27

Mexico (INEGI, 1983; Servicio Mete- to 55 Ma (Johnston & Stephen, 2004). The reologico´ Nacional, 2017). There are two uplift of the Sierra de Jua´rez, including the annual seasons in the region: a rainy season Sierra Mazateca, is thought to have begun 16 from June to October, and a dry season from Ma, in the middle Miocene (Centeno- November to May. Different types of soils Garcıa,´ 2004). The largest structures within are found in the highlands, including the Sierra de Jua´rez are thrust faults overlain umbrisols, luvisols, cambisols, acrisols, reg- by different rock types. Three of them are osols and phaezems (INEGI, 2007). associated with the Sierra Mazateca: (1) the In terms of its substrate composition, Cuicateco Fault in the western part of the the Sierra Mazateca can be divided into a sierra, near Teotitla´n de Flores Magon,´ western noncarbonate rock area composed characterized by rocks of the Cuicatla´n of allochthonous Jurassic rocks (sandstones Complex consisting of metavolcanics, schists and shales), and an eastern area composed (Charleston, 1980) and lens of serpentines of Cretaceous karstic limestone. The divid- (Carfantan, 1981; Campa & Coney, 1983); ing line between the two areas is east of the (2) the Huautla Fault in the central part of town of Huautla de Jimenez´ and runs in a the sierra, east of Huautla de Jimenez,´ N–SW direction from the village of San characterized by allochthonous Jurassic Andres´ to the Santo Domingo River. In the sandstones, shales, mudstones, and thin western noncarbonate areas, limestone out- bedded limestones (Moreno, 1980); and crops sometimes also occur. These outcrops (3) the Cerro Rabon´ Fault in the eastern are Lower Cretaceous limestones that have part of Sierra, characterized by coraline been exposed because of erosion through the limestones from the Lower to Middle thrust sheet caused by the Petlapa River Cretaceous (Smith, 1994; 2002) of the (Smith, 1994). In the karstic part of the Cordoba Platform (Gonza´lez, 1976). Sierra Mazateca there are many large cave The Sierra Mazateca is located within systems. One of them is located just east of the Papaloapan River basin, the second most Huautla de Jimenez,´ in the area where the important fluvial system in Mexico—after region’s two different rock substrates come the Grijalva-Usumacinta system— in terms into contact. This cave system, known as of its streamflow (Pereyra-Dıaz´ et al., 2010; Sistema Huautla, is 78.3 km (256,890 ft) CONAGUA, 2016). Within the region, the long and 1,560 m (5,118 ft) deep. It is the two largest rivers are: 1) the Santo Domingo deepest known cave in the Western Hemi- River, in the southern part of the sierra, sphere and the ninth deepest cave in the which eventually becomes the Papaloapan world. This cave system has been explored River and discharges into the Gulf of and mapped since the 1960s, but a new long- Mexico, and 2) the Petlapa River, in the term expedition project, Proyecto Espe- northern part of the sierra, which discharges leologico´ Sistema Huautla (PESH), is work- into the Miguel Alema´n dam in the eastern ing to extend its known length and depth lowlands. Throughout the region there are (Steele, 2017). perennial and intermittent streams. In the The geological history of the Sierra western part of the Sierra Mazateca, where Mazateca, like that of south central Mexico the substrate is composed of mostly imper- in general, is complex. The region belongs to meable noncarbonate rocks, several springs the Sierra de Jua´rez Geological Subprovince, and waterfalls are present. However, in the characterized by flat-lying, NE-directed dry season, their flow decreases dramatically overthrusts and fold-structures that have and some even disappear completely. In the complex internal structures. Geological eastern karstic part of the sierra, there are no studies indicate that the northern mountain significant surface streams, as all the rain- range of Oaxaca was formed by folding and water percolates into the karst. In the Cerro faulting processes during the Laramide Rabon´ massif area, the water collects near Orogeny that took place from the late the base level creating a number of subter- Cretaceous to Paleocene, approximately 80 ranean rivers (Bitterli & Jeanin, 1996). 28 LUNDELLIA DECEMBER, 2017

In the western part of the Sierra cultivation was the main source of work and Mazateca there is a large mountain range, income for Mazatec people. However, due to known as the Sierra de los Frailes, that lies the sharp decline in the international coffee diagonally in a N–SE direction. In the prices of this crop in the 1990s, most of the central part of the region there is a major plantations have been abandoned, and gorge formed by a tributary of the Petlapa current coffee production is primarily for River that runs northward from a locality local consumption. known as Puente de Fierro. Maps refer to it The oldest reference to the highland as the Puente de Fierro River (INEGI, 2015) vegetation of the Sierra Mazateca is found in although locals rarely refer to it by a specific the Relacion´ de Teotitla´n, a 1581 description name. In the eastern part of the sierra, the of the town of Teotitla´n de Flores Magon´ karstic massif known as Cerro Rabon´ , rises (also known as Teotitla´n del Camino) and prominently from the lowlands. The SE its adjacent areas. This is part of a series of portion of this massif has a spectacular rock documents written in the 16th century to cliff that rises from the ground almost inform King Phillip II about the character- vertically for at least 1,000 m (3,300 ft). istics of the territories in New Spain. The Smaller, exposed rock cliffs are frequent in Relacion´ says that: the central portion of the region. Guauhtla [referring to present-day Like other areas of Oaxaca, the Sierra Huautla de Jimenez]´ is cold and Mazateca has a rich cultural history. Its humid, and it rains there most of population is composed of indigenous the year. . .it has few grasses, as most Mazatec people with different degrees of of the land is covered with thick acculturation to modern Mexican society. forests. . .corn, beans, and squashes Their tonal language—Mazatec—is the third abound. . .it has large hills of pines, most widely spoken in the state of Oaxaca cedars, cypresses, oaks, and madro- after Zapotec and Mixtec. Mazatecs repre- n˜os. . .[the people] live from agri- sent the ninth largest indigenous group in culture, hunting, and the sale of Mexico, based on the number of individuals ocote [pine wood used for making who speak Mazatec (INEGI, 2010). Nahuatl torches and as fuel wood] (Acun˜a, and Mixtec communities are also found in 1984, pp. 206–207). the region. Some of the earliest evidence of In January 1906, Mexican geologist human settlements in the Sierra Mazateca is Manuel Villada visited the region to explore found in tombs, archaeological sites, and and document the cave of ‘‘Nindo-Da-G´ e´’’ caves scattered throughout the region. Based (Mazatec for: Cerro del Agua Crecida or on the artifacts found in these sites, there is Mountain of High Water), located in the evidence of habitation going back to 500– central area of the Sierra Mazateca, near the 750 CE (Hapka & Rouvinez, 1997; Munn, town of San Antonio Eloxochitla´n. In 2014), although further studies are required addition to presenting information about to establish a detailed chronology of the pre- the geology of the area, he also provided Hispanic history of the region. While many brief descriptions of the topography and residents of the Sierra Mazateca live in the weather, and he identified some of the most region’s large towns and urban centers ‘‘notable’’ plants he saw during his visit. He where there is access to a wide selection of began his expedition in the town of goods and services, a substantial number live ‘‘Teotitla´n del Camino’’, and as he made in smaller, rural communities where they his way into the sierra, he traveled through grow different crops including maize, beans, the Sierra de Los Frailes, which he described squash, and a variety of edible greens, and as follows: raise livestock including chickens, turkeys, As one advances, an interminable pigs, and goats. The Sierra Mazateca was, series of eminent peaks appear, of until recently, a major coffee-growing region different altitudes and shapes, sim- in Oaxaca. Since the early 1900s, coffee ulating the waves of an agitated NUMBER 20 MUNN-ESTRADA: FLORA OF THE SIERRA MAZATECA OF OAXACA, MEXICO 29

ocean. After the small town of San occidentalis L., or Alamo. . .I will Bernardino, one arrives at the enumerate some of the species highest point of the sierra. . .known found among the shrubs that as ‘‘Cumbre de los frailes’’ which emerge from the stream, and that has an altitude of approximately are also found on the mountain 2,470 m. Unfortunately, we made sides. The one that stands out above the trip on a bad day. . .a sky dark as all others is known in horticulture lead hovered over our heads. . .an by the names of Monte de oro or impetuous wind was blowing in- Pluma de oro, because of the color cessantly. . .layers of ice covered the of its flowers, which rise in tight tree tops. . .the temperature was two racemes: it is Jacobinia aurea of or three degrees below zero. . .mak- Hemsl. in the beautiful Acanthaceae ing our stay in the area very family. . .Aphelandra schiedeana Ch. painful. . .so we hurried our depar- y Sch. is another representative of ture. . .(Villada, 1906, pp. 487–488). the same family that lives here, but He then described the plants he saw in not in the water. Of the Gesneriad the highest elevations of the region and in family, there is the Isoloma deppea- the area near the cave he explored: na. . .of the Bignon family there is The always magnificent forests of Tecoma stans Juss., or Nextamax- oaks and conifers crown the highest ochitl. . .of the Verbenaceae family altitudes, although they have been there is the Lantana camara L. so diminished by immoderate cutting. well-known in our gardens by the Of the first [oaks] I will only same genus name. The most notable mention Quercus repanda K. in of the legumes is Cassia multiflora H.B. . .On the branches of the sec- Mart. and Gal., or Retama, which is ond [conifers] grow a few false another ornamental plant, with parasites, such as that called flowers disposed in racemes of an Soluche or Tillandsia recurvata L., intense yellow. Of Iresine celosioides as well as others of the same genus. L., in the Amaranthaceae, of Peper- Sedum dendroideum Moc. y Ses., omia edule L., in the Piperaceae, and Penstemones, Salvias, Senecios, etc., of a beautiful terrestrial orchid in and various grasses are interspersed the genus Laelia, I conserve vivid at the base of the trees, beautifying memories of my peregrination those places with their showy bou- through those mountains...It quets of red, blue, and yellow would be strange not to mention flowers (Villada, 1906, p. 503). an interesting group of trees that are . . .in the canyon of Nindo-Da-G´ e,´ typically found in certain regions of which has a warmer and humid the country, such as the one we are climate...there is a noticeable currently discussing. I am referring change in the aspect of the flora, to the Amates or Higuerones, trees which to a certain degree becomes that produce adventitious roots in exuberant. I will first mention a their branches, which descend ver- voluptuous tree of short stature tically until they embed themselves with elegant panicles of white flow- in the ground, and which from their ers. . .that grows along the edges of exterior aspect appear to be other the stream at the entrance to the trunks. I was only able to ascertain canyon. . .It is Saurauja villosa... the presence of one species, yet known as Pipicho. . .Another tree unidentified, that has a certain of much greater stature and corpu- affinity with Ficus padiaefolia K. in lence than the latter, and which is H.B, known as Cozahui- also a lover of water, is Platanus que. . .Because of its beautiful fo- 30 LUNDELLIA DECEMBER, 2017

liage it is a truly ornamental tree, of so poorly preserved that Schultes was unable medium, elliptic, thin, and some- to identify them, but the news that they what rigid leaves, a bit lustrous and existed sparked his interest so much that he of a pleasant green; that when headed to Huautla de Jimenez´ to investigate shaken by the wind, produces a the matter personally. During his stay in the strong noise that stands out from Sierra Mazateca, Schultes confirmed the use the others of its kind (Villada 1906, of psychoactive mushrooms by the Maza- pp. 503–504). tecs, but he erroneously identified them as a Villada’s account is significant, not only species of Panaeolus (Schultes, 1939). Nev- because it is one of the earliest records of ertheless, his publications on this topic geological research in the Sierra Mazateca, prompted others to visit the Sierra Mazateca, but also because it provides an indication and in the late 1950s, French mycologist that the region’s forests were already being Roger Heim properly identified the mush- impacted by wood harvesting activities. rooms as a species in the genus Psilocybe In the late 1930s, Richard Evans (Guzma´n, 2008). To this day, the region is Schultes conducted a botanical study in the famous worldwide for these mushrooms Sierra Mazateca as part of his Ph.D. research which have attracted both Mexicans and at Harvard University. In his dissertation foreigners interested in consuming them for (Schultes, 1941), entitled The Economic either medicinal or recreational use. Aspects of the Flora of Northeastern Oaxaca, As the title of Schultes’ thesis implies, Schultes wrote: his research focused on economically useful To be sure, the survival of many plants. Therefore, many of the species he culture-traits was the most con- includes in his work are cultivated plants. vincing argument in favor of inves- For the Mazatec region, Schultes listed 164 tigating north-eastern Oaxaca species of vascular plants (130 dicots, 26 ethnobotanically. But it was not monocots, 2 conifers, and 6 ferns) that he the only argument. A glance at any found to be of economic importance flora of southern Mexico or at any (Schultes, 1941). monograph which includes species In 1993, the Sociedad para el Estudio de from southern Mexico reveals at los Recursos Bioticos´ de Oaxaca (SERBO) once that the mountainous districts wrote a brief report on the botanical of the northeastern part of Oaxaca observations their team made on a visit to are of peculiar botanical interest. the plateau of the Cerro Rabon´ massif. The Floristically, this region is the report has not been published by SERBO, richest part of the state, and it is but it was used to write a brief description of probably more varied than any the Cerro Rabon´ forests for a speleological other area of equal size in Mexico publication (Bitterli et al., 1996). In the (Schultes, 1941: xiii). latter, the authors note the presence of One of the most important culture- Cupressus lusitanica var. benthamii (Endl.) traits that attracted Schultes’ attention to Carriere` and Oreomunnea mexicana northeastern Oaxaca was the use of psycho- (Standl.) J.F.Leroy in the region, two species active mushrooms by Mazatec people. Early with limited distributions in Mexico. colonial documents noted the use of psy- Oreommunea is a relict species from the choactive mushrooms by the Aztecs as part Miocene, and its presence points to the of their religious and healing traditions, but ancient origin of the region’s cloud forests this practice subsequently went unreported (Rzedowski & Palacios-Cha´vez, 1977; Pala- until the 1930s, when it became evident that cios-Cha´vez & Rzedowski, 1993; Rzedowski, some communities in Oaxaca were still 1996, Herrera et al., 2014). using fungi as part of ritual ceremonies. In In 1994, Carlos Ruız-Jim´ enez´ made a 1937, Schultes received specimens of these structural analysis of the vegetation found in fungi for identification. The specimens were a portion of the Sierra de los Frailes as part NUMBER 20 MUNN-ESTRADA: FLORA OF THE SIERRA MAZATECA OF OAXACA, MEXICO 31

of his undergraduate studies at Mexico’s cation of vouchers. The exploration of the National Autonomous University. In his Sierra Mazateca focused on three study areas thesis, Ana´lisis Estructural del Bosque which were selected because they present Mesofilo´ de la Region de Huautla de Jimenez´ dense vegetation cover and have distinct (Oaxaca), Mexico´ , Ruız-Jim´ enez´ listed 178 geological, altitudinal, and landscape char- species (3 lycophytes, 25 ferns, 3 gymno- acteristics. The study areas selected were: the sperms, 147 angiosperms) for an area known Sierra de los Frailes, the canyon area as Puerto Soledad (Ruiz-Jimenez,´ 1995; between the towns of Huautla de Jimenez´ Ruiz-Jimenez´ et al., 2000). The present study and Chilchotla, and the Cerro Rabon´ plateau includes 75 of the species in Ruiz-Jimenez’´ (Fig. 1). The collection sites in these three work. However, a careful analysis of synon- areas are described in Appendix 1. ymy has not been made of the rest of the STUDY AREA NUMBER 1. SIERRA DE LOS species. FRAILES: This area is the westernmost moun- The biological diversity of the Tehua- tain range of the Sierra Mazateca, located in ca´n-Cuicatla´n Valley, located west of the the noncarbonate portion of the region (Fig. sierra, has been studied extensively. This 1). Its highest peak (and that of the region in mostly semi-arid region, encompassing ap- general) is the Cerro Pelon´ at 2,750 m (9,000 proximately 10,000 km2 (1,000,000 ha), ft). Five general localities (see Appendix 1, maintains 36 different plant communities Localities 1–5) were surveyed in this area, and its flora represents 10–11.4% of Mex- ranging in altitude from 2,020–2,750 m. The ico’s national plant diversity. In the region’s tree canopy in the Sierra de los Frailes higher elevations there are oak forests, pine- generally reaches heights of 20–30 m, as oak forests, and to a smaller extent, cloud observed in this study. forests, particularly in the southernmost part The Sierra de los Frailes is readily of the valley (Da´vila et al., 2002; Ruiz- accessible by a major two-lane road— Jimenez,´ 2003; Vaiente-Banuet et al. 2009; Mexican Federal Highway 182—that pro- Canseco-Ma´rquez & Gutierrez-May´ en,´ 2010; vides access to the Sierra Mazateca, and by SEMARNAT, 2013). other unpaved roads. If approached from Beyond these studies, botanists from Teotitla´n de Flores Magon´ in the Tehuaca´n various Mexican and international institu- Valley, the Sierra de los Frailes will be tions have made collections in the Sierra encountered after making an ascent of Mazateca, but no concerted efforts, apart approximately 1,000 m (3,280 ft) from that from those already outlined, have been made town. As the altitude increases, the climate to study the flora of the Sierra Mazateca. becomes noticeably cooler and the vegeta- I am a native of the Sierra Mazateca and tion more lush. As one approaches 2,000 m undertook this study to better document its (6,561 ft), the landscape changes dramati- floristic diversity and to provide baseline cally and epiphyte-covered trees of 15–20 m data that could support the development of (50–65 ft) in height become a common educational materials about the region’s sight. This change in landscape and climatic biodiversity. The present study does not conditions marks the beginning of the Sierra attempt to synthesize previous work by de los Frailes. Continuing along highway 182 others in the area; rather, it aims to toward the community of Plan de Guada- contribute to the knowledge of the region lupe, at about 2,300 m (7,545 ft), one finds by enumerating the plants collected and extensive stands of mature cloud forests in identified during the project. the north-facing slopes of the area. Some of the trees present in these forests are: Quercus METHODS corrugata, Q. ocoteifolia, Q. scytophylla, Podocarpus matudae, Clethra hartwegii, This study was conducted in two phases: Cleyera integrifolia, Styrax ramirezii, Drimys the first focused on the collection of granadensis var. mexicana, Ocotea betazensis, specimens, and the second on the identifi- Cinnamomum effusum, Osmanthus ameri- 32 LUNDELLIA DECEMBER, 2017

depressa. A few herbs such as Osbertia stolonifera, Ottoa oenanthoides, Lycopodium clavatum, and L. thyoides, as well as a Calamagrostis grass and a terrestrial orchid can be found growing among the stunted shrubs. Whether this vegetation is natural or a result of human activities is unknown. Toward the southeastern part of the Sierra de los Frailes, along the road to Agua Duende, one can find large specimens of Clethra lanata, as well as Myrsine juergense- FIG. 2. Aspect of Study Area 1. View from nii, Pinus patula, Ternstroemia tepezapote, ´ Cerro Pelon, Sierra de los Frailes. Several species and Citharexylum mocinnoi. Further south, of oaks are common in this area. Photo by Diana near the communities of Agua de Cerro and X. Munn-Estrada. San Pedro de los Encinos (between Agua Duende and Soyaltitla), an excursion into canus, Phyllonoma laticuspis, Prunus brachy- the forested slopes leads to similar forests botrya, and Ternstroemia tepezapote, among such as those in the previous site, but others. These forests have a canopy of 20–30 populated with Quercus aff. corrugata, Oreo- m(65–100ft)anddonotpresenta panax liebmannii, Chiococca sp., and many significant understory except for the areas specimens of Cyathea fulva, among others. where they may have been disturbed by An interesting and quite particular aspect of human activities, or where there are gaps in the areas near the community of San Pedro the vegetation due to natural events (i.e., de los Encinos is the presence of dense falling of trees). In the fringes of the forests stands of oaks—5–7 m tall—with very and along the highway, many Asteraceae narrow trunks. The inhabitants of San Pedro herbs can be found including Dahlia aus- de los Encinos (named after the oak trees tralis and Senecio callosus. described, encino being the Spanish term for In comparison to the other areas studied oak) have indicated that these stands grew in the Mazateca region, these forests present after a major fire devastated parts of the area a much lower degree of human disturbance. many years ago (the exact date is unknown). At the Plan de Guadalupe one can The vegetation found in the Sierra de los continue along highway 182 toward Huautla Frailes study area is predominantly ever- de Jimenez,´ ascend the Cerro Pelon´ by foot, green, although some of its elements are or travel to the southeastern portion of the deciduous (e.g., Alnus). study area. STUDY AREA 2. HUAUTLA–CHILCHOTLA: On the slopes of the Cerro Pelon´ one This area is located in the central part of can find: Arbutus xalapensis, Clethra hartwe- the Sierra Mazateca, between the noncar- gii, Cornus excelsa, Ocotea betazensis, Quer- bonate and the karstic portions of the cus ocoteifolia, Ugni myricoides,and region. Its geographical reference point is a Weinmannia pinnata, among other trees. large river gorge that begins at a location In the higher parts of the Cerro Pelon,´ known as Puente de Fierro—near the town starting at about 2,600 m (8530 ft), trees on of Huautla de Jimenez—and´ continues the slopes exposed to the northeastern north to the town of Chilchotla. This area winds, such as Clethra hartwegii, present presents mosaics of vegetation with different many morphological deformations in their degrees of human disturbance, and different trunks and branches. At the very top of the successional stages. Eleven general localities mountain there are no trees, only dwarf (Appendix 1, Localities 6–16) were surveyed shrubs measuring 40–70 cm (1.3–2.3 ft), in this area, with altitudes ranging from including Gaultheria erecta, Pernettya pros- 1,069–2,151 m. Because of its large extension trata, Vaccinium confertum, and Quercus and distinct physical characteristics, the river NUMBER 20 MUNN-ESTRADA: FLORA OF THE SIERRA MAZATECA OF OAXACA, MEXICO 33

gorge can best be described if divided into the exposed rock faces in the road cuts of the three sub-parts: (1) the areas immediately road from Puente de Fierro to San Antonio adjacent to the river, (2) the mountain in the Eloxochitla´n. These rocks generally harbor western part of the gorge (known among many ferns, a wide variety of herbs, and Mazatecs as Nindo-Ntahe, or Mountain of small shrubs. At higher altitudes (ca.1,430 the High Water, located near the town of m; 4,691 ft), an intermittent stream that San Antonio Eloxochitla´n, and (3) the travels from San Antonio toward the river mountain range in the eastern part of the below supports a riparian forest with gorge, which is the location of many large Platanus mexicana var. mexicana, Ficus sp., villages and communities, including Huautla Deppea grandiflora, and Cornutia pyramida- de Jimenez´ in the south, and Santa Marıa´ ta. Chilchotla in the north. In the winter, many trees on this Starting from the Puente de Fierro, and mountain range lose their leaves, making following the river—a tributary of the the vegetation appear quite different from Petlapa River—to the north, the vegetation that found in the Sierra de los Frailes, where is best described as riparian forest with the vegetation tends to be nearly evergreen. deciduous elements, interspersed with sec- The presence of Heliocarpus on the slopes of ondary vegetation. Among the trees found the mountain range is an indicator of the along the river and its adjacent areas are: secondary nature of the vegetation. Ulmus mexicana, Platanus mexicana var. The mountain range east of the gorge mexicana, Cinnamomum effusum, Ficus sp., also has highly disturbed areas, which is Glossostipula concinna, Litsea glaucescens, expected given that it is the most populated Ocotea bernoulliana, Damburneya salicifolia, part of the Sierra Mazateca. Wherever one Mollinedia viridiflora, Oecopetalum mexica- goes in this mountain range, one is sure to num, Quercus sartorii, Saurauia aff. villosa, find remnants of coffee plantations inter- Symplocos limoncillo, Deppea erythrorhiza, spersed with cloud forest. Therefore, its Citharexylum mocinnoi, Juglans aff. mollis, vegetation is best described as secondary. Capparidastrum mollicellum, Telanthophora Altitudes in this area range from 1,200– grandifolia, and Liquidambar styraciflua in 2,150 m (3,937–7,050 ft.). The highest peak the nearby slopes. In the understory of the is the Cerro de la Adoracion,´ located riparian forest, Chamaedorea palms are southeast of Huautla de Jimenez´ at 2,150 common, as well as many begonias and m (7,053 ft). ferns such as Llavea cordifolia and Niphi- Trees found in this range include: dium crassifolium. Among the epiphytes, Liquidambar styraciflua, Inga vera, Alchornea species of Peperomia abound, as well as latifolia, Myrsine coriacea, Quercus sartorii, many orchids. Persea americana, Solanum aphyodendron, The slopes of the Nindo-Ntahe, west of Frangula capreifolia, Miconia sylvatica, Bud- the river (1,060–1,450 m; 3,477–4,757 ft), dleja americana, Berberis gracilis, and Clusia present mosaics of disturbed cloud forests, sp. In the southern parts of this area, near areas of subsistence agriculture, and second- the Cerro de la Adoracion´ , the following ary vegetation. Despite being highly dis- species were found: Clethra hartwegii, Mico- turbed, one still finds many large specimens nia hemenostigma, Phyllonoma laticuspis, of oaks in this area including: Quercus Alnus acuminata subsp. arguta, Vaccinium candicans, Q. eugeniifolia, Q. lancifolia, Q. leucanthum, and Solanum nigricans, among ocoteifolia, and Q. polymorpha, and other others. trees such as Turpinia sp., Wimmeria To the northeast of Huautla, towards bartletti, Hedyosmum mexicanum, Inga acro- the town of Santa Cruz de Jua´rez, one can cephala, Liquidambar styraciflua, Ocotea encounter: Dendropanax arboreus, Garrya bernoulliana, Palicourea padifolia, and the aff. laurifolia, Persea americana, Prunus tree fern Sphaeropteris horrida. An interest- brachybotrya, Saurauia leucocarpa, Solanum ing aspect of this area is the flora growing in nigricans, Vaccinium leucanthum, Deppea 34 LUNDELLIA DECEMBER, 2017

has a rugged landscape composed of cone karst and large dolines. The highest peak in the region is Cerro Caballero at 2,150 m (7,054 ft). This area is the most difficult to access, as there are no roads that go close to the plateau. One can enter the area by foot from three localities: the town of San Jose´ Tenango, the community of Rancho Aven- dan˜o which is east of Cerro Central, and the town of Jalapa de Dıaz.´ The altitudinal range of the five localities (Appendix 1, Localities FIG. 3. Study Area 3. View of Cerro Rabon´ 17–21) explored in the northeastern and from Jalapa de Dıaz´ where a member of the southeastern portions of the plateau is walnut family is among the dominant trees. 1,100–1,645 m. Photo by Diana X. Munn-Estrada. Although the Cerro Rabon´ is difficult to access and has no surface streams because of grandiflora, Myriocarpa longipes, and Psycho- its karstic, porous substrate, its plateau is tria fruticetorum. quite populated, contrary to the popular belief among many Mazatecs that no one Along the road toward Santa Marıa´ lives on this mountain. Chilchotla (1,070–1,230 m; 3,510–4,035 ft), The vegetation in the populated por- north of Puente de Fierro, the vegetation is tions of the plateau is generally disturbed, also mostly secondary. Some of the trees but it is well conserved in the less populated found here are: Persea schiedeana, Acalypha areas. The mature cloud forests found in the cf. longipes, Dendropanax arboreus, Juglans explored areas are very distinct in floristic mollis, Alchornea latifolia, Tonduzia long- composition from others found in the Sierra ifolia, Wimmeria bartletti, and Coccoloba Mazateca, particularly because one of their hirtella. The black limestone outcrops that dominant elements is Oreomunnea mexica- border the road, harbor many locality- na, a large tree not found anywhere else in restricted species and many ferns and this sierra. The specimens of Oreomunnea gesneriads. observed were 25–30 m (82–100 ft) in East of Marıa´ Luisa, and near the height, and grew in close association with community of Agua de Gancho, there are a trees of Sloanea cruenta, also reaching the few patches of forest growing on a limestone same heights. Other species living among the substrate that contain: Ardisia verapazensis, Oreomunnea trees are: Weinmannia pinnata, A. liebmannii, Cojoba arborea, Glossostipula Pinus sp., Ardisia liebmannii, Randia matu- concinna, Psychotria fruticetorum, Sommera dae, Psychotria galeottiana, P. sarapiquiensis, arborescens, Deppea grandiflora, Hoffmannia Parathesis leptopa, and Sommera arborescens. nicotianifolia, Psychotria mexiae, Coccoloba In the understory, especially in areas with sp., many Chamaedorea palms, ferns, or- light gaps, numerous Chamaedorea species chids, and other epiphytes such as Marc- thrive, in conjunction with many species of gravia stonei. North of these forests, at an Acanthaceae, and other herbs. Epiphytic altitude of 1,700 m (5,577 ft) there are areas ferns and orchids are a common component that have been highly disturbed by fires and of these forests. Other species present in this consequently, their vegetation is secondary area but not as abundant as Oreomunnea in nature. forests are: Saurauia villosa, Clethra con- STUDY AREA 3. CERRO RABON´ : This area is zattiana, Fuchsia paniculata, Glossostipula located in the easternmost karstic front concinna, Myriocarpa longipes, Oreopanax range of the Sierra Mazateca. It is the xalapensis, and Arachnothryx heteranthera. plateau of the Cerro Rabon´ massif that rises Many begonias, ferns, mints, and asters west of the Miguel Alema´n dam. The plateau thrive in the open area of the limestone NUMBER 20 MUNN-ESTRADA: FLORA OF THE SIERRA MAZATECA OF OAXACA, MEXICO 35

trails found in the Cerro Rabon´ plateau. In sented following the classification of Smith the southern portion of the plateau one can et al. (2006) and Christenhusz et al. (2011), find: Clethra conzattiana, Ardisia liebmannii, which are congruent with each other except and Hedyosmum mexicanum. for the recognition of Nephrolepidaceae and Preliminary collecting expeditions to the Athyriaceae, which follows Christenhusz. study areas were carried out in September Gymnosperms are classified according to 1999 and the summer of 2000. Intensive Christenhusz et al. (2011). Angiosperms collecting began in March 2001 and contin- follow the classification system of the ued through April 2002. A short expedition Angiosperm Phylogeny Group IV (APG IV, to the region was also made in December 2016; Stevens, 2017). Scientific names were 2002. In total, 21 general localities distrib- confirmed in journals and database systems uted in the three study areas were selected, (The Plant List, 2013; Tropicos.org, 2017; and frequent expeditions were made to these Villasen˜or, 2010). areas (see Appendix 1). Collections were Species names are followed by a descrip- carried out with permits DOO.02.1463 and tion of the plant’s general habit, the altitu- SGDPA/DGVS/3312 issued by Mexico’s dinal range represented by the specimens Secretariat of Environment and Natural collected, and the localities where they were Resources (SEMARNAT, 2010), and addi- collected (see localities, Appendix 1), with tional permits were provided by local voucher numbers in parentheses. Species are authorities. The coordinates of localities terrestrial unless otherwise stated. The local- were recorded with a GPS system. Speci- ities of species recognized as endangered, mens were pressed in the field and dried in threatened, or protected by Mexican law are Huautla de Jimenez,´ using standard meth- not disclosed (SEMARNAT, 2010). ods (Bridson & Forman, 1992). All of the In Mexico, specimens were deposited at collections were made by the author with the the Instituto de Ecologıa,´ A.C. (XAL), and the assistance of local guides, colleagues, and National Autonomous University (MEXU). family. The focus, throughout the study, was In the US they were deposited at The on collecting fertile vouchers. The identifi- University of Texas at Austin (TEX) and the cation of specimens began in June 2002 at Smithsonian’s National Museum of Natural The University of Texas at Austin. Many History (US); there are additional partial sets specialists contributed to the final identifi- to be distributed. Vouchers sent to specialists cation of specimens (see Acknowledgments). for identification may be found in select SPECIES CHECKLIST: The species checklist herbaria in the US (see Acknowledgments). (Appendix 2) is based entirely on the There are several collections that remain collections made by the author and collab- unidentified even to the rank of family. As orators during the present study; collections new determinations are received, and when made by other collectors, whether or not a critical mass of newly identified material is cited in previous published works (Schultes, 1941; Ruiz-Jimenez,´ 1995; Ruiz-Jimenez´ et available, the author will publish an update al. 2000, 2012) are not included. The species of the checklist. checklist thus does not attempt to reflect current knowledge of the flora of the study RESULTS AND DISCUSSION areas but rather the contributions of the current study, the largest undertaken in the A total of 1,200 collections made by the Sierra Mazateca. Species are presented al- author are included in this study. These phabetically according to four major cate- collections represent 648 species distributed gories: Lycophytes, Ferns, Gymnosperms, among 136 families and 389 genera (Ap- and Angiosperms. Authors of plant names pendix 2). Twenty-nine of the species are are cited after Brummit and Powell (1992). only determined to genus, but are recog- The classification of lycophytes follows nized as distinct taxa and counted as Christenhusz et al. (2011). Ferns are pre- individual species. 36 LUNDELLIA DECEMBER, 2017

TABLE 1. Distribution of taxa according to tion of remaining material collected during vascular plant groups. this study as well as and further fieldwork in the areas mentioned above. In addition to Families Genera Species supporting a better understanding of cloud Lycophytes 239forests in Mexico, the relevance of this study Ferns 18 39 85 is its potential to contribute to the conser- Gymnosperms 444vation and sustainable management of Angiosperms 112 343 550 vegetation in the Sierra Mazateca, as well Total 136 389 648 as to the development of educational materials for the communities in this region.

In terms of the number of species they ACKNOWLEDGEMENTS contain, angiosperms represent the largest of the vascular plant groups (84%), followed by This study was carried out under the ferns (14%), lycophytes (1.5%), and gym- auspices of the Plant Biology Graduate nosperms (0.5%) (Table 1). Program of The University of Texas at Based on the records included in this Austin (UT-Austin) under the supervision paper, the five largest families of angio- of Dr. Jose´ Panero. In Mexico, the project sperms found in the Sierra Mazateca, in was sponsored by the Instituto de Ecologıa,´ terms of their number of genera and species, A.C. in Xalapa, Veracruz, under the super- are: Asteraceae, Orchidaceae, Rubiaceae, vision of Dr. Francisco Lorea. Without the Melastomataceae, and Piperaceae. The Poly- backing of Drs. Panero and Lorea, this podiaceae is the largest family of ferns. project could not have been carried out. Twenty-one of the recorded species are Key support was received from the late Dr. protected by Mexican law because they are Mario Sousa Sa´nchez and Rafael Torres- endangered or threatened with extinction, or Colın´ at the National Herbarium (MEXU) very close to becoming threatened (SEM- of the Universidad Nacional Autonoma´ de ARNAT, 2010). These species are noted, Mexico´ (UNAM) in Mexico City. The study without specific locality, in Appendix 2. was made possible by research grant U-028 The 648 species collected through this received from the Mexican Consejo Nacional study give a positive indication of the para el Uso y Conocimiento de la Biodiversi- biological richness found in the Sierra dad (CONABIO). Additional funding was Mazateca, but further research in herbaria obtained from the Tinker Foundation and exploration of the region will surely (through the Institute of Latin American increase the checklist included in this work. Studies at UT-Austin), the Linda Escobar The Sierra de los Frailes and the Cerro Fellowship for Tropical Studies (Plant Biol- Rabon´ plateau, in particular, merit further ogy Graduate Program at UT), and Bacardi study. &Cıa.´ Plant identifications were generously Apreliminaryreportofthisstudy provided by the following botanists (families available online (Lorea-Herna´ ndez & identified and herbaria affiliations in paren- Munn-Estrada, 2005) was one of the data theses): Pedro Acevedo (Sapindaceae; US), sets used in a study to determine the floristic Salvador Acosta (Acanthaceae; ENCB), affinities among Mexican cloud forests. The Frank Almeda (Melastomataceae, Symplo- findings indicate that the flora of the Sierra caceae; CAS), Paul Berry (Onagraceae; Mazateca has close affinities to other cloud MICH), Kathleen Burt-Utley (Begoniaceae; forests in Oaxaca including: Chinantla, USF), Lynn Clark (Poaceae; ISC), Thomas Cerro Salomon,´ Sierra de Jua´rez, Tiltepec, Croat (Araceae; MO), Thomas Daniel El Rincon´ and Santa Cruz Tepetotutla (Acanthaceae; CAS), Ricardo de Santiago (Ruiz-Jimenez´ et al., 2012). This author (Melastomataceae; FCME), Alfonso Delgado feels that any further analysis of the data Salinas (Fabaceae; MEXU), Robert Dressler should await completion of the identifica- (Orchidaceae; JBL), Adolfo Espejo (Mono- NUMBER 20 MUNN-ESTRADA: FLORA OF THE SIERRA MAZATECA OF OAXACA, MEXICO 37

cots; UAMIZ), Vicki Funk (Asteraceae; US), housed me in different communities, the Peter Fritsch (Styracaceae; CAS), Abisaı´ many individuals who guided me through Garcıa´ Mendoza (Agavaceae; MEXU), James their land, and my own family which Henrickson (Rosaceae; TEX), Hugh Iltis provided key contacts, and guidance in the (Capparaceae; WIS), Rolando Jimenez´ (Or- region, and financial support. chidaceae; AMO), Lawrence Kelly (Actini- daceae, Aristolochiaceae, Symplocaceae; LITERATURE CITED NY), Blanca Leon´ (Ferns; TEX), Ana Rosa Lopez´ Ferrari (Monocots; UAMIZ), Fran- Acun˜a, R. (ed.). 1984. Relaciones Geogra´ficas del siglo cisco Lorea (Lauraceae; XAL), David Lor- XVI: Antequera. Volumenes´ I & II. Universidad ence (Rubiaceae; PTBG), Lucio Lozada Nacional Autonoma´ de Mexico,´ Mexico City. (Apocynaceae/Asclepiadoideae; FCME), APG IV.2016.AnupdateoftheAngiosperm Phylogeny Group classification for the orders and James Luteyn (Ericaceae; NY), Andrew families of flowering plants: APG IV. Bot. J. Linn. McDonald (Convolvulaceae; UT-Rio Soc. 181: 1–20. Grande Valley), John Mickel (Ferns; NY), Arriaga, L., J.M. Espinoza, C. Aguilar, E. Martınez,´ L. Juan Carlos Montero (Solanaceae; MEXU), Gomez,´ and E. Loa (eds.). 2000. Regiones terrestres Mike Nee (Solanaceae; NY), Guy Nesom prioritarias de Mexico.´ Comision´ Nacional para el Conocimiento y Uso de la Biodiversidad, Mexico. (Asteraceae; TEX), Kevin Nixon (Fagaceae; http://www.conabio.gob.mx. BH), Jose´ Panero (Asteraceae; TEX), Her- Bitterli, T. and P. Jeanin. 1996. Hydrology. In: T. Bitterli milo Quero Rico (Arecaceae; MEXU), Jon (ed.). Proyecto Cerro Rabon´ 1990–1994, Oaxaca, Ricketson (Rubiaceae; MO), Lourdes Rico- Mexico. Speleo Projects, Basel, Switzerland, pp. 48–50. Arce (Fabaceae; K), Gerardo Salazar (Or- ——— and K. Meyers. 1996. Geologic Overview. In: T. Bitterli (ed.). Proyecto Cerro Rabon´ 1990–1994, chidaceae; MEXU), Elizabeth Skendzic (Poa- Oaxaca, Mexico. Speleo Projects, Basel, Switzerland, ceae; Kutztown University), Lawrence Skog pp. 40–41. (Gesneriaceae; US), Mario Sousa Sa´nchez Bridson, D. and L. Forman (eds.).1992.The (Fabaceae; MEXU), Charlotte Taylor (Ru- Herbarium Handbook. Royal Botanic Gardens, Kew. biaceae; MO), Rafael Torres-Colın´ (Faba- Brummit, R.K. & C.E. Powell (eds.). 1992. Author of ceae; MEXU), Leticia Torres-Colın´ Plant Names. Royal Botanic Gardens, Kew. Campa, M. F. and P. J. Coney. 1983. Tectono- (Fabaceae; MEXU), Billie Turner (Astera- stratigraphic terranes and mineral resource distri- ceae; TEX), Luz Marıa´ Villareal (Clethra- butions in Mexico. Can. J. Earth Sci. 20: 1040–1051. ceae; Universidad de Guadalajara), Grady Canseco-Ma´rquez, L. and G. Gutierrez-May´ en.´ 2010. Webster (Euphorbiaceae; DAV), Tom Anfibios y Reptiles del Valle de Tehuaca´n-Cuicatla´n. Wendt (various taxa; TEX), Justin Williams Comision´ Nacional para el Conocimiento y Uso de la Biodiversidad (CONABIO), Fundacion´ para la Reser- (Apocynaceae; Sam Houston State Univer- va de la Biosfera Cuicatla´n, A. C, y la Benemerita´ sity), and George Yatskievych (Ferns; TEX). Universidad Autonoma´ de Puebla, Mexico.´ Celia del Carmen Zun´ ˜iga Rıos´ Zertuche Carfantan, J. C. 1981. Evolucion´ estructural del sureste produced the map of the region, and Carlos de Mexico;´ paleogeografıa´ e historia tectonica´ de las E. Arroyo Cruz (CONABIO) provided key zonas internas mesozoicas. Univ. Natl. Auton. Mexico, Inst. Geol. Revista 5(2): 207–216. information on soils. The following individ- Centeno-Garcıa,´ E. 2004. Configuracion´ geologica´ del uals provided academic guidance and logis- estado. In: A.J. Garcıa-Mendoza,´ M.J. Ordon´ ˜ez y tics support: USA: Jose´ Panero, Thomas M. Briones-Salas (eds). Biodiversidad de Oaxaca. Wendt, Beryl Simpson, Billie Turner, Robert Instituto de Biologıa,´ UNAM-Fondo Oaxaquen˜o Nicholson, Gustavo Romero, Ernie Garza; para la Conservacion´ de la Naturaleza-World Wildlife Fund, Mexico, pp. 29–42. MEXICO: Francisco Lorea, Claudia Gallar- Charleston, S. 1980. Stratigraphy and tectonics of the do, Mario Sousa Sa´nchez, Rafael Torres- Rio Santo Domingo Area, State of Oaxaca, Mexico: Colın;´ SIERRA MAZATECA, OAXACA: 26th Congres Geologique International, v. I, Estrada-Pineda Family, Francisco Mendoza, Sections 1 a 5, Juillet, Resumes, 324 p. Jua´rez Family, Cerqueda Family, Renato Christenhusz, M.J.M, X.C. Zhang, and H. Schneider. 2011. A linear sequence of extant families and Garcıa.´ I thank the various municipal genera of lycophytes and ferns. Phytotaxa 19: 7–54 presidents and authorities who provided ———, J.L. Reveal, A. Farjon, M.F. Gardner, R.R. permits to work, the Mazatec families who Mill and M.W. Chase. 2011. A new classification 38 LUNDELLIA DECEMBER, 2017

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APPENDIX 1 Mature cloud forest presenting slight human disturbance (compared to other localities). COLLECTING LOCALITIES WITHIN THE THREE STUDY AREAS OF Locality 4: Agua de Cerro – San Pedro de los Encinos THE SIERRA MAZATECA Municipio: Mazatla´n Villa de Flores. Location: Between the communities of Agua de Cerro and San Pedro de los STUDY AREA NUMBER 1. SIERRA DE LOS FRAILES: Encinos (SE of Agua Duende, and before Soyaltitla); (Western portion of the Sierra Mazateca) accessible from the road that connects Plan de Guadalupe Locality 1: Puerto de la Soledad with Palo de Marca, and continues towards Huautla de Municipio: Teotitla´n de Flores Magon.´ Location: 29– Jimenez.´ Altitude: 2,020–2,345 m (6,627–7,693 ft). 32 km (18–20 mi) from Teotitla´n de Flores Magon,´ Vegetation: Mature cloud forest that presents a higher along highway Mex 182 to Huautla de Jimenez.´ degree of disturbance than that found in the previous Altitude: 2,300–2,340 m (7,545–7,677 ft) Vegetation: localities (mainly from wood–cutting for fuel). The Mature cloud forest presenting slight human forests of this locality have also been disturbed by major disturbance (compared to other localities). fires, but details about their last occurrence is unknown.

Locality 2: Plan de Guadalupe–Cerro Pelon´ Locality 5: San Juan la Union´ Zoquia´pan Municipio: San Jeronimo´ Tecoatl.´ Location: Plan de Municipio: San Lucas Zoquia´pan. Location: Areas Guadalupe; approximately 40 km (25 mi) from surrounding the community of San Juan la Union.´ ´ ´ Teotitlan de Flores Magon, along highway Mex 182 Accessible by the terracerıa´ from Huautla de Jimenez´ ´ to Huautla de Jimenez. Altitude: 2,242–2,742 m to Palo de Marca. Altitude: 2,183–2,333 m (7,162– (7,355–8,996 ft). Vegetation: Mature cloud forest 7,654 ft). Vegetation: Cloud forests highly disturbed by presenting slight human disturbance (compared to fire in the lower elevations, mature cloud forests in the other localities). higher elevations. Locality 3: Plan de Guadalupe – San Mart´ın Zoquiapan STUDY AREA 2. HUAUTLA–CHILCHOTLA:(Central Municipio: San Lucas Zoquia´pam. Location: Along the portion of the Sierra Mazateca) road from Plan de Guadalupe toward Agua Duende. Altitude: 2,167–2,259 m (7,109–7,411ft). Vegetation: Locality 6: Puente de Fierro 40 LUNDELLIA DECEMBER, 2017

Municipio: Huautla de Jimenez.´ Location: Areas Municipio: Huautla de Jimenez.´ Location: adjacent to the Puente de Fierro (approximately 8 Approximately 3–4 km north of Puente de Fierro, km (5 mi) from Huautla de Jimenez´ on the road to along the terracer´ıa to the town of Santa Marıa´ Teotitla´n de Flores Magon),´ alongside the Petlapa Chilchotla; west facing mountain slopes approaching River, and along the terracer´ıa to Santa Marıa´ the town of Aguacatitla. Altitude: 1,188–1,590 m Chilchotla. Altitude: 1,150–1,270 m (3,772– 4,166 ft). (3,897–5,216 ft). Vegetation: The area is highly Vegetation: Riparian forest along the river. disturbed by agricultural activities, but it still maintains elements of cloud forest vegetation. The Locality 7: Puente de Fierro – San Antonio Eloxochitla´n Municipio: San Antonio Eloxochitla´n. Location: East area had until recently been used to grow coffee, but facing mountain slopes of gorge. Altitude: 1,069–1,450 given the current low profits of this crop, most m (3,507–4,757 ft). Vegetation: Cloud forest disturbed plantations have been abandoned. This has given way by agricultural activities. to the regrowth of the natural vegetation. The area presents mosaics of vegetation at different successional Locality 8: San Antonio Eloxochitla´n – San Jose´ stages. Buenavista Municipio: San Antonio Eloxochitla´n. Location: Locality 13: Agua de Fierro – Santa Mar´ıa Chilchotla Locality begins at the bridge on the outskirts of the Municipio: Santa Marıa´ Chilchotla. Location: 5–7 km village of San Antonio Eloxochitla´n that leads to the (3–4.5 mi) N from Puente de Fierro, along the road to terracer´ıa to San Jose´ Buenavista. Altitude: 1,316–1,436 the town of Santa Marıa´ Chilchotla, and below it, m (4,317–4,711 ft). Vegetation: Cloud forest and towards the Petlapa River. Altitude: 1,070–1,230 m riparian forest disturbed by human activities. (3,510–4,035 ft). Vegetation: The area has also been highly disturbed by agricultural activities, mainly Locality 9: Huautla de Jimenez´ – Puente de Fierro coffee plantations. The talus of black limestone that Municipio: Huautla de Jimenez.´ Location: Trail borders some parts of the road is extremely interesting: between the outskirts of Huautla de Jimenez´ and the many locality-restricted species grow on this substrate. area of Puente de Fierro (see Locality 11). Altitude: 1,229–1,673 m (4,032–5,488 ft). Vegetation: Disturbed Locality 14: Mar´ıa Luisa – Agua de Gancho cloud forest, secondary vegetation. Municipio: Santa Marıa´ Chilchotla. Location: Approximately 8 km (5 mi) N of Puente de Fierro, Locality 10: Huautla de Jimenez´ – Cerro de la Adoracion´ along the road to the town of Santa Marıa´ Chilchotla; Municipio: Huautla de Jimenez.´ Location: Urban area west facing mountain slopes approaching the of Huautla de Jimenez´ and the adjacent mountain, community of Agua de Gancho. Altitude: 1,289– Cerro de la Adoracion.´ Altitude: 1,650–1,800 m 1,483 m (4,229–4,865 ft).Vegetation: In the populated (5,413–5,905 ft) in town area, 1,800–2,150 m (5,905– parts of this locality the vegetation is highly disturbed 7,053 ft) in adjacent mountain. Vegetation: Huautla de Jimenez´ has been inhabited for a very long time and its by agricultural activities. Because of the presence of vegetation has been decimated for the most part. pre-Hispanic ruins in the area, it is likely that the area However, throughout the area one can find elements of has been heavily impacted by human activities for a cloud forest vegetation. The Cerro de la Adoracion´ is long period of time. However, towards the NE of Agua considered a sacred place for the Mazatec people of the de Gancho, there are forested areas that are peculiar in highlands, and to this day it remains unpopulated. that their substrate is composed of large limestone Nevertheless, the vegetation is highly disturbed and blocks. only at its very top can one find a small patch of forest Locality 15: Zongolica Chilchotla composed of oak trees. Municipio: San Jose´ Tenango. Location: Altitude: Locality 11: Huautla de Jimenez´ – Santa Cruz de Jua´rez 1,709 m (5,606 ft). Vegetation: Highly disturbed by Municipios: Huautla de Jimenez´ and Santa Cruz de fire. Cloud forest remnants include an Ulmus tree that Jua´rez. Location: Path between Huautla de Jimenez´ is at least 40 m (131 ft) in height. and the town of Santa Cruz de Jua´rez. Altitude: 1,554– Locality 16: San Miguel Nuevo 1,819 m (5,098–5,967 ft). Vegetation: Disturbed cloud Municipio: San Jose´ Tenango. Location: San Miguel forest. Nuevo. Altitude: Approximately 1,100–1,200 m Locality 12: Agua de Fierro – Aguacatitla (3,608–3,937 ft). Vegetation: Disturbed cloud forest. NUMBER 20 MUNN-ESTRADA: FLORA OF THE SIERRA MAZATECA OF OAXACA, MEXICO 41

STUDY AREA 3. CERRO RABON´ : (Eastern portion of LYCOPODIACEAE the Sierra Mazateca) Huperzia pringlei (Underw. & F.E.Lloyd) Holub Locality 17: San Mart´ın Caballero Epiphytic herb; 2242 m; 2 (2323). Municipio: San Jose´ Tenango. Location: Cerro Rabon´ Huperzia taxifolia (Sw.) Trevis plateau. Accessible by paths from San Jose´ Tenango or Epiphytic herb; 1458–1515 m; 14 (621A), 18 Jalapa de Dıaz.´ Altitude: 1,250–1,645 m (4,101–5,396 (1775). ft). Vegetation: Mature cloud forest in non-populated Lycopodium clavatum L. areas; disturbed cloud forest in populated areas. Herb; 1589–2742 m; 2 (1719), 11 (1006). Lycopodium thyoides Humb. & Bonpl. ex Willd. Locality 18: San Mart´ın Caballero – Cerro Caballero Herb; 2223–2742 m; 2 (1714), 3 (2077), 4 (2282), ´ Municipio: San Jose Tenango. Location: Along paths 5(1628). east of the town of San Martın´ Caballero. Altitude: 1,360–1,515 m (4,461–4,970 ft). Vegetation: Mosaic of SELAGINELLACEAE areas with mature cloud forest and areas with Selaginella guatemalensis Baker disturbed vegetation. Herb; 1600 m; 17 (1754). Locality 19: San Mart´ın Caballero – San Jose´ Tenango Selaginella martensii Spring Municipio: San Jose´ Tenango. Location: Along paths Herb; 1530 m; 12 (1002A). leading to San Martın´ Caballero. Altitude: 1,200–1,424 Selaginella oaxacana Spring m (3,937–4,672 ft). Vegetation: Mosaic of areas with Herb; 1560 m; 17 (532). mature cloud forest and areas with disturbed Selaginella silvestris Aspl. vegetation. Herb; 2321 m; 4 (2300). Selaginella stellata Spring Locality 20: Rancho Avendan˜o – San Mart´ın Caballero Herb; 1233 m; 6 (1115). Municipio: San Jose´ Tenango. Location: Path from the Rancho Avendan˜o, accessible from Cerro Central, FERNS toward San Martın´ Caballero. Altitude: ANEMIACEAE Approximately 1,100–1,450 m (3,608–4,757 ft). Vegetation: Highly disturbed cloud forest. Anemia phyllitidis (L.) Sw. Herb; 1352 m; 7 (1963). Locality 21: Cerro Rabon´ – Cerro Alamo Municipio: San Jose´ Tenango. Location: Between the ASPLENIACEAE communities of Cerro Rabon´ and Cerro Alamo. Asplenium auriculatum Sw. Because of its remoteness from the field base and the Epiphytic herb; 1645 m; 17 (1759). costs involved in reaching the site, this locality was not Asplenium cuspidatum Lam. explored in detail. The one trip made to the site Epiphytic herb; 1323 m; 7 (1928). revealed that the locality has cloud forests on limestone Asplenium fragrans Sw. substrates that deserve more study. Altitude: 1,100– Epiphytic herb; 1600 m; 17 (1746). 1,555 m (3,608-5,101 ft). Vegetation: Mature cloud Asplenium monanthes L. forest in non-populated areas; disturbed cloud forest in Herb; 2300–2340 m; 1 (1939), 4 (1676), 21 (762). populated areas. Asplenium nesioticum Maxon Herb; 7 (1191).

APPENDIX 2 ATHYRIACEAE

Diplazium cf. urticifolium Christ SPECIES CHECKLIST Herb; 1180 m; 16 (705).

See ‘‘Methods’’ for explanation of format and content. BLECHNACEAE Families, genera and species are listed alphabetically by Blechnum appendiculatum Willd. family etc. within the four major groups: Lycophytes, Herb; 1352 m; 7 (1966). Ferns, Gymnosperms, and Angiosperms. Blechnum falciforme (Liebm.) C.Chr. LYCOPHYTES Herb; 2340 m; 4 (1681). 42 LUNDELLIA DECEMBER, 2017

Blechnum polypodioides Raddi Epiphytic herb; 1600 m; 17 (1742). Herb; 1197 m; 12 (1650). Phanerophlebia macrosora (Baker) Underw. Blechnum schiedeanum (Schltdl. ex C.Presl) Hieron. Herb; 2300 m; 1 (1478). Herb; 1418 m; 9 (2368). Phanerophlebia remotispora E.Fourn. Blechnum wardiae Mickel & Beitel Herb; 1200 m; 7 (1111). Herb; 1560 m; 17 (1730). EQUISETACEAE Woodwardia spinulosa M.Martens & Galeotti, vel aff. Herb; 1426 m; 9 (2367). Equisetum myriochaetum Schltdl. & Cham. Herb; 1200 m; 7 (1104). CYATHEACEAE GLEICHENIACEAE Alsophila firma (Baker) D.S.Conant Tree fern; 2 (2201). Sticherus bifidus (Willd.) Ching Cyathea divergens Kunze Herb; 1632 m; 10 (802). Tree fern; 1233 m; 6 (1177). Sticherus palmatus (W.Schaffn. ex E.Fourn.) Copel Cyathea fulva (M.Martens & Galeotti) Fee´ Herb; 1418 m; 9 (2369). Tree fern; 1515–2345 m; 4 (2287), 5 (1620), 18 HYMENOPHYLLACEAE (1783). Sphaeropteris horrida (Liebm.) R.M.Tryon Hymenophyllum fucoides (Sw.) Sw. Tree fern; 1300 m; 7 (1421). Epiphytic herb; 2611 m; 2 (1704). Hymenophyllum myriocarpum Hook. DICKSONIACEAE Epiphytic herb; 2242 m; 2 (2328), 2330). Lophosoria quadripinnata (J.F.Gmel.) C.Chr. Hymenophyllum polyanthos (Sw.) Sw. Herb; 2242 m; 2 (2235, 2334). Epiphytic herb; 1515 m; 18 (1776). Trichomanes collariatum Bosch DRYOPTERIDACEAE Epiphytic herb; 21 (761). Arachniodes denticulata (Sw.) Ching LINDSAEACEAE Herb; 1600–2345 m; 4 (2286), 10 (1079), 17 (1755). Odontosoria schlechtendalii (C.Presl) C.Chr. Dryopteris wallichiana (Spreng.) Hyl. Herb; 1460 m; 12 (1381). Herb; 2300–2611 m; 1 (1481), 2 (1609, 1712), 4 MARATTIACEAE (2294). Elaphoglossum erinaceum (Fee)´ T.Moore var. Marattia laxa Kunze erinaceum Herb; 2345 m; 2 (2237), 4 (2283). Epiphytic herb; 1560–1645 m; 17 (536, 1726, NEPHROLEPIDACEAE 1758). Elaphoglossum glaucum T.Moore Nephrolepis cordifolia (L.) C.Presl. Epiphytic herb; 2328 m; 4 (1670). Huautla–Chilchotla Area, specific locality not Elaphoglossum leebrowniae Mickel disclosed; species is endangered and protected Herb; 2487 m; 2 (1688). under Mexican law. Elaphoglossum paleaceum (Hook. & Grev.) Sledge Nephrolepis pectinata (Willd.) Schott Epiphytic herb; 2320–2473 m; 1 (1456), 2 (1690). Herb; 1185–1474 m; 6 (875), 14 (1288). Elaphoglossum peltatum (Sw.) Urb. POLYPODIACEAE Epiphytic herb; 1600 m; 17 (1753). Elaphoglossum sartorii (Liebm.) Mickel Campyloneurum amphostenon (Kunze ex Klotzsch) Epiphytic herb; 2340–2487 m; 2 (1683), 4 (1674, Fee´ 1677). Epiphytic herb; 1197–2487 m; 2 (1682), 12 Elaphoglossum squamipes (Hook.) T.Moore (1643). Epiphytic herb; 2611 m; 2 (1703). Campyloneurum angustifolium (Sw.) Fee´ Elaphoglossum vestitum (Schltdl. & Cham.) T.Moore Epiphytic herb; 1184 m; 6 (922). Herb; 1197–1515 m; 12 (1648), 18 (1785). Campyloneurum ensifolium (Willd.) J. Sm. Elaphoglossum viride (E.Fourn.) C.Chr. Herb; 1185–1203 m; 6 (854, 931). NUMBER 20 MUNN-ESTRADA: FLORA OF THE SIERRA MAZATECA OF OAXACA, MEXICO 43

Campyloneurum serpentinum (Christ) Ching Epiphytic herb; 1197–2340 m; 2 (2347), 3 (2095), Herb; 1226 m; 13 (2142). 4(1679), 5 (1622), 10 (1082), 12 (1653), 13 Campyloneurum xalapense Fee´ (2178), 19 (1834). Epiphytic herb; 1197–2313 m; 5 (1623), 7 (1128, Polypodium plesiosorum Kunze 1932, 1965), 12 (1644). Epiphytic herb; 1323–2300 m; 1 (1480), 7 (1929). Cochlidium linearifolium (Desv.) Maxon ex C.Chr. Polypodium pleurosorum Kunze ex Mett. Epiphytic herb; 2604 m; 2 (1695). Epiphytic herb; 2313–2328 m; 4 (1669), 5 (1621). Lellingeria prionodes (Mickel & Beitel) A.R.Sm. & Polypodium polypodioides (L.) Watt R.C.Moran Herb; 1197–1226 m; 7 (1192), 12 (1638), 13 Epiphytic herb; 2313 m; 5 (1631A). (2150). Melpomene leptostoma (Fee)´ A.R.Sm. & R.C.Moran Polypodium puberulum Schltdl. & Cham. Epiphytic herb; 1515–2313 m; 5 (1631), 18 Epiphytic herb; 2300–2340 m; 1 (1479), 4 (1680). (1779). Terpsichore delicatula (M.Martens & Galeotti) Niphidium crassifolium (L.) Lellinger A.R.Sm. Epiphytic herb; 1184–1560 m; 6 (921), 7 (1129), 17 Epiphytic herb; 2333 m; 5 (1632). (535). PTERIDACEAE Pecluma alfredii (Rosenst.) M.G.Price var. cupreolepis (A.M.Evans) A.R.Sm. Adiantum andicola Liebm. Herb; 1290–2223 m; 3 (2072), 7 (1408), 7 (1969), Herb; 1197–1515 m; 9 (1038), 12 (1649), 13 7(1408). (2148), 18 (1790). Adiantum capillus-veneris L. Phlebodium areolatum (Humb. & Bonpl. ex Willd.) Herb; 1185–1327 m; 6 (1656), 8 (2064). J.Sm. Adiantum poiretii Wikstr. Herb; 1197–1375 m; 6 (1262), 12 (1645), 13 Herb; 1637 m; 11 (1994). (2145), 19 (1822). Cheilanthes bonariensis (Willd.) Proctor Pleopeltis angusta Humb. & Bonpl. ex Willd. var. Herb; 2313 m; 5 (1634). stenoloma (Fee)´ Farw. Llavea cordifolia Lag. Herb; 1197–1226 m; 12 (1641), 13 (2144). Herb; 1185–1266 m; 6 (814, 855). Pleopeltis crassinervata (Fee)´ T.Moore Mildella fallax (M.Martens & Galeotti) G.L.Nesom Herb; 1197–1352 m; 7 (1970), 8 (2063), 9 (1059), Herb; 1197 m; 12 (1640). 12 (1642), 13 (2149). Mildella intramarginalis (Kaulf. ex Link) Trevis. Pleopeltis fallax (Schltdl. & Cham.) Mickel & Beitel Epiphytic herb; 2242 m; 2 (2344). Epiphytic herb; 1229 m; 9 (1033). Myriopteris lendigera (Cav.) J.Sm. Pleopeltis polylepis (Roem. ex Kunze) T.Moore var. Herb; 2242 m; 2 (2346). interjecta (Weath.) E.A.Hooper Pteris longifolia L. Epiphytic herb; 2151–2151 m; 2 (1693), 10 Herb; 1290 m; 7 (1413). (1088). Pteris orizabae M.Martens & Galeotti Polypodium echinolepis Fee´ Herb; 2345 m; 4 (2284). Herb; 1226 m; 13 (2139). Pteris podophylla Sw. Polypodium furfuraceum Schltdl. & Cham. Herb; 1197 m; 12 (826). Epiphytic herb; 1226–1352 m; 6 (1115A), 7 Vittaria graminifolia Kaulf. (1972), 13 (2140). Epiphytic herb; 1184–2604 m; 2 (1696), 6 (906), Polypodium hartwegianum Hook. 17 (533, 1745A). Epiphytic herb; 2611 m; 2 (1706). TECTARIACEAE Polypodium longepinnulatum E.Fourn. Epiphytic herb; 1645 m; 17 (1766). Tectaria heracleifolia (Willd.) Underw. Polypodium loriceum L. Herb; 1184–1226 m; 6 (918), 13 (2141). Herb; 21 (773). THELYPTERIDACEAE Polypodium munchii Christ Epiphytic herb; 2611–2640 m; 2 (1160, 1708). Thelypteris atrovirens (C.Chr.) C.F.Reed Polypodium plebeium Schltdl. & Cham. Herb; 1637 m; 11 (1993). 44 LUNDELLIA DECEMBER, 2017

Thelypteris concinna (Willd.) Ching Saurauia pedunculata Hook. Herb; 1197 m; 12 (1651). Tree; 1323 m; 8 (1240). Thelypteris paucipinnata (Donn.Sm.) C.F.Reed Saurauia villosa DC. Herb; 1200 m; 7 (1113). Tree; 1513 m; 18 (1806). Saurauia aff. villosa DC. GYMNOSPERMS Tree; 1184–1300 m; 6 (828, 1428), 7 (1410, 1422, CUPRESSACEAE 1899).

Cupressus lusitanica Mill. ADOXACEAE Huautla–Chilchotla and Cerro Rabon´ Areas, Viburnum hartwegii Benth specific locality not disclosed; species is Treel 1262–1424 m; 7 (1400, 1402); 19 (462). protected under Mexican law. ALSTROEMERIACEAE PINACEAE Bomarea acutifolia (Link & Otto) Herb Pinus patula Schiede ex Schltdl. & Cham. Woody vine; 1424–1458 m; 14 (631), 19 (459), 21 Tree; 2183–2223 m; 1 (2231A), 3 (2100), 5 (1615). (746).

PODOCARPACEAE ALTINGIACEAE

Podocarpus matudae Lundell Liquidambar styraciflua L. Sierra de los Frailes Area, specific locality not Tree; 1229–1412 m; 7 (2037), 9 (1066). disclosed; species is protected under Mexican law. AMARANTHACEAE TAXACEAE Alternanthera lanceolata (Benth.) Schinz Taxus globosa Schltdl. Herb; 1185 m; 6 (2124), 17 (435). Sierra de los Frailes Area, specific locality not Iresine diffusa Humb. & Bonpl. ex Willd. disclosed; species is protected under Mexican law. Herb; 1185–1637 m; 6 (834), 11 (1995), 14 (1494). ANGIOSPERMS Iresine hebanthoides Suess Shrub; m; 13 (2183). ACANTHACEAE ANACARDIACEAE Aphelandra schiedeana Schltdl. & Cham. Shrub; 1180–1350 m; 7 (1505), 13 (943). Toxicodendron radicans (L.) Kuntze Woody vine; 1184 m; 6 (926). Dicliptera sumichrastii Lindau Herb; 1266 m; 6 (819, 1875). APIACEAE Justicia aurea Schltdl. Eryngium carlinae F.Delaroche Shrub; 1184–1434 m; 6 (900, 1655), 9 (1861). Herb; 2636 m; 2 (1347). Justicia fimbriata (Nees) V.A.W.Graham Ottoa oenanthoides Kunth Shrub; 1180 m; 16 (713). Herb; 2725 m; 2 (1154). Odontonema callistachyum (Schltdl. & Cham.) Sanicula liberta Cham. & Schltdl. Kuntze Herb; 1350 m; 7 (1254), 17 (565). Herb; 1185–1226 m; 6 (869), 13 (2152). APOCYNACEAE Stenandrium chameranthemoideum Oerst. Herb; 1250–1560 m; 14 (969), 17 (437, 566, 1720), Alstonia longifolia (A.DC.) Pichon 18 (1773). Tree; 1226 m; 13 (2165). Stenostephanus haematodes (Schltdl.) T.F.Daniel Asclepias curassavica L. Shrub; 1185–1560 m; 6 (2126), 14 (954, 1531), 17 Herb; 1184 m; 6 (1289) (1721), 19 (441). Mandevilla oaxacana (A.DC.) Hemsl. Woody vine; 1226 m; 13 (1306). ACTINIDIACEAE Matelea velutina (Schltdl.) Woodson Saurauia leucocarpa Schltdl. Herb; 1184 m; 6 (1291) Tree; 1589 m; 11 (1007). Stemmadenia litoralis (Kunth) L.Allorge NUMBER 20 MUNN-ESTRADA: FLORA OF THE SIERRA MAZATECA OF OAXACA, MEXICO 45

Tree; 1233 m; 6 (1175). Epiphtic; 1233–1460 m; 6 (1181), 8 (1442), 12 (1387A). ARACEAE Agave obscura Schiede Anthurium lucens Standl. Epiphtic; 1226 m; 13 (1308). Herb; 1323–1350 m; 8 (1298), 17 (570, 571). Maianthemum macrophyllum (M.Martens & Anthurium podophyllum (Schltdl. & Cham.) Kunth Galeotti) LaFrankie Huautla–Chilchotla Area, specific locality not Herb; 1100–1200 m; 16 (663). disclosed; species is endangered and protected Maianthemum paniculatum (M.Martens & Galeotti) under Mexican law. LaFrankie Monstera deliciosa Liebm. Epiphytic herb; 1350 m; 17 (567). Herb; 1180–1645 m; 16 (681), 17 (1761). Maianthemum scilloideum (M.Martens & Galeotti) Philodendron smithii Engl. LaFrankie Herb; 1474 m; 14 (1286). Herb; 2300 m; 1 (1468). Xanthosoma robustum Schott Yucca sp. Herb; 1323 m; 7 (1923). Tree; 1418 m; 9 (2379). Xanthosoma sagittifolium (L.) Schott ASTERACEAE Epiphytic herb; 1600 m; 17 (1750). Achyrocline sp. ARALIACEAE Shrub; 1650 m; 11 (1991). Acmella oppositifolia (Lam.) R.K.Jansen Dendropanax arboreus (L.) Decne. & Planch. Herb; 1184–1415 m; 6 (891). Tree; 1184–1589 m; 6 (896), 11 (2009), 13 (2166). Acmella cf. oppositifolia (Lam.) R.K.Jansen Oreopanax capitatus (Jacq.) Decne. & Planch. Herb; 1250 m; 17 (412). Tree; 1323 m; 8 (1235). Ageratina glauca (Sch. Bip. ex Klatt) R.M.King & Oreopanax liebmannii Marchal H.Rob Tree; 2321–2335 m; 1 (1919, 2229), 4 (2297). Shrub; 2473–2742 m; 2 (1613, 1717). Oreopanax xalapensis (Kunth) Decne. & Planch. Ageratina grandifolia (Regel) R.M.King & H.Rob. Tree; 1360–2320 m; 1 (1463), 18 (1812). (Ageratina conspicua (Kunth & Bouche)´ R.M.King & ARECACEAE H.Rob.) Shrub; 1658 m; 11 (1987). Chamaedorea oblongata Mart. Ageratina isolepis (B.L. Rob.) R.M.King & H.Rob. Herb; 1323 m; 8 (1236). Herb; 1412 m; 7 (2044). Chamaedorea oreophila Mart. Ageratina ligustrina (DC.) R.M.King & H.Rob. Huautla–Chilchotla Area, specific locality not Shrub; 1515–2473 m; 1 (1452), 2 (1614), 4 (1667), disclosed; species is endangered and protected 18 (1789). under Mexican law. Ageratina malacolepis (B.L.Rob.) R.M.King & H.Rob. Chamaedorea rojasiana Standl. & Steyerm. Herb; 1458 m; 14 (993). Cerro Rabon´ Area, specific locality not disclosed; Ageratina mazatecana B.L.Turner species is endangered and protected under Herb; 1466 m; 14 (981). Mexican law. Ageratina ovilla (Standl. & Steyerm.) R.M.King & Chamaedorea tepejilote Liebm. H.Rob. Herb; 1560 m; 17 (1731). Woody vine; 2335 m; 1 (1914). Chamaedorea sp. Ageratina pichinchensis (Kunth) R.M.King & H.Rob. Herb; 1424–1560 m; 17 (541, 542, 549), 19 (488). Herb; 1466 m; 14 (980). Ageratina pringlei (B.L.Rob. & Greenm.) R.M.King & ARISTOLOCHIACEAE H.Rob. Aristolochia tricaudata Lem. Herb; 1185–1229 m; 6 (881), 9 (1030). Shrub; 1069–1375 m; 7 (1214, 1250), 19 (1824). Ageratina prunellifolia (Kunth) R.M.King & H.Rob. Shrub; 2640 m; 2 (1156). ASPARAGACEAE Ageratina rubricaulis (Kunth) R.M.King & H.Rob. Agave ellemeetiana K.Koch subsp. ellemeetiana Tree; 1203 m; 6 (930). 46 LUNDELLIA DECEMBER, 2017

Ageratina vernalis (Vatke & Kurtz) R.M.King & Fleischmannia pycnocephala (Less.) R.M.King & H.Rob. H.Rob. Shrub; 1632 m; 10 (801). Herb; 1229 m; 9 (1020). Ageratum houstonianum Mill. Fleischmannia seleriana (B.L.Rob.) R.M.King & Herb; 1412–1673 m; 7 (2046), 9 (1856), 10 (799), H.Rob. 11 (2002), 14 (979). Herb; 1185–1327 m; 6 (883, 887), 8 (2061). Aldama dentata La Llave & Lex. Fleischmanniopsis mendax (Standl. & Steyerm.) Herb; 1184–1458 m; 6 (789), 14 (1488). R.M.King & H.Rob. Alloispermum integrifolium (DC.) H.Rob. Herb; 2 (2202). Herb; 1300–1626 m; 7 (1952, 2045), 10 (804), 14 Galinsoga quadriradiata Ruiz & Pav. (982), 14 (994). Herb; 21 (749). Archibaccharis schiedeana (Benth.) J.D.Jacks. Gnaphalium sp. Herbaceous vine; 1184 m; 6 (1430). Herb; 1300 m; 7 (1951A). Baccharis conferta Kunth Jaegeria hirta (Lag.) Less. Shrub; 1709–2725 m; 2 (1146, 2213), 3 (2088), 10 Herb; 1185–1426 m; 6 (873), 9 (2354), 19 (442), (1093), 11 (1014), 15 (950). 20 (584). Baccharis trinervis (Lam.) Pers. Koanophyllon pittieri (Klatt) R.M.King & H.Rob. Shrub; 1229–2335 m; 4 (2270), 7 (1125), 9 (1029, Tree; 1184 m; 6 (907). 1060), 11 (1999). Lagascea helianthifolia Kunth Bartlettina karwinskiana (DC.) R.M.King & H.Rob. Herb; 1229–1261 m; 7 (1887), 9 (1069). Shrub; 2223–2300 m; 1 (1937), 3 (2099). Leiboldia serrata (D.Don) Gleason Bartlettina oresbia (B.L.Rob.) R.M.King & H.Rob. Shrub, 1266 m, 6 (815) Shrub; 2223 m; 3 (2096). Lepidaploa tortuosa (L.) H. Rob. Bartlettina sordida (Less.) R.M.King & H.Rob. (Vernonia tortuosa (L.) S.F.Blake) Shrub; 1185–2220 m; 3 (2028), 6 (868, 888), 14 Shrub; 1266 m; 7 (1890). (966), 18 (1796), 21 (771). Melampodium divaricatum (Rich. ex Rich.) DC. Bartlettina tuerckheimii (Klatt) R.M.King & H.Rob. Herb; 1184–1458 m; 6 (1426), 7 (1117), 14 (1489), Shrub; 2325–2328 m; 4 (2262). 19 (405). Bidens triplinervia Kunth Microspermum debile Benth. Herb; 2151 m; 10 (1091). Herb; 2314 m; 2 (1331). Calea urticifolia (Mill.) DC. Mikania cordifolia (L.f.) Willd. Herb; 1458 m; 14 (985). Woody vine; 1229–1631 m; 9 (1048, 1858). Chionolaena salicifolia (Bertol.) G.L.Nesom Mikania pyramidata Donn.Sm. (Gnaphaliothamnus salicifolius (Bertol.) G.L. Nesom) Woody vine; 1513 m; 18 (1799). Herb; 1632–2742 m; 2 (1716), 3 (2076), 10 (803). Montanoa speciosa (DC.) Sch.Bip. ex C.Koch Cirsium sp. Tree; 20 (574). Herb; 1300 m; 7 (1951). Neomirandea araliifolia (Less.) R.M.King & H.Rob. Critonia daleoides DC. Tree; 1185 m; 6 (1877A). Tree; 1466 m; 14 (984). Osbertia stolonifera (DC.) Greene Critonia hospitalis (B.L.Rob.) R.M.King & H.Rob. Herb; 2725 m; 2 (1153). Tree; 1515 m; 18 (1780). Oxylobus oaxacanus S.F.Blake Dahlia australis (Sherff) P.D.Sorensen Herb; 2742 m; 2 (1717A). Herb; 2300 m; 1 (1467). Pentacalia parasitica (Hemsl.) H.Rob. & Cuatrec. Erechtites valerianifolia (Link ex Wolf) Less. ex DC. (Pentacalia wilburii H.Rob.) Herb; 1250 m; 17 (434). Shrub; 2604 m; 2 (1699). Erigeron karvinskianus DC. Perymeniopsis ovalifolia (A. Gray) H.Rob. Herb; 1185–1474 m; 6 (813, 885), 14 (636), 19 Herbaceous vine; 1184 m; 6 (795, 1368). (448), 21 (748). Perymenium gracile Hemsl. Erigeron longipes DC. Shrub; 2203 m; 4 (1659). Herb; 1229 m; 9 (1052). Piqueria trinervia Cav. NUMBER 20 MUNN-ESTRADA: FLORA OF THE SIERRA MAZATECA OF OAXACA, MEXICO 47

Herb; 1434–1658 m; 9 (1860), 11 (1983), 14 Herb; 1184–1658 m; 6 (788), 6 (1290), 9 (1019), 9 (975). (2374), 11 (1985). Podachaenium eminens (Lag.) Sch.Bip. Verbesina hypoglauca Sch.Bip. ex Klatt Tree; 1285 m; 7 (1900). Shrub; 2242–2611 m; 2 (1710, 2338). Roldana jurgensenii (Hemsl.) H.Rob. & Brettell Vernonia arctioides Less. Shrub; 1278–2335 m; 1 (1903), 7 (1896, 1921). Tree; 1185–1570 m; 6 (815, 886), 9 (1866), 11 Roldana lanicaulis (Greenm.) H.Rob. & Brettell (2011). Herb; 2223 m; 3 (2098). Vernonia heydeana J.M.Coult. Roldana mazatecana B.L.Turner Shrub; 1380 m; 7 (1974). Shrub; 2320 m; 1 (1947) Vernonia jonesii B.L.Turner Roldana mexicana (McVaugh) H.Rob. & Brettell Shrub; 1589–1645 m; 11 (1009), 17 (1770). Herb; 2611 m; 2 (1713). Vernonia patens Kunth Roldana schaffneri (Sch.Bip. ex Klatt) H.Rob. & Tree; 1300–1415 m; 6 (2310), 7 (1126). Brettell Viguiera cordata (Hook. & Arn.) D’Arcy Herb; 1229–1466 m; 9 (1065), 14 (974). Shrub; 1185 m; 6 (880). Schistocarpha bicolor Less. Villasenoria orcutti (Greenm.) B.L.Clark Herb; 1412–1412 m; 7 (2042). Huautla–Chilchotla Area, specific locality not Schistocarpha pedicellata Klatt disclosed; species is endangered and protected Herb; 2327–2604 m; 2 (1702), 4 (1671). under Mexican law. Senecio callosus Sch.Bip. Youngia japonica (L.) DC. Herb; 2223–2335 m; 1 (1901), 3 (2103), 4 (1663). Herb; 21 (747). Senecio salignus DC. BALSAMINACEAE Tree; 2170 m; 3 (2022). Sigesbeckia jorullensis Kunth Impatiens walleriana Hook.f. Herb; 2242 m; 2 (2327). Herb; 1290 m; 7 (1416), 20 (582). Sinclairia andromachioides (Less.) Sch.Bip. ex Rydb. BEGONIACEAE Shrub; 1180–1226 m; 13 (2128), 13 (2169). Sinclairia deppeana (Less.) Rydb. Begonia caroliniifolia Regel Shrub; 1180–2335 m; 1 (1913), 13 (2127), 18 Herb; 1185 m; 6 (865, 2112). (1798). Begonia crassicaulis Lindl. Sinclairia discolor Hook. & Arn. Herb; 1460 m; 12 (1390). Shrub; 1185–2204 m; 3 (2034), 6 (871). Begonia glabra Aubl. Smallanthus uvedalia (L.) Mack. Herb; 7 (1185). Shrub; 1226–1290 m; 7 (1419), 13 (1318). Begonia heracleifolia Schltdl. & Cham. Stevia jorullensis Kunth Herb; 1346 m; 19 (1840). Herb; 2604 m; 2 (1701). Begonia ludicra A.DC. Stevia monardifolia Kunth Herb; 1474 m; 14 (1283). Herb; 2604 m; 2 (1700). Begonia manicata Brongn. ex F.Cels Tagetes filifolia Lag. Herb; 1150–1327 m; 6 (820, 864, 1869, 2109), 8 Herb; 1460 m; 12 (1382). (2056). Tanacetum parthenium (L.) Sch.Bip. Begonia nelumbiifolia Schltdl. & Cham. Herb; 1601 m; 9 (1171). Epiphytic herb; 1203 m; 6 (934). Telanthophora grandifolia (Less.) H.Rob. & Brettell Begonia pustulata Liebm. Shrub; 1184–1360 m; 6 (879, 1433), 18 (1815). Herb; 1180 m; 16 (674). Tetrachyron manicatum Schltdl. Begonia sartorii Liebm. Herb; 1180–1709 m; 6 (807), 6 (889, 1878), 8 Herb; 1323 m; 7 (1926). (2060), 9 (1026), 11 (1980), 12 (1002), 13 (942), BERBERIDACEAE 15 (946). Tithonia diversifolia (Hemsl.) A.Gray Berberis gracilis Hartw. ex Benth. Tree; 1266–1266 m; 6 (808), 7 (1889), 7 (1889). (Berberis gracilis Hartw. ex Benth. var. madrensis Trigonospermum melampodioides DC. Marroq.) 48 LUNDELLIA DECEMBER, 2017

Tree; 1229 m; 9 (1018). Tree; 1330 m; 12 (1374).

BETULACEAE CACTACEAE

Alnus acuminata Kunth subsp. arguta (Schltdl.) Disocactus martianus (Zucc. ex Pfeiff.) Barthlott Furlow (Aporocactus conzattii Britton & Rose) Tree; 2151–2340 m; 1 (1447, 1911), 3 (2019, Epiphytic succulent; 2242–2450 m; 2 (1164, 2020), 10 (617, 1076). 2236). Carpinus tropicalis (Donn.Sm.) Lundell Disocactus sp. Sierra de los Frailes and Huautla–Chilchotla (Nopalxochia sp.) Areas, specific locality not disclosed; species is Epiphytic succulent; 1069–2340 m; 1 (2226), 7 endangered and protected under Mexican law. (1245). Rhipsalis baccifera (J.S.Muell.) Stearn BORAGINACEAE Epiphytic succulent; 1184–1185 m; 6 (1321, Cynoglossum amabile Stapf & J.R.Drumm. 2122). Herb; 1589 m; 12 (1179). CALCEOLARIACEAE Tournefortia acutiflora M.Martens & Galeotti Tree; 2220 m; 2 (2194), 3 (2030). Calceolaria mexicana Benth. Wigandia urens (Ruiz & Pav.) Kunth Herb; 1185–1226 m; 6 (860), 13 (2163), 16 (716). Tree; 1626 m; 10 (798). Calceolaria tripartita Ruiz & Pav. BROMELIACEAE Herb; 1673 m; 9 (1857).

Pitcairnia recurvata (Scheidw.) K.Koch CAMPANULACEAE Epiphytic herb; 1289 m; 14 (1274). Lobelia cardinalis L. Tillandsia butzii Mez Herb; 1500 m; 17 (555), 20 (401). Epiphytic herb; 1474 m; 14 (970). Lobelia laxiflora Kunth Tillandsia filifolia Schltdl. & Cham. (Lobelia laxiflora Kunth subsp. laxiflora) Epiphytic herb; 1270–1350 m; 6 (1273A), 7 Herb; 1229–1604 m; 9 (1042), 11 (2004). (1257). Lobelia sartorii Vatke Tillandsia grandis Schltdl. Herb; 1185–2450 m; 2 (1162), 3 (2091), 6 (847), 6 1474–1589 m; 12 (1182), 14 (958). (2125). Tillandsia gymnobotrya Baker Lobelia xalapensis Kunth Epiphytic herb; 2328 m; 4 (1660). Herb; 1226 m; 13 (2175). Tillandsia imperialis E. Morren ex Roezl Sierra de los Frailes Area, specific locality not CANNABACEAE disclosed; species is endangered and protected Trema micrantha (L.) Blume under Mexican law. Tree; 1323 m; 8 (1228). Tillandsia leiboldiana Schltdl. Epiphytic herb; 1323–1375 m; 8 (1237), 19 CAPPARACEAE (1829). Capparidastrum mollicellum (Standl.) Cornejo & Iltis Tillandsia multicaulis Steud. Tree; 1184–1350 m; 6 (935), 7 (1249). Epiphytic herb; 1315 m; 12 (1371). Tillandsia punctulata Schltdl. & Cham. CAPRIFOLIACEAE Epiphytic herb; 1375–1515 m; 18 (1774), 19 Sambucus sp. (1821). Shrub; 9 (605). Tillandsia tricolor Schltdl. & Cham. Valeriana candolleana Gardner Huautla–Chilchotla Area, specific locality not Herb; 1185–1226 m; 6 (850), 13 (2180). disclosed; species is endangered and protected Valeriana scandens L. under Mexican law. Herbaceous vine; 1350–1424 m; 17 (560), 19 BRUNELLIACEAE (467).

Brunellia mexicana Standl. CARYOPHYLLACEAE NUMBER 20 MUNN-ESTRADA: FLORA OF THE SIERRA MAZATECA OF OAXACA, MEXICO 49

Arenaria lanuginosa (Michx.) Rohrb. Vine; 2314 m; 19 (1837). Herb; 1658–2223 m; 3 (2092), 11 (1981). Dichondra repens J.R.Forst & G.Forst Drymaria cordata (L.) Willd. ex Schult. Herb; 1226 m; 13 (2143). Herb; 1185 m; 6 (2105). Dichondra sericea Sw. Herb; 1426 m; 9 (2360). CELASTRACEAE Ipomoea batatas (L.) Lam. Celastrus liebmannii Standl. Woody vine; 1233 m; 7 (1395). Shrub; 2321 m; 4 (2299). Ipomoea batatoides Choisy Maytenus sp. Herbaceous vine; 1226 m; 13 (1311). Tree; 1474 m; 14 (961) Ipomoea cholulensis Kunth Wimmeria bartletti Lundell Herbaceous vine; 1229–1458 m; 9 (1053), 14 Tree; 1226–1352 m; 7 (1958), 13 (2129, 2130). (1491). Zinowiewia integerrima Turcz. Ipomoea funis Schltdl. & Cham. Tree; 2167–2220 m; 3 (2025, 2027). Woody vine; 1380 m; 7 (1973). CHLORANTHACEAE Ipomoea mairetii Choisy Herbaceous vine; 1436 m; 8 (2053). Hedyosmum mexicanum C.Cordem. Merremia tuberosa (L.) Rendle Tree; 1330–1431 m; 7 (1243), 12 (1376, 756), 21 Herbaceous vine; 20 (578). (782). CORNACEAE CLETHRACEAE Cornus excelsa Kunth Clethra conzattiana L.M.Gonza´lez Tree; 2 (2190). Tree; 1513–1515 m; 18 (1781, 1804, 1817). Clethra hartwegii Britton CRASSULACEAE Tree; 2300–2604 m; 1 (1460, 1485), 2 (1344, Echeveria rosea Lindl. 1698). Succulent; 2328 m; 1 (1666). Clethra lanata M.Martens & Galeotti Kalanchoe pinnata (Lam.) Pers. Tree; 1100–2223 m; 3 (2080), 10 (1085), 21 (752). Succulent; 1200–1604 m; 6 (933), 11 (2006). CLUSIACEAE Sedum sp. Succulent; 1226 m; 13 (2181). Clusia guatemalensis Hemsl. Tree; 1226–1600 m; 9 (612), 13 (2170), 14 (973), CUCURBITACEAE 17 (1747), 19 (1841) Clusia sp. Hanburia mexicana Seem. Tree; 1226–1474 m; 8 (1233), 14 (514) Herbaceous vine; 1100–1555 m; 21 (743).

COMMELINACEAE CUNONIACEAE

Matudanthus nanus (M.Martens & Galeotti) Weinmannia pinnata L. D.R.Hunt Tree; 1560–2520 m; 2 (1345), 4 (2273), 17 (509). Herb; 2400 m; 2 (1340). CYPERACEAE Tradescantia zanonia (L.) Sw. Herb; 1184–1474 m; 6 (916, 2118), 14 (639, 963), Cyperus hermaphroditus (Jacq.) Standl. 17 (546). Herb; 1424 m; 19 (495). Tripogandra purpurascens (Schauer) Handlos Rhynchospora aristata Boeck. Herb; 1100–1450 m; 20 (579). Herb; 1100–1200 m; 16 (677, 679). Tripogandra serrulata (Vahl) Handlos Rhynchospora radicans (Schltdl. & Cham.) H.Pfeiff Herb; 1100–1300 m; 6 (1266), 7 (1127), 13 (2146), Herb; 1424 m; 19 (447). 16 (676), 17 (433). DIPENTODONTACEAE CONVOLVULACEAE Perrottetia longistylis Rose Cuscuta sp. Tree; 1180–1460 m; 12 (1392), 16 (710), 19 (451). 50 LUNDELLIA DECEMBER, 2017

ELAEOCARPACEAE Calliandra houstoniana (Mill.) Standl. Shrub; 1184 m; 6 (1431). Sloanea cruenta Lundell Canavalia glabra (M.Martens & Galeotti) J.D.Sauer Tree; 1505–1600 m; 17 (504, 1745, 1748), 18 Herbaceous vine; 1290 m; 7 (1409). (1773A). Centrosema sp. ERICACEAE Herbaceous vine; 1262 m; 7 (1399). Cojoba arborea (L.) Britton & Rose Arbutus xalapensis Kunth Tree; 1474 m; 14 (962). Tree; 2473 m; 2 (1689). Crotalaria sp. Comarostaphylis discolor (Hook.) Diggs Tree; 1300 m; 7 (1954). Sierra de los Frailes Area, specific locality not Desmodium helleri Peyr. disclosed; species is protected under Mexican law. Herbaceous vine; 1185–1278 m; 6 (831, 2114), 7 Gaultheria acuminata Schltdl. & Cham. (1898). Tree; 1185 m; 6 (849). Desmodium intortum (Mill.) Urb. Gaultheria erecta Vent. Herbaceous vine; 1233 m; 7 (1398). Tree; 2170–2725 m; 1 (1904), 2 (1145, 1161), 3 Desmodium strobilaceum Schltdl. (2017, 2067), 4 (2246, 2288). Herb; 1233 m; 7 (1397). Lyonia squamulosa M.Martens & Galeotti Erythrina sp. Shrub; 2151–2325 m; 4 (2253), 10 (616, 1095). Tree; 7 (1207). Macleania insignis M.Martens & Galeotti Indigofera thibaudiana DC. Epiphytic woody vine; 1184–2321 m; 4 (2290), 6 Tree; 1233 m; 7 (1396). (919), 13 (2156), 14 (630), 15 (947), 19 (1830). Inga acrocephala Steud. Pernettya prostrata (Cav.) DC. Tree; 1320 m; 7 (1510). Shrub; 2725–2742 m; 2 (1152, 1718). Inga tuerckeimii Pittier Vaccinium confertum Kunth Tree; 1315 m; 12 (1372). Shrub; 2725 m; 2 (1148). Inga vera Willd. Vaccinium leucanthum Schltdl. Tree; 1229 m; 9 (1021). Tree; 1737–2325 m; 3 (2074), 4 (2255, 2291), 5 Leucaena diversifolia (Schltdl.) Benth. (1619), 10 (620, 1073), 11 (1012). Tree; 1570 m; 11 (2013). Vaccinium matudae Lundell Lupinus sp. Epiphytic shrub; 2321 m; 4 (2293). Shrub; 2203 m; 4 (1658). Lysiloma auritum (Schltdl.) Benth. EUPHORBIACEAE Tree; 1320 m; 8 (1440). Acalypha cf. longipes S.Watson Mimosa albida Humb. & Bonpl. ex Willd. Herb; 1185–1226 m; 6 (846, 848), 13 (2132), 16 Shrub; 1270 m; 6 (1270). (661). Phaseolus chiapasanus Piper Alchornea costaricensis Pax & K.Hoffm. Woody vine; 1290 m; 7 (1415). Tree; 1250 m; 6 (1263). Phaseolus glabellus Piper Alchornea latifolia Sw. Woody vine; 1184–1290 m; 6 (1551), 7 (1414). Tree; 1070–1415 m; 6 (2318), 9 (1041), 12 (996), Senna pallida (Vahl) H.S.Irwin & Barneby var. 13 (1213). trichocraspedon (Sandwith) H.S.Irwin & Barneby Croton draco Schltdl. Tree; 1233–1320 m; 7 (1393), 8 (1437). Tree; 6 (608), 7 (1888). FAGACEAE Euphorbia nutans Lag. Quercus candicans Nee´ Herb; 1229 m; 9 (1045). Tree; 1300 m; 7 (1420). Euphorbia xalapensis Kunth Quercus corrugata Hook. Herb; 1226 m; 13 (1363, 2138). Tree; 2300 m; 1 (1471). Gymnanthes riparia (Schltdl.) Klotzsch Quercus aff. corrugata Hook. Tree; 1226 m; 13 (2159). Tree; 2340 m; 4 (1675). FABACEAE Quercus depressa Bonpl. NUMBER 20 MUNN-ESTRADA: FLORA OF THE SIERRA MAZATECA OF OAXACA, MEXICO 51

Shrub; 2725 m; 2 (1150). HYPERICACEAE Quercus elliptica Nee´ Hypericum sp. Tree; 1930 m; 10 (615). Herb; 1424 m; 19 (449). Quercus eugeniifolia Liebm. Vismia baccifera (L.) Triana & Planch. Tree; 7 (1189). Tree; 1250 m; 17 (430). Quercus lancifolia Schltdl. & Cham. Tree; 1300 m; 7 (1141). HYPOXIDACEAE Quercus ocoteifolia Liebm. Hypoxis decumbens L. Tree; 1450–2450 m; 1 (1450), 7 (1434), 12 (1391). Herb, 1185–1424 m; 6 (2107), 19 (446). Quercus aff. ocoteifolia Liebm. Tree; 1184 m; 6 (920). IRIDACEAE Quercus polymorpha Schltdl. & Cham. Tree; 1320–1320 m; 7 (1187), 8 (1438, 1441). Crocosmia 3 crocosmiiflora (Lemoine) N.E.Br. Quercus sartorii Liebm. Herb, 1100–1850 m, 10 (613), 20 (404). Tree; 1229–1250 m; 6 (1268), 9 (1043). JUGLANDACEAE Quercus scytophylla Liebm. Tree; 2320 m; 1 (1457). Juglans mollis Engelm. Tree; 1226–1589 m; 12 (1180), 13 (2187). GARRYACEAE Oreomunnea mexicana (Standl.) J.F.Leroy Garrya laurifolia Hartw. ex Benth. Tree; 1560 m; 17 (508). Shrub; 2223–2223 m; 3 (2086, 2087). LAMIACEAE Garrya aff. laurifolia Hartw. ex Benth. Tree; 1570 m; 11 (2012). Cornutia pyramidata L. Tree; 1323–1408 m; 8 (1210, 1297). GENTIANACEAE Holmskioldia sanguinea Retz. Lisianthius nigrescens Schltdl. & Cham. Herb; 1180 m; 16 (703). Herb; 1226 m; 13 (1307). Hyptis lantanifolia Poit. Lisianthius quichensis Donn.Sm. Herb; 1229 m; 9 (1323). Shrub; 1275–1709 m; 7 (1893), 14 (977), 15 (945). Hyptis mutabilis (Rich.) Briq. Herb; 1184–1458 m; 6 (796), 14 (1490), 19 (407). GERANIACEAE Salvia sp. Geranium sp. Herb; 1229–1460 m; 7 (1123), 9 (1057), 12 (1383). Herb; 2242 m; 2 (2343). Scutellaria sp. Herb; 2400 m; 2 (1341). GESNERIACEAE Stachys sp. Achimenes grandiflora (Schiede) DC. Herb; 2151–2640 m; 2 (1157), 10 (1087). Herb; 1184 m; 6 (1369). LAURACEAE Columnea schiedeana Schltdl. Epiphytic woody vine; 1070–1226 m; 6 (929), 13 Cinnamomum effusum (Meisn.) Kosterm. (1211), 13 (2137). Tree; 1184 m; 1 (2224), 6 (923). Moussonia deppeana (Schltdl. & Cham.) Klotzsch ex Damburneya salicifolia (Kunth) Trofimov & Rohwer Hanst. Tree; 1233 m; 6 (1097). Shrub; 1180–2328 m; 3 (2079), 4 (1661), 5 (1624), Litsea glaucescens Kunth 7(2048), 16 (658), 17 (431), 19 (1846), 20 (402). Tree; 1185 m; 6 (853). Smithiantha multiflora (M.Martens & Galeotti) Ocotea bernoulliana Mez Fritsch Tree; 1184–1323 m; 6 (911), 7 (1933). Herb; 1184–1226 m; 6 (1370), 13 (1361), 20 (400). Ocotea betazensis (Mez) van der Werff Tree; 1404–2335 m; 1 (1912, 2222), 2 (2212, HELICONIACEAE 2238), 19 (1839). Heliconia tortuosa Griggs Ocotea disjuncta Lorea-Hern. Herb; 1100–1200; 16 (673). Tree; 1 (2217). 52 LUNDELLIA DECEMBER, 2017

Persea americana Mill. Tree; 1275 m; 7 (1895). Tree; 1589–1673 m; 9 (1853), 11 (2008). Malvaviscus achanioides (Turcz.) Fryxell Persea pallescens (Mez) Lorea-Hern. Shrub; 1424 m; 19 (472). Shrub; 2170 m; 3 (2018). Malvaviscus arboreus Cav. Persea schiedeana Nees Herb; 1180 m; 16 (695). Tree; 1180–1375 m; 13 (940), 19 (1820). Pavonia schiedeana Steud. Herb; 1424–1560 m; 17 (539), 19 (469). LENTIBULARIACEAE Pavonia uniflora (Sesse´ & Moc.) Fryxell Pinguicula macrophylla Kunth Shrub; 1226 m; 13 (1308A). Herb; 1229 m; 9 (1070), 10 (614). Sida glabra Mill. Herb; 1300 m; 7 (1142). LINACEAE Sida rhombifolia L. Linum nelsonii Rose Herb; 1185–1229 m; 6 (2111), 9 (1322), 16 (694). Herb; 1185 m; 6 (856). Trichospermum galeottii (Turcz.) Kosterm. Tree; 1180 m; 16 (717) LORANTHACEAE Triumfetta bogotensis DC. Psittacanthus ramiflorus (Moc. & Sesse´ ex DC.) Herb; 1184–1424 m; 6 (1433B), 17 (563), 19 (468) G.Don Triumfetta grandiflora Vahl Shrub; 1474 m; 14 (648). Tree; 1180 m; 13 (941)

LYTHRACEAE MARCGRAVIACEAE

Cuphea sp. Marcgravia stonei Utley Herb; 1185 m; 6 (835). Shrub; 1424–1560 m; 17 (531), 19 (463).

MAGNOLIACEAE MELASTOMATACEAE

Magnolia dealbata Zucc. Arthrostemma ciliatum Pav. ex D.Don Sierra de los Frailes Area, specific locality not Herb; 1458 m; 14 (989). disclosed; species is endangered and protected Centradenia grandifolia (Schltdl.) Endl. under Mexican law. Herb; 1474 m; 14 (1278). Magnolia schiedeana Schltdl. Conostegia arborea Steud. Sierra de los Frailes Area, specific locality not Tree; 1589 m; 11 (1004). disclosed; species is endangered and protected Conostegia xalapensis (Bonpl.) D.Don ex DC. under Mexican law. Tree; 1458 m; 14 (990). Heterocentron subtriplinervium (Link & Otto) MALPIGHIACEAE A.Braun & C.D.Bouche´ Bunchosia lindeniana A.Juss. sens. lat. Shrub; 1250 m; 6 (1265). Shrub; 1360 m; 18 (1813), 21 (766) Miconia anisotricha (Schltdl.) Triana Shrub; 2223–2300 m; 3 (2093). MALVACEAE Miconia costaricensis Cogn. Anoda cristata (L.) Schltdl. Shrub; 1645 m; 17 (1756). Herb; 1185 m; 6 (2113), 20 (581). Miconia glaberrima (Schltdl.) Naudin Anoda cf. cristata (L.) Schltdl. Tree; 1330–1381 m; 12 (1375). Herb; 1184 m; 6 (915). Miconia globulifera Naudin Hampea integerrima Schltdl. Shrub; 1381 m; 12 (997). Tree; 1460 m; 12 (1380). Miconia hemenostigma Naudin Heliocarpus americanus L. Tree; 2151–2300 m; 1 (1476), 3 (2068), 10 (1077), Tree; 1342–1380 m; 7 (1955), 9 (1864). 10 (1094). Heliocarpus appendiculatus Turcz Miconia lonchophylla Naudin Tree; 1233–1513 m; 6 (812, 818), 7 (1883), 9 Tree; 1645 m; 17 (1765). (1863), 18 (1809), 19 (1842). Miconia mazatecana de Santiago Heliocarpus sp. Tree; 1709 m; 15 (944). NUMBER 20 MUNN-ESTRADA: FLORA OF THE SIERRA MAZATECA OF OAXACA, MEXICO 53

Miconia sylvatica (Schltdl.) Naudin Tree; 1483 m; 14 (1540). Shrub; 1229 m; 9 (1049). Fuchsia microphylla Kunth Monochaetum floribundum (Schltdl.) Naudin Shrub; 2325–2640 m; 2 (1159), 4 (1662, 2256). Shrub; 1200–1424 m; 9 (2370), 19 (1832, 1835, Fuchsia paniculata Lindl. 1847). Tree; 1513 m; 18 (1808), 21 (755). Tibouchina longifolia (Vahl) Baill Lopezia racemosa Cav. Shrub; 1604 m; 11 (2005). Herb; 1185–1658 m; 6 (840, 2120), 11 (1984), 17 Tibouchina scabriuscula (Schltdl.) Cogn. (421), 19 (471). Shrub; 2223 m; 3 (2084). ORCHIDACEAE Topobea laevigata (D.Don) Naudin Tree; 1184–1513 m; 6 (903, 869A), 18 (1807). Aulosepalum pyramidale (Lindl.) M.A.Dix & M.W.Dix MELIACEAE Terrestrial; 1226 m; 13 (2182). Trichilia havanensis Jacq. Beloglottis costaricensis (Rchb.f.) Schltr. Tree; 1185–1637 m; 2 (2239), 6 (837, 2051), 11 Epiphytic herb; 1200 m; 7 (1098). (1997). Brassia verrucosa Lindl. Epiphytic herb; 1450 m; 7 (1199). MENISPERMACEAE Calanthe calanthoides (A.Rich. & Galeotti) Hamer & Cissampelos pareira L. Garay Woody vine; 1424 m; 19 (487). Terrestrial; 1474 m; 14 (632). METTHENIUSACEAE Campylocentrum schiedei (Rchb.f.) Benth ex Hemsl. Epiphytic herb; 1184–1200 m; 6 (793), 7 (1110). Oecopetalum mexicanum Greenm. & C.H.Thomps. Coelia macrostachya Lindl. Tree; 1184 m; 6 (910). Epiphytic herb; 21 (764). MONIMIACEAE Coelia triptera (Sm.) G.Don ex Steud. Epiphytic herb; 1226 m; 13 (2147). Mollinedia viridiflora Tul. Comparettia falcata Poepp. & Endl. Tree; 1184–1474 m; 6 (909, 912), 14 (1280). Epiphytic herb; 1474 m; 9 (607), 14 (633). MORACEAE Dichaea glauca (Sw.) Lindl. Epiphytic herb; 7 (1200). Ficus sp. Dichaea muricatoides Hamer & Garay Tree; 1270–1350 m; 6 (1269), 7 (1258). Epiphytic herb; 1180 m; 16 (666). Trophis mexicana (Liebm.) Bureau Dichaea suaveolens Kraenzl. Herb; 1250–1300 m; 17 (411), 7 (1131). Epiphytic herb; 1474 m; 14 (1287). MYRTACEAE Elleanthus cynarocephalus (Rchb.f.) Rchb.f. Epiphytic herb; 1180–1424 m; 16 (659), 19 Eugenia sp. (1850). Tree; 1184 m; 6 (899). Epidendrum laucheanum Bonhof ex Rolfe Ugni myricoides (Kunth) O.Berg Epiphytic herb; 21 (744). Tree; 2611 m; 2 (1705). Epidendrum polyanthum Lindl. NYCTAGINACEAE Epiphytic herb; 1458 m; 14 (622). Epidendrum cf. polyanthum Lindl. Mirabilis longiflora L. Epiphytic herb; 1474 m; 14 (1287A). Herb; 1320 m; 8 (1439). Epidendrum radicans Pav. ex Lindl. Epiphytic herb; 1229–1380 m; 7 (1975), 9 (1055). OLEACEAE Epidendrum ramosum Jacq. Osmanthus americanus (L.) A.Gray Epiphytic herb; 1424 m; 19 (473). Tree; 2302–2340 m; 1 (1454, 1465). Habenaria cf. distans Griseb. Terrestrial; 1250 m; 17 (427). ONAGRACEAE Isochilus oaxacanus Salazar & Soto Arenas Fuchsia arborescens Sims Epiphytic herb; 2450 m; 2 (1163). 54 LUNDELLIA DECEMBER, 2017

Jacquiniella leucomelana (Rchb.f.) Schltr. Rhynchostele rossii (Lindl.) Soto Arenas & Salazar Epiphytic herb; 1560 m; 17 (543). Sierra de los Frailes Area, specific locality not Lepanthes disticha Garay & R.E.Schult. disclosed; species is endangered and protected Epiphytic herb; 1100–1555 m; 21 (745). under Mexican law. Lepanthes rekoi R.E.Schult. Sobralia macrantha Lindl. Epiphytic herb; 2242 m; 2 (2322). Terrestrial; 1229–1673 m; 9 (611). Lycaste aromatica (Graham) Lindl. Stanhopea tigrina Bateman ex Lindl. Epiphytic herb; 1069 m; 7 (1218). Huautla–Chilchotla Area, specific locality not Malaxis histionantha (Link) Garay & Dunst. disclosed; species is endangered and protected Epiphytic herb; 1350 m; 17 (492). under Mexican law. Maxillaria cucullata Lindl. Stelis argentata Lindl. Epiphytic herb; 21 (763). (Stelis endresii Rchb.f) Maxillaria densa Lindl. Epiphytic herb; 1184–1474 m; 6 (654A), 14 (644). Epiphytic herb; 1226–1352 m; 7 (1959), 13 Stelis cobanensis (Schltr.) Pridgeon & M.W.Chase (2158). Huautla–Chilchotla Area, specific locality not Maxillaria variabilis Bateman ex Lindl. disclosed; species is protected under Mexican law. Epiphytic herb; 1226–1226 m; 13 (2155), 21 Stelis ornata (Rchb.f.) Pridgeon & M.W.Chase (751). Epiphytic herb; 1184 m; 6 (792). Nidema boothii (Lindl.) Schltr. Stelis purpurascens A.Rich. & Galeotti Epiphytic herb; 1226–1352 m; 6 (904), 7 (1960), Epiphytic herb; 1150–1226 m; 6 (1872), 13 13 (2167). (2157). Oestlundia luteorosea (A.Rich. & Galeotti) Stelis rubens Schltr. W.E.Higgins Epiphytic herb; 1184 m; 6 (654). Epiphytic herb; 1240 m; 6 (1261). Stelis veracrucensis Solano Platanthera vulcanica Lindl. Epiphytic herb; 2151 m; 10 (1081). (Platanthera limosa Lindl.) Trichocentrum pachyphyllum (Hook.) R.Jimenez´ & Terrestrial; 2313–2742 m; 2 (1715), 5 (1627). Carnevali Pleurothallis cardiothallis Rchb.f. Epiphytic herb; 1327 m; 8 (2055). Epiphytic herb; 1424–1560 m; 17 (1733), 19 (465). OROBANCHACEAE Prosthechea cochleata (L.) W.E.Higgins Castilleja integrifolia L.f. Epiphytic herb; 1069–1458 m; 7 (1217, 1972A), 14 Shrub; 1819 m; 2 (2192), 11 (1015). (627). Conopholis alpina Liebm. Prosthechea ochracea (Lindl.) W.E.Higgins Epiphytic herb; 7 (1198). Herb; 2223–2345 m; 1 (1941), 2 (2208), 3 (2071), Prosthechea pseudopygmaea (Finet) W.E.Higgins 4(2274). Epiphytic herb; 1424–1474 m; 14 (645), 19 (439). Lamourouxia dasyantha (Cham. & Schltdl.) Prosthechea varicosa (Bateman ex Lindl.) W.E.Higgins W.R.Ernst Epiphytic herb; 2313–2473 m; 2 (1687), 5 (1630). Herb; 1226 m; 13 (1317). Prosthechea vitellina (Lindl.) W.E.Higgins Lamourouxia xalapensis Kunth Sierra de los Frailes Area, specific locality not Herb; 2328 m; 4 (1664). disclosed; species is protected under Mexican law. Seymeria mazatecana B.L. Turner Rhynchostele beloglossa (Rchb.f.) Dressler & Herb; 2640 m; 2 (1351). N.H.Williams OXALIDACEAE Sierra de los Frailes Area, specific locality not disclosed; species is endangered and protected Oxalis latifolia Kunth under Mexican law. Herb; 2369 m; 2 (1338). Rhynchostele cordata (Lindl.) Soto Arenas & Salazar PAPAVERACEAE Sierra de los Frailes Area, specific locality not disclosed; species is endangered and protected Bocconia frutescens L. under Mexican law. Tree; 1290 m; 7 (1418). NUMBER 20 MUNN-ESTRADA: FLORA OF THE SIERRA MAZATECA OF OAXACA, MEXICO 55

PASSIFLORACEAE Peperomia tetraphylla (G.Forst.) Hook. & Arn. Epiphytic herb; 1226–1645 m; 11 (1008), 13 Passiflora sp. (2177), 17 (1768), 18 (1778). Woody vine; 1100–1426 m; 1 (2257), 7 (1248, Piper amalago L. 1403, 1411, 1423), 9 (2351), 16 (699). Shrub; 1184–1327 m; 6 (901), 8 (2058), 9 (1062), PENTAPHYLACACEAE 16 (685). Piper hispidum Sw. Cleyera integrifolia (Benth.) Choisy Shrub; 1229–1515 m; 9 (1058), 18 (1786). Tree; 2320–2340 m; 1 (1449, 1944). Piper martensianum C.DC. Ternstroemia tepezapote Schltdl. & Cham. Shrub; 1184 m; 6 (913). Tree; 2167–2640 m; 1 (1455, 1945), 2 (1332, Piper obliquum Ruız´ & Pav. 1348), 3 (2023, 2078), 4 (2295), 5 (1618). Tree; 1474 m; 14 (1279). PHYLLANTHACEAE Piper umbellatum L. Herb; 1560 m; 17 (547). Phyllanthus niruri L. Piper yzabalanum C.DC. ex Donn.Sm. Herb; 1250–1290 m; 7 (1407), 17 (432). Shrub; 1184–1424 m; 6 (995), 7 (1920), 19 Phyllanthus aff. purpusii Brandegee (476). Tree; 1560–1645 m; 17 (520, 1764). PLANTAGINACEAE PHYLLONOMACEAE Lophospermum erubescens D.Don Phyllonoma laticuspis (Turcz.) Engl. Herb; 1290–1458 m; 7 (1190, 1412), 8 (1238), 14 Tree; 2151–2340 m; 1 (1448), 2 (2215), 10 (1080). (629). PHYTOLACCACEAE Plantago australis Lam. subsp. hirtella (Kunth) Rahn Herb; 1185–1458 m; 6 (2106), 14 (991), 17 (413). Phytolacca rugosa A.Braun & C.D.Bouche´ Shrub; 2151 m; 10 (1086). PLATANACEAE

PICRAMNIACEAE Platanus mexicana Moric. var. mexicana Tree; 1185–1412 m; 6 (890, 2308), 7 (2039), 8 Picramnia antidesma Sw. (719). Tree; 1323–1375 m; 7 (1927), 19 (1831). POACEAE PIPERACEAE Agrostis semiverticillata (Forssk.) C.Chr. Peperomia angustata Kunth Herb; 1150 m; 6 (1871). (Polypogon viridis (Gouan) Breistr.) Peperomia dendrophila Schltdl. & Cham. Herb; 1226 m; 13 (2161). Herb; 1184–1300 m; 6 (859, 925), 7 (1130, Arundinella deppeana Ness 1111A). Herb; 1185–1300 m; 6 (844, 851, 878, 2119), 7 Peperomia galioides Kunth (1144). Epiphytic herb; 2328–2487 m; 2 (1684), 4 (1665). Cynodon dactylon (L.) Pers. Peperomia hernandiifolia (Vahl) A.Dietr. Herb; 1229 m; 9 (1051). Herb; 1600 m; 17 (1744). Dichanthelium laxiflorum (Lam.) Gould Peperomia liebmannii C.DC. Herb; 2223 m; 3 (2085). Herb; 1460 m; 12 (1384). Isachne arundinacea (Sw.) Griseb. Peperomia magnoliifolia (Jacq.) A.Dietr. Herbaceous vine; 1229–1466 m; 7 (1922), 9 Herb; 1460–1560 m; 12 (1385), 17 (1737). (1027), 14 (976). Peperomia obtusifolia (L.) A.Dietr. Lasiacis divaricata (L.) Hitchc. Herb; 16 (684). Herb; 21 (772). Peperomia rotundifolia (L.) Kunth Lasiacis procerrima (Hack.) Hitchc. Epiphytic herb; 1233 m; 6 (1176). Herb; 1300 m; 7 (1424). Peperomia tenerrima Schltdl. & Cham. Muhlenbergia sp. Epiphytic herb; 2220 m; 2 (2206), 3 (2029). Herb; 1197 m; 12 (1637). 56 LUNDELLIA DECEMBER, 2017

Oplismenus hirtellus (L.) P.Beauv. subsp. setarius Parathesis rekoi Standl. (Lam.) Mez ex Ekman Tree; 2300 m; 1 (1477). Herb; 1424 m; 19 (444). RANUNCULACEAE Otatea aztecorum (McClure & E.W.Sm.) C.E. Calderon´ ex Soderstr. Anemone mexicana Kunth Herb; 2333 m; 5 (1629). Herb; 21 (780). Paspalum convexum Humb. & Bonpl. ex Flugg¨ e´ Clematis grossa Benth. Herb; 1229 m; 9 (1054). Woody vine; 1323–1372 m; 7 (1924), 9 (1862). Paspalum langei (E.Fourn.) Nash Ranunculus petiolaris Humb., Bonpl. & Kunth ex DC. Herb; 1184 m; 6 (1365). Herb; 1601 m; 9 (1166). Sporobolus indicus (L.) R.Br. RHAMNACEAE Herb; 1229 m; 9 (1032). Frangula capreifolia (Schltdl.) Grubov POLEMONIACEAE Tree; 1229–2345 m; 4 (2280), 6 (1264), 9 (1067), Cobaea biaurita Standl. 12 (1000), 12 (1183). Herbaceous vine; 21 (783). ROSACEAE POLYGALACEAE Alchemilla pectinata Kunth Monnina xalapensis Kunth Herb; 2640 m; 2 (1158). Shrub; 2151–2313 m; 3 (2075), 5 (1625), 10 Cercocarpus pringlei (C.K.Schneid.) Rydb. (1083). Tree; 1270 m; 6 (1271), 7 (1203). Polygala paniculata L. Prunus brachybotrya Zucc. Herb; 1233–1424 m; 7 (1394), 19 (445). Tree; 1571–2320 m; 1 (1943), 11 (2001). Securidaca diversifolia (L.) S.F.Blake Prunus lundelliana Standl. Shrub; 1323 m; 8 (1300). Tree; 1323 m; 8 (1241). Prunus matudae Lundell POLYGONACEAE Tree; 1 (2219). Coccoloba hirtella Lundell Rubus fagifolius Schltdl. & Cham. Tree; 1226–1515 m; 13 (2174), 14 (964), 18 Woody vine; 1513 m; 18 (1802). (1772). Rubus glaucus Benth. Woody vine; 1513 m; 18 (1805). PRIMULACEAE Rubus sapidus Schltdl. Anagallis arvensis L. Woody vine; 1229 m; 9 (1056). Herb; 1185 m; 6 (845). RUBIACEAE Ardisia liebmannii Oerst. Shrub; 1474–1645 m; 14 (650, 1538), 17 (517, Arachnothryx heteranthera (Brand.) Borhidi 1760). Tree; 1515 m; 18 (1771). Ardisia verapazensis Donn.Sm. Arachnothryx ovadensis (Lundell) Borhidi Tree; 1323–1474 m; 8 (1301), 14 (640, 1282), 19 Shrub; 6 (1294), 18 (1794) (454). Bouvardia ternifolia (Cav.) Schltdl. Myrsine coriacea (Sw.) R.Br. ex Roem. & Schult. Herb; 1229 m; 9 (1063). Tree; 1229 m; 9 (606), 9 (1050). Chiococca alba (L.) Hitchc. Myrsine juergensenii (Mez) Ricketson & Pipoly Herbaceous vine; 1180 m; 16 (670). Tree; 2223–2320 m; 1 (1459), 2 (2198), 3 (2082). Chiococca phaenostemon Schltdl. Parathesis leptopa Lundell Tree; 1 (2227). Tree; 1505 m; 17 (503). Chomelia brachypoda Donn.Sm. Parathesis macronema Bullock Tree; 1300 m; 7 (1137). 1320–1350 m; 7 (1259, 1508), 17 (497). Crusea calocephala DC. Parathesis melanosticta (Schtdl.) Hemsl. Herb; 1180–1601 m; 9 (1168, 2357), 14 (988), 16 Tree; 1320–2335 m; 1 (1472, 1918, 1938), 2 (692), 17 (416). (2234), 3 (2073), 17 (1727). Crusea coccinea DC. NUMBER 20 MUNN-ESTRADA: FLORA OF THE SIERRA MAZATECA OF OAXACA, MEXICO 57

Herb; 2151–2325 m; 4 (2258), 10 (1078). Rogiera edwardsii (Standl.) Borhidi Deppea erythrorhiza Schltdl. & Cham. Tree; 1350 m; 7 (1253). Tree; 1200 m; 7 (1101). Rogiera stenosiphon (Hemsl.) Borhidi Deppea grandiflora Schltdl. Tree; 1185 m; 6 (832). Shrub; 1327–2335 m; 1 (1908, 2221), 8 (2062), 11 Sommera arborescens Schltdl. (1010, 1992), 14 (967), 15 (948). Tree; 1184–1483 m; 6 (908), 7 (1102, 1247), 14 Deppea scoti (J.H.Kirkbr.) Lorence (1535). (Bellizinca scoti (J.H.Kirkbr.) Borhidi) Spermacoce assurgens Ruız´ & Pav. Shrub; 2223 m; 3 (2097). (Spermacoce remota Lam.) Galium hypocarpium (L.) Endl. ex Griseb. Herb; 1184 m; 6 (1427). Herb; 2320 m; 1 (1461). Spermacoce remota Lam. Glossostipula concinna (Standl.) Lorence Shrub; 1226 m; 13 (2151). Tree; 1184–1513 m; 6 (895), 7 (1132), 14 (971), 16 RUTACEAE (704). Hamelia calycosa Donn.Sm. Citrus sp. Tree; 1226 m; 13 (1314) Tree; 1184 m; 6 (928). Hedyotis exigula W.H.Lewis Amyris aff. attenuata Standl. (Mexotis latifolia (M.Martens & Galeotti) Terrell Tree; 1180 m; 16 (706). & H.Rob.) Herb; 1185–1327 m; 6 (839), 8 (2057), 9(610, SALICACEAE 1040), 13 (2168). Hasseltiopsis dioica (Benth.) Sleumer Hoffmannia nicotianifolia (M.Martens & Galeotti) Tree; 1184 m; 6 (917) L.O.Williams Salix paradoxa Kunth Shrub; 1270–1474 m; 2 (2196), 7 (1499), 14 (641), Shrub; 2223 m; 3 (2101), 3 (2102). 18 (1801), 19 (452, 453, 1845), 21 (767). Xylosma quichensis Donn.Sm. Hoffmannia psychotrifolia (Benth.) Griseb. Tree; 2242–2742 m; 2 (2193). Shrub; 1323 m; 8 (1305). Hoffmannia aff. psychotrifolia (Benth.) Griseb. SAPINDACEAE Tree; 1560 m; 17 (515). Serjania flaviflora Radlk. Notopleura hondurensis C.M.Taylor Woody vine; 1185–1203 m; 6 (932), 6 (2108). Herb; 1424–1560 m; 17 (1722), 19 (457). Palicourea padifolia (Willd. ex Roem. & Schult.) SAPOTACEAE C.M.Taylor & Lorence Sideroxylon persimile (Hemsl.) T.D.Penn subsp. Tree; 1300–1560 m; 7 (1143), 14 (983), 17 (518, persimile 527), 18 (1787). Tree; 1323 m; 8 (1303). Posoqueria coriacea M.Martens & Galeotti Tree; 1316 m; 8 (1445). SCHOEPFIACEAE Psychotria elata (Sw.) Hammel Schoepfia schreberi J.F.Gmel. Tree; 1180 m; 16 (667). Tree; 1233 m; 7 (1885), 7 (1886). Psychotria fruticetorum Standl. Schoepfia vacciniiflora Planch. ex Hemsl. Shrub; 1483–1589 m; 11 (1005), 14 (1528). Tree; 1184–1352 m; 6 (927), 7 (1962). Psychotria galeottiana (M.Martens) C.M.Taylor & Lorence SCROPHULARIACEAE Tree; 1600–2320 m; 1 (1464), 17 (1751). Buddleja americana L. Psychotria mexiae Standl. Tree; 1673 m; 9 (1854). Tree; 1320–1560 m; 7 (1251, 1511), 12 (1378), 14 Russelia coccinea (L.) Wettst. (1533), 17 (1728), 18 (1788, 1795). Herb; 1266–1270 m; 6 (810), 6 (1273). Psychotria sarapiquiensis Standl. Russelia ternifolia Kunth Tree; 1300–1560 m; 7 (1136), 17 (1724, 1736). Herb; 1185 m; 6 (867). Randia matudae Lorence & Dwyer Tree; 1560 m; 17 (505). SMILACACEAE 58 LUNDELLIA DECEMBER, 2017

Smilax domingensis Willd. Symplocos coccinea Bonpl. Herbaceous vine; 1226–2325 m; 4 (2268), 13 Sierra de los Frailes Area, specific locality not (2153). disclosed; species is protected under Mexican Smilax glauca Walter law. Herbaceous vine; 1229 m; 9 (1064). Symplocos limoncillo Bonpl. Smilax subpubescens A.DC. Tree; 1069–1280 m; 6 (1260), 7 (1204), 7 (1219), 7 Herbaceous vine; 1229–1300 m; 7 (1135), 9 (1401). (1072). ULMACEAE SOLANACEAE Trema micrantha (L.) Blume Cestrum elegans (Brongn.) Schltdl. Tree; 1380 m; 7 (1957). Tree; 1460–1530 m; 12 (1003, 1387). Ulmus mexicana (Liebm.) Planch. Cestrum fasciculatum (Schltdl.) Miers Tree; 1185 m; 6 (882, 884, 2117). Shrub; 2335 m; 1 (1917). URTICACEAE Cestrum laxum Benth. Shrub; 2327 m; 4 (1672). Cecropia obtusifolia Bertol. Cestrum nocturnum L. Tree; 7 (1186A). Tree; 1184 m; 6 (1320). Myriocarpa longipes Liebm. Lycianthes anomala Bitter Shrub; 1185–1658 m; 6 (829, 1880), 11 (1990), 18 Tree; 1226 m; 13 (1315). (1792, 1803, 1816). Physalis sp. Pilea hyalina Fenzl Herb; 1184 m; 6 (914). Herb; 1323 m; 8 (1304). Schraderanthus viscosus (Schrad.) Averett Pilea microphylla (L.) Liebm. Tree; 2325 m; 4 (2263). Herb; 1250 m; 7 (1193), 17 (438). Solanum aligerum Schltdl. Pilea pubescens Liebm. Tree; 2300 m; 1 (1474). Herb; 1250 m; 17 (425), 21 (781). Solanum americanum Mill. Pilea tridentata Killip Herb; 1185 m; 6 (841). Herb; 1424 m; 19 (458). Solanum aphyodendron S.Knapp Urera caracasana (Jacq.) Gaudich. ex Griseb. Shrub; 1185–1431 m; 6 (838), 7 (1244), 9 (1061). Tree; 1226 m; 13 (1312). Solanum appendiculatum Dunal VERBENACEAE Woody vine; 1184 m; 6 (1546A). Solanum chrysotrichum Schltdl. Citharexylum mocinnoi D.Don Tree; 1069–1184 m; 6 (894), 7 (1216). Tree; 1266–2259 m; 3 (1486), 6 (811, 816), 8 Solanum lanceolatum Cav. (1239), 9 (1852), 19 (1818). Tree; 2300 m; 1 (1475). Lantana camara L. Solanum nigricans M.Martens & Galeotti Herb; 1229–1460 m; 9 (1024), 12 (1379). Tree; 1381–2314 m; 2 (1333), 10 (1092), 11 Lippia myriocephala Schltdl. & Cham. Tree; 1184–1460 m; 6 (794), 9 (1037), 12 (1389), (1011), 12 (1001B). 14 (1492), 19 (406). STAPHYLEACEAE Verbena carolina L. Herb; 1229–1350 m; 9 (1047), 17 (559). Turpinia insignis (Kunth) Tulasne Tree; 1278–1323 m; 7 (1897, 1925). VIOLACEAE Turpinia occidentalis (Sw.) G.Don Viola scandens Humb. & Bonpl. ex Schult. Tree; 2204 m; 3 (2033), 4 (2275). Herb; 2151 m; 10 (1084). STYRACACEAE VITACEAE Styrax ramirezii Greenm. Vitis popenoei Fennell Tree; 2300–2320 m; 1 (1458, 1473, 1935, 1936). Woody vine; 1290–1320 m; 7 (1417, 1506). SYMPLOCACEAE Vitis tiliifolia Humb. & Bonpl. ex Roem. & Schult. NUMBER 20 MUNN-ESTRADA: FLORA OF THE SIERRA MAZATECA OF OAXACA, MEXICO 59

Woody vine; 1229 m; 7 (2047), 9 (1046).

WINTERACEAE

Drimys granadensis L.f. var. mexicana (DC.) A.C.Sm. Tree; 2300 m; 1 (1482).

ZINGIBERACEAE

Renealmia mexicana Klotzsch ex Petersen Herb; 1180–1350 m; 7 (1504), 16 (675).