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Botanical Sciences

On-line version ISSN 2007-4476Print version ISSN 2007-4298

Bot. sci vol.95 n.3 México Jul./Sep. 2017 

Taxonomy and floristics

Floristic composition of the cloud forest of the Bicentenario Reserve, Zongolica, Veracruz, México

Luis Alfonso Castillo-Hernández1 

Hilda Flores-Olvera1  * 

1 Departamento de Botánica. Instituto de Biología. Universidad Nacional Autónoma de México, Ciudad de México, México



The Bicentenario Reserve (BR) located in Sierra de Zongolica, Veracruz, includes 63 hectares of cloud forest (cf) which lacks of systematic floristic studies, but the Sierra proposed as an area for bird conservation.


i) Is the floristic composition of the BR taxonomical rich? ii) How the growth forms are represented in this Reserve ? iii) Has the BR endemic or threatened species?

Studied species:

Vascular plants.

Study site and years of study:

The BR was explored from March 2011 to October 2012.


Botanical samples from each species considered different were collected and processed according to conventional procedures to be identified with the use of taxonomic tools, and consults with specialist. Analysis of the richness, life forms, endemism and threatened species were made.


We recorded 401 species, distributed in 272 genera and 102 families, being the most diverse Orchidaceae, Asteraceae, Fabaceae and Piperaceae; whilst Peperomia, Tillandsia, Polypodium, Quercus and Solanum are the genera with the highest number of species. Sixty-nine species are endemic to Mexico, but six are restricted to Veracruz. We found 23 new records for the municipality of Zongolica, Quercus ghiesbreghtii in the cf for the first time, and Q. furfuraceae for Veracruz; 86 species are threatened.


The BR has a high floristic richness compared to other areas with cf in Mexico, revealing the importance to complete the floristic studies along the Sierra de Zongolica. Due the high number of threatened species in the BR its long-term conservation acquires a greater importance.

Key words: conservation; endemism; growth forms; threatened species



La Reserva del Bicentenario (BR) ubicada en la Sierra de Zongolica, Veracruz, contiene 63 hectáreas de bosque mesófilo (bm) que carece de estudios florísticos sistemáticos, mientras que la sierra se ha propuesto como área para la conservación de las aves.


i) ¿La composición florística de la BR es rica taxonómicamente? ii) ¿Cómo están representadas las formas de crecimiento en esta Reserva? iii) ¿La RB tiene especies endémicas o en peligro de extinción?

Especies en estudio:

Plantas Vasculares.

Sitio de estudio y fechas:

La BR fue explorada de marzo de 2011 a octubre de 2012.


Muestras botánicas de cada especie considerada diferente se colectaron y procesaron con procedimientos convencionales para ser identificadas con herramientas taxonómicas y consulta con especialistas. Se realizaron análisis de la riqueza de especies, formas de crecimiento, endemismo y especies en riesgo.


Registramos 401 especies, distribuidas en 272 géneros y 102 familias; Orchidaceae, Asteraceae, Fabaceae y Piperaceae son las más diversas, mientras que los géneros Peperomia, Tillandsia, Polypodium, Quercus y Solanum tienen el mayor número de especies. Sesenta y nueve especies son endémicas de México y seis están restringidas a Veracruz. Encontramos 23 registros nuevos para el municipio de Zongolica, Quercus ghiesbreghtii por primera vez en bm, Q. furfuraceae en Veracruz y 86 especies amenazadas.


La RB tiene una alta riqueza florística comparada con otras áreas de México con bm, revelando la importancia de completar los estudios florísticos de la Sierra de Zongolica. Debido a alto número de especies amenazadas en la RB su conservación a largo plazo adquiere gran importancia.

Palabras clave: conservación; endemismo; especies amenazadas; forma de crecimiento

The cloud forest (cf) (sensu Rzedowski 1978a) is a set of plant communities distributed along the mountains, having diverse structures, floristic affinities and species compositions (CONABIO 2010). They have a discontinuous distribution, but they occupy an area of 0.5 to 1 % of the national territory (González-Espinosa et al. 2011, Gual-Díaz & González-Medrano 2014). The cf concentrate a high floristic richness which goes from 2,500 to 2,822 species; the highest per unit of surface among all vegetation types in Mexico (Rzedowski 1996, Villaseñor 2010). The largest extension of cf is located in the Sierra Madre Oriental, the Sierra Norte de Oaxaca, the Sierra Madre del Sur, the Northern Chiapas mountains and the Sierra Madre de Chiapas (González-Espinosa et al. 2011). This vegetation is severely threatened due to land use change, which has reduced its original cover by half (Challenger 1998) and due to only 12 % of the cover in Mexico being protected (Ponce-Reyes et al. 2012).

Studies on cf have focused on both floristic and structural aspects in at least 83 localities in the country (Ruiz-Jiménez et al. 2012), but specially in Chiapas (Ramírez-Marcial 2001), State of Mexico (López-Pérez et al. 2011), Guerrero (Meave et al. 1992; Catalán-Everástio et al. 2003), Hidalgo (Luna et al. 1994, Alcántara-Ayala & Luna-Vega 2001, Ponce-Vargas et al. 2006), Jalisco (Sánchez-Rodríguez et al. 2003), Oaxaca (Acosta-Castellanos 1997, Mejía-Domínguez et al. 2004, Ruiz-Jiménez et al. 2000, Lorea & Munn 2005), Morelos (Luna-Vega et al. 1989), Querétaro (Cartujano et al. 2002) and Tamaulipas (Cavazos-Camacho 2000).

In the state of Veracruz, the cf covers approximately 135,271 ha, with approximately 2,028 plant species (Castillo-Campos et al. 2011). The majority of the 19 studies on this vegetation type focus on the central zone of the state (Ruiz-Jiménez et al. 2012). The Sierra de Zongolica, located in the central zone of the state, is part of the physiographic province of the Sierra Madre del Sur (Medina-Chena et al. 2010). The biological exploration of the zone has been focused mainly on the knowledge of birds being proposed as an Important Area for Bird Conservation (AICA 15 Sierra de Zongolica) (Arizmendi & Márquez-Valdemar 2000).

Regarding the botanical studies of the Sierra de Zongolica, Vázquez-Torres (1977) identified six vegetation types and contributed a list of 290 species. The floristic studies have mainly focused on the study of the epiphytes of the cloud forest in La Quinta Ecological Reserve, Zongolica (Flores-Méndez 2008, Viccon-Esquivel 2009). Etnobotanical studies of the region emphasize the cultural richness in Tlaquilpa (Pérez-Pacheco 1992, López & Reyes 1997), Astacinga (Navarro-Pérez & Avendaño-Reyes 2002), Soledad Atzompa (Trueba-Sánchez 2009) and Tequila (Mata-Labrada 2011, Martínez-Bolaños 2012). Montoy-Koh (2010) recorded 94 plant species in the Sierra de Zongolica useful for bees (Apis mellifera L.) 69 of them from the cf.

In the study of the cf of Mexico, CONABIO (2010) classifies the forests of the Huautla-Zongolica subregion of Sierra Norte de Oaxaca, where the area of this study is located, as forests of low quality and under a high threat, since the area is highly affected by degradation processes and by deforestation due to the demand of agricultural lands.

As a contribution to evaluate the plant richness of the Bicentenario Reserve, and to know the value for conservation we aim: i) to know the floristic composition of the BR, ii) to know the growth forms of the area, and iii) to document if there are endemic and/or threatened species to the area.

Material and methods

Area of study. The Bicentenario Reserve (Figure 1) is located northeast of the municipality of Zongolica (Figure 1B), between 18° 39’ 01.89” and 18° 39’ 27.36” North and 97° 00’ 35.6” and 97° 0’ 55.62” West, along an altitudinal interval of 1,313 to 1,560 m. It belongs to the Instituto Tecnológico Superior of Zongolica (Figure 2A) and was decreed as an Área Privada de Conservación on March 30th, 2009; it benefits about 2,000 inhabitants (Rodríguez-Luna et al. 2011). The Reserve has an extension of 63 ha and is divided in three zones: Core Area, Multiple Use Zone and Buffer Zone (Figure 1). The Reserve borders the ejido Olla Chica to the East and Northeast, whilst the rest adjoins private terrains.

Figure 1 Area of study. A) Location of the Bicentenario Reserve. B) Zonification (map modified from the one from the ITZ). 

Figure 2 Area of study. A) View of the Instituto Tecnológico and the Reserve surroundings. B) La Compañía stream. C) Agave gomez-pompae, endemic species to Amatlán and Zongolica, Veracruz. 

Physiographically, the Bicentenario Reserve is located in the province of the Sierra Madre del Sur, within the subprovince of the Sierras Orientales, whose northern portion is known as Sierra de Zongolica, which is less abrupt than the rest of the subprovince and where Cretaceous calcareous rocks predominate, giving it affinity to the Sierra Madre Oriental (Medina-Chena et al. 2010). The soil is a luvisol (INEGI 2009).

According to data from the Zongolica meteorological station (18° 40’ 00” N, 97° 00’ 00” W; 1,294 meters above sea level), the climate is humid template with rains along the year. The mean annual precipitation is 2,770 mm, and the mean annual temperature is 17.5 ºC, being May the warmest month (26.4 ºC) and February the coldest (7.7 ºC). The mean annual number of days with mist is 48.4 (SMN, 2014) and the climate formula is C(fm)b(e) (García 1973). Regarding hydrography, the La Compañía stream is located in the southeastern limit of the area of study (Figure 2B).

Field- and deskwork. We explored the Bicentenario Reserve from March 2011 to October 2012 during nine field walks, lasting two to four days each. All the paths of the Reserve were walked in each visit. The entire plants in the case of herbs or branches of shrubs and trees, were collected from species considered distinctively different along the paths, selecting the individuals in the best state of flowering and/or fructification. The botanical samples collected were pressed in the field and dried out later at the Herbario Nacional de Mexico (MEXU). The specimens were identified in the Departamento de Botánica, in the Instituto de Biología, UNAM, with the available tools, such as electronic keys; Milliken et al. 2009 onwards, Nixon 2001-2002; regional floras, especially the Flora de Veracruz (Gómez-Pompa & Sosa 1978-1991, Sosa 1992-2005; Castillo-Campos 2006-2013), Flora of Guatemala (McVaugh 1963, Nash & Williams 1976, Standley & Steyermark 1946-1958, Standley & Williams 1961-1962, 1975. Standley et al. 1970-1974,), Flora del Bajío y de Regiones Adyacentes (Rzedowski & Calderón de Rzedowski 1985-2013), Flora Mesoamericana (Davidse et al. 1994, 1995, 2009, 2012), Flora de Nicaragua (Stevens et al. 2009), and specialized literature such as The Pteridophytes of Mexico (Mickel & Smith 2004), Trees and Shrubs of Mexico (Standley 1920-1926); species revisions and plant descriptions (e.g. Rzedowski 1978b, Zavala-Chávez 1989, Valencia 1995, González-Villarreal 2005, McMillan et al. 2006, Ibarra-Manríquez et al. 2012). Determinations were checked against specimens at MEXU, and images of mainly type specimens from electronic pages such as and JSTOR Global Plants (2000-2013). In the cases of problematic specimens for determination experts in several groups were consulted (see acknowledgements).

In order to know the areas of distribution of the species founded in the BR, the same references used for specimen determination were consulted, as well as Luna-Vega et al. (2004), Villaseñor (2010), González-Espinosa et al. (2011), Viccon-Esquivel & Krömer (2012), Vergara-Rodríguez (2013). In order to know the risk categories of the species in the Bicentenario Reserve, the Norma Oficial Mexicana-NOM 059 (SEMARNAT 2010) and The Red List of Mexican Cloud Forest Trees (González-Espinosa et al. 2011) were consulted.

The first duplicates of the voucher specimens were deposited in MEXU. Duplicates of these vouchers were deposited in the Instituto Tecnológico Superior de Zongolica (ITZ), the Herbario del Instituto de Ecología A.C. (XAL), and the Missouri Botanical Garden (MO).

The list of the species identified follows APG III (Stevens 2001 onwards) for the Angiosperm families, and Mickel & Smith (2004) for pteridophytes. The species are listed alphabetically by the groups Monilophyta, Lycopodiophyta, Pinophyta and Magnoliophyta. The species names were standardized according to The International Plant Name Index.


Floristic Composition. We collected 641 botanical specimens which represent 102 families, 272 genera and 401 species (Appendix 1 and Table 1). Among the groups of vascular plants, Magnoliophyta is the most numerous, followed by Monilophyta and Lycopodiophyta, whilst Pinophyta is represented by only one species (Table 1). Regarding the species number, the most diverse families are Orchidaceae with 36 (9 %), Asteraceae with 30 (7.5 %), Fabaceae with 22 (5.5 %), Rubiaceae and Polypodiaceae both with 17 each (4.2 %), Bromeliaceae with 16 (4 %), Piperaceae with 15 (3.7 %), Malvaceae and Solanaceae both with 13 each (3.2 %) and Fagaceae with 11 (2.7 %). The rest of the families includes 211 species, corresponding to 52.6 % (Table 2). The most rich families in number of genera are Asteraceae with 27 (9.9 %), Orchidaceae with 23 (8.5 %), Fabaceae with 18 (6.6 %), Rubiaceae with 16 (5.9 %), Malvaceae with 7 (2.6 %), Polypodiaceae and Poaceae both with 6 each (2.2 %), Lamiaceae, Melastomataceae, Pteridaceae and Urticaceae all with 5 each (1.8 %); the remaining families have 149 genera, representing 54.8 % (Table 2). The genera with the largest number of species are Peperomia, Quercus, Tillandsia, Polypodium and Solanum (Table 2). The predominant growth form is herbaceous with 135 species (33.67 %), followed by trees with 82 (20.45 %), epiphytes with 79 (19.70 %), shrubs with 72 (17.96 %), climbers with 29 (7.23 %), arborescent with 2 (0.50 %), holoparasites and hemiparasites both with 1 (0.25 %) species each (Table 3).

Table 1 Vascular plant richness (percent in parenthesis) in the Bicentenario Reserve, Zongolica, Veracruz. 

Group Number of species Number of genera Number of families
Monilophyta 42 (10.47) 26 (9.56) 12 (11.77)
Lycopodiophyta 3 (0.75) 2 (0.74) 2 (1.96)
Pinophyta 1 (0.25) 1 (0.37) 1 (0.98)
Magnoliophyta 355 (88.53) 243 (89.34) 87 (85.29)
TOTAL 401 272 102

Table 2 Families and genera with the highest number of species in the Bicentenario Reserve, Zongolica, Veracruz. 

Family Number of
Genus (Family) Number of
Asteraceae 27/30 Peperomia (Piperaceae) 11
Orchidaceae 23/36 Quercus (Fagaceae) 11
Fabaceae 18/22 Tillandsia (Bromeliaceae) 11
Rubiaceae 16/17 Polypodium (Polypodiaceae) 9
Malvaceae 7/13 Solanum (Solanaceae) 7
Polypodiaceae 6/17 Salvia (Lamiaceae) 5
Poaceae 6/8 Miconia (Melastomataceae) 4
Lamiaceae 5/10 Epidendrum y Prosthechea (Orchidaceae) 4/4
Melastomataceae 5/8 Smilax (Smilacaceae) 4
Pteridaceae 5/6 Piper (Piperaceae) 4
Urticaceae 5/6 Ipomoea (Convolvulaceae) 4
Bromeliaceae 4/16 Maxillaria y Jacquiniella (Orchidaceae) 3/3
Solanaceae 4/13 Saurauia (Actinidiaceae) 3
Piperaceae 2/15 Cestrum (Solanaceae) 3
Fagaceae 1/11 Desmodium (Fabaceae) 3
Total 134/227 Total 93

Table 3 Life forms in the Bicentenario Reserve, Zongolica, Veracruz. 

Growth forms Number of species
Herb 135
Tree 82
Epiphyte 79
Shrub 72
Climber 29
Arborescent 2
Hemiparasite 1
Holoparasite 1
Total 401

Forest characterization. Trees of Liquidambar styraciflua up to 20 meters height are better represented in the core area, while in the lowest part of the Reserve and along the La Compañía stream (Figure 2B), Platanus mexicana is the tallest tree. In the buffer zone Pinus pseudostrobus var. apulcensis is among the tallest trees with approximately 18 m. This layer is composed by species of Carpinus, Citharexylum, Clethra (Figure. 3I), Clusia (Figure 3F), Cornus, Hedyosmum, Myriocarpa, Myrsine, Ocotea, Oreopanax, Ostrya, Phyllonoma (Figure 3E), Picramnia, Prunus, Quercus, Rhamnus, Saurauia, Symplocos, Ternstroemia, Trema, Turpinia, Viburnum, etc. Species of Bunchosia, Deppea, Erythrina, Hampea, Heliocarpus, Ilex, Juglans, Litsea, Nectandra (Figure 3A), Nyssa, and Tilia can also be found. Arborescent ferns species of Alsophila and Cyathea up to 3 or 4 meters height can be found. Towards the lowest parts of the Reserve, species of Ficus are present while in the southeastern slope close to the La Compañia stream (Figure 2B) we find Arachnothryx aff. buddleioides, Myrcianthes fragans, Perrottetia longistylis, Platanus mexicana, Siparuna thecaphora and Sommera arborescens.

Figure 3 Some species of the flora of the Bicentenario Reserve. A) Nectandra salicifolia, near threatened. B) Iresine hebanthoides, endemic to México. C) Phyllonoma laticuspis, vulnerable. D) Passiflora adenopoda. E) Moussonia deppeana. F) Clusia flava. G) Begonia barkeri, endemic to Veracruz and Puebla. H) Gelsemium sempervirens. I) Clethra macrophylla, endemic to México. Photographies A-F by Helga Ochoterena Booth, G. by Alejandro Torres Montúfar, and H by Luis Alfonso Castillo-Hernández. 

In the more pronounced slopes, rupicolous species, such as Agave ellemeetiana, A. gomezpompae (Figure 2C), Echeveria rubromarginata and Tillandsia grandis are found. The epiphytes are very rich, growing in dense groups on the trunks and branches of the trees; among them there are many orchids, bromeliads, ferns and Piperaceae and Araceae species from the genera Asplenium, Brassia, Campyloneurum, Catopsis, Elaphoglossum, Epidendrum, Huperzia, Jacquiniella, Maxillaria, Monstera, Nidema, Peperomia, Phlebodium, Polypodium, Sedum, Sobralia and Tillandsia. The holoparasites and hemiparasites found in the Reserve are Conopholis alpina and Phoradendron nervosum, respectively, the latter being better represented in the zones where the Reserve borders with coffee plantations and paths.

The buffer zone and borders of the Reserve, have species of shrubs of the genera Aphelandra, Ardisia, Bartlettina, Cestrum, Chiococca, Conostegia, Gaultheria, Lepidaploa, Lyonia, Miconia, Mimosa, Moussonia (Figure 3E), Palicourea, Piper, Psychotria, Rogiera, Xylosma and Zapoteca. Among the climbers, both the herbaceous and woody ones, the best represented are Archibaccharis schiedeana, Cologania broussonetii, Clitoria mexicana, Gelsemium sempervirens (Figure 3H), Helicotropis linearis, Iresine hebanthoides (Figure 3B), Passiflora adenophora (Figure 3D), Rhynchosia longeracemosa, Selenicereus hamathus, Smilax spp, Toxicodendron radicans, Valeriana candolleana and Vitis spp. The herbaceous plants are species of Achimenes, Anthurium, Begonia (Figure 3G), Chamaedorea, Coccocypselum, Cuphea, Desmodium, Elaphoglossum, Govenia, Habenaria, Lisianthus, Oldenlandia, Peperomia, Polypodium, Salvia, Sticherus, Thelypteris, Tradescantia and Tripogandra in addition to species of Asteraceae.

Species under risk category. Out of the 401 species collected in the Reserve, 86 of them (21.44 %) are included in some of the categories in the NOM 059 (SEMARNAT 2010) or in The Red List of Mexican Cloud Forest Trees (González-Espinosa et al. 2011); four of them are cited in both lists: Litsea glaucescens, Juglans pyriformis, Carpinus caroliniana and Ostrya virginiana. Ten species are listed under some of the risk categories proposed in the NOM 059 (SEMARNAT 2010); two of them are in risk of extinction (P): Litsea glaucescens and Tilia americana var. mexicana; five are threatened (A): Psilotum complanatum, Juglans pyriformis, Chamaedorea schiedeana, Carpinus carolianiana and Tillandsia tricolor, whilst three are under special protection: (Pr) Cyathea divergens var. tuerckheimii, Alsophila firma and Ostrya virginiana. Out of the 80 species found in The Red List of Mexican Cloud Forest Trees categories (González-Espinosa et al. 2011), seven of them are Endangered (EN), 19 Vulnerable (VU), 10 Near Threatened (NT), 43 Least Concern (LC) and one Data Deficient (DD) (see Appendix 1).

Endemism. The Bicentenario Reserve contains 69 species that distributes only within the borders of Mexico and represent the 17.2 % of the total found species (see Appendix 1). Six of them are known only from some parts of the state of Veracruz: Agave gomez-pompae (Figure 2C) from Amatlán and Zongolica; Clethra tuxtlensis from Los Tuxtlas and Zongolica; Erythrina berenices from Chiconquiaco, Yecuatla, Xalapa, San Andrés Tuxtla and Zongolica; Quercus ghiesbreghtii from Yecuatla, Ixhuatlán de los Reyes and Zongolica, as well as Hampea integerrima and Cuphea nitidula which are known from the central zone of the state.

New records. In the Bicentenario Reserve, were found 23 species that extend its distribution into the municipality of Zongolica. For example, Quercus furfuraceae is a new report for the state of Veracruz, since its distribution was considered previously only for Michoacán, Puebla (Valencia-A 2004), Oaxaca and San Luis Potosí (Villaseñor 2010). Quercus dysophylla was only known from the municipalities of Acultzingo, Huayacocotla and Tlalnelhuayocan, Veracruz; Quercus ghiesbreghtii, was only known from Yecuatla and Ixhuatlan de los Reyes, Veracruz, and it is the first record in a cloud forest, since it is not cited for this type of vegetation by Villaseñor (2010) or Valencia-A & Gual-Díaz (2014). Other species that extend their distribution into the Zongolica municipality in the present contribution are Cornus excelsa known from Huayacocotla, Orizaba and Xalapa, Gelsemium semprevirens from Acajete, Atzalan, Banderilla, Calcahualco, Chiconquiaco, Coatepec, Huayacocotla, Huatusco, Jilotepec, Naolinco, Rafael Lucio, Soteapan, Tlacolulan, Tlalnelhuayocan, Xalapa and Xico, Nyssa sylvatica from Atzalan, Peperomia consoquitlana from Atoyac, Coatepec, Ixtaczoquitlan, Jilotepec, Naolinco, Orizaba, San Andrés Tuxtla and Xalapa, P. epidendron from Orizaba, Tenochtitlan, Tlacotepec de Mejia and Totula, Ponthieva rinconii from Coetzala, Racinaea ghiesbreghtii from Coatepec, Huatusco, Jilotepec, Tlalnelhuayocan, Xalapa and Xico.


In the Bicentenario Reserve a group of families that Rzedowski (1996) considered as thriving preferentially in the cloud forest, or whose genera can thrive exclusively or almost exclusively in the cloud forest, is found. Among the former, there are Actinidiaceae, Aquifoliaceae, Begoniaceae, Clethraceae, Cornaceae, Cyatheaceae, Gesneriaceae, Gleicheniaceae, Lauraceae, Lycopodiaceae, Marattiaceae, Monimiaceae, Myrsinaceae, Orchidaceae, Piperaceae, Selaginellaceae, Staphyleaceae, Symplocaceae and Theaceae (Pentaphylacaceae). And among the ones whose genera thrive in the cloud forest are Chloranthaceae, Hamamelidaceae (=Altingiaceae) and Nyssaceae (=Cornaceae). Some of the tree genera with a quantitatively importance in the cloud forest (Rzedowski 1996) found in the Bicentenario Reserve are Alnus, Carpinus, Clethra, Cornus, Juglans, Liquidambar, Nyssa, Oreopanax, Prunus, Quercus, Symplocos and Ternstroemia.

The cf of the Bicentenario Reserve contains a rich floristic composition with at least 401 species of vascular plants, representing the 16.4 % of the floristic richness estimated for this vegetation type in Mexico by Rzedowski (1996), or 14.21 % of the richness estimated by Villaseñor (2010) and the 19.77 % out of the total species number recorded in the cf in the state of Veracruz (Castillo-Campos et al. 2011). This richness is high, taking into account the reduced sampling area (63 ha = 0.63 km2) compared to other areas studied in the state of Veracruz, such as Teocelo, where Luna et al. (1988) recorded 271 species in approximately 21 km2 (= 2,100 ha); Huayacocotla, where Juárez-Medina (2008) recorded 494 species from the information of three previous works and their samplings in 20 km2 (= 2,000 ha), or La Cortadura, Coatepec, where García-Franco et al. (2008) recorded 258 species in 3,000 m2 (= 0.3 ha, = 0.003 Km2). In Hidalgo, 336 species were recorded in Tlanchinol (Luna et al. 1994), 359 in Lolotla (Ponce-Vargas et al. 2006), 287 in Eloxochitlan and 233 in Tahuelompa (Alcántara-Ayala & Luna-Vega 2001). In the cf of Landa de Matamoros (Querétaro), Cartujano et al. (2002) recorded 774 species while studying an area of approximately 54 km2 and a greater sampling effort, with 14 monthly field walks, from March 1998 to April 2000, each lasting four to five days.

The high number of fern species (pteridophytes and related) found in the Bicentenario Reserve can be an indicator that the forest is conserved, since ferns require very particular environmental conditions for their growth, such as intense shade and high humidity (Williams-Linera 2012). These are the reasons for which ferns are represented by a larger number of species in the cf than in any other vegetation type (Challenger 1998). The presence of arborescent ferns in the Reserve shows low perturbation conditions for a long time period, in addition to them being a distinctive characteristic of the cf, due to their absence in any other type of primary vegetation (Challenger 1998, Luna et al. 2001).

Regarding growth forms, the highest numbers of species are herbs, which are better represented in the buffer zone, since logging and transit of people are common in this part of the Reserve. The borders of the Reserve favor the diversification of herbs, whereas in the core area a lower number of herbs was found. Luna et al. (2001) consider that the low amount of light penetrating the cf prevents the development and diversification of herbs, whereas its development is favored in the margins and they diversify as the disturbance increases.

The sampling of Quercus furfuracea for the first time in the state of Veracruz, of Quercus ghiesbreghtii for the first time in a cf, as well as of Peperomia consoquitlana, P. epidendron, Quercus dysophylla, Q. ghiesbreghtii, and other species which extend their known distribution into the Zongolica municipality exemplifies the need to explore the zone, which would remarkably improve the phytogeographical analyses.

It is interesting to emphasize that in the taxonomical treatments of some families of the Flora de Veracruz the presence of some species is not recorded for Zongolica, so they also extend their distribution into this municipality (e.g., Cornus excelsa, Nyssa sylvatica, Racinaea ghiesbreghtii and Gelsemium semprevirens). In this work, species of epiphytes not recorded by Flores-Méndez (2008), Viccon-Esquivel (2009), or Viccon-Esquivel & Krömer (2012) for Zongolica were found. Among them Asplenium praemorsum, Catopsis morreniana, Dryopteris patula, Echeveria rosea, Mainthemum macrophyllum, Maxillaria cucullata, Myoxanthus congestus, Peperomia galioides, P. pililimba, P. rhexiifolia, Polypodium fraternum, P. plesiosorum, Prosthechea cochleata, P. chondylobulbon, Sedum botterii, Stelis emarginata, Tillandsia leiboldiana and Xylobium sulfurinum, increasing the epiphyte diversity in the municipality, even though the mentioned authors reported a greater number of species not found in the area of study.

Due to the deterioration of the vegetation in a large part of the Sierra de Zongolica and to the Reserve terrain currently being private property, it is common to find species that according to literature thrive in perturbed places, such as Bidens squarrosa, Cyclospermum leptophyllum, Elephantopus mollis, Fleischmannia pycnocephala, Gaultheria acuminata, Lantana camara, Miconia mexicana, Psidium guajava, Sida rhombifolia, Triumfetta bogotensis, Verbena carolina, Witheringia solanacea and Viguiera cordata. The presence of other species such as Conostegia xalapensis indicate that a secondary succession is under process, whereas other such as Epidendrum radicans and Erigeron karvinskianus are ruderals. Nonetheless, these are mainly found in the paths of the buffer zone, in the multiple-use zone and in the borders of the Reserve, where there is a larger transit of people and so the amount of light penetrating the forest is higher. In the Bicentenario Reserve there are seven introduced species Hypoestes phyllostachya, Ocimum campechianum, Ricinus communis and Trifolium repens, from the Old World (Villaseñor & Espinosa-García 2004), Impatiens walleriana from South America, Eriobotrya japonica native from China (Pankhurst 2009) and Kalanchoe pinnata introduced from Africa and Madagascar (Brunner 2009). These species are still scarce and are found in the borders of the Reserve or in the buffer zone.

The presence in the Bicentenario Reserve of 86 species included in the NOM-059 (SEMARNAT 2010) and in the The Red List of Mexican Cloud Forest Trees (González-Espinosa et al. 2011) stress the importance of the reservation as a biodiversity refuge, protecting species of great importance in the Mexican flora. As Martínez-Camilo et al. (2012) mention, the role performed by a protected area, the Bicentenario Reserve in this case, in the conservation of biodiversity acquires a greater importance when the species under threat, in danger of extinction or in another risk category, are known. Based on this information it is possible to promote more appropriate conservation policies, focused on the species that need them, that is why these species need special attention and why actions for their conservation and preservation in the ecosystem should be taken.


This first contribution to the knowledge of the flora of the cf in the Bicentenario Reserve in particular and in the Zongolica municipality in general, emphasizes its high floristic richness, with 401 species, 272 genera and 102 families of vascular plants, representing the 19.77 % of the total species number recorded in the cf of the state of Veracruz and between 14.21 and 16.04 % of the floristic richness estimated for this vegetation type in Mexico.

Despite the cf of the region having been cataloged as of low quality, the remnant flora, according to the results of this work, contains a high species richness, with 69 species endemic to Mexico, out of which eight are endemic to the state of Veracruz, as well as species in some risk category and species never before recorded for the zone. The conservation of the Reserve should be a priority, nonetheless because it has ten species considered in the NOM 059 (SEMARNAT, 2010) and 80 in The Red List of Mexican Cloud Forest Trees (González-Espinosa et al. 2011).


To Miguel Ángel de la Torre Loranca, General Director of the Instituto Tecnológico Superior de Zongolica, for facilitating the making of this study. Particularly, we thank Fortino Corona Jiménez and Yavé Carrera Quirino of the ITSZ, as well as Helga Ochoterena Booth, Alejandro Torres Montufar and Nidia Mendoza Díaz, for their support and company in the field work. To the experts Oscar Hinojosa Espinoza and José Luis Villaseñor (Asteraceae), Carlos Gómez Hinostrosa (Cactaceae), Helga Ochoterena and Alejandro Torres Montufar (Rubiaceae), Alfonso Delgado (Fabaceae), Angélica Ramírez Roa (Gesneriaceae), Gerardo Salazar Chávez (Orchidaceae), Susana Valencia Ávalos (Quercus), Daniela Vergara (Peperomia), Alberto Reyes and Emily Lott, for the determination of some species or the corroboration of the identifications. We thank David Gernandt and Angélica Ramírez Roa for the support of the MEXU in mounting duplicates donated to the ITSZ. We also thank José Luis Caballero for his help in the making of Figure 1.

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Appendix 1

Floristic composition of the Bicentenario Reserve, Zongolica, Veracruz. Voucher number refers to the collections of the first author (LA Castillo-Hernández), except those of the Instituto Tecnológico de Zongolica marked as ITS. Categories in the NOM 059-SEMARNAT (2010): P (risk of extinction), A (threatened), Pr (special protection). The Red List of Mexican Cloud Forest Trees Categories (González-Espinosaet al. 2011): EN (endangered), VU (vulnerable), NT (near threatened), LC (least concern), DD (data deficient). * = endemic to Mexico. 

Taxa Voucher
The Red List
of Mexican
Cloud Forest
Trees Categories
Asplenium cuspidatum Lam. 322 Epiphyte
Asplenium miradorense Liebm. 416 Herb
Asplenium praemorsum Sw. 160 Epiphyte
Cyathea divergens var. tuerckheimii (Maxon) R.M.Tryon 337 Arborescent Pr
Alsophila firma (Baker) D.S. Conant 336 Arborescent Pr
Pteridium aquilinum (L.) Kuhn ITSZ Herb
Dryopteris patula (Sw.) Underw. 562 Epiphyte
*Elaphoglossum sartorii (Liebm.) Mickel 581, 584, 612 Herb
*Elaphoglossum vestitum (Schltdl. & Cham.) T. Moore 578 Epiphyte
Phanerophlebia remotispora E.Fourn. 102, 280, 545 Herb
Polystichum distans E.Fourn. 526 Herb
Sticherus palmatus (W.Schaffn. ex E. Fourn.) Copel 100, 308 Herb
Marattia laxa Kunze 534 Herb
Campyloneurum angustifolium Fée 284, 605 Epiphyte
Campyloneurum xalapense Fée 604, 614 Epiphyte
Niphidium crassifolium (L.) Lellinger 99 Epiphyte
Pecluma alfredii (Rosenst.) M.G.Price 603 Epiphyte
Pecluma plumula (Humb. & Bonpl. ex Willd.) M.G.Price 716 Herb
Phlebodium areolatum (Humb. & Bonpl. ex Willd.) J.Sm. 46, 48, 243 Epiphyte
Pleopeltis angusta var. stenoloma Farw. 393 Herb
Pleopeltis crassinervata T.Moore 162-A, 244 Epiphyte
Polypodium conterminans Liebm. 307 Herb
Polypodium fraternum Schltdl. & Cham. 585 Epiphyte
*Polypodium lepidotrichum (Fée) Maxon 525 Herb
Polypodium plebeium Schltdl. & Cham. 51, 606 Epiphyte
Polypodium plesiosorum Kunze 162-B, 576 Epiphyte
Polypodium polypodioides (L.) Watt 552 Herb
Polypodium rhodopleuron Kunze 564, 721 Epiphyte
Polypodium subpetiolatum Hook. 613 Epiphyte
*Polypodium villagranii Copel. 728 Herb
Psilotum complanatum Sw. 350 Epiphyte A
Adiantum andicola Liebm. 101 Herb
Llavea cordifolia Lag. 129 Herb
Pityrogramma ebenea (L.) Proctor ITSZ Herb
Pteris cretica L. 563 Herb
Pteris longifolia L. 601 Herb
Vittaria bradeorum Rosenst. 481 Epiphyte
Anemia phyllitidis (L.) Sw. 161 Herb
Tectaria heracleifolia(Willd.) Underw. 717 Herb
Thelypteris oligocarpa (Humb. & Bonpl. ex Willd.) Ching 586 Herb
Thelypteris hispidula (Decne.) C.F.Reed 524 Herb
Thelypteris ovata var. lindheimeri (C.Chr.) A.R.Sm. 24 Herb
Huperzia pithyoides (Schltdl. & Cham.) Holub 357-A Epiphyte
Huperzia taxifolia (Sw.) Trevis. 357-B Epiphyte
Selaginella stellata Spring 16, 96 Epiphyte
Pinus pseudostrobus var. apulcensis (Lindl.) Shaw 78, 79, 500 Tree
Aphelandra schiedeana Schltdl. & Cham. 108, 270 Shrub
Hypoestes phyllostachya Baker 175 Herb
Justicia pectoralis Jacq. 22 Herb
Odontonema callistachyum (Schltdl. & Cham.) Kuntze 487 Shrub
Saurauia leucocarpa Schltdl. 453 Tree VU
*Saurauia pedunculata Hook. 277, 468, 544 Tree VU
*Saurauia scabrida Hemsl. 588 Tree NT
Sambucus nigra subsp. canadensis (L.) Bolli 472 Shrub LC
*Viburnum tiliifolium (Oerst.) Hemsl. 83, 171, 230 Tree VU
Bomarea edulis (Tussac) Herb. 267 Herb
Liquidambar styraciflua L. 455 Tree LC
Iresine diffusa Humb. & Bonpl. ex Willd. 19, 288, 289, 295 Herb
*Iresine hebanthoides Suess. 296, 328 Climber
Rhus terebinthifolia Schltdl. & Cham. 278 Shrub
Toxicodendron radicans (L.) Kuntze 434 Climber
Cyclospermum leptophyllum (Pers.) Sprague ex Britton & P.Wilson 443 Herb
Sanicula liberta Cham. & Schltdl. 348 Herb
Apocynaceae (Asclepiadaceae)
Asclepias curassavica L. 436 Herb
*Ilex condensata Turcz. 516 Tree
Ilex sp. 315 Tree
Anthurium scandens (Aubl.) Engl. subsp. scandens 557 Epiphyte
Anthurium schlechtendalii Kunth 182, 406 Herb
Monstera deliciosa Liebm. 149 Epiphyte
*Syngonium sagittatum G.S.Bunting 346 Epiphyte
Hydrocotyle mexicana Schltdl. & Cham. 163 Herb
Oreopanax capitatus (Jacq.) Decne. & Planch. 252, 394, 499, 561 Tree NT
*Chamaedorea schiedeana Mart. 148, 299, 356 Herb A
*Agave ellemeetiana Jacobi 360 Herb
*Agave gomezpompae Cházaro & Jimeno-Sevilla 359 Herb
Maianthemum macrophyllum (M.Martens & Galeotti) LaFrankie 505 Epiphyte
Acourtia veracruzana B.L.Turner 425 Herb
Ageratina ligustrina (DC.) R.M.King & H.Rob. 273, 386 Shrub
Ageratum houstonianum Mill. 20 Herb
Aldama dentata La Llave 118, 730 Herb
Alloispermum integrifolium (DC.) H.Rob. 6, 382, 489 Herb
Archibaccharis hirtella (DC.) Heering 313 Climber
Archibaccharis schiedeana (Benth.) J.D.Jacks. 145, 146, 297, 301 Climber
Bartlettina tuerckheimii (Klatt) R.M.King & H.Rob. 398, 400 Shrub
Bidens odorata Cav. var. odorata 150 Herb
Bidens squarrosa Kunth 168, 729 Climber
Calyptocarpus wendlandii Sch.Bip. 113, 542 Herb
Critonia hospitalis (B.L.Rob.) R.M.King & H.Rob. 383, 495 Tree
Dahlia coccinea Cav. 139 Herb
Decachaeta incompta (DC.) R.M.King & H.Rob. 247 Shrub
Elephantopus mollis Kunth 14, 384 Herb
Erigeron karvinskianus DC. 13, 426 Herb
Fleischmannia pycnocephala (Less.) R.M. King & H.Rob. 11, 261, 316, 404, 497 Herb
Fleischmanniopsis leucocephala (Benth.) R.M.King & H.Rob. 475 Shrub
Hymenostephium cordatum (Hook. & Arn.) S.F.Blake 287, 549, 722 Herb
Lagascea helianthifolia Kunth 366 Shrub
*Leiboldia serrata (D.Don) Gleason 362, 402 Shrub
Lepidaploa tortuosa (L.) H.Rob. 268, 401 Shrub
Piqueria trinervia Cav. 200, 428 Herb
*Roldana oaxacana (Hemsl.) H.Rob. & Brettell 409 Herb
Roldana schaffneri (Sch.Bip. ex Klatt) H.Rob. & Brettell 8, 496 Shrub
*Schistocarpha bicolor Less. 407 Shrub
Tagetes filifolia Lag. 179 Herb
Telanthophora grandifolia (Less.) H.Rob. & Brettell 338 Tree
Tetrachyron manicatum Schltdl. 387, 501 Shrub
Verbesina turbacensis Kunth 724 Shrub
Impatiens walleriana Hook.f. 574 Herb
*Begonia barkeri Knowles & Westc. 335, 392 Herb
Begonia sp. 345 Herb
Alnus acuminata Kunth 537 Tree LC
Carpinus caroliniana Walter 71 Tree A NT
Ostrya virginiana (Mill.) K.Koch 73 Tree Pr NT
Catopsis morreniana Mez 228 Epiphyte
Catopsis sessiliflora (Ruiz & Pav.) Mez 53, 566 Epiphyte
Pitcairnia heterophylla (Lindl.) Beer 460 Epiphyte
*Racinaea adscendens (L.B.Sm.) M.A. Spencer & L.B.Sm. ITSZ Epiphyte
*Racinaea ghiesbreghtii (Baker) M.A. Spencer & L.B.Sm. 98 Epiphyte
Tillandsia butzii Mez 464 Epiphyte
Tillandsia grandis Schltdl. 583 Herb
*Tillandsia heterophylla E.Morren 522 Herb
Tillandsia juncea (Ruiz & Pav.) Poir. ITSZ Epiphyte
*Tillandsia kirchhoffiana Wittm. 97 Epiphyte
Tillandsia leiboldiana Schltdl. 70 Epiphyte
Tillandsia multicaulis Steud. 188-B Epiphyte
Tillandsia punctulata Schltdl. & Cham. 363 Epiphyte
Tillandsia schiedeana Steud. ITSZ Epiphyte
Tillandsia tricolor Schltdl. & Cham. 58 Epiphyte A
Tillandsia viridiflora (Beer) Baker 593 Epiphyte
*Selenicereus hamatus (Scheidw.) Britton & Rose 521 Herb
Lobelia laxiflora Kunth 5, 306 Herb
Lobelia xalapensis Kunth 17 Herb
Cannabaceae (Ulmaceae)
Trema micrantha (L.) Blume var. micranta 555, 600 Tree LC
Caprifoliaceae (Valerianaceae)
Valeriana candolleana Gardner 135, 424, 490 Climber
Drymaria gracilis Schltdl. & Cham. 590 Herb
Hedyosmum mexicanum C.Cordem. 210, 532 Tree LC
*Clethra macrophylla M.Martens & Galeotti 246, 385 Tree LC
*Clethra tuxtlensis L.M.González 61, 310, 395, 523 Tree LC
Clusia flava Jacq. 35, 198 Tree LC
Callisia monandra (Sw.) Schult. & Schult.f. 177 Herb
Commelina diffusa Burm.f. 442 Herb
Tradescantia zanonia (L.) Sw. 45, 323 Herb
Tripogandra serrulata (Vahl) Handlos 44, 255 Herb
Ipomoea batatas (L.) Lam. 180, 317 Climber
*Ipomoea funis Schltdl. & Cham. var. funis 342 Climber
Ipomoea indica (Burm.) Merr. 181 Climber
Ipomoea mairetii Choisy 372 Climber
Cornus excelsa Kunth 281, 515 Tree LC
Nyssa sylvatica Marshall 528 Tree LC
*Echeveria rosea Lindl. 463 Epiphyte
*Echeveria rubromarginata Rose 103 Herb
Kalanchoe pinnata (Lam.) Pers. 21 Herb
*Sedum botterii Hemsl. 293, 432 Epiphyte
Carex aff. standleyana Steyerm. 520 Herb
Carex polystachya Sw. ex Wahlenb. 304 Herb
Cyperus tenuis Sw. 176, 616 Herb
Rhynchospora aristata Boeckeler 190, 567 Herb
Rhynchospora radicans (Schltdl. & Cham.) H.Pfeiff. subsp. radicans 60 Herb
Dioscorea mexicana Scheidw. 548 Climber
Perrottetia longistylis Rose 351, 599 Tree LC
Chimaphila maculata (L.) Pursh 509 Herb
Gaultheria acuminata Schltdl. & Cham. 80, 476 Shrub LC
Lyonia squamulosa M.Martens & Galeotti 314, 456 Shrub NT
Vaccinium leucanthum Schltdl. 530 Tree EN
Acalypha sp. 242, 291 Herb
Cnidoscolus multilobus (Pax) I.M.Johnst. 462 Tree LC
Euphorbia graminea Jacq. 312 Herb
Ricinus communis L. 591 Shrub
Acacia angustissima (Mill.) Kuntze var. angustissima 551 Tree
Canavalia villosa Benth. 380, 570 Climber
Centrosema pubescens Benth. 158 Climber
Chamaecrista rufa Britton & Rose 106 Herb
Clitoria mexicana Link 167, 207, 379 Climber
Cologania pulchella Kunth 157 Climber
Crotalaria sagittalis L. 253 Herb
Desmodium caripense (Kunth) G.Don 164 Herb
*Desmodium helleri Peyr. 271, 365, 367, 368, 422 Herb
Desmodium psilophyllum Schltdl. 110 Herb
Diphysa floribunda Peyr. 361-B Tree NT
*Erythrina berenices Krukoff & Barneby 361-A Tree
Helicotropis linearis (Kunth) A.Delgado Inga vera Willd. 334 Climber
Mimosa albida Humb. & Bonpl. ex Willd. var. albida 151, 199, 378 Shrub
*Phaseolus glabellus Piper 119, 405 Climber
Rhynchosia longeracemosa M.Martens & Galeotti 109, 238, 572 Climber
Senna hirsuta (L.) H.S.Irwin & Barneby 569 Herb
*Senna foetidissima var. grandiflora (Benth.) H.S.Irwin & Barneby 156, 374 Shrub
Senna septemtrionalis (Viv.) H.S.Irwin & Barneby 503 Shrub
Trifolium repens L. 465 Herb
Zapoteca portoricensis (Jacq.) H.M.Hern. subsp. portoricensis 41, 282, 446, 715 Shrub LC
*Quercus aff. glabrescens Benth. 104 Tree VU
*Quercus aff. sartorii Liebm. 609 Tree
Quercus candicans Née 130, 414 Tree VU
Quercus castanea Née 169, 220, 272, 507 Tree
*Quercus dysophylla Benth. 269 Tree
Quercus elliptica Née 92, 49 Tree VU
*Quercus furfuracea Liebm. 187 Tree
*Quercus ghiesbreghtii M.Martens & Galeotti 568 Tree
*Quercus obtusata Bonpl. 77 Tree LC
*Quercus polymorpha Schltdl. & Cham. 239, 610, 611 Tree EN
*Quercus sartorii Liebm. 607 Tree EN
Gelsemiaceae (Loganiaceae)
Gelsemium sempervirens (L.) J.St.-Hil. 276, 364 Climber
*Lisianthius nigrescens Schltdl. & Cham. var. nigrescens 94 Herb
Achimenes antirrhina (DC.) C.V.Morton 183, 565-B Herb
Achimenes grandiflora (Schltdl.) DC. 565-A Herb
Columnea schiedeana Schltdl. 471 Epiphyte
Moussonia deppeana (Schltdl. & Cham.) Klotzsch ex Hanst. 132, 262 Shrub
Heliconia adflexa (Griggs) Standl. 538 Herb
Hypericum hypericoides (L.) Crantz 88, 403 Shrub
Vismia baccifera (L.) Triana & Planch. 554 Tree LC
Hypoxis decumbens L. 111, 215 Herb
*Sisyrinchium schaffneri S.Watson 18 Herb
*Juglans pyriformis Liebm. 533 Tree A EN
Hyptis atrorubens Poit. 331 Herb
Hyptis mutabilis (Rich.) Briq. 127, 339 Herb
Ocimum campechianum Mill. 3, 93, 122, 125 Herb
Prunella vulgaris L. 539 Herb
*Salvia filipes Benth. 486 Shrub
Salvia karwinskii Benth. 435 Shrub
*Salvia longispicata M.Martens & Galeotti 126 Herb
Salvia mocinoi Benth. 38, 131, 258 Herb
Salvia purpurea Cav. 222, 423, 726 Shrub
Stachys lindenii Benth. 594 Herb
Litsea glaucescens Kunth 251, 473 Tree P VU
Nectandra salicifolia (Kunth) Nees 283 Tree NT
Ocotea helicterifolia (Meisn.) Hemsl. 461 Tree VU
*Ocotea psychotrioides Kunth 413 Tree EN
Cuphea hyssopifolia Kunth 353, 354 Shrub
*Cuphea nitidula Kunth 42, 196 Herb
Bunchosia lindeniana A.Juss. 279, 411, 553-A Tree LC
Bunchosia biocellata Schltdl. 735 Shrub
Anoda cristata (L.) Schltdl. 10, 373 Herb
*Hampea integerrima Schltdl. 33, 300 Tree NT
Heliocarpus americanus L. 420 Tree LC
Heliocarpus appendiculatus Turcz. 469 Tree LC
Heliocarpus mexicanus (Turcz.) Sprague 128 Shrub LC
Pavonia schiedeana Steud. 154 Shrub
*Pavonia uniflora (Sessé & Moc.) Fryxell 285, 369 Shrub
Sida glabra Mill. 1, 136 Herb
Sida rhombifolia L. 399, 491 Shrub
*Tilia americana var. mexicana (Schltdl.) Hardin 575 Tree P
Triumfetta bogotensis DC. 112, 485 Shrub
Triumfetta grandiflora Vahl 28, 311, 457 Shrub
Triumfetta semitriloba Jacq. 30 Shrub
Arthrostemma ciliatum Pav. ex D.Don 559-A Shrub
Conostegia xalapensis (Bonpl.) D.Don ex DC. 433, 466 Shrub LC
Heterocentron subtriplinervium (Link & Otto) A.Braun & C.D.Bouché 134, 508 Shrub
Miconia affinis DC. 39, 89 Shrub
Miconia desmantha Benth. 72, 492 Shrub
Miconia glaberrima (Schltdl.) Naudin 410 Shrub LC
Miconia mexicana (Bonpl.) Naudin 32, 90, 206, 234 Tree
*Tibouchina galeottiana (Naudin) Cogn. 527, 559-B Herb
Mollinedia viridiflora Tul. 529 Tree VU
Dorstenia contrajerva L. 719, 720 Herb
Ficus aurea Nutt. 558 Tree LC
Ficus velutina Humb. & Bonpl. ex Willd. 560 Tree LC
Pseudolmedia glabrata(Liebm.) C.C.Berg 454 Tree LC
Eugenia sp. 347 Tree
Myrcianthes fragrans (Sw.) McVaugh 504, 615 Tree
Psidium guajava L. 540 Tree
Lopezia racemosa Cav. 2, 4, 115, 178 Herb
Oenothera rosea L’Hér. ex Aiton 173, 543 Herb
Brassia verrucosa Lindl. 512 Epiphyte
Coelia triptera (Sm.) G.Don ex Steud. 470 Epiphyte
Dichaea glauca (Sw.) Lindl. 69 Epiphyte
Dichaea neglecta Schltr. 63 Epiphyte
Elleanthus cynarocephalus (Rchb.f.) Rchb.f. 66, 227 Epiphyte
Epidendrum radicans Pav. ex Lindl. 91, 211 Herb
Epidendrum ramosum Jacq. 249, 480, 571 Epiphyte
Epidendrum repens Cogn. 235, 303 Epiphyte
Epidendrum veroscriptum Hágsater 166 Epiphyte
*Gongora galeata (Lindl.) Rchb.f. 598 Epiphyte
Govenia mutica Rchb.f. 54, 186 Herb
Habenaria odontopetala Rchb.f. 320 Herb
Isochilus major Schltdl. & Cham. 59 Epiphyte
Jacquiniella equitantifolia (Ames) Dressler 459 Epiphyte
*Jacquiniella leucomelana (Rchb.f.) Schltr. 333 Epiphyte
Jacquiniella teretifolia (Sw.) Britton & P.Wilson 202 Epiphyte
Leochilus carinatus (Knowles & Westc.) Lindl. 52 Epiphyte
Lycaste aromatica (Graham) Lindl. 531 Epiphyte
Lycaste deppei (Lodd.) Lindl. 546 Epiphyte
Malaxis histionantha (Link, Klotzsch & Otto) Garay & Dunst. 547 Herb
Maxillaria cucullata Lindl. 329-A Epiphyte
Maxillaria meleagris Lindl. 329-B Epiphyte
Maxillaria variabilis Bateman ex Lindl. 159, 189, 205 Epiphyte
Myoxanthus congestus (A.Rich. & Galeotti) Soto Arenas 332 Epiphyte
Nemaconia graminifolia Knowles & Westc. 142, 327 Epiphyte
Nidema boothii (Lindl.) Schltr. 427, 478 Epiphyte
*Ponthieva rinconii Salazar 114, 573 Herb
Prosthechea chondylobulbon (A.Rich. & Galeotti) W.E.Higgins 292 Epiphyte
Prosthechea cochleata (L.) W.E.Higgins 107, 219, Epiphyte
Prosthechea ochracea (Lindl.) W.E.Higgins 57, 75, 274 Epiphyte
Prosthechea pseudopygmaea (Finet) W.E.Higgins 580 Epiphyte
Sobralia macrantha Lindl. 65 Epiphyte
Stelis emarginata (Lindl.) Soto Arenas & Solano 47 Epiphyte
Stelis platystylis (Schltr.) Solano & Soto Arenas 204, 233, 479 Epiphyte
*Trichocentrum pachyphyllum (Hook.) R. Jiménez & Carnevali 412 Herb
Xylobium sulfurinum (Lem.) Schltr. 597 Epiphyte
Castilleja arvensis Schltdl. & Cham. 12 Herb
Conopholis alpina Liebm. 452 Holoparasite
Biophytum dendroides (Kunth) DC. 361-C Herb
Passiflora adenopoda DC. 377, 725 Climber
*Passiflora conzattiana Killip 62 Climber
*Ternstroemia sylvatica Schltdl. & Cham. 82, 217 Tree VU
Phyllanthus compressus Kunth 438 Herb
Phyllonoma laticuspis (Turcz.) Engl. 26-B, 302 Tree VU
Phytolacca purpurascens A.Braun & C.D.Bouché 592 Shrub
Phytolacca rivinoides Kunth & C.D.Bouché 397 Shrub
Picramniaceae (Simaroubaceae)
Picramnia antidesma Sw. 165, 488, 608 Tree
*Peperomia consoquitlana C.DC. 55,85,86, 121, 213, 550 Epiphyte
Peperomia dendrophila Schltdl. & Cham. 56, 321 Epiphyte
Peperomia donaguiana C.DC. 216, 294 Herb
*Peperomia epidendron C.DC. 477, 553-B, 723 Herb
Peperomia galioides Kunth 236 Epiphyte
*Peperomia liebmannii C.DC. 141, 318 Herb
Peperomia obtusifolia (L.) A.Dietr. 120 Epiphyte
Peperomia quadrifolia (L.) Kunth 140 Epiphyte
Peperomia rhexiifolia Moritz ex C.DC. 358 Epiphyte
Peperomia sanjoseana C.DC. 214 Herb
Peperomia tetraphylla Hook. & Arn. 325 Epiphyte
Piper aduncum L. 587 Shrub LC
Piper amalago L. 37, 40, 123 Shrub
Piper glabrescens (Miq.) C.DC. 144, 275 Shrub
Piper umbellatum L. 579 Shrub
Plantago australis subsp. hirtella (Kunth) Rahn 15 Herb
Russelia coccinea (L.) Wettst. 248, 381 Shrub
Platanus mexicana Moric. 418 Tree NT
Arundinella deppeana Nees ex Steud. 431 Herb
Dichanthelium laxiflorum (Lam.) Gould 203 Herb
Lasiacis nigra Davidse 241 Herb
Lasiacis scabrior Hitchc. 518 Herb
Lasiacis sp. 286 Herb
*Muhlenbergia gigantea (E. Fourn.) Hitchc. 172 Herb
Oplismenus hirtellus (L.) P.Beauv. 143 Herb
Panicum polygonatum Schrad. 519 Herb
Loeselia glandulosa (Cav.) G.Don 371 Herb
Polygala paniculata L. 184, 192 Herb
Monnina sylvatica Schltdl. & Cham. 449 Shrub
*Ardisia liebmannii Oerst. subsp. Liebmannii 34, 326 Shrub VU
Ardisia sp. 341 Tree
Myrsine coriacea (Sw.) R.Br. subsp. coriacea 209, 429 Tree LC
Parathesis aff. donnell-smithii Mez 582 Tree NT
Clematis grossa Benth. 212, 419 Climber
Thalictrum pubigerum Benth. 174, 218 Herb
*Rhamnus pompana M.C.Johnst. & L.A.Johnst. 43, 439, 451 Tree VU
Eriobotrya japonica (Thunb.) Lindl. 556 Tree
Prunus brachybotrya Zucc. 84, 376 Tree VU
Rubus coriifolius Liebm. 467 Shrub
Arachnothryx buddleioides (Benth.) Planch. 535 Tree VU
Bouvardia ternifolia (Cav.) Schltdl. 185, 223, 245, 263, 265 Shrub
Chiococca phaenostemon Schltdl. 74,237 Climber DD
Coccocypselum hirsutum Bartl. ex DC. 81, 232 Herb
Crusea calocephala DC. 117, 483 Herb
Deppea grandiflora Schltdl. 444 Tree VU
Galianthe brasiliensis subsp. angulata (Benth.) E.L. Cabral & Bacigalupo 152 Shrub
*Galium seatonii Greenm. 254 Herb
Hoffmannia nicotianifolia (M.Martens & Galeotti) L.O. Williams 266,355 Shrub
*Mexotis latifolia (M.Martens & Galeotti) Terrell & H.Rob. 155 Herb
*Oldenlandia microtheca (Schltdl. & Cham.) DC. 95, 250,319 Herb
Palicourea padifolia (Humb. & Bonpl. ex Schult.) C.M.Taylor & Lorence 23, 27,260 Shrub LC
Psychotria galeottiana (M.Martens) C.M. Taylor & Lorence 31 Shrub VU
Psychotria trichotoma M.Martens & Galeotti 26-A, 224 Shrub LC
*Rogiera ligustroides (Hemsl.) Borhidi 67, 256 Shrub
Sommera arborescens Schltdl. 352, 536 Tree EN
Spermacoce remota Lam. 153, 484 Herb
Zanthoxylum melanostictum Schltdl. & Cham. 513 Tree LC
Zanthoxylum petenense Lundell 718 Tree
Xylosma flexuosa (Kunth) Hemsl. 29, 208, 502 Shrub LC
Phoradendron nervosum Oliv. 194, 421 Hemiparasite
Siparuna thecaphora (Poepp. & Endl.) A.DC. 437, 595 Tree LC
Smilacaceae (Liliaceae)
Smilax bona-nox L. 448, 498 Climber
Smilax moranensis M.Martens & Galeotti 7, 441, 447 Climber
Smilax mollis Humb. & Bonpl. ex Willd. 64, 240, 324, 391 Climber
Smilax subpubescens A.DC. 493 Climber
*Cestrum elegans (Brongn.) Schltdl. 133, 195, 231, 259, 290, 408 Shrub
Cestrum nocturnum L. 226 Shrub LC
Cestrum tomentosum L.f. 197, 298 Tree
Physalis gracilis Miers 445 Herb
*Physalis stapelioides (Regel) Bitter 390, 482 Herb
Solanum aphyodendron S.Knapp 375, 450 Tree LC
Solanum chrysotrichum Schltdl. 430 Shrub LC
Solanum lanceolatum Cav. 25, 494 Shrub LC
Solanum myriacanthum Dunal 50 Shrub
Solanum nigrescens M.Martens & Galeotti 370 Herb
Solanum schlechtendalianum Walp. 221 Shrub LC
Solanum sp. 343 Shrub
Witheringia solanacea L’Hér. 36, 514 Shrub
Turpinia insignis (Kunth) Tul. Symplocaceae 76, 458 Tree EN
Symplocos limoncillo Bonpl. 68, 511 Tree VU
Boehmeria caudata Sw. 440 Tree LC
Myriocarpa longipes Liebm. 344, 388 Tree LC
Phenax hirtus (Sw.) Wedd. 589 Shrub
Pilea microphylla (L.) Liebm. 87, 188 Herb
Pilea pubescens Liebm. 147, 517 Herb
Urera caracasana (Jacq.) Gaudich. ex Griseb. 602 Tree LC
Citharexylum mocinnoi D.Don 305, 396, 389 Tree LC
Lantana camara L. 9 Shrub
Lantana hirta Graham 137 Shrub
Lippia myriocephala Schltdl. & Cham. 105, 193 Shrub LC
Verbena carolina L. 541 Herb
Vitis popenoei J.L.Fennell 510, 596 Climber
Vitis tiliifolia Humb. & Bonpl. ex Schult. 474 Climber

Received: February 11, 2017; Accepted: April 17, 2017

* Corresponding author: Hilda Flores-Olvera, e-mail:

Author Contributions. This paper is derived from the bachelor’s thesis of L. A. Castillo-Hernández guided by H. Flores-Olvera as the main adviser. Both conducted fieldwork and contributed to the content and revisions of the paper.

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