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 293 Acta Botanica Mexicana 100: 293-315 (2012) 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. 294 Sosa y De-Nova: Endemic angiosperm lineages in Mexico 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 295 Acta Botanica Mexicana 100: 293-315 (2012) (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. 296 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. 297 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). 298 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). 299 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, 300 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). 301 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). 302 Sosa y De-Nova: Endemic angiosperm lineages in Mexico 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 303 Acta Botanica Mexicana 100: 293-315 (2012) and Diego Angulo. 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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