Solanaceae diversity in the state of Jalisco, Mexico
Diversidad de la familia Solanaceae en el estado de Jalisco, México
Carmen Teresa Cuevas–Arias, Ofelia Vargas and Aarón Rodríguez*
Instituto de Botánica, Departamento de Botánica y Zoología, Centro Universitario de Ciencias Biológicas y Agropecuarias, Universidad de Guadalajara, Apartado postal 139, 45110 Zapopan, Jalisco, México
]]> * Correspondent:
Recibido: 08 noviembre 2006
Aceptado: 13 septiembre 2007
Abstract
Mexico is a center of diversity for Solanaceae. Our objective is to analyze the species diversity and geographical distribution of the Solanaceae in Jalisco. The data come from 3 405 herbarium specimens. An analysis of these specimens indicates that the Solanaceae in Jalisco are represented by 20 genera, 138 species and 140 taxa. Four genera, Solanum (55 species), Physalis (35), Cestrum (10) and Lycianthes (9) represent 79% of the total number of species. In contrast, Brachistus, Browallia, Chamaesaracha, Jaltomata, Juanulloa, Lycium, Nectouxia, Nicandra and Nierenbergia have only 1 species each. In Jalisco, the Solanaceae are widely distributed throughout the state occurring at altitudes ranging from sea level to 3 400 m. Mostly, they grow in conifer and oak forest (81 species) followed by tropical subdeciduous forest (57), tropical deciduous forest (54), and cloud forest (43). Fifty–one species are commonly found in disturbed and ruderal areas. Lycianthes jalicensis, Physalis lignesens, P. longipedicellata, P. longiloba and P. tamayoi are endemic to the state. These results indicate that Jalisco ranks fourth in species diversity for Solanaceae after the states of Oaxaca, Chiapas and Veracruz.
Key words: Solanaceae, distribution, species richness, Jalisco, Mexico.
]]> Resumen
México es un centro de diversificación de la familia Solanaceae. El objetivo de este trabajo es analizar la riqueza y distribución de las especies de Solanaceae en Jalisco. Se examinaron 3 405 ejemplares de herbario y como resultado se registra la presencia de 138 especies y 140 taxones agrupadas en 20 géneros. Los géneros con el mayor número de especies son Solanum (55 especies), Physalis (35), Cestrum (10) y Lycianthes (9). Estos representan el 79% de las especies. En contraste, Brachistus, Browallia, Chamaesaracha, Jaltomata, Juanulloa, Lycium, Nectouxia, Nicandra y Nierenbergia están representados por una especie. En Jalisco, las solánaceas crecen desde el nivel del mar hasta los 3 400 m. Las especies habitan con más frecuencia en el bosque de pino y encino (81 especies), seguido de bosque tropical subcaducifolio (57), bosque tropical caducifolio (54) y bosque mesófilo de montaña (43). En vegetación secundaria o como ruderales crecen 51 especies, Lycianthes jalicensis, Physalis lignesens, P. longipedicellata, P. longiloba y P. tamayoi son endémicas de Jalisco. Basados en estos resultados, Jalisco constituye la cuarta entidad más rica en especies de Solanaceae en México después de Oaxaca, Chiapas y Veracruz.
Palabras clave: Solanaceae, distribución, riqueza de especies, Jalisco, México.
Introduction
Solanaceae are almost worldwide in distribution, however, the majority of genera and species are neotropical. The family encompases 96 genera and almost 2 300 species (D'Arcy, 1991). The genera with most species are Solanum L. (1,000), Lycianthes (Dunal) Hassl. (200), Cestrum L. (175), Nicotiana L. (95), Physalis L. (80), and Lycium L. (75), (D'Arcy, 1991; Martínez, 1999; Nee, 2001; Fukuda et al., 2001, Knapp et al., 2004). Floristic studies (Rodríguez and Vargas, 1994; Vargas and Rodríguez, 1993, 1995; Vargas, 1998; Vargas et al., 1998, 1999) have shown the presence of numerous species of Solanaceae in the state of Jalisco.
Jalisco is located in western Mexico between 22°45' and 18°55'N latitude and 101°28' and 105°42'W longitude (Fig. 1). Altitudes range from 0 to 4 300 meters above sea level. The state has a surface area of 80 137 km2 which represents 4% of the Mexican territory. It includes 5 morphotectonic provinces: Northwestern Plains and Sierras, Sierra Madre Occidental, Central Plateau, Trans–Mexican Volcanic Belt, and Sierra Madre del Sur (Ferrusquía–Villafranca, 1993). As a consequence its flora includes phytogeographic elements from all 5 provinces.
Jalisco has several river basins with the most notable from the Lerma–Santiago River, which drains the northern and northeastern parts of the state. The Ameca river basin drains Central and Western Jalisco. The Mascota, Tomatlán, San Nicolás, Purificación, Marabasco–Minatitlán, Ayuquila, Tuxcacuesco, Armería, and Tuxpan rivers flow almost perpendicular to the Pacific Ocean and drain the coastal area. The southeastern corner is part of the Balsas River drainage basin. Lake Chapala, the largest freshwater lake in Mexico, is located at the eastern border of Jalisco and Michoacán. In addition, in south–central Jalisco, there are a series of seasonal and salty lakes forming the Zacoalco–Sayula land–locked system. A temperate climate with summer humidity and the tropical climate with summer humidity dominate the state (García, 1989; Villalpando and García, 1993). Rainfall is strongly seasonal extending from June to October. The northeastern corner and the coastal plains of Tomatlán in Jalisco are the driest areas, with less than 500 mm annually. In contrast, rainfall reaches its maximum, 1 600 mm in the uplands of the Sierras of Manantlán, Cacoma, Cuale and Mascota near the coastal plains (García et al., 1997a). Mean temperature ranges from 22° to 26°C in the coastal plains and 18° to 20°C at elevations of 1 600 m. Lastly, the mean temperature is less than 18°C in the highlands (García et al., 1997b).
]]> Thirteen plant communities are present in Jalisco (Fig. 2; Rzedowski and McVaugh, 1966; Rzedowski, 1978). Forty–five to 50% of the state is characterized by deciduous and subdeciduous forests. They occur along the coastal plains as well as in canyons in the central part of the state from sea level to 1 600 m. Some areas, scattered within the tropical subdeciduous forest along the coastal plains, are dominated by palms of the genus Orbignya Mart. ex Endl. Conifer and oak forests are most common in the highlands between 800 and 3 400 m and account for approximately one fourth of the state's surface area. Lower elevational pine–dominated areas are found only in the western corner of the state. Cloud and fir–dominated forests are restricted to the ravines and protected steep slopes within the conifer and oak forest zones. Savannas are found between 400 and 800 m on the Pacific slope and represent a transition zone between the tropical subdeciduous forest and the oak forest. The thorn forest includes an area of the coastal plains in the western part of the state as well as a mezquite community within the tropical deciduous forest. Grasslands are restricted to the northeastern corner interspersed with the xerophilous scrub vegetation. The mangrove community follows the ocean shoreline in areas with low–energy waves. In contrast, beach and frontal dune vegetation extend along the coastline. The beach habitat occupies sandy substrate adjacent to open ocean that extends from mean tide line to the top of the frontal dune. Finally, freshwater lakes, where the aquatic vegetation is found, are generally located in the central part of the state. The objective of this paper is to report and analyze the species diversity and distribution of Solanaceae in Jalisco.
Material and methods
The data come from 3 405 herbarium specimens held at the following herbaria: CHAPA, ENCB, F, GUADA, IBUG, IEB, MEXU, MICH, TEX, WIS, and ZEA. A database can be obtained upon request from the corresponding author. The determination of the specimens in herbaria was carried out using the identification keys in Bitter (1920), Standley (1924), Francey (1935, 1936), Correll (1962), Martínez (1966), Roe (1967, 1972), Waterfall (1967a,b,c), Hunziker (1969), D'Arcy (1973, 1978), D'Arcy and Eshbaugh (1974), Gentry and Standley (1974), Nee (1979, 1986, 1993), Whalen (1979, 1984), Davis (1980), Schilling (1981), Whalen et al. (1981), Knapp (1985, 2002), Bernardello and Hunziker (1987), Vargas and Rodríguez (1993), Bohs (1994), Rodríguez and Vargas (1994), Dean (1995), Vargas et al. (1998), and Spooner et al. (2004). Lastly, the species of Physalis and Solanum section Petota were verified by consultation of type specimens.
Results
The Solanaceae in Jalisco are represented by 20 genera (Table 1), 138 species and 140 taxa (Table 2). Solanum (55 species), Physalis (35), Cestrum (10) and Lycianthes (9) are the most diverse genera and include 79% of the taxa. Datura (6), Nicotiana (4), Bouchetia (2), Brugmansia (2), Capsicum (2), Petunia (2), and Solandra (2) group only few species. In contrast, Brachistus Miers, Browallia L., Chamaesaracha (A. Gray) Benth., Jaltomata Schltdl., Juanulloa Ruiz et Pavón, Lycium, Nectouxia H.B.K., Nicandra Adans, and Nierenbergia Ruiz et Pavón are represented by only 1 species each (Table 1).
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Discussion
The Solanaceae are widespread in Jalisco. They grow from the sea level to 3 400 m in the Nevado de Colima and are found in 11 out of the 13 vegetation communities described for Jalisco (Fig. 2, Table 2). They have not been collected in either savanna or palm forest. Eighty one taxa grow in conifer and oak forests whereas the tropical subdeciduous forest hosts 57 species. Fifty–four and 43 species have been reported from the tropical deciduous forest and the cloud forest, respectively. Furthermore, 28 species grow in grasslands and 11 species in thorn forests. Twelve species of Solanaceae are elements of the fir–dominated forest and 11 species prefer seasonally flooded areas. Seventeen species occur in the xerophilous scrub and mangrove is the habitat for Physalis acutifolia, Solanum diphyllum and S. tampicense. Finally, P. minuta grows only in the beach strand and frontal dune vegetation. Various species grow in more than 1 vegetational community (Fig. 3; Table 2).
Some species are frequent in conifer and oak forest, tropical subdeciduous forest, and tropical deciduous forest. These include Solanum americanum, S. nigrescens, and S. ferrugineum. Other less common species but recorded throughout the state are Cestrum confertiflorum, C. tomentosum, C. terminale, Jaltomata procumbens, Lycianthes moziniana, Physalis angulata, P. lagascae, P. nicandroides, P. waterfallii, Solanum dulcamaroides, S. elaeagnifolium, S. erianthum, S. grayi var. grandiflorum, S. lycopersicun var. cerasiforme, S. nigrescens, S. pseudocapsicum, S. refractum, and S. umbellatum. Less frequently, we can find Brachistus stramoniifolius, Capsicum annuum var. aviculare, Cestrum nitidum, C. thyrsoideum, Nicotiana plumbaginifolia, Petunia parviflora, Physalis orizabae, P. pruinosa, P. pubescens, P. sulphurea, Solanum adscendens, S. appendiculatum, S. bulbocastanum, S. candidum, S. chrysotrichum, S. hougasii, S. iopetalum, S. nigricans, S. pubigerum, S. stoloniferum, S. trifidum, and S. verrucosum. Lastly, Cestrum anagyris, C. aurantiacum, Chamaesaracha cernua, Datura quercifolia, Lycianthes pringlei, L. stephanocalyx, L. surotatensis, Physalis aggregata, P. angustiphysa, P. cinerascens, P. cordata, P. coztomatl, P. chenopodifolia, P. glutinosa, P. solanaceus, P. volubilis, Solanum aligerum, S. angustifolium, S. aphyodendron, S. brevipedicellatum, S. cardiophyllum, S. fructo–tecto, S. guerreroense, S. hazenii, S. pinnatisectum, S. stenophyllidium, and S. tridynamum have been collected in only a few areas.
Other species are known only from areas with particular environmental conditions. For instance, Lycium carolinianum grows exclusively around salty and periodically flooded lakes in south–central Jalisco. Similarly, Datura ceratocaula is a subacuatic species found only in seasonal ponds. Physalis acutifolia, Solanum diphyllum, and S. tampicense occur only in the mangroves while Physalis minuta grows only in the beach and frontal dune plant community. Juanulloa mexicana is an epiphytic shrub growing on Lysiloma sp. (Vargas–Rodríguez et al. 2006). Solanum verrucosum reaches the highest altitudinal range at 3 400 m.
The endemic species of Solanaceae in Jalisco are Lycianthes jalicensis, Physalis lignescens, P. longipedicellata, P. longiloba, and P. tamayoi. Other species are endemic to Jalisco and neighboring states. For instance, Physalis volubilis and P. waterfallii are endemic to Jalisco and Michoacán whereas P. hastatula has been collected only on the border between the states of Jalisco, Aguascalientes, and Guanajuato. Physalis aggregata and Solanum guerreroense were known only from the type locality in Oaxaca and Guerrero, respectively. Therefore, the collections for both of these species from Jalisco represent disjunct populations or range extensions. No genus is endemic to Jalisco. Nor are any species of Solanaceae listed in the Mexican Endangered Species Act (NOM–059–ECOL–2001, 2002).
In Jalisco, 51 native species occur in disturbed areas and ruderal vegetation. Datura stramonium, Jaltomata procumbens, Physalis nicandroides, Solanum americanum, S. elaeagnifolium, S. lycopersicum var. cerasiforme, and S. rostratum are favored by these disturbed ecological conditions. Two South American species introduced here, Nicandra physalodes and Nicotiana glauca, are very common in disturbed areas. In contrast, Solanum marginatum and S. sisimbriifolium, other introduced species in Mexico, are rare in the state of Jalisco. Solanum marginatum occurs as an invasive ruderal in marginal areas of the conifer and oak forest whereas S. sisymbriifolium grows in salty environments and tropical deciduous forest.
]]> Solanaceae are important vegetable crops and others are cultivated as ornamentals. The economies of the counties of Mazamitla and Tapalpa depend on the cultivation of potato (Solanum tuberosum). Similarly, the tomato (Solanum lycopersicum) is an important crop in the counties of Sayula and Autlán de Navarro. Peppers (Capsicum annuum var. annuum) and husk tomatillo (Physalis philadelphica) are extensively cultivated in Yahualica and Cuquío. Brugmansia candida and B. suaveolens are commonly planted in the uplands as ornamentals and as living fencepost while Cestrum nocturnum and Petunia hybrida are used extensively as ornamentals throughout the state. Likewise, Solandra grandiflora is a preferred ornamental along the coast and in the vicinity of Lake Chapala. Lycianthes rantoneii, Solanum seafortianum, S. wendlandi and S. wrightii are ornamentals common in the Guadalajara metropolitan area (the second largest city in Mexico). Finally, tobacco (Nicotiana tabacum) is extensively cultivated in Jalisco.Some wild species of Solanaceae represent sources of food in Jalisco. The fruits of Physalis philadelphica and the tubers of Solanum cardiophyllum and S. ehrenbergii are collected and consumed locally. Similarly, but less frequently, the fruits of Jaltomata procumbens, Solanum americanum, and S. lycopersicum var. cerasiforme are eaten in Jalisco. Lastly, the tubers of Solanum stenophyllidium also represent sources of food in the northern parts of the state.
As expected, the areas with the highest number of species correspond to nature preserves, national parks, and biosphere reserves in Jalisco. Comprehensive floristic studies have been published for these areas that include the Chamela Biological Station (Lott, 1993), Sierra de la Primavera Natural Reserve (Rodríguez and Reynoso, 1992), and the Manantlán Biosphere Reserve (Vázquez et al., 1995). The Guadalajara and Zapopan counties in central Jalisco, also evince a high number of species. Most likely, this result is correlated with the intensity of botanical exploration of the area considering that the botanical research centers are located there. These 2 counties include most of the Guadalajara metropolitan area. In contrast, we did not find any herbarium specimens documenting the presence of species of Solanaceae in the counties of Cuautla, Chimaltitán, Degollado, Guachinango, Manuel M. Diéguez, Mexticacán, Totatiche, Unión de San Antonio, and Villa Hidalgo. Certainly, it is an artifact of botanical collection.
The presence of 5 morphotectonic provinces in Jalisco could contribute to the species diversity of Solanaceae in the state. The south and the southeastern parts of Jalisco share floristic affinities with the states of Michoacán, Guerrero and Oaxaca through the Sierra Madre del Sur. Physalis aggregata, Solanum guerreroense, and S. iopetalum show this geographical distribution pattern. Lycianthes manantlanensis, P. gracilis, P. minuta, S. morelliforme, S. brevipedicellatum, and S. bulbocastanum are elements of the flora of Chiapas and Central America that have been found in Jalisco. Similarly, the Trans–Mexican Volcanic Belt, which spans Mexico from coast to coast, allows some floristic affinities with central and eastern Mexico. Physalis coztomatl, P. subrepens, P. volubilis, S. polyadenium, and S. verrucosum are examples of this geographical pattern. The Central Plateau extends to the northeastern corner of Jalisco as well as in other regions of Mexico. As a result, xeric taxa such as P. cinerascens, P. glutinosa, P. sordida, S. corymbosum, S. dasyadenium, S. elaeagnifolium, and S. fructo–tecto, typical of northern Mexico, are found there. The southern boundary of the Sierra Madre Occidental touches northern Jalisco and therefore these 2 regions share P. hederifolia and S. stenophyllidium. Finally, Solanum grayi is a common species of the Northwestern Plains and Sierras and reaches its southernmost geographical distribution at the northwestern part of Jalisco.
The results of this analysis show that the Solanaceae flora of Jalisco is the fourth most species diverse in Mexico, after Oaxaca, Chiapas, and Veracruz. Rodríguez (2004) reported the presence of 19 genera and 165 species in the state of Oaxaca with 7 species endemic to that state. Similarly, 24 genera and 164 species haven been reported for Chiapas, in southeastern Mexico (Breedlove, 1986). Four species are endemic to Chiapas. The Solanaceae in Veracruz, on the Gulf of Mexico slope, are represented by 138 species in 21 genera (Nee, 1986, 1993). Four species are endemic to Veracruz. These numbers are similar to those reported in the floras of Guatemala (27 genera and 182 species; Gentry and Standley, 1974; Knapp et al. 2006), Nicaragua (22 genera and 117 species; D'Arcy, 2001), and Panama (21 genera and 140 species; D'Arcy, 1973).
Acknowledgements
We thank Mollie Harker and Julie Villand for comments on an earlier draft of the manuscript. We also thank Christian Briseño and Arturo Castro for artwork and the staff at CHAPA, ENCB, F, GUADA, H, IBUG, IEB, MEXU, MICH, TEX, WIS, and ZEA for their kindnesses during our visit and for specimen loans. This paper is part of the B.S. thesis of C. T. Cuevas–Arias (2000). Distribución y riqueza de la familia Solanaceae en Jalisco. Centro Universitario de Ciencias Biológicas y Agropecuarias, Universidad de Guadalajara.
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