Acta Botanica Mexicana 100: 293-315 (2012)
ENDEMIC ANGIOSPERM LINEAGES IN MEXICO: HOTSPOTS FOR
CONSERVATION
Victoria SoSa1,4 and J. arturo de-noVa2,3
Instituto de Ecología A.C., Biología Evolutiva,
Apdo. postal 63, 91070 Xalapa, Veracruz, Mexico.
2
Universidad Autónoma de San Luis Potosí, Instituto de Investigación en Zonas
Desérticas, Altair 200, 78377 San Luis Potosí, Mexico.
3
Universidad Autónoma de San Luis Potosí, Facultad de Agronomía, km 14.5
carretera San Luis Potosí-Matehuala, 78321 San Luis Potosí, Mexico.
4
Autor para la correspondencia: victoria.sosa@inecol.edu.mx
1
ABSTRACT
As a megadiverse country, Mexico harbors 4 to 8% of the lora of the world and
of this, 51% is endemic. There is concern because several factors are impeding its
conservation. In this paper, areas of endemism for the lowering plants of Mexico are
identiied to prioritize regions for conservation. To categorize zones for preservation, the
approach followed takes biodiversity, weighted endemism and evolutionary history into
account. Lineages of angiosperms, families, genera, and formal or informal groups within
genera previously retrieved as monophyletic are selected to represent evolutionary history
in equivalent spatial units. A database with 9416 entries based on specimens of species
belonging to 259 monophyletic groups of angiosperms from Mexico was compiled, and their
presence-absence recorded for every unit area. Species richness and weighted endemism
index was calculated for each of these units. The results indicate that the majority of the
regions with the highest indices of endemism have a dry climate with xeric vegetation,
with the exception of two areas of tropical and temperate forests. They are: the northeastern
rosette scrub in Nuevo León and Coahuila, gypsum grasslands in San Luis Potosí, the Sierra
Gorda in Queréraro, Tolantongo in Hidalgo, the Tehuacán-Cuicatlán Valley in Puebla and
Oaxaca, El Salto in Durango, Sierra de Quila in Jalisco, a western portion of the Balsas
River Basin in Michoacán, Guerrero, Morelos and State of Mexico, the Tehuantepec area in
Oaxaca, the Central Depression of Chiapas and El Triunfo in Chiapas. Some of the areas of
endemism in the Chihuahuan Desert, Balsas River Basin, the Central Depression of Chiapas
and the southern area of Oaxaca are not suficiently protected. Approximately 340 species
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were microendemic, i.e. restricted to a single quadrat, and the Cactaceae account for the
majority of the species on the Mexican Red List.
Key words: Cactaceae, Chihuahuan Desert, endemism, Mega-Mexico, xeric
vegetation.
RESUMEN
México está considerado como uno de los países megadiversos y en su territorio
se encuentran entre 4 y 8% del número de total de especies de plantas del mundo, de las
cuales 51% son endémicas. Existe una gran preocupación sobre la conservación de la lora
mexicana, ya que se han detectado varias actividades y factores que la amenazan. En este
trabajo se identiican áreas de endemismo para las angiospermas de México con el objetivo
de priorizar regiones para conservación. Para categorizar estas zonas se sigue el enfoque
que toma en cuenta la biodiversidad, el índice de endemismo ponderado y la historia
evolutiva. Se identiicaron los linajes de angiospermas, ya sean familias, géneros, o grupos
infragenéricos con o sin estatus taxonómico que previamente se habían determinado como
monoiléticos para representar la historia evolutiva en unidades espaciales equivalentes.
Se construyó una base de datos de 9416 registros de especies de 259 grupos monoiléticos
de angiospermas restringidas a México y se registró su presencia en estas áreas. Para cada
una se calculó la riqueza de taxones y el índice de endemismo ponderado. Los resultados
muestran que la mayoría de las zonas de más alto endemismo están en climas secos, con
vegetación xérica, con dos excepciones de vegetación tropical y templada. Los índices de
endemismo ponderado más altos se localizaron en: el área norte de matorral rosetóilo
en Nuevo León y Coahuila, matorrales gipsóilos en San Luis Potosí, la Sierra Gorda en
Querérato, Tolantongo en Hidalgo, el Valle de Tehuacán-Cuicatlán en Puebla y Oaxaca, El
Salto en Durango, la Sierra de Quila en Jalisco, la zona oeste de la Depresión del Balsas en
Michoacán, Guerrero, Morelos y el Estado de México, la zona de Tehuantepec en Oaxaca
y El Triunfo en Chiapas. Algunas áreas de endemismo en el Desierto Chihuahuense, en
la Cuenca del Balsas y en la Depresión de Chiapas, así como del sur de Oaxaca no están
suicientemente protegidas. Se registraron aproximadamente 340 especies con distribución
restringida a un solo cuadrante y de éstas la mayoría de las que se incluyen en la lista de
taxones amenazados de México pertenecen a las Cactaceae.
Palabras clave: Cactaceae, Desierto Chihuahuense, endemismo, Mega-México,
vegetación xérica.
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INTRODUCTION
Different criteria have been applied to prioritize areas for conservation and
undoubtedly the most widely used is the concentration of endemic species undergoing increased habitat loss. The areas defined this way are known as “biodiversity
hotspots” (Myers et al., 2000). Globally, thirty-five biodiversity hotspots, comprising 44% of the total number of species of vascular plants, have been identified and
Mesoamerica is one of them (Myers et al., 2000; Mittermeier et al., 2011).
Other approaches for prioritizing areas for conservation have added the element of evolutionary history, because diversity and evolutionary history are unequally distributed in different areas of the world (Vane-Wright et al., 1991). Among
these approaches, phylogenetic endemism uses phylogenetic diversity and weighted
endemism as measures to identify areas for conservation based on equivalent spatial units (Rosauer et al., 2009). Additionally, attributes such as scarcity (Cadotte &
Davies, 2010), local and global rarity of taxa (e.g. Crain et al., 2011), phylogenetic
distinctiveness and isolation (e.g. Collen et al., 2011), phylogenetic diversity (Forest
et al., 2007) and functional diversity (e.g. Devictor et al., 2010; Pio et al., 2011) have
variously been incorporated to the evolutionary history approach.
In this paper, we identify areas of endemism for the flowering plants of Mexico
to prioritize areas for conservation. The concept followed here for an area of endemism is that of a geographic region that includes the distributions of two or more
monophyletic taxa with phylogenetic and distributional congruence (Harold & Mooi,
1994). Areas of endemism have several attributes: they have a single history, they are
smaller than the entire study area, they do not overlap with other areas of endemism,
they host at least two taxa with ranges restricted to the area and they are maximally
congruent (Linder, 2001; Szumik et al., 2004; Ebach et al., 2008).
Thus, to categorize areas for preservation we follow an approach that combines biodiversity, weighted endemism and evolutionary history. Lineages of Mexican angiosperms, families, genera or formal and informal groups within genera that
have been previously recognized as monophyletic were selected to represent evolutionary history in equivalent spatial units.
As a megadiverse country, Mexico houses 4 to 8% of the flora of the world and
there is concern because several factors are impeding its conservation. The greatest
threats to the flora of Mexico are intensification of habitat loss, the adverse effects of
climate change and the overexploitation of the majority of habitats (Dávila et al., 2011).
Angiosperms were chosen because they are one of the most diverse groups of
organisms in Mexico. Their diversity has been estimated at 24,500-29,000 species
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(Villaseñor, 2003; Espejo-Serna et al., 2004) and more than 50% are endemic to the
country (Rzedowski, 1993). Furthermore, the genera distributed in Mexico have
been documented (Villaseñor, 2004), and the floristic knowledge of the country has
been recently summarized (Anonymous, 2009). The groups of angiosperms distributed in the area known as Mega-Mexico were used in this study. This biogeographic
province was proposed by Rzedowski (1993) and includes, in addition to Mexico’s
current territory, the areas of the Sonoran Desert, the Chihuahuan Desert and the
Tamaulipan scrub that lie in the United States of America, as well as those portions
of Central America as far south as northern Nicaragua.
The families with the largest number of endemic Mexican genera are Cactaceae and Asteraceae (Turner, 1996-2010; Guzmán et al., 2003; Hernández & GómezHinostrosa, 2011a,b). Setchellanthaceae, a monotypic family, only grows in Mexico
(Iltis, 1999). In the monocots, a clade of geophyte genera in the Asparagaceae, the
Milla clade, grows in Mega-Mexico (Gándara et al., 2009) and a group in the Crassulaceae, the Acre clade includes several genera exclusive to Mexico (Acevedo-Rosas
et al., 2004; Carrillo-Reyes et al., 2010). Three related genera, Morkilia, Sericodes
and Viscainoa in the Zygophyllaceae (Sheahan & Chase, 2006), and two genera
in the Anacardiaceae, Bonetiella and Pseudosmodingium (Aguilar-Ortigoza et al.,
2004) are found in this biogeographic province. In the Acanthaceae, nine genera and
a clade within Ruellia are endemic to Mexico (Daniel, 1993; Tripp, 2010). In addition,
among the more remarkable endemic groups of Mexico are clades of Bursera (Rzedowski et al., 2005; De-Nova et al., 2012), Agave and groups nested within this genus
such as Manfreda, Polianthes and Prochnyanthes (García-Mendoza, 1995; Rocha et
al., 2006, Good-Avila et al., 2006), the section Physodium in Melochia (Dorr & Barret, 1989), a clade in the Zea diploperennis group (Poaceae) (Buckler & Holtsford,
1996), and a clade of Yucca within the Sarcocarpa group (Pellmyr et al., 2007).
Fouquieria and Leucophyllum are arid land groups in Mega-Mexico (Henrickson &
Flyr, 1985; Schultheis & Baldwin, 1999). Enigmatic genera like Velascoa (Crossosomataceae) (Sosa & Chase, 2003), Chiangiodendron (Achariaceae) (Sosa et al., 2005),
Enriquebeltrania (Euphorbiaceae) (De-Nova et al., 2006), Cerdia (Caryophyllaceae)
(Sosa et al., 2006), Olmeca (Bambusoideae, Poaceae) (Dávila-Aranda et al., 2004;
Ruiz-Sanchez et al., 2011), Peltophorum (Leguminosae) (Sousa, 2005), the parasitic
Eremitilla (Orobanchaceae) (Yatskievych & Contreras-Jiménez, 2009), Echinopterys (Malpighiaceae) (Davis et al., 2001), Nowickea (Phytolaccaceae) (Martínez
& McDonald, 1989), and Mexipedium (Orchidaceae) (Albert & Chase, 1992) are
endemic to Mexico, to mention just a few examples. We recorded 259 monophyletic
angiosperm groups endemic to Mega-Mexico.
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Sosa y De-Nova: Endemic angiosperm lineages in Mexico
The objectives of this paper are: 1) to identify the areas of endemism of the
angiosperms of Mexico, using monophyletic groups to prioritize areas for conservation, and 2) to detect species from these natural groups with a restricted distribution
to highlight the threatened taxa.
MATERIALS AND METHODS
Taxa
Mexican angiosperm lineages, families, genera, and infrageneric groups
with or without formal taxonomic status were compiled based on the literature
(Rzedowski, 1993; Villaseñor, 2004; Anonymous, 2009). Distribution records were
obtained from herbarium specimens in ANSM, ENCB, HCIB, IBUG, IEB, MEXU,
MO, NY, TEX, UAMIZ, US and XAL, and by consulting the Mexican Biodiversity
Database (REMIB) (www.conabio.org).
Study area
The study area includes the entire country of Mexico. Even though the distribution of some groups extends into the south of the United States of America and
northern Central America in Mega-Mexico, only the localities within Mexico were
used. A system of land quadrats based on one degree squares was used to define
arbitrary area units, resulting in a set of 237 area units with records of endemic taxa.
The occurrence of every specimen of each monophyletic group in each quadrat was
recorded. The data matrix had a total of 9416 georeferenced records. Quadrats with
no records were eliminated. Species restricted to a single quadrat were identified as
microendemics.
Areas of endemism
First, the number of species was added up for each quadrat to estimate its
diversity (unweighted species richness, Pearson & Juliano, 1993; Kershaw et al.,
1995). Then, the weighted endemism index, a method that weights species inversely
to their distribution areas was also calculated (Linder, 2001).
Microendemic species
The species with a restricted distribution, i.e., those only found in a single
quadrat, were recorded and of these the taxa on the Mexican Red List (Anonymous,
2010) were identified.
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Acta Botanica Mexicana 100: 293-315 (2012)
RESULTS
Areas of endemism
The data matrix included the presence/absence data for 878 species belonging to 259 monophyletic groups for 237 area units. The highest unweighted species
richness values for each quadrat are shown in Table 1 and Fig. 1. The areas with the
highest number of endemic species are in Tehuacán-Cuicatlán, in the eastern of the
Balsas River Basin, in Tolantongo and Tepeapulco, Hidalgo and in the Sierra Gorda.
The weighted endemism values are listed in Table 1 and shown in Fig. 2.
Eleven areas with the highest weighted endemism values (10.657-34.819) were identified: 1) A northeastern area of rossette scrub in Nuevo León and Coahuila (Ramos
Arizpe, Aramberri, Galeana and Zaragoza); 2) an area of gypsum grasslands in San
Luis Potosí, 3) the Sierra Gorda, Querétaro (extending to San Luis Potosí); 4) Tolantongo in Hidalgo, 4) the area of Tehuacán-Cuicatlán, Puebla and Oaxaca; 5) El Salto,
Durango; 6) the Sierra de Quila in Jalisco; 7) the western area of the Balsas River
Basin (Michoacán, Guerrero, Morelos, State of Mexico); 8) the Tehuantepec area,
Unweighted endemic
species richness
1-7
8 - 17
18 - 33
34 - 57
500
58 - 108
0
500
1000
km
Fig. 1. Distribution map of the Mexican lineages of angiosperms (unweighted richness).
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Sosa y De-Nova: Endemic angiosperm lineages in Mexico
Table 1. Areas of endemism of the Mexican lineages of angiosperms with the highest species
richness (number of species 23-108) and the highest indices of weighted endemism (6.6134.81). Values for each land quadrat for every area of endemism are included.
Areas of endemism
Tehuacán-Cuicatlán
Balsas River Basin
Northeastern rosette scrub
Sierra Gorda
Northeastern rosette scrub
Tolantongo
Balsas River Basin
Tehuacán-Cuicatlán
Balsas River Basin
Sierra de Quila
Central Depression of Chiapas
Tehuantepec Region
Northeastern rosette scrub
El Triunfo
Northeastern rosette scrub
Balsas River Basin
El Salto
Gypsum grasslands
Sierra de Órganos
Baja California Sur
Unweighted endemic
species richness
108
94
64
66
61
71
64
53
55
74
67
47
39
57
39
50
39
25
44
52
38
40
33
31
24
23
23
Weighted
endemism
34.81888723
30.53009253
20.95559334
19.20541089
18.9554685
18.76557059
16.97099585
16.89719931
16.79210378
16.68154352
16.08813242
15.86234919
15.67449119
14.40864616
15.67449119
14.27457894
13.88012541
13.71388889
13.52460031
12.92819513
11.6864493
11.59780087
8.122629758
8.805300868
7.578488054
6.938598987
6.618010751
Oaxaca; 9) the Central Depression of Chiapas; 10) El Triunfo, Chiapas. Among the
areas with high weighted endemism indices is the southern area of Baja California
and the Sierra de Órganos, Zacatecas (Fig. 2).
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Acta Botanica Mexicana 100: 293-315 (2012)
N
Northeastern rosette scrub
Baja California Sur
Gypsum grasslands
Sierra Gorda
Tolantongo
Weighted endemism
6.11 - 10.667
10.667 - 34.819
Elevation
0 - 544
545 - 1163
1164 - 1783
1784 - 2402
2403 - 3022
3023 - 3641
3642 - 4261
4262 - 4880
4881 - 5500
El Salto
Central Depression of Chiapas
Sierra de Órganos
Sierra de Quila
Balsas River Basin
Tehuacán-Cuicatlán
Tehuantepec
El Triunfo
500
0
500
1000
km
Fig. 2. Areas of endemism for the Mexican lineages of angiosperms (weighted endemism).
Microendemics
Appendix lists the 340 species whose distribution is restricted to a single
quadrat, with their threatened status indicated when applicable.
DISCUSSION
Rzedowski (1993) pointed out that the distribution of areas with endemic species for the flora of Mexico does not coincide with the distribution of biodiversity.
He indicated that the endemic taxa are concentrated in areas of dry climate and this
conclusion was reached based on species richness alone, without taking the historical element into account. The latter has been included in this paper by identifying
areas of endemism shared by at least two monophyletic groups. Our results indicate
that majority of the areas: the northeastern rosette scrub, the gypsum grasslands,
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Sosa y De-Nova: Endemic angiosperm lineages in Mexico
the Sierra Gorda, the southern portion of the Chihuahuan Desert, the area of Tehuacán-Cuicatlán, the Sierra de Quila, the western area of the Balsas River Basin,
the Tehuantepec area and the Central Depression of Chiapas, have a dry climate,
corroborating Rzedowski’s hypothesis. The endemic groups occur at low to middle
elevations, in xeric vegetation. The only area with a tropical climate and a high
weighted endemism index is El Triunfo in Chiapas, and El Salto in Durango has a
temperate climate.
The Sierra Gorda was previously recognized as an area with significant endemism and it is included in the Mexican System of Natural Protected Areas, the
SINAP (Arriaga-Cabrera et al., 2000). Additionally, the Tehuacán-Cuicatlán area is
comprised of arid vegetation and is perhaps the most important biosphere reserve
in Mexico (Arriaga-Cabrera et al., 2000). It is a floristic province, and an ecological
island given the high number of endemics, estimated at 365 species (Méndez-Larios
et al., 2004; Dávila et al., 2002).
The area of Metztitlán-Tolantongo was previously known for having high
endemism and it was decreed as a biological reserve in 2000 (Hiriart-Valencia &
González-Medrano, 1983; Arriaga-Cabrera et al., 2000). Furthermore, Sierra de
Quila was earlier identified as a hotspot for conservation based on mammal distributional predictions as biodiversity surrogates (Sánchez-Cordero et al., 2005). In
addition, the semiarid gypsum karstlands in north central Mexico characterized by a
mosaic of shrubby communities and endemic gypsophile grasslands were formerly
acknowledged as areas with elevated endemism and important to preserve (Henrickson & Johnston, 1986; Meyer et al., 1992; Huerta-Martínez & García-Moya, 2004).
The western area of the Balsas River Basin in Michoacán, Guerrero, Morelos
and State of Mexico, is another region with high indices of endemism. RodríguezJiménez et al. (2005) have identified 337 endemic species of vascular plants in this
biogeographic province. Cañón del Zopilote and Infiernillo are two proposed areas
for conservation in this province (Arriaga-Cabrera et al., 2000), yet they represent
only a small area within the Balsas River Basin.
Several regions in Nuevo León harbor extremely large numbers of endemisms
in the Cactaceae (Juárez et al., 2009), and a high concentration of narrowly distributed Asteraceae (González-Zamora et al., 2007; Alanís-Flores et al., 2011). Moreover,
these regions coincided with one of the areas of high endemism identified in this
study: the Northeastern rosette scrub.
Our results detected the Sierra La Laguna in Baja California Sur as a territory with high endemism. Plant diversity and endemism on the entire Baja California Peninsula have previously attracted attention (Riemann & Ezcurra, 2007).
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Acta Botanica Mexicana 100: 293-315 (2012)
The notable endemism in the Sierra La Laguna, Baja California, had also been
pointed out (León de la Luz & Breceda, 2006) and resulted in the Sierra La Laguna reserve being decreed one of Mexico’s biosphere reserves (Arriaga-Cabrera
et al., 2000).
El Triunfo is a biosphere reserve with elevated endemism and richness, for
which approximately 1000 species of vascular plants have been reported. The area
includes several habitats such as cloud, oak and tropical forest (Martínez-Meléndez
et al., 2008; Pérez-Farrera et al., 2012). It was decreed as a reserve because it is considered to be a Pleistocene refugium for several tropical species (Arriaga-Cabrerra
et al., 2000). As well, the pine and oak forests in El Salto have provided suitable
habitats for several gymnosperm relict species (Valenzuela-Núñez & GranadosSánchez, 2009).
The Central Depression of Chiapas has been earlier identified as an area with
seasonally dry tropical forests and tropical oak forests where approximately 3.4% of
the total number of vascular plant species in Mexico are distributed (Reyes-García
& Sousa, 1997). Furthermore, among the habitats of Tehuantepec, the tropical dry
forests harbor the largest diversity including several taxa of endemic angiosperm
(Acosta et al., 2003; Pérez-García et al., 2010).
It is noteworthy that the majority of the areas with the highest endemism indices, such as El Salto (Durango), the Central Depression of Chiapas, Tehuantepec,
(Oaxaca), and Tolantongo in the southern area of the Chihuahuan Desert are not
protected under the SINAP scheme (Arriaga-Cabrera et al., 2000).
Microendemic species
It is crucial to take the rarity of species into account when setting conservation priorities (Mooers & Redding, 2009). It has been mentioned that in areas of
endemism the species with restricted distributions are usually on the red lists (e.g.,
Argentina, Szumik et al., in press). The same happens in Mexico where we found
that a large number of the species whose distribution is restricted to a single quadrat
in our study area are included on the Mexican List of Threatened Species (Anonymous, 2010).
Most of the species on this list with a limited distribution are cacti. More than
900 species of Cactaceae are present throughout Mexico (Ortega-Baes & GodínezÁlvarez, 2006). This is one of the groups that are most used as ornamental plants
and so have been continuously extracted from their habitats, with the result that they
are now the most threatened group in Mexico (Gómez-Hinostrosa & Hernández,
2000; Hernández & Gómez-Hinostrosa, 2011a,b).
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CONCLUSIONS
Future research should examine the probable causes of diversification for the
angiosperm lineages in the areas of endemism in Mega-Mexico. For other areas with
high degrees of endemism, such as the Andes, isolation caused diversification, similar in many respects to the floras of remote oceanic islands (Särkinem et al., in press)
or along elevational gradients (Kessler, 2000). Climate was the factor that promoted
speciation in Australia’s areas of endemism (Ladiges et al., 2011), while tectonic stability in central and southern China influenced the permanence of areas of plant
endemism (López-Pujol et al., 2011). In Sub-Saharan Africa, elevation range and low
seasonality were core environmental predictors for centers of endemism (Jetz et al.,
2004). Diversification in hotspots of biodiversity and endemism in Brazil were attributed to the effect of fire on vegetation (Simon et al., 2009), while serpentine soils
and a benign climate favored endemism in California (Anacker & Harrison, in press).
Our results suggest that various causes promoted the diversification of several
groups of plants in the areas of endemism, and a dry climate together with isolation are probably the most remarkable. Nine areas of endemism have a dry climate.
Furthermore, the Central Depression of Chiapas and the Balsas River Basin are two
areas that remained isolated, bordered by mountain ranges. In contrast, El Triunfo in
Chiapas probably acted as a refugium for angiosperm lineages that remained there
throughout the Pleistocene. Gypsum soils probably favored endemism in the northeastern rosette scrub and the grasslands of San Luis Potosí. However investigation is
needed to corroborate these hypotheses.
It should be emphasized that hotspots do not necessarily coincide with species
richness, the degree of threat or areas of endemism (Orme et al., 2005). The areas
of endemism identified in our study do not coincide with the areas with elevated
diversity of the flora of Mexico, as Rzedowski (1993) pointed out, and some of the
areas of endemism in the Chihuahuan Desert, Balsas River Basin and the southern
area of Oaxaca are not sufficiently protected.
ACKNOWLEDGEMENTS
We are grateful to Patricia Dávila and Jorge Meave del Castillo for their useful comments that improved the manuscript. We are grateful to Manuel Cuéllar and
Ismael G. Valdivieso for their help designing the database, and to Rosario Landgrave for her invaluable help producing the maps, as well as to Manuel Cuéllar
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Acta Botanica Mexicana 100: 293-315 (2012)
and Diego Angulo. Jerzy Rzedowski provided important information on the endemism of the Mexican flora; Carolina Calviño provided information on Apiaceae
(Eryngium), Etelvina Gándara on Asparagaceae (Milla clade) and Scrophulariaceae
(Leucophyllum); John Bain, Jose Panero, Edward Schilling and José Luis Villaseñor
on Asteraceae; Salvador Arias and Héctor Hernández on Cactaceae; Mark Simmonds on Celastraceae; Pablo Carrillo Reyes and Raúl Acevedo-Rosas on Crassulaceae; Rafael Lira on Cucurbitaceae; Eduardo Estrada, Lourdes Rico and Jenny
Sotuyo on Fabaceae; Susana Valencia on Fagaceae; Paul Peterson, María Elena
Siqueiros and J. Travis Columbus on Poaceae; Rafael Fernández Nava on Rhamnaceae and Lynn Boss and Aarón Rodríguez on Solanaceae.
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Recibido en abril de 2012.
Aceptado en junio de 2012.
309
Acta Botanica Mexicana 100: 293-315 (2012)
APPENDIX
Microendemic species of the Mexican lineages of angiosperms. These are
restricted to a single quadrat. Species that are on the Mexican Red List (Anonymous,
2010) are indicated with the following designations in parentheses: A = threatened,
P = endangered, Pr = under special protection.
Acanthaceae
Gypsacanthus nelsonii E.J. Lott, V. Jaram. &
Rzed.
Holographis anisophylla T.F. Daniel
Holographis argyrea (Leonard) T.F. Daniel (Pr)
Holographis caput-medusae T.F. Daniel
Holographis hintonii (Leonard) T.F. Daniel
Holographis pallida Leonard & Gentry
Holographis tamaulipica T.F. Daniel
Holographis tolantongensis T.F. Daniel
Ixtlania acicularis M.E. Jones
Mexacanthus mcvaughii T.F. Daniel
Mirandea andradenia T.F. Daniel
Mirandea huastecensis T.F. Daniel
Mirandea hyssopus (Nees) T.F. Daniel
Ruellia conzattii Standl.
Ruellia guerrerensis T.F. Daniel
Ruellia laslobasensis E.A. Tripp
Ruellia sarukhaniana Ramamoorthy
Ruellia sororia Standl.
Achariaceae
Chiangiodendron mexicanum T. Wendt
Achatocarpaceae
Phaulothamnus spinescens A. Gray
Amaryllidaceae
Sprekelia clintiae Traub
Anacardiaceae
Pseudosmodingium andrieuxii Engl.
Apiaceae
Eryngium humile Cav.
Eryngium mexicanum S. Watson
Apocynaceae
Thenardia gonoloboides Woodson
310
Asparagaceae
Beaucarnea purpusii Rose
Beschorneria tubilora Kunth (Pr)
Dasylirion inerme S. Watson
Hemiphylacus mahindae L. Hern.
Hemiphylacus novogalicianus L. Hern.
Jaimehintonia gypsophila B.L. Turner
Milla magniica H.E. Moore
Milla rosea H.E. Moore
Nolina humilis S. Watson
Nolina lindheimeriana S. Watson
Nolina pliabilis (Baker) Lundell
Nolina pumila Rose
Yucca baccata Torr.
Yucca capensis L.W. Lenz
Asteraceae
Ageratum albidum (DC.) Hemsl.
Ageratum conyzoides L.
Ageratum maritimum Kunth
Ageratum microcephalum Hemsl.
Ageratum munaense R.M. King & H. Rob.
Ageratum paleaceum (Gay ex DC.) Hemsl.
Ageratum tomentosum (Benth.) Hemsl.
Alomia hintonii R.M. King & H. Rob.
Alvordia angusta S.F. Blake
Amauria carterae A.M. Powell
Arnicastrum glandulosum Greenm.
Axiniphyllum pinnatisectum (Paul G. Wilson)
B.L. Turner
Axiniphyllum sagittalobum B.L. Turner
Axiniphyllum tomentosum Benth.
Baeriopsis guadalupensis J.T. Howell
Bahiopsis carterae (E.E. Schill.) E.E. Schill. &
Panero
Bahiopsis chenopodina (Greene) E.E. Schill. &
Panero
Bahiopsis laciniata (A. Gray) E.E. Schill. &
Panero
Sosa y De-Nova: Endemic angiosperm lineages in Mexico
Appendix. Continuation.
Bahiopsis lanata Kellogg
Bahiopsis tomentosa (A. Gray) E.E. Schill. &
Panero
Brickellia adenolepis (B.L. Rob.) Shinners
Brickellia adontophylla A. Gray
Brickellia amblyoleopsis (B.L. Rob.) R.M. King
& H. Rob.
Brickellia aramberrana B.L. Turner
Brickellia cardiophylla B.L. Rob.
Brickellia coahuilensis (A. Gray) Harc. & Beaman
Brickellia loribunda A. Gray
Brickellia frutescens A. Gray
Brickellia glabrata (Rose) B.L. Rob.
Brickellia glutinosa A. Gray
Brickellia hastata Benth.
Brickellia hebercarpa (DC.) A. Gray
Brickellia kellermanii Greenm.
Brickellia pedunculosa (DC.) Harc. & Beaman
Brickellia peninsularis Brandegee
Brickellia rusbyi A. Gray
Brickellia simplex A. Gray
Brickellia urolepis S.F. Blake
Brickellia vernicosa B.L. Rob.
Brickellia wislizeni A. Gray
Calanticaria brevifolia (Greenm.) E.E. Schill. &
Panero
Conoclinium mayieldii T.F. Patterson
Correllia montana A.M. Powell
Eryngiophyllum pinnatisectum Paul G. Wilson
Eryngiophyllum rosei Greenm.
Eupatoriastrum triangulare (DC.) B.L. Rob.
Faxonia pusilla Brandegee
Gonzalezia hypargyrea (Greenm.) E.E. Schill. &
Panero
Gonzalezia rosei (Greenm.) E.E. Schill. & Panero
Gymnolaena serratifolia Rydb.
Gymnolomia scaposa Brandegee
Henricksonia mexicana B.L. Turner
Hofmeisteria gayleana B.L. Turner
Hybridella anthemidifolia (B.L. Rob. & Greenm.)
Olsen
Hydropectis aquatica Rydb.
Jaliscoa goldmanii (B.L. Rob.) R.M. King & H.
Rob.
Jaliscoa paleacea (Cronquist) R.M. King & H.
Rob.
Jaliscoa pappifera S.F. Blake
Jefea gnaphalioides (A. Gray) Strother
Jefea pringlei (Greenm.) Strother
Lundellianthus breedlovei (B.L. Turner) Strother
Lundellianthus guatemalensis (Donn. Sm.)
Strother
Lundellianthus salvinii (Hemsl.) Strother
Marshalljohnstonia gypsophila Henrickson
Mexerion sarmentosum (Klatt) G.L. Nesom
Nesomia chiapensis B.L. Turner
Otopappus acuminatus S. Watson
Otopappus pittieri (Greenm.) B.L. Turner
Paneroa stachyofolia (B.L. Rob.) E.E. Schill.
Perymenium ovalifolium (A. Gray) B.L. Turner
Philactis zinnioides Schrad.
Pittocaulon bombycophole (Bullock) H. Rob. &
Brettell
Pleurocoronis gentryi (Wiggins) R.M. King &
H. Rob.
Pleurocoronis pluriseta (A. Gray) R.M. King &
H. Rob.
Psacaliopsis purpusii (Greenm. ex Brandegee) H.
Rob. & Brettell
Psacalium brachycomum (S.F. Blake) H. Rob. &
Brettell
Psacalium calvum (Brandegee) Pippen
Psacalium decompositum (A. Gray) H. Rob. &
Brettell
Psacalium globosum (B.L. Rob. & Fernald) H.
Rob. & Brettell
Psacalium hintonii (Pippen) H. Rob. & Brettell
Psacalium hintoniorum B.L. Turner
Psacalium pachyphyllum (Sch. Bip.) Rydb.
Psacalium paucicapitatum (B.L. Rob. &
Greenm.) H. Rob. & Brettell
Psacalium peltigerum (B.L. Rob. & Seaton) Rydb.
Psacalium radulifolium (Kunth) H. Rob. &
Brettell
Psacalium tussilaginoides (Kunth) H. Rob. &
Brettell
Robinsonecio porphyresthes (T.M. Barkley) T.M.
Barkley & Janovec
Squamopappus skutchii (S.F. Blake) R.K. Jansen,
N.A. Harriman & Urbatsch
Sidneya tenuifolia (A. Gray) E.E. Schill. & Panero
Stenocarpha ritovegana B.L. Turner
311
Acta Botanica Mexicana 100: 293-315 (2012)
Appendix. Continuation.
Stephanodoria tomentella Greene
Stevia chilapensis Soejima & Yahara
Stevia coahuilensis Soejima & Yahara
Stevia crassifolia Soejima & Yahara
Stevia ecatepecana Soejima, Yahara & K. Watan.
Stevia ilodecaballoana Soejima, Yahara & K.
Watan.
Stevia mascotensis Soejima & Yahara
Stevia mexicana Soejima, Yahara & K. Watan.
Stevia oaxacana Soejima & Yahara
Stevia oligophylla Soejima & Yahara
Stevia potosina Soejima, Yahara & K. Watan.
Stevia rotundifolia Soejima, Yahara & K. Watan.
Stevia scabrelloides Soejima & Yahara
Stevia viejoana Soejima, Yahara & K. Watan.
Steviopsis adenosperma (Sch. Bip.) B.L. Turner
Steviopsis amblyolepis (B.L. Rob.) R.M. King &
H. Rob.
Steviopsis nesomii B.L. Turner
Steviopsis squamulosa (A. Gray) B.L. Turner
Steviopsis vigintiseta (DC.) R.M. King & H. Rob.
Stuessya apiculata (S.F. Blake) B.L. Turner &
F.G. Davies
Stuessya perennans B.L. Turner & F.G. Davies
Tetrachyron chimalapanum B.L. Turner
Tetrachyron grayi (Klatt) Wussow & Urbatsch
Tetrachyron orizabensis (Klatt) Wussow &
Urbatsch
Tuxtla pittieri (Greenm.) Villaseñor & Strother
Wamalchitamia appressipila (S.F. Blake) Strother
Wamalchitamia aurantiaca (Klatt) Strother
Wamalchitamia dionysi Strother
Zexmenia virgulta Klatt
Boraginaceae
Lasiarrhenum confundum B.L. Turner
Lashiarrhenum pinetorum I.M. Johnst.
Mimophytum omphalodoides Greenm.
Brassicaceae
Lexarzanthe mexicana (Iltis & Al-Shehbaz)
Diego & Calderón
Raphanorhyncha crassa Rollins
Burseraceae
Beiselia mexicana Forman
312
Bursera rzedowskii C.A. Toledo
Cactaceae
Astrophytum asterias Lem. (P)
Aztekium hintonii (Glass & W.A. Fitz Maur.) (Pr)
Aztekium ritteri Boed. (A)
Cumarinia odorata (Boed.) Buxb. (Pr)
Geohintonia mexicana Glass & W.A. Fitz Maur.
(Pr)
Leuchtenbergia principis Hook. (A)
Neobuxbaumia multiareolata (Daws.) Bravo,
Scheinvar & Sánchez-Mej.
Obregonia denegrii Frič & A. Berger (A)
Pachycereus tepamo S. Gama-López & S. Arias
Pelecyphora aselliformis Ehrenb. (Pr)
Pelecyphora strobiliformis (Werderm.) Frič &
Schelle ex Kreuz. (A)
Thelocactus hastifer (Werderm. & Boed.) F.M.
Knuth (A)
Turbinicarpus alonsoi Glass & S. Arias
Turbinicarpus hoferi Lüthy & A.B. Lau (A)
Turbinicarpus lophophoroides (Werderm.) Buxb.
& Backeb. (Pr)
Turbinicarpus pseudopectinatus (Backeb.) Glass
& R.A. Foster (Pr)
Caryophyllaceae
Cerdia virescens Moc. & Sessé
Convolvulaceae
Ipomoea decemcornuta O’Donell
Crassulaceae
Cremnophila linguifolia (Lem.) Moran
Cremnophila nutans (Rose) Rose
Graptopetalum amethystinum E. Walther
Graptopetalum bartramii Rose
Pachyphytum amethystinum Rose
Pachyphytum brachetii J. Reyes, O. González &
A. Gut.
Pachyphytum brevifolium Rose
Pachyphytum caesium Kimnach & Moran
Pachyphytum coeruleum J. Meyrán
Pachyphytum contrerasii Pérez-Calix, I. García
& Cházaro
Pachyphytum ittkaui Moran
Sosa y De-Nova: Endemic angiosperm lineages in Mexico
Appendix. Continuation.
Pachyphytum garciae Pérez-Calix & Glass
Pachyphytum hookeri A. Berger
Pachyphytum kimnachii Moran
Pachyphytum longifolium Rose
Pachyphytum machucae I. García, Glass &
Cházaro
Pachyphytum oviferum J.A. Purpus
Pachyphytum rzedowskii I. García, Pérez-Calix
& J. Meyrán
Pachyphytum saltense Brachet, J. Reyes &
Mondragón
Pachyphytum werdermannii Poelln.
Thompsonella garcia-mendozae P. Carrillo &
Pérez-Calix
Thompsonella mixtecana J. Reyes & L. López
Thompsonella spathulata Kimnach
Thompsonella xochipalensis M. Gual Diaz, S.
Peralta & Pérez-Calix
Crossosomataceae
Velascoa recondita Calderón & Rzed.
Cucurbitaceae
Cucurbita fraterna L.H. Bailey
Cucurbita pedatifolia L.H. Bailey
Apatzingania arachoidea I.M. Johnston
Vaseyanthus brandegeei Rose
Acaciella igualensis Britton & Rose
Acaciella sotoi L. Rico
Calliandropsis nervosus (Britton & Rose) H.M.
Hern. & P. Guinet
Dalea laniceps Barneby
Dalea parrasana Brandegee
Hesperothamnus ehrenbergii (Harms) Harms
Hesperothamnus littoralis (Brandegee) Brandegee
Hesperothamnus purpusii (Harms) Harms
Marina brevis León de la Luz
Marina capensis Barneby
Marina catalinae Barneby
Marina divaricata (Benth.) Barneby
Marina interstes Barneby
Marina oculata (Rydb.) Barneby
Mariosousa acatlensis (Benth.) Seigler & Ebinger
Fagaceae
Quercus clivicola Trel. & C.H. Mull.
Quercus radiata Trel.
Quercus tarahumara Spellenb., J.D. Bacon &
Breedlove
Quercus verde C.H. Mull.
Fouquieriaceae
Fouquieria leonilae Miranda (Pr)
Fouquieria purpusii Brandegee (P)
Gentianaceae
Cyperaceae
Cypringlea evadens (C.D. Adams) Reznicek & S. Geniostemon atarjanus B.L. Turner
Geniostemon rotundifolius Rzed. & Calderón
González
Euphorbiaceae
Euphorbia coalcomanensis (Croizat) V.W.
Steinm. (A)
Euphorbia cyri V.W. Steinm. (E)
Euphorbia dressleri V.W. Steinm. (E)
Euphorbia inkii (Boiss.) V.W. Steinm. (A)
Euphorbia peritropoides (Millsp.) V.W. Steinm.
Euphorbia personata (Croizat) V.W. Steinm.
Euphorbia tehuacana (Brandegee) V.W. Steinm. (A)
Euphorbia tithymaloides L.
Gesneriaceae
Achimenes candida Lindl.
Achimenes hintoniana Ramírez Roa & L.E. Skog
Achimenes nayaritensis L.E. Skog
Achimenes occidentalis C.V. Morton
Achimenes pedunculata Benth.
Smithiantha aurantiaca Wiehler
Fabaceae
Acaciella barrancana (Gentry) L. Rico
Acaciella goldmanii Britton & Rose
Iteaceae
Pterostemon bravoanus J. Jiménez Ram. & M.
Martínez
Iridaceae
Ainea conzattii (R.C. Foster) Ravenna (A)
313
Acta Botanica Mexicana 100: 293-315 (2012)
Appendix. Continuation.
Lamiaceae
Salvia canescens C.A. Mey.
Salvia dolichantha E. Peter
Salvia univerticillata Ramamoorthy ex Klitg.
Lauraceae
Mocinnodaphne cinnamomoidea Lorea-Hern.
Liliaceae
Hesperaloe tenuifolia G.D. Starr
Loasaceae
Schismocarpus matudae Steyerm.
Schismocarpus pachypus S.F. Blake
Malphighiaceae
Lasiocarpus multilorus Nied.
Lasiocarpus ovalifolius Nied.
Malvaceae
Bastardiastrum tarasoides Fryxell
Bastardiastrum tricarpellatum (B.L. & Rob. &
Greenm.) D.M. Bates
Periptera lobelioides Fryxell & S.D. Koch
Periptera trichostemon Bullock
Nyctaginaceae
Grajalesia fasciculata (Standl.) Miranda
Grajalesia ferruginea Miranda
Oleaceae
Hesperelaea palmeri A. Gray (P)
Onagraceae
Lopezia clavata Brandegee
Lopezia lopezioides (Hook. & Arn.) Plitmann,
P.H. Raven & Breedlove
Lopezia ovata (Plitmann, P.H. Raven &
Breedlove) Plitmann, P.H. Raven & Breedlove
Lopezia smithii Rose
Lopezia suffrutescens Munz
Megacorax gracielanus S. González & W.L. Wagner
Orchidaceae
Hagsatera rosilloi R. González
314
Mexipedium xerophyticum (Soto Arenas, Salazar
& Hágsater) V.A. Albert & M.W. Chase (P)
Nezahualcoyotlia gracilis (L.O. Williams) R.
González
Physogyne garayana R. González & Szlach.
Physogyne sparsilora (C. Schweinf.) Garay
Svenkoeltzia luzmariana R. González
Svenkoeltzia pamelae Szlach., Rutk. & Mytnik
Orobanchaceae
Castilleja ililora G.L. Nesom
Castilleja hidalgensis J.M. Egger
Castilleja macrostigma B.L. Rob.
Castilleja ornata Eastw.
Castilleja perelegans G.L. Nesom
Castilleja sphaerostigma Eastw.
Castilleja stipifolia G.L. Nesom
Castilleja tancitaroana G.L. Nesom
Eremitilla mexicana Yatsk. & J.L. Contr.
Lamourouxia brachyantha Greenm.
Lamourouxia macrantha M. Martens &
Galeotti
Lamourouxia nelsonii B.L. Rob. & Greenm.
Phytolaccaceae
Nowickea glabra J. Martínez & J.A. McDonald
Nowickea xolocotzii J. Martínez & J.A.
McDonald
Poaceae
Muhlenbergia brevis C.O. Goodd.
Muhlenbergia majalcensis P.M. Peterson
Olmeca clarkiae (Davidse & R.W. Pohl) RuizSanchez, Sosa & Mejía-Saules
Olmeca zapotecorum Ruiz-Sanchez, E., Sosa &
Mejía Saules
Otatea glauca L.G. Clark & G. Cortés
Otatea ramirezii Ruiz-Sanchez
Otatea transvolcanica Ruiz-Sanchez & L.G.
Clark
Otatea ximenae Ruiz-Sanchez & L.G. Clark
Rhamnaceae
Karwinskia calderonii Urb.
Karwinskia johnstonii R. Fernandez
Sosa y De-Nova: Endemic angiosperm lineages in Mexico
Appendix. Continuation.
Rubiaceae
Carterella alexanderae (A.M. Carter) Terrell
Habroneuron radicans (Wernham) S.P. Darwin
Omiltemia parvifolia Borhidi & K.Velasco
Placocarpa mexicana Hook. f.
Stenotis gracilenta (I.M. Johnst.) Terrell
Stenotis peninsularis (Brandegee) Terrell
Stylosiphonia glabra Brandegee
Leucophyllum hintoniorum G.L. Nesom
Leucophyllum langmaniae Flyr
Leucophyllum lanosum Flyr
Leucophyllum mojinense Henrickson & T. Van
Devender
Leucophyllum ultramonticola Flyr
Leucophyllum virescens I.M. Johnst.
Sapindaceae
Balsas guerrerensis Cruz Durán & K. Vega
Solanaceae
Physalis heterophylla Nees
Physalis virginiana Mill.
Physalis walteri Nutt.
Solanum johnstonii Whalen
Solanum morelliforme Bitter & Münch
Solanum tribulosum S. Schauer
Tzeltalia amphitricha (Bitter) E. Estrada & M.
Martínez
Tzeltalia calidaria (Standl. & Steyerm.) E.
Estrada & M. Martínez
Scrophulariaceae
Leucophyllum alejandrae G.L. Nesom
Leucophyllum lyrii B.L. Turner
Zygophyllaceae
Morkillia acuminata Rose & Painter
Viscainoa pinnata Gentry
Rutaceae
Ptelea baldwinii Torr. & A. Gray
Ptelea coninis Greene
Ptelea megacarpa Rose ex Greene
Ptelea obscura Greene
Ptelea obtusata Greene
Ptelea subintegra Greene
315