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Endemic angiosperm lineages in Mexico: hotspots for conservation.

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 (Davila 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 (Villasenor, 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 (Villasenor, 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; Guzman et al., 2003; Hernandez & Gomez-Hinostrosa, 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 (Gandara 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 (Garcia-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) (Davila-Aranda et al., 2004; Ruiz-Sanchez et al., 2011), Peltophorum (Leguminosae) (Sousa, 2005), the parasitic Eremitilla (Orobanchaceae) (Yatskievych & Contreras-Jimenez, 2009), Echinopterys (Malpighiaceae) (Davis et al., 2001), Nowickea (Phytolaccaceae) (Martinez & 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.

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; Villasenor, 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.

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 Tehuacan-Cuicatlan, 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 Leon and Coahuila (Ramos Arizpe, Aramberri, Galeana and Zaragoza); 2) an area of gypsum grasslands in San Luis Potosi, 3) the Sierra Gorda, Queretaro (extending to San Luis Potosi); 4) Tolantongo in Hidalgo, 4) the area of Tehuacan-Cuicatlan, Puebla and Oaxaca; 5) El Salto, Durango; 6) the Sierra de Quila in Jalisco; 7) the western area of the Balsas River Basin (Michoacan, Guerrero, Morelos, State of Mexico); 8) the Tehuantepec area, 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 Organos, Zacatecas (Fig. 2).

[FIGURE 1 OMITTED]

[FIGURE 2 OMITTED]

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, the Sierra Gorda, the southern portion of the Chihuahuan Desert, the area of Tehuacan-Cuicatlan, 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 Tehuacan-Cuicatlan 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 (Mendez-Larios et al., 2004; Davila et al., 2002).

The area of Metztitlan-Tolantongo was previously known for having high endemism and it was decreed as a biological reserve in 2000 (Hiriart-Valencia & Gonzalez-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 (Sanchez-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-Martinez & Garcia-Moya, 2004).

The western area of the Balsas River Basin in Michoacan, Guerrero, Morelos and State of Mexico, is another region with high indices of endemism. Rodriguez-Jimenez et al. (2005) have identified 337 endemic species of vascular plants in this biogeographic province. Canon 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 Leon harbor extremely large numbers of endemisms in the Cactaceae (Juarez et al., 2009), and a high concentration of narrowly distributed Asteraceae (Gonzalez-Zamora et al., 2007; Alanis-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). The notable endemism in the Sierra La Laguna, Baja California, had also been pointed out (Leon 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 (Martinez-Melendez et al., 2008; Perez-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-Nunez & Granados Sanchez, 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-Garcia & 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; Perez-Garcia 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 & Godinez-Alvarez, 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 (Gomez-Hinostrosa & Hernandez, 2000; Hernandez & Gomez-Hinostrosa, 2011a,b).

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 (Sarkinem 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 (Lopez-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 Potosi. 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.

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

Asparagaceae

Beaucarnea purpusii Rose

Beschorneria tubiflora Kunth (Pr)

Dasylirion inerme S. Watson

Hemiphylacus mahindae L. Hern.

Hemiphylacus novogalicianus L. Hern.

Jaimehintonia gypsophila B.L. Turner

Milla magnifica 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.

Ageratumpaleaceum (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

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 floribunda 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.

Brickelliapedunculosa (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 mayfieldii 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.

Jaliscoapappifera 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

Otopappuspittieri (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.

Psacaliopsispurpusii (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.

Psacaliumpaucicapitatum (B.L. Rob. & Greenm.) H. Rob. & Brettell

Psacaliumpeltigerum (B.L. Rob. & Seaton) Rydb.

Psacalium radulifolium (Kunth) H. Rob. & Brettell

Psacalium tussilaginoides (Kunth) H. Rob. & Brettell

Robinsonecioporphyresthes (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

Stephanodoria tomentella Greene

Stevia chilapensis Soejima & Yahara

Stevia coahuilensis Soejima & Yahara

Stevia crassifolia Soejima & Yahara

Stevia ecatepecana Soejima, Yahara & K. Watan.

Steviafilodecaballoana 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

Stuessyaperennans B.L. Turner & F.G. Davies

Tetrachyron chimalapanum B.L. Turner

Tetrachyron grayi (Klatt) Wussow & Urbatsch

Tetrachyron orizabensis (Klatt) Wussow & Urbatsch

Tuxtla pittieri (Greenm.) Villasenor & 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 & Calderon

Raphanorhyncha crassa Rollins

Burseraceae

Beiselia mexicana Forman

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 & Sanchez-Mej.

Obregonia denegrii Fric & A. Berger (A)

Pachycereus tepamo S. Gama-Lopez & S. Arias

Pelecyphora aselliformis Ehrenb. (Pr)

Pelecyphora strobiliformis (Werderm.) Fric & Schelle ex Kreuz. (A)

Thelocactus hastifer (Werderm. & Boed.) F.M. Knuth (A)

Turbinicarpus alonsoi Glass & S. Arias

Turbinicarpus hoferi Luthy & A.B. Lau (A)

Turbinicarpus lophophoroides (Werderm.) Buxb. & Backeb. (Pr)

Turbinicarpus pseudopectinatus (Backeb.) Glass & R.A. Foster (Pr)

Caryophyllaceae

Cerdia virescens Moc. & Sesse

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. Gonzalez & A. Gut.

Pachyphytum brevifolium Rose

Pachyphytum caesium Kimnach & Moran

Pachyphytum coeruleum J. Meyran

Pachyphytum contrerasii Perez-Calix, I. Garcia & Chazaro

Pachyphytum fittkaui Moran

Pachyphytum garciae Perez-Calix & Glass

Pachyphytum hookeri A. Berger

Pachyphytum kimnachii Moran

Pachyphytum longifolium Rose

Pachyphytum machucae I. Garcia, Glass & Chazaro

Pachyphytum oviferum J.A. Purpus

Pachyphytum rzedowskii I. Garcia, Perez-Calix & J. Meyran

Pachyphytum saltense Brachet, J. Reyes & Mondragon

Pachyphytum werdermannii Poelln.

Thompsonella garcia-mendozae P. Carrillo & Perez-Calix

Thompsonella mixtecana J. Reyes & L. Lopez

Thompsonella spathulata Kimnach

Thompsonella xochipalensis M. Gual Diaz, S. Peralta & Perez-Calix

Crossosomataceae

Velascoa recondita Calderon & Rzed.

Cucurbitaceae

Cucurbita fraterna L.H. Bailey

Cucurbitapedatifolia L.H. Bailey

Apatzingania arachoidea I.M. Johnston

Vaseyanthus brandegeei Rose

Cyperaceae

Cypringlea evadens (C.D. Adams) Reznicek & S. Gonzalez

Euphorbiaceae

Euphorbia coalcomanensis (Croizat) V.W. Steinm. (A)

Euphorbia cyri V.W. Steinm. (E)

Euphorbia dressleri V.W. Steinm. (E)

Euphorbia finkii (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.

Fabaceae

Acaciella barrancana (Gentry) L. Rico

Acaciella goldmanii Britton & 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 Leon 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

Geniostemon atarjanus B.L. Turner

Geniostemon rotundifolius Rzed. & Calderon

Gesneriaceae

Achimenes candida Lindl.

Achimenes hintoniana Ramirez Roa & L.E. Skog

Achimenes nayaritensis L.E. Skog

Achimenes occidentalis C.V. Morton

Achimenes pedunculata Benth.

Smithiantha aurantiaca Wiehler

Iridaceae

Ainea conzattii (R.C. Foster) Ravenna (A)

Iteaceae

Pterostemon bravoanus J. Jimenez Ram. & M. Martinez

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.

Schismocarpuspachypus S.F. Blake

Malphighiaceae

Lasiocarpus multiflorus 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. Gonzalez & W.L. Wagner

Orchidaceae

Hagsatera rosilloi R. Gonzalez

Mexipedium xerophyticum (Soto Arenas, Salazar & Hagsater) V.A. Albert & M.W . Chase (P)

Nezahualcoyotlia gracilis (L.O. Williams) R. Gonzalez

Physogyne garayana R. Gonzalez & Szlach.

Physogyne sparsiflora (C. Schweinf.) Garay

Svenkoeltzia luzmariana R. Gonzalez

Svenkoeltziapamelae Szlach., Rutk. & Mytnik

Orobanchaceae

Castilleja filiflora G.L. Nesom

Castilleja hidalgensis J.M. Egger

Castilleja macrostigma B.L. Rob.

Castilleja ornata Eastw.

Castillejaperelegans 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. Martinez & J.A. McDonald

Nowickea xolocotzii J. Martinez & J.A. McDonald

Poaceae

Muhlenbergia brevis C.O. Goodd.

Muhlenbergia majalcensis P.M. Peterson

Olmeca clarkiae (Davidse & R.W. Pohl) Ruiz Sanchez, Sosa & Mejia-Saules

Olmeca zapotecorum Ruiz-Sanchez, E., Sosa & Mejia Saules

Otatea glauca L.G. Clark & G. Cortes

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

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

Rutaceae

Ptelea baldwinii Torr. & A. Gray

Ptelea confinis Greene

Ptelea megacarpa Rose ex Greene

Ptelea obscura Greene

Ptelea obtusata Greene

Ptelea subintegra Greene

Sapindaceae

Balsas guerrerensis Cruz Duran & K. Vega

Scrophulariaceae

Leucophyllum alejandrae G.L. Nesom

Leucophyllum flyrii B.L. Turner

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.

Solanaceae

Physalis heterophylla Nees

Physalis virginiana Mill.

Physalis walteri Nutt.

Solanum johnstonii Whalen

Solanum morelliforme Bitter & Munch

Solanum tribulosum S. Schauer

Tzeltalia amphitricha (Bitter) E. Estrada & M. Martinez

Tzeltalia calidaria (Standl. & Steyerm.) E. Estrada & M. Martinez

Zygophyllaceae

Morkillia acuminata Rose & Painter

Viscainoa pinnata Gentry

ACKNOWLEDGEMENTS

We are grateful to Patricia Davila and Jorge Meave del Castillo for their useful comments that improved the manuscript. We are grateful to Manuel Cuellar 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 Cuellar and Diego Angulo. Jerzy Rzedowski provided important information on the endemism of the Mexican flora; Carolina Calvino provided information on Apiaceae (Eryngium), Etelvina Gandara on Asparagaceae (Milla clade) and Scrophulariaceae (Leucophyllum); John Bain, Jose Panero, Edward Schilling and Jose Luis Villasenor on Asteraceae; Salvador Arias and Hector Hernandez on Cactaceae; Mark Simmonds on Celastraceae; Pablo Carrillo Reyes and Raul Acevedo-Rosas on Crassulaceae; Rafael Lira on Cucurbitaceae; Eduardo Estrada, Lourdes Rico and Jenny Sotuyo on Fabaceae; Susana Valencia on Fagaceae; Paul Peterson, Maria Elena Siqueiros and J. Travis Columbus on Poaceae; Rafael Fernandez Nava on Rhamnaceae and Lynn Boss and Aaron Rodriguez on Solanaceae.

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Victoria Sosa (1,4) and J. Arturo De-Nova (2,3)

(1) Instituto de Ecologia A.C., Biologia Evolutiva, Apdo. postal 63, 91070 Xalapa, Veracruz, Mexico.

(2) Universidad Autonoma de San Luis Potosi, Instituto de Investigacion en Zonas Deserticas, Altair 200, 78377 San Luis Potosi, Mexico.

(3) Universidad Autonoma de San Luis Potosi, Facultad de Agronomia, km 14.5 carretera San Luis Potosi-Matehuala, 78321 San Luis Potosi, Mexico.

(4) Autor para la correspondencia: victoria.sosa@inecol.edu.mx
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 (6.61-34.81). Values for each land quadrat
for every area of endemism are included.

Areas of endemism               Unweighted endemic     Weighted
                                 species richness      endemism

Tehuacan-Cuicatlan                     108           34.81888723
                                        94           30.53009253
Balsas River Basin                      64           20.95559334
                                        66           19.20541089
Northeastern rosette scrub              61            18.9554685
Sierra Gorda                            71           18.76557059
                                        64           16.97099585
Northeastern rosette scrub              53           16.89719931
Tolantongo                              55           16.79210378
Balsas River Basin                      74           16.68154352
Tehuacan-Cuicatlan                      67           16.08813242
Balsas River Basin                      47           15.86234919
                                        39           15.67449119
Sierra de Quila                         57           14.40864616
Central Depression of Chiapas           39           15.67449119
Tehuantepec Region                      50           14.27457894
Northeastern rosette scrub              39           13.88012541
El Triunfo                              25           13.71388889
Northeastern rosette scrub              44           13.52460031
Balsas River Basin                      52           12.92819513
El Salto                                38            11.6864493
Gypsum grasslands                       40           11.59780087
Sierra de Organos                       33           8.122629758
Baja California Sur                     31           8.805300868
                                        24           7.578488054
                                        23           6.938598987
                                        23           6.618010751
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Author:Sosa, Victoria; De-Nova, J. Arturo
Publication:Acta Botanica Mexicana
Date:Jul 1, 2012
Words:7608
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