Bromeliad flora of Oaxaca, Mexico: richness and distribution

Abstract

The current knowledge of the bromeliad flora of the state of Oaxaca, Mexico is presented. Oaxaca is the Mexican state with the largest number of bromeliad species. Based on the study of 2,624 herbarium specimens corresponding to 1,643 collections, and a detailed bibliographic revision, we conclude that the currently known bromeliad flora for Oaxaca comprises 172 species and 15 genera. All Mexican species of the genera Bromelia, Fosterella, Greigia, Hohenbergiopsis, Racinaea, and Vriesea are represented in the state. Aechmea nudicaulis, Bromelia hemisphaerica, Catopsis nitida, C. oerstediana, C. wawranea, Pitcairnia schiedeana, P. tuerckheimii, Racinaea adscendens, Tillandsia balbisiana, T. belloensis, T. brachycaulos, T. compressa, T. dugesii, T. foliosa, T. flavobracteata, T. limbata, T. maritima, T. ortgiesiana, T. paucifolia, T. pseudobaileyi, T. rettigiana, T. utriculata, T. x marceloi, Werauhia pycnantha, and W. nutans are recorded for the first time from Oaxaca. Collections from 226 (of 570) municipalities and all 30 districts of the state were studied. Among the vegetation types occurring in Oaxaca, oak forest is the richest with 83 taxa, followed by tropical deciduous forest with 74, and cloud forest with 73 species. Species representation and distribution in Oaxaca are analyzed in detail. We also provide a comparison with bromeliad floras of the states of Chiapas, Guerrero, Puebla and Veracruz. The analysis of the species and collections by altitudinal intervals shows that the highest numbers of both ocurre between 1,500 and 2,000 m, with the number of species markedly decreasing above 2,500 m.

Full text

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Acta Botanica Mexicana 81: 71-147 (2007)
BROMELIAD FLORA OF OAXACA, MEXICO: RICHNESS AND
DISTRIBUTION
Adolfo EspEjo-sErnA1, AnA rosA lópEz-fErrAri1,nAncy MArtínEz-corrEA1 And
VAlEriA AngélicA pulido-EspArzA2
1Universidad Autónoma Metropolitana-Iztapalapa,
División de Ciencias Biológicas y de la Salud, Departamento de Biología,
Herbario Metropolitano, Apdo. postal 55-535, 09340
México, D.F., México. aes@xanum.uam.mx
2El Colegio de la Frontera Sur - San Cristóbal de las Casas,
Laboratorio de Análisis de Información Geográfica y Estadística,
Chiapas, México. feliscatus@yahoo.com
ABSTRACT
The current knowledge of the bromeliad flora of the state of Oaxaca, Mexico
is presented. Oaxaca is the Mexican state with the largest number of bromeliad species.
Based on the study of 2,624 herbarium specimens corresponding to 1,643 collections,
and a detailed bibliographic revision, we conclude that the currently known bromeliad
flora for Oaxaca comprises 172 species and 15 genera. All Mexican species of the genera
Bromelia, Fosterella, Greigia, Hohenbergiopsis, Racinaea, and Vriesea are represented in
the state. Aechmea nudicaulis, Bromelia hemisphaerica, Catopsis nitida, C. oerstediana,
C. wawranea, Pitcairnia schiedeana, P. tuerckheimii, Racinaea adscendens, Tillandsia
balbisiana, T. belloensis, T. brachycaulos, T. compressa, T. dugesii, T. foliosa, T.
flavobracteata, T. limbata, T. maritima, T. ortgiesiana, T. paucifolia, T. pseudobaileyi, T.
rettigiana, T. utriculata, T. x marceloi, Werauhia pycnantha, and W. nutans are recorded for
the first time from Oaxaca. Collections from 226 (of 570) municipalities and all 30 districts
of the state were studied. Among the vegetation types occurring in Oaxaca, oak forest is the
richest with 83 taxa, followed by tropical deciduous forest with 74, and cloud forest with 73
species. Species representation and distribution in Oaxaca are analyzed in detail. We also
provide a comparison with bromeliad floras of the states of Chiapas, Guerrero, Puebla and
Veracruz. The analysis of the species and collections by altitudinal intervals shows that the
highest numbers of both ocurre between 1,500 and 2,000 m, with the number of species
markedly decreasing above 2,500 m.
Key words: Bromeliaceae, distribution, diversity, endemism, flora, Mexico, Oaxaca.

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Acta Botanica Mexicana 81: 71-147 (2007)
72
RESUMEN
Se presenta el estado actual del conocimiento de la flora bromeliológica del estado
de Oaxaca, México. La entidad ocupa el primer lugar en el país en cuanto a número de
especies de Bromeliaceae se refiere. Los resultados obtenidos de la revisión de 2,624
ejemplares herborizados, correspondientes a 1,643 colectas, así como la revisión de
bibliografía especializada, muestran que en el estado están presentes 172 especies
agrupadas en 15 géneros. Bromelia, Fosterella, Greigia, Hohenbergiopsis, Racinaea
y Vriesea tienen representados a todos sus taxa mexicanos. Se registran por primera
vez para el estado: Aechmea nudicaulis, Bromelia hemisphaerica, Catopsis nitida, C.
oerstediana, C. wawranea, Pitcairnia schiedeana, P. tuerckheimii, Racinaea adscendens,
Tillandsia balbisiana, T. belloensis, T. brachycaulos, T. compressa, T. dugesii, T. foliosa,
T. flavobracteata, T. limbata, T. maritima, T. ortgiesiana, T. paucifolia, T. pseudobaileyi,
T. rettigiana, T. utriculata, T. x marceloi, Werauhia pycnantha y W. nutans. Se registraron
colectas para 226 municipios de los 570 y para el total de los distritos (30) en los que
está dividido políticamente el estado. Se hizo una comparación de la flora bromeliológica
de Oaxaca con la de Chiapas, Guerrero, Puebla y Veracruz. De los tipos de vegetación
presentes, el Bosque de Quercus es el que presenta mayor riqueza de taxa (83), seguido por
el Bosque Tropical Caducifolio (74) y el Bosque Mesófilo de Montaña (73). El análisis del
número de especies y de colecciones por intervalo altitudinal muestra que las cantidades
más altas, tanto de colectas como de especies, se concentran entre los 1,500 y los 2,000 m
s.n.m., disminuyendo claramente por arriba de los 2,500 m.
Palabras clave: Bromeliaceae, distribución, diversidad, endemismo, flora, México,
Oaxaca.
INTRODUCTION
The Bromeliaceae contain approximately 3,086 species in 56 genera (Luther,
2006); in Mexico there are 18 genera and 342 species (Espejo et al., 2004). The fami-
ly is well distributed across the country, as well as in all the vegetation types sensu
Rzedowski (1978). It is a valuable component of many different plant associations,
not only for the shear abundance of its species but also for the ecological role they
play.
The family is important on many levels. Ecologically they are central epiphy-
tic components in various ecosystems, possessing various physiological adaptations
for atmospheric stress and serving as unique habitats for many animal species. Eco-
nomically, bromeliads are valuable because of their ornamental and edible species.

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Espejo-Serna et al.: Bromeliad flora of Oaxaca
Their floristic importance relates to their wide distribution throughout the Neotro-
pics and the high number of narrowly endemic species. Despite these factors, the
family has received relatively little attention with regard to floristic studies.
Although there are various regional treatments for the Mexican Bromeliaceae
(e.g., McVaugh (1989), Utley and Burt-Utley (1994), Victoria (2001), Arellano Mijan-
gos (2002), Pulido-Esparza et al. (2004), Espejo et al. (2005), and Ramírez-Morillo
et al. (2004)), the only two contributions of national scope are those of Smith and
Downs (1974; 1977; 1979) and Espejo et al. (2004). The purpose of this article is to
provide an updated list of the members of the family in Oaxaca, with emphasis on
aspects of endemism and geographic distribution.
Study area
Oaxaca is located in the southern portion of Mexico between 15°39' and
18°09' N latitude and 93°52' and 98°32' W longitude and covers an area of 93,343
km2, 4.8% of the country’s surface (Anonymous, 2004) (Fig. 1). The state is poli-
tically divided into 30 districts and 570 municipios that include more than 10,000
towns and villages (Anonymous, 2004). It also is divided economically into eight
regions: Cañada, Costa, Istmo, Mixteca, Papaloapan, Sierra Norte, Sierra Sur, and
Valles Centrales (Anonymous, 2004).
Five physiographic regions are distinguished in Oaxaca (Anonymous, 2004):
1) Sierra Madre del Sur, 2) Cordillera Centroamericana, 3) Llanura Costera del
Golfo sur, 4) Eje Neovolcánico, and 5) Sierras de Chiapas y Guatemala. Elevation
varies from sea level along the coastal plains at the Pacific Ocean to 3,000 m in the
“Sierra Madre de Oaxaca” (Ortíz-Pérez et al., 2004).
Also, five climatic regions are distinguished in Oaxaca (Trejo, 2004): 1)
Cuenca del Balsas, 2) Sierra Madre del Sur y Llanura Costera, 3) Mixteca y Valles
de Oaxaca, 4) Sierra Madre de Chiapas y Llanura Costera, and 5) Sierra Madre
Oriental.
The climatic and physiographic variation is reflected in the diversity of ve-
getation communities that exists in Oaxaca, and these communities possess the
greatest biological diversity of all Mexican states. Within the state the following
types of vegetation (sensu Rzedowski, 1978) have been reported: Bosque Tropi-
cal Perennifolio, Bosque Tropical Subcaducifolio, Bosque Tropical Caducifolio,
Bosque Espinoso, Matorral Xerófilo, Bosque de Quercus, Bosque de Coníferas,
Bosque Mesófilo de Montaña, Palmar, Bosque de Galería,Vegetación Acuática y
Subacuática.

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Acta Botanica Mexicana 81: 71-147 (2007)
74
Fig. 1. Oaxaca State location in Mexico and known collection sites of the bromeliad species within the state.

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Espejo-Serna et al.: Bromeliad flora of Oaxaca
METHODS
From June of 1997 until June of 2006, Oaxacan specimens from the follo-
wing herbaria were examined, identified, annotated, and captured in a database: B,
BM, BR, C, CHAP, CICY, CIIDIR, CODAGEM, ECON, ENCB, F, FCME, FI, GH,
GOET, HEID, IBUG, IEB, K, LG, LL, M, MA, MEXU, MICH, MO, NY, OXF, P,
SEL, SERO, TEX, UAMIZ, UC, UNICACH, US, VT, WU, XAL, and Z. The infor-
mation from these specimens was enhanced by a carefull revision of pertinent bi-
bliographic material (Baker, 1889; Mez, 1896; Smith, 1961; Smith & Downs, 1974;
1977; 1979; Grant, 1995 a; b; Palací, 1997; Till, 1990, 1992).
In order to analyze the distribution of the species, geographic coordinates
were obtained from the labels of the examined specimens. These were checked and
corrected if necessary, and subsequently, the data were processed in the program
Arc View GIS 8 (Anonymous, 2003). In instances where such information was lac-
king, the location was obtained from topographic 1:250,000 scale maps (Instituto
Nacional de Estadística, Geografía e Informática (INEGI) maps D14-3, E14-6, E14-
8, E14-9, E14-12, E15-7 and E15-10), and from the web site http://mapserver.inegi.
gob.mx/dsist/ahl2003/index.html?c=424.
Two data matrices for species presence-absence were constructed, one relati-
ve to the vegetation types and other relative to the adjacent states (Guerrero, Chia-
pas, Puebla, and Veracruz). These matrices were used to obtain Jaccard indices of
similarity. Also, cluster analyses were done in order to determine the affinities of
the Oaxacan bromeliad flora with respect to that of the adjacent states and to eva-
luate the similarity of the flora with respect to the types of vegetation present in
the state. These analyses were conducted using the program NTSYSpc2.0 (Rohlf,
1998).
A species accumulation curve was generated to assess the knowledge and
extent of exploration of the state´s bromeliad flora. The collection year information
used to generate this curve was obtained directly from herbarium labels, using as a
sample the total number of species recorded each five years. The starting point was
1840, the year of the first known collection of Bromeliaceae in Oaxaca. In addition,
a rarefaction curve, taking into account the abundance of each taxon, was genera-
ted (Colwell & Coddington, 1994) to compare the data obtained in the accumula-
tion curve with that expected using Chao1 and Jacknife 2 estimators (Chao, 1984;
Colwell & Coddington, 1994). The algorithms were calculated with the program
EstimatesS Ver. 7 (Colwell, 2004).

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Acta Botanica Mexicana 81: 71-147 (2007)
76
RESULTS AND DISCUSSION
After examining 2,624 herbarium sheets, corresponding to 1,643 collections,
we can say now that the known bromeliad flora of the state is composed by 172
species and 15 genera (appendix). This figure represents 50.29% of the 342 species
of the family registered for the country (Espejo et al., 2004). In contrast, the two
previous publications that provide integral information about the bromeliad Flora of
Oaxaca, Flora Neotropica (Smith & Downs, 1974, 1977, 1979) and the Checklist of
Mexican Bromeliaceae (Espejo et al., 2004), reported 14 genera, 88 species based on
213 collections and 15 genera, 135 species based on 1482 collections, respectively.
The genera with the greatest number of species in the area are Tillandsia (101
spp.) and Hechtia (20 spp.). In contrast, Billbergia, Fosterella, Guzmania and Ho-
henbergiopsis each possess a single species. Bromelia, Fosterella, Greigia, Hohen-
bergiopsis, Racinaea, and Vriesea have all their Mexican taxa represented in Oaxa-
ca. The following species are reported from the state for the first time: Aechmea
nudicaulis, Bromelia hemisphaerica, Catopsis nitida, C. oerstediana, C. wawranea,
Pitcairnia schiedeana, P. tuerckheimii, Racinaea adscendens, Tillandsia balbisia-
na, T. belloensis, T. brachycaulos, T. compressa, T. dugesii, T. foliosa, T. flavobrac-
teata, T. limbata, T. maritima, T. ortgieseana, T. paucifolia, T. pseudobaileyi, T.
rettigiana, T. utriculata, T. x marceloi, Werauhia nutans, and W. pycnantha.
Oaxaca, with 43 narrowly endemic species (Appendix; Figs. 10, 11, 12, 13, 14,
15, 16, 17, 18, 19, 20, 21, and 22), possesses a higher number of endemics than any
other state in the country. These endemics belong to four genera: Hechtia (13 spp.),
Pitcairnia (1 sp.), Tillandsia (28 spp.), and Viridantha (1 sp.). Of all species present
in the state, 95 (55.23%) are endemic to Mexico and represent eight genera: Brome-
lia (2 spp.), Catopsis (1 sp.), Greigia (2 spp.), Hechtia (20 spp.), Pitcairnia (6 spp.),
Tillandsia (59 spp.), Viridantha (4 spp.) and Werauhia (1 sp.).
It is worth pointing out that the majority of the Bromeliaceae collections are
concentrated in the north, north-west and center portions of the state along its prin-
cipal roadways (Fig. 1). Collections are known from 226 (39.64%) of the 570 muni-
cipalities and from all of the districts (Table 1).
The analysis of species number by altitude (Fig. 2) shows that 79 species, or
45.93 % of the total, ocurr between 1,750 and 2,000 m. The altitudinal average varies
between 30 and 45 species in the other altitudinal intervals, and decreases suddenly
above 2,500 m. A similar analysis for the collections is showed in figure 3. The gra-
ph shows two peaks, one between 0 and 250 m and another between 1,750 and 2,000
m. The distribution of the collections between intervals is heterogeneous.

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Espejo-Serna et al.: Bromeliad flora of Oaxaca
District
Number of
collections
193
174
135
109
85
88
84
83
63
54
57
53
49
46
44
39
41
28
32
29
28
21
19
16
10
Km2
Collection index
Standarized
collection index
24.11
100.00
36.98
32.57
64.16
38.79
44.69
59.33
38.89
40.10
60.87
17.74
24.69
19.14
17.02
55.30
36.34
27.33
20.65
17.27
13.13
12.70
57.34
7.25
9.15
7.01
9.07
1.16
1.69
1.92
Juchitán
Ixtlán de Juárez
Tehuantepec
Tuxtepec
Cuicatlán
Pochutla
Tlacolula
Teotitlán
Tlaxiaco
Etla
Teposcolula
Mixe
Huajuapam
Miahuatlán
Yautepec
Villa Alta
Juxtlahuaca
Coixtlahuaca
Putla
Nochixtlán
Juquila
Choapam
Centro
Sola de Vega
Silacayoapam
Ejutla
Zaachila
Jamiltepec
Zimatlán
Ocotlán
Without data
13189
2867
6015
5515
2183
3738
3097
2305
2669
2219
1543
4922
3270
3959
4259
1162
1859
1688
2553
2767
3514
2725
546
3635
1800
940
545
4276
975
858
1.46
6.07
2.24
1.98
3.98
2.35
2.71
3.60
2.36
2.43
3.69
1.08
1.50
1.16
1.03
3.36
2.21
1.66
1.25
1.05
0.80
0.77
3.48
0.44
0.56
0.43
0.55
0.07
0.10
0.12
-
4
3
3
1
1
43 --
Table 1. Number of collections and indices of collection density fot each district.

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Acta Botanica Mexicana 81: 71-147 (2007)
78
Fig. 2. Altitudinal distribution of the bromeliad species in Oaxaca.
Fig. 3. Altitudinal distribution of the bromeliad collections in Oaxaca.
0-250 251-500501-750 751-1000 1001-12501251-15001501-1750 1751-20002001-22502251-2500 2501-27502751-3000
Altitude (m)
Number of species
0
10
20
30
40
50
60
70
80
90
250.0
300.0
225.0
150.0
75.0
0.0
20.0
15.0
10.0
5.0
0.0
0.0
500.0750.0
1000.0 1250.01500.0 1750.02000.0 2250.02500.02750.03000.0
Altitude (m)
Collection number
Collection percentage

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Espejo-Serna et al.: Bromeliad flora of Oaxaca
Concerning genera (Fig. 4), their distribution is more or less homogeneous,
the average varying between 7 and 11 genera per altitudinal interval, and, as with
species, they decrease suddenly above 2,500 m. All the Oaxacan species of Aech-
mea, Billbergia, Bromelia, Fosterella, Guzmania, and Vriesea, grow below 1,500 m.
The genus Tillandsia is well represented from the sea level to 3,000 m.
The species accumulation curve (Fig. 5) shows that even though there has
been a substantial increase in the number of Bromeliaceae species and collections
reported, the asymptote had not been reached. This suggests that the state has not
been sufficiently explored. The rarefaction curve, and the Chao and Jacknife esti-
mates show that the predicted number of Bromeliaceae species for the state varies
between 219 and 243, and thus between 50 and 74 species remain to be encountered
yet.
These results can be explained by the fact that there is a tendency to co-
llect along highways and roads, and this has led to a disproportionate amount of
collecting in relatively few areas. In contrast, there are many areas that have not
been explored at all. For example, of the 570 municipalities, only in 226 are known
collections of Bromeliaceae. In order to increase our knowledge about the floristics
and biogeography of the family in the state, it is necessary to undertake a much more
extensive sampling in areas that are far from the principal roads. However, it should
be noted that a large portion of Oaxaca (and Mexico in general) has suffered severe
loss of primary vegetation, and it is therefore possible that even with an extensive
collecting effort, few new records may be found.
Fig. 4. Altitudinal distribution of the bromeliad genera in Oaxaca.
0
2
4
6
8
10
12
0-250251-500 501-750751-1000 1001-1250 1251-15001501-17501751-20002001-2250 2251-25002501-27502751-3000
Altitude (m)
Number of genera

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Acta Botanica Mexicana 81: 71-147 (2007)
80
With respect to the comparison between the bromeliad flora of Oaxaca with
that of the neighbour states, the dendrogram shows that the Floras of Chiapas and
Veracruz are very similar and both, as a group, are highly similar to that of Oaxaca
(Fig. 6).
The similarity of the Oaxacan bromeliad flora with that of Chiapas-Veracruz
is mainly attributed to the Mesoamerican influence in these states. Another factor
is the continuous extension of vegetation between these three states, that is repre-
sented by tropical forest that extends from northern South America through Central
America. The southern states of Mexico represent the northernmost limit of the
distribution of various taxa, and in these states the geographic and biological con-
ditions have generated a particular flora. Puebla also shares some elements with the
Oaxaca-Chiapas-Veracruz group, but is clearly separate from them (Jaccard index =
0.22). Puebla appears to possess a much stronger influence from the Eje Neovolcáni-
co Transmexicano and the Sierra Madre Oriental. The bromeliad flora of Guerrero
has the least similarity in comparison that of the other states analyzed (Jaccard index
= 0.21).
Although Oaxaca shares many floristic elements with adjacent regions, it re-
presents a special case, due to its great ecological diversity, ranging from the arid re-
gions of the north to the coastal regions and central zones, the latter of which include
cloud forests and a grand extension of coniferous forests (Table 2).
Fig. 5. Rarefaction and species accumulation curves.
Lustrum
Species number
Accumulation
Chao 1 mean
Jack 1 mean
Jack 2 mean
Cole rarefaction
0
50
100
150
200
250
1357911 13 1517 1921 23 2527 29 3133 35

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Espejo-Serna et al.: Bromeliad flora of Oaxaca
With regard to the comparison of bromeliads among the types of vegetation
present in the state, the dendrogram (Fig. 7) shows that if we exclude from the analy-
sis the Palmares (because it is generally found distributed in small patches within
other types of vegetation where it is associated with humid and generally warm cli-
mates or secondary vegetation), there are two main groups (Jaccard Index = 0.04).
One contains the majority of the vegetation types (seven in total). Within this graph
Bosque de Coníferas and the Bosque de Quercus are the most similar (Jaccard In-
dex = 0.39) and are grouped with the Bosque Mesófilo de Montaña (Jaccard Index
= 0.32). The second main group encompasses two subgroups: one conformed by the
Vegetación Acuática (mainly Manglares) and the Bosque Espinoso which are always
found in the southeast coastal region of the state, and the other represented by the
Bosque Tropical Subcaducifolio. Factors such as soil type, temperature and humi-
dity microhabitats and other environmental conditions where these communities
are found could be why the bromeliad floras are so similar within these vegetation
types.
Of the 11 types of vegetation (sensu Rzedowski, 1978) present in the state,
the Bosque de Quercus is the richest in number of species (83), followed by Bosque
Tropical Caducifolio with 74 species, and Bosque Mesófilo de Montaña with 73
taxa. Five species are restricted to Bosque de Coníferas: Tillandsia festucoides, T.
leucolepis T. quaquaflorifera, T. x marceloi, and T. wuelfinghoffi. Thirteen species
Fig. 6. Similarity of the bromeliad floras between Oaxaca and adjacent states.
Oaxaca
Chiapas
Veracruz
Puebla
Guerrero
0.210.25 0.29
Jaccard
0.340.38

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Acta Botanica Mexicana 81: 71-147 (2007)
82
Table 2. Total number of known species by state, shared species with Oaxaca and similarity
index (Jaccard).
State
Total number of
known species
123
88
57
89
Number shared
with Oaxaca
75
55
44
69
Similarity index (Jaccard)
Chiapas
Guerrero
Puebla
Veracruz
34.42
27.5
24.44
36.9
are known only from Bosque de Encino: Bromelia hemisphaerica, Hechtia nuusvio-
rum and two undescribed species of Hechtia, Pitcairnia schiedeana, P. tuerckhei-
mii, Tillandsia atroviolacea, T. lampropoda, T. rettigiana, T. rubrispica, T. ulrici,
T. yerbasantae, and Viridantha penascoenesis. Thirteen species are exclusive to
Bosque Mesófilo de Montaña: Catopsis nitida, C. oerstediana, Greigia vanhynin-
gi, Racinaea adscendens, Tillandsia belloensis, T. copalaensis, T. kirchhoffiana, T.
kolbii, T. laui, T. lautneri, T. leiboldiana, T. mixtecorum, and Werauhia nocturna.
Seventeen species are confined to Bosque Tropical Caducifolio: Hechtia caudata,
H. fosteriana, H. fragilis, H. lanata, H. lyman-smithii, H. marnier-lapostollei, and
three additional undescribed species, Tillandsia atenangoensis, T. califanii, T. gla-
brior, T. ortgiesiana, T. pseudobaileyi, T. schatzlii, T. schusteri, and T. vernardoi.
Twelve species are limited to Bosque Tropical Perennifolio: Aechmea magdalenae,
A. nudicaulis, Catopsis wawranea, Pitcairnia undulata, Tillandsia flavobractea-
ta, T. foliosa, T. limbata, T. pruinosa, Vriesea heliconioides, V. malzinei, Werauhia
gladioliflora, and W. nutans. Tillandsia socialis grows only in Bosque Tropical Sub-
caducifolio. Two species are restricted to Matorral Xerófilo: Hechtia confusa, and
Viridantha mauryana. Tillandsia balbisiana occurs solely in Bosque Espinoso.
ACKNOWLEDGMENTS
We thank the following herbaria for providing specimens and data used in
this work: B, BM, BR, C, CHAP, CICY, CIIDIR, CODAGEM, ECON, ENCB, F,
FCME, FI, GH, GOET, HEID, IBUG, IEB, K, LG, LL, M, MA, MEXU, MICH,
MO, NY, OXF, P, SEL, SERO, TEX, UAMIZ, UC, UNICACH, US, VT, WU, XAL,
and Z; the Elizabeth Bascom Fellowship for Latinoamerican Women and the Mis-

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Espejo-Serna et al.: Bromeliad flora of Oaxaca
Fig. 7. Similarity of the bromeliad flora between the types of vegetation in Oaxaca. BC:
Bosque de Coníferas (Coniferous forest); BQ: Bosque de Quercus (Oak forest); BMM:
Bosque Mesófilo de Montaña (Cloud forest); BTC: Bosque Tropical Caducifolio (Tropical
deciduous forest); BTP: Bosque tropical perennifolio (Tropical evergreen forest); MX:
Matorral xerófilo (Xerophytic scrub); BG: Bosque de Galería (Gallery forest); BE: Bosque
Espinoso (Thorn forest); VA: Vegetación Acuática (Aquatic vegetation); BTSC: Bosque
tropical subcaducifolio (tropical subdeciduous forest); PAL: Palmar (Palm savanna).
souri Botanical Garden for a scholarship granted to the second author; Jacqueline
Ceja Romero and Aniceto Mendoza Ruiz for their invaluable and constant help in
the field work; Victor Steinmann for his help with the English translation; and Javier
García-Cruz suggestions and improvements to the manuscript.
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Recibido en julio de 2006.
Aceptado en julio de 2007.