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Journal of Species Lists and Distribution 



 

ISSN 1809-127X (online edition) 

www.checklist.org.br 

© 2009 Check List and Authors 

 

 

LISTS OF SPECIES 

 

 

Check List, Campinas, 5(4): 787–799, December, 2009. 

                    

787


 

 

Angiosperms, Los Tuxtlas Biosphere Reserve, Veracruz, Mexico 



 

 

Víctor Arroyo-Rodríguez,

1 *

 Jacob C. Dunn,



2

 Julieta Benítez-Malvido,

1

 and Salvador Mandujano



3

 

 



1

 Universidad Nacional Autónoma de México (UNAM), Centro de Investigaciones en Ecosistemas. 



Antígua Carretera a Pátzcuaro 8701, Ex Hacienda de San José de la Huerta, Morelia 58190. Michoacán, Mexico. 

 

2



 

Universitat de Barcelona, Centre Especial de Recerca en Primats. 

Vall Hebron 171. Barcelona 08035, Spain. 

 

3



 Departamento de Biodiversidad y Ecología Animal, Instituto de Ecología A. C. 

Km 2.5 Carretera Antigua a Coatepec 351, Congregacion del Haya, Xalapa 91070. Veracruz, Mexico. 

 

*



 Corresponding author e-mail: victorarroyo_rodriguez@hotmail.com 

 

 



Abstract: 

The Los Tuxtlas Reserve has been heavily deforested and fragmented since the 1970’s. Although the flora 

of Los Tuxtlas has been described previously, most floristic lists come from the large forest reserve of the Los Tuxtlas 

field station. Here we present a check list of Angiosperms recorded in 45 rainforest fragments (< 1 to 266 ha) located in 

three landscapes with different levels of deforestation. We sampled all trees, shrubs, lianas, palms and herbs with 

diameter at breast height (dbh)   2.5 cm within ten 50 m x 2 m plots per fragment. We recorded 9,435 plants belonging 

to 73 families and 372 species. Fabaceae, Rubiaceae, and Moraceae were best represented. Eight species are classified 

as Endangered by the Mexican government, and five are human-introduced species. We conclude that the conservation 

and restoration of all the remaining rainforest fragments are necessary to effectively preserve the plant diversity of this 

region. 


 

Introduction 

Accelerated deforestation and fragmentation of 

primary forest in tropical regions (Achard et al. 

2002) are threatening global biodiversity at an 

alarming rate (FAO 2006). These two processes 

modify the spatial pattern of the remaining forest 

(e.g. reduced patch area, increased patch isolation, 

and increased proportion of forest edges; Andrén 

1994), potentially affecting ecological processes 

such as pollination, seed dispersal, recruitment, 

competition, migration, herbivory and extinction 

(Turner et al. 1996; Benítez-Malvido 1998; 

Benítez-Malvido et al. 1999; Cordeiro and Howe 

2001; Wright and Duber 2001; Chacoff et al. 

2004; Aguirre and Dirzo 2008). As a 

consequence, habitat fragmentation can not only 

decrease the number of plant species, but also lead 

to significant changes in composition and 

vegetation structure (Turner et al. 1996; Laurance 

et al. 1998; Hill and Curran 2003; Arroyo-

Rodríguez and Mandujano 2006; Chazdon et al. 

2007; Santos et al. 2008; Arroyo-Rodríguez et al. 

2009; Dirzo et al. 2009).  

 

In Mexico deforestation has led to the loss of 



approximately 90 % of the tropical rainforest 

(Flores-Villela and Gerez 1994), particularly 

affecting the Los Tuxtlas Biosphere Reserve 

(Dirzo and García 1992; Guevara et al. 2004). 

This region represents the northern limit of 

tropical rainforest distribution in the Neotropics 

(Dirzo and Miranda 1991), and 95 % of the 

original rainforest here has already disappeared 

(Castillo-Campos and Laborde 2004; Guevara et 

al. 2004). While several studies have analyzed the 

plant communities in this region (e.g. Bongers et 

al. 1988; Ibarra-Manríquez et al. 1995; 1996a; b; 

1997a; b), most were carried out in the large forest 

reserve (700 ha) of the Los Tuxtlas biological 

field station of the National Autonomous 

University of Mexico (UNAM), with little 

attention being paid to the changes in plant 

communities that arise from deforestation and 

forest fragmentation (but see Arroyo-Rodríguez 

and Mandujano 2006; Arroyo-Rodríguez et al. 

2009; Dirzo et al. 2009).  

 

To contribute to the understanding of how the loss 



and fragmentation of the rainforest affect the 

vegetation in Los Tuxtlas, and to assess the 

potential conservation value of forest fragments, 

we sampled vegetation in 45 rainforest fragments 



Lists of Species 

 

Check List, Campinas, 5(4): 787–799, December, 2009. 

 

788


(< 1 to 266 ha) located in three landscapes with 

different levels of deforestation (24 %, 11 % and 4 

% of remaining forest cover; Arroyo-Rodríguez et 

al. 2007; 2009). Here we present a species list of 

all the plants recorded in the study landscapes and 

suggest some conservation priorities. 

 

Materials and Methods 

Study Site  

The Los Tuxtlas region is located in the southeast 

of the state of Veracruz, Mexico (18°8' - 18°45' N, 

94°37' - 95°22' W; Figure 1). The climate is warm 

and humid, with a mean annual temperature of 25 

ºC, and annual rainfall between 3,000 and 4,600 

mm. This region covers an area of 155,122 ha, 

with elevation ranging from 0 to 1,780 m above 

sea level (a.s.l.). Los Tuxtlas was decreed a 

Biosphere Reserve in 1998 owing to its 

exceptional biodiversity (CONABIO 2000). The 

original dominant vegetation type (below 700 m 

a.s.l.) was tropical rainforest, but the reserve was 

heavily deforested and fragmented between 1972 

and 1993, and the remaining rainforest is 

surrounded by a matrix of pastures and croplands 

(Castillo-Campos and Laborde 2004; Guevara et 

al. 2004; Figure 2). 

We selected three landscape fragmentation units 

(Figure 1) considering that: (1) they represent a 

gradient of rainforest deforestation; (2) they are 

all situated between 0 and 400 m a.s.l. (to avoid 

changes in vegetation associated with altitude; see 

Castillo-Campos and Laborde 2004); and (3) they 

each occupy a similar area (ca. 5,000 ha). 

Elsewhere we have presented a full description of 

the methods used to digitize the landscapes, and 

detailed the differences in spatial attributes 

between landscapes (Arroyo-Rodríguez et al. 

2007; 2009). Only a brief overview is given here. 

The three landscapes have been highly deforested

but there were notable differences in the degree of 

deforestation. The landscape with the lowest 

deforestation level (LDL) covered 5,356 ha, 24 % 

of which was rainforest distributed among 75 

patches ranging from 0.5 to 700 ha; the landscape 

with intermediate deforestation level (IDL) 

covered 4,965 ha, 11 % of which was rainforest 

distributed among 88 patches ranging from 0.5 to 

76 ha; and the landscape with the highest 

deforestation level (HDL) covered 5,046 ha, 4 % 

of which was rainforest distributed among 46 

patches ranging from 0.5 to 68 ha. 

 

 

 

Figure 1

Location of the three landscapes studied in the Los Tuxtlas Biosphere Reserve, southeastern Veracruz, 

Mexico. Black polygons represent studied patches (LDL, lowest level of deforestation, 24 % of remaining forest 

cover; IDL, intermediate deforestation level, 11 %; HDL, highest deforestation level, 4 %).

  


Lists of Species 

 

Check List, Campinas, 5(4): 787–799, December, 2009. 

 

789


 

 

Figure 2

The remaining rainforest fragments in Los Tuxtlas (A-B) are surrounded by a matrix of pastures and 

croplands, where isolated trees and live fences (i.e. several strands of barbed wire held up by a line of trees) are 

very common (C-D). Vegetation within forest fragments is highly variable in composition and structure (E-G), 

with larger fragments dominated by large trees, with a closed canopy (E-F), and smaller fragments dominated by a 

higher density of smaller trees (G). 

 

————————————————



 

 

 


Lists of Species 

 

Check List, Campinas, 5(4): 787–799, December, 2009. 

 

790


Data Collection 

Vegetation was sampled in 45 randomly selected 

rainforest fragments (15 per landscape) using the 

Gentry (1982) protocol. Within each fragment we 

randomly located ten 50 m x 2 m plots. All trees, 

shrubs, lianas, palms and herbs with dbh   2.5 cm 

were recorded. Lianas were measured at the base, 

not at dbh (Gentry 1982). Species not identified in 

the field were collected for identification in the 

MEXU (Institute of Biology, UNAM, Mexico 

City) and XAL (Institute of Ecology A.C., 

Xalapa, Veracruz) herbaria (see further details in 

Arroyo-Rodríguez and Mandujano 2006; Arroyo-

Rodríguez et al. 2007; 2009). 

 

Results and Discussion 

In total, we recorded 9,435 plants belonging to 73 

families and 372 species (Table 1). We identified 

88.4 % (n = 329) of the species and 98.8 % of the 

stems sampled. Of the 329 identified species, 320 

were dicotyledonous and 9 monocotyledonous 

(i.e. families Arecaceae, Heliconiaceae, and 

Smilacaceae) (Table 1). The 320 identified 

dicotyledonous species represent ca. 51 % of the 

dicotyledonous species reported for the Los 

Tuxtlas biology field station (627 spp.; Ibarra-

Manríquez et al. 1995; 1996a; b), and ca. 17 % of 

all the plant species (including epiphytes) reported 

for the Los Tuxtlas Biosphere Reserve (1,873 

spp.; Castillo-Campos and Laborde 2004). As two 

well-known ecological theories predict (island 

biogeography theory and metapopulation theory: 

MacArthur and Wilson 1967; Hanski 1999) we 

found that the number of species was higher in the 

landscape with lowest deforestation level (LDL = 

253 species), than in the other two landscapes 

(IDL = 160 species; HDL = 180 species) (Table 

1). 

 

The families with the highest number of species 

were Fabaceae (31 species), Rubiaceae (19), and 

Moraceae (19), together representing 21 % of all 

the identified species (Table 1). These are also the 

best represented families in the Los Tuxtlas 

biological field station (Bongers et al. 1988; 

Ibarra-Manríquez et al. 1997a). In general, the 

most common species were Astrocaryum 

mexicanum (Arecaceae), Siparuna andina 

(Monimiaceae), 



Croton schiedeanus 

(Euphorbiaceae), 



Vochysia guatemalensis 

(Vochysiaceae) and Stemmadenia donnell-smithii 

(Apocynaceae) (together representing ca. 19 % of 

all stems) (Table 1). All these species (except A. 



mexicanum) are light-demanding; a functional 

group that is common in forest gaps and close to 

forest edges (Benítez-Malvido 1998). As the 

majority of the study fragments were small (60 % 



< 5 ha), and therefore, highly affected by edge 

effects (e.g. increases in light, temperature and 

wind intensity; Saunders et al. 1991), we expected 

light-demanding species to be relatively dominant 

in terms of abundance of stems. However, as we 

have previously reported, most of the identified 

species are old-growth forest species (Arroyo-

Rodríguez et al. 2009).  

 

Eight of the 372 species sampled (2.2 %) are 



classified as Endangered by the Mexican 

government (Calophyllum brasiliense



Chamaedorea alternans,  Geonoma oxycarpa

Mortoniodendron guatemalense

Spondias 

radlkoferi,  Talauma mexicana,  Tetrorchidium 

rotundatum and Vatairea lundellii; SEMARNAT 

2002). In the LDL, we found 82 stems of 7 of 

these species, in the IDL 119 stems of 5 species, 

and in the HDL only 35 stems of 5 species. 

Furthermore, of the 148 plant species reported as 

useful for commerce (i.e. timber, fuelwood, 

ornamental, artwork, and others) in the Los 

Tuxtlas rainforest (not including epiphytes; 

Ibarra-Manríquez et al. 1997b), 113 species (76 

%) were sampled in our study fragments (Table 

1). These species made up 34 % of all identified 

species, and 52 % of all identified stems. Of the 

249 species identified in the LDL, 95 (38 %) were 

useful species. Of the 145 species identified in the 

IDL, 63 (44 %) were useful species, and of the 

154 species identified in the HDL, 65 (42 %) were 

useful species.  

 

Although evidence indicates that fragmentation 



may favor the invasion of exotic plant species in 

forest fragments (Turner et al. 1996; Dislich and 

Pivello 2002), in our sample most of the species 

(369 species, 99 %) were native to the region, and 

only five (1 %) were human-introduced species 

(Citrus sp., Coffea arabica,  Psidium guajava, 



Theobroma cacao, and Manguifera indica

representing only 0.4 % of the stems sampled 

(Table 1). This finding is similar to that reported 

by Dirzo et al. (2009) in the same region, and 

could be caused by, on the one hand, a relatively 

short amount of isolation time for the fragments 

(Turner et al. 1996; Santos et al. 2008), and, on 

the other hand, the lack of environmental 

conditions for the natural dispersion, 

establishment and development of these cultivated 

species. 

 

In conclusion, our results demonstrate that forest 



fragments may serve as reservoirs of diverse 

native plant communities, including endangered 



Lists of Species 

 

Check List, Campinas, 5(4): 787–799, December, 2009. 

 

791


and economically important plant species. In spite 

of the small size of most of the fragments, in a 

previous paper we demonstrated that the smallest 

fragments present a similar species density to the 

biggest fragments, and also that the species 

turnover (beta diversity) among fragments and 

landscapes is very high (Arroyo-Rodríguez et al. 

2009). Therefore, and in accordance with Dirzo et 

al. (2009), we believe that the conservation and 

restoration of all of the remaining forest fragments 

is necessary in order to effectively preserve the 

plant biodiversity in Los Tuxtlas region. 



 

————————————————

 

 

Table 1. 

Check list  of the Angiosperms sampled in 45 rainforest fragments located in Los Tuxtlas, Veracruz, 

Mexico. Plant nomenclature was used according to the Missouri Botanical Garden nomenclatural update database 

(Anonymous 2009). The life form (LF) and stem abundances in each fragmented landscape (LDL = lowest 

deforestation level; IDL = intermediate deforestation level; HDL = highest deforestation level) are also indicated. 

Species marked with an asterisk (*) are native species reported by Ibarra-Manríquez et al. (1997b) as useful for 

commerce (i.e. timber, fuel wood, ornamental, artwork, and others). 

 

Family Species 



LF 

LDL 

IDL 

HDL 

Total 

Actinidiaceae 



Saurauia scabrida 

Hemsl. 


Tree 

 19 


 19 

 

Saurauia 

sp. 

Tree 


 22 

 22 


 

Saurauia yasicae 

Loes. 


Tree 

15 21 7  43 

Amaranthaceae 

Iresine arbuscula Uline et W. L. Bray 

Tree 


 



Anacardiaceae 



Mangifera indica 

L. 


Tree 

  3 


 

Mosquitoxylum jamaicense Krug and Urb. 

Tree 

 

43 



 

43 


 

Spondias mombin 

L.* 


Tree 

3 22 


9 34 

 

Spondias radlkoferi 

Donn. 

Sm.* 


Tree 

33 16 22 71 

 

Tapirira mexicana 

Marchand 

Tree 

2 163 


6 171 

Annonaceae 



Cymbopetalum baillonii R. E. Fr. 

Tree 


41 

45 


49 

135 


 

Cymbopetalum penduliflorum (Dunal) Baill. 

Tree 


 

 



 

Desmopsis trunciflora var. glabra G.E. Schatz 

Tree 

10 


 

 

10 



 

Guamia sp. 

Tree 


 

20 



22 

 

Guatteria amplifolia Triana and Planch. 

Tree 

 

 



 



Malmea depressa (Baill.) R. E. Fr.  

Tree 


 

 



 

Rollinia mucosa 

Baill.* 

Tree 


27 23 95 145 

 

Tridimeris hahniana 

Baill. 

Tree 


  1 


 

Xylopia frutescens 

Aubl. 


Tree 

 4 


 4 

Apocynaceae 



Aspidosperma megalocarpon Müll. Arg.* 

Tree 


 

 



 

Forsteronia viridescens S. F. Blake 

Liana  15 

 

 



15 

 

Stemmadenia donnell-smithii (Rose) Woodson 

Tree 

29 


44 

178 


251 

 

Stemmadenia galeottiana (A. Rich.) Miers*  

Tree 



 



 

 



Tabernaemontana alba 

Mill. 


Tree 

42 20 6  68 

 

Tabernaemontana arborea 

Rose 


Tree 

7  36 38 81 

Aquifoliaceae 

Ilex quercetorum I.M. Johnst.* 

Tree 


13 


21 


 

Ilex valerioi Standl.* 

Tree 


 

11 



16 

Araliaceae 



Dendropanax arboreus (L.) Decne. and Planch.* 

Tree 


53 

65 


74 

192 


 

Oreopanax obtusifolius L. O. Williams 

Tree 


 

 



Arecaceae 



Astrocaryum mexicanum 

Liebm.* 


Palm 216 233 90  539 

 

Bactris mexicana 

Mart. 

Palm 


25 9  16 50 

 

Chamaedorea alternans H. Wendl.* 

Palm  13 

 

 



13 

 

Chamaedorea tepejilote Liebm. ex Mart.* 

Palm  23 

41 



65 

 

Desmoncus ferox 

Bartlett* 

Palm 


  4 


 

Geonoma oxycarpa 

Mart 


Palm 

  3 



Aristolochiaceae 

Aristolochia grandifolia 

Salisb. 


Liana 

  1 



 

Aristolochia ovalifolia 

Duch. 


Liana 

  1 


Asteraceae 



Eupatorium galeotti B. L. Rob* 

Shrub  45 

117 

14 


176 

 

Eupatorium quadrangulare 

DC. 

Shrub 


  1 

 



Mikania aromatica 

Oerst. 


Liana 

  1 



 

Neurolaena lobata (L.) Cass. 

Herb  2 


 

 



Lists of Species 

 

Check List, Campinas, 5(4): 787–799, December, 2009. 

 

792


Family Species 

LF 

LDL  IDL HDL  Total 

 

Tuxtla pittieri (Greenm.) Villaseñor and Strother 

Liana  4 

 

 



 

Vernonia deppeana 

Less. 

Shrub 


  1 

 



Vernonia patens 

Kunth 


Shrub 

  1 



Bignoniaceae 

Amphitecna tuxtlensis A. H. Gentry 

Tree 




13 

 

Anemopaegma chrysanthum 

Dugand 

Liana 


  1 


 

Arrabidaea verrucosa (Standl.) A. H. Gentry 

Liana  5 

 

 



 

Callichlamys latifolia (Rich.) K. Schum. 

Liana  2 

 

 



 

Mansoa hymenaea (DC.) A. H. Gentry 

Liana  3 

25 

 

28 



 

Mansoa verrucifera (Schltdl.) A. H. Gentry  

Liana  2 

 

 



 

Paragonia pyramidata (Rich.) Bureau 

Liana  6 

 

 



 

Stizophyllum riparium (Kunth) Sandwith 

Liana  1 

 

 



 

Tabebuia rosea (Bertol.) A. DC.* 

Tree 


 

 



Bombacaceae 



Bernoullia flammea 

Oliv.* 


Tree 

  7 


 

Ceiba pentandra (L.) Gaertn.* 

Tree 





12 

16 


 

Pachira aquatica 

Aublet 


Tree 

  9 



 

Quararibea funebris (La Llave) Vischer* 

Tree 


10 

 

 



10 

 

Quararibea yunckeri 

Standl. 

 

Tree 



  2 


Boraginaceae 

Cordia alliodora (Ruiz and Pav.) Oken*  

Tree 


17 

29 



49 

 

Cordia dodecandra 

DC. 

Tree 


 10 

 10 


 

Cordia megalantha S. F. Blake*  

Tree 


17 



24 

 

Cordia stellifera I. M. Johnst.* 

Tree 



 



 

 



Cordia stenoclada I. M. Johnst. 

Shrub  18 

 



24 



 

Rochefortia lundelli 

Camp* 


Tree 

  6 



Burseraceae 

Bursera simaruba 

(L.) 


Sarg.* 

Tree 


36 62 36 134 

Capparaceae 



Capparis baduca 

L. 


Tree 

14 


  14 

 

Capparis mollicella 

Standl. 

Tree 


  3 


 

Crataeva tapia 

L. 


Tree 

  7 



Caricaceae 

Carica papaya 

L. 


Tree 

3   3 6 


 

Jacaratia dolichaula (Donn. Sm.) Woodson 

Tree 


 

 



Cecropiaceae 



Cecropia obtusifolia 

Bertol.* 

Tree 

50 45 17 112 



 

Coussapoa purpusii 

Standl. 


Tree 

  2 



Celastraceae 

Crossopetalum parviflorum (Hemsl.) Lundell  

Shrub  2 

 

 



 

Perrottetia longistylis 

Rose 


Tree 

  1 



 

Wimmeria bartletti 

Lundell 


Tree 

1 6 1 8 


Chrysobalanaceae  Couepia polyandra (Kunth) Rose* 

Tree 


13 


 

14 


 

Hirtella triandra (Standl.) 

Prance* 


Tree 

 

71 14 85 



Clethraceae 

Clethra macrophylla M. Martens and Galeotti* 

Tree 


 

 



Clusiaceae 



Calophyllum brasiliense var. rekoi 

(Standl.) 

Standl.* 

Tree 


5 35 

3 43 


 

Rheedia edulis (Seem.) Planch. and Triana* 

Tree 


17 

50 


71 


 

Vismia baccifera (L.) Triana and Planch. 

Tree 


 

 



Cochlospermaceae  Cochlospermum vitifolium (Milld.) Spreng. 

Tree 

 

16 



 

16 


Combretaceae 

Combretum laxum 

Jacq. 


Liana 

  2 



 

Terminalia amazonia (J. F. Gmel) Exell 

Tree 


 

45 


 

45 


Connaraceae 

Connarus schultesii 

Standl. 


Liana 

  4 



Convolvulaceae 

Ipomoea batatas (L.) Lam. 

Liana  2 

 

 



 

Ipomoea philomega 

(Vell.)House 

 

Liana 


3   1 4 

Cyclanthaceae 



Carludovica gracilis Liebm. ex. Matuda 

Herb   




Dilleniaceae 

Tetracera volubilis L. 

Liana  1 

12 

 

13 



Ebenaceae 

Diospyros digyna 

Jacq.* 


Tree 

  6 



Elaeocarpaceae 

Sloanea medusula K. Schum. and Pittier* 

Tree 


 

34 


 

34 


Erythroxylaceae 

Erythroxylum panamense 

Turcz. 


Tree 

  2 



Euphorbiaceae 

Acalypha diversifolia 

Jacq.* 


Shrub 

21 10 14 45 

 

Adelia barbinervis Schltdl. and Cham.* 

Tree 


 

 



 

Alchornea latifolia 

Sw.* 

Tree 


5  56 16 77 

Lists of Species 

 

Check List, Campinas, 5(4): 787–799, December, 2009. 

 

793



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