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The Chaparral Vegetation in Mexico Under Nonmediterranean Climate: the Convergence and Madrean-Tethyan Hypotheses Reconsidered1

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The Chaparral Vegetation in Mexico Under Nonmediterranean Climate: the Convergence and Madrean-Tethyan Hypotheses Reconsidered1 American Journal of Botany 85(10): 1398±1408. 1998. THE CHAPARRAL VEGETATION IN MEXICO UNDER NONMEDITERRANEAN CLIMATE: THE CONVERGENCE AND MADREAN-TETHYAN HYPOTHESES RECONSIDERED1 ALFONSO VALIENTE-BANUET,2,4 NOEÂ FLORES-HERNAÂ NDEZ,2 MIGUEL VERDUÂ ,3 AND PATRICIA DAÂ VILA3 2Instituto de EcologõÂa, Universidad Nacional AutoÂnoma de MeÂxico, Apartado Postal 70±275, UNAM, 04510 MeÂxico, D.F.; and 3UBIPRO, ENEP-Iztacala, Universidad Nacional AutoÂnoma de MeÂxico, Apartado Postal 314, MeÂxico, 54090, Tlalnepantla, MeÂxico A comparative study between an unburned evergreen sclerophyllous vegetation located in south-central Mexico under a wet-summer climate, with mediterranean regions was conducted in order to re-analyze vegetation and plant characters claimed to converge under mediterranean climates. The comparison considered ¯oristic composition, plant-community struc- ture, and plant characters as adaptations to mediterranean climates and analyzed them by means of a correspondence analysis, considering a tropical spiny shrubland as the external group. We made a species register of the number of species that resprouted after a ®re occurred in 1995 and a distribution map of the evergreen sclerophyllous vegetation in Mexico (mexical) under nonmediterranean climates. The TehuacaÂn mexical does not differ from the evergreen sclerophyllous areas of Chile, California, Australia, and the Mediterranean Basin, according to a correspondence analysis, which ordinated the TehuacaÂn mexical closer to the mediter- ranean areas than to the external group. All the vegetation and ¯oristic characteristics of the mexical, as well as its distribution along the rain-shadowed mountain parts of Mexico, support its origin in the Madrean-Tethyan hypothesis of Axelrod. Therefore, these results allow to expand the convergence paradigm of the chaparral under an integrative view, in which a general trend to aridity might explain ¯oristic and adaptive patterns detected in these environments. Key words: chaparral; convergence; evergreen; mexical; Mexico; sclerophyll; Tehuacan Valley; vegetation. The evergreen sclerophyllous vegetation consists most- tensively in a comparative manner for more than three ly of low-stature shrubs and trees (1±3 m high) and decades. Originally, all the studies were focused on test- broad-leafed sclerophyllous species with 40±100% cov- ing plant community-level convergence, assuming that erage by woody vegetation. This ecosystem has been as- under a similar climate these communities will evolve sociated typically with mediterranean climates with towards convergent solutions, including a maximum ef- warm, dry summers and cool, wet winters (Griesebach, ®ciency ¯ow of energy and nutrients (Barbour and Min- 1872; Cain, 1950; Naveh, 1967; Specht, 1969; Thrower nich, 1990). Since then, a signi®cant number of studies and Bradbury, 1977; Cody and Mooney, 1978; Di Castri, testing convergence have been published in the literature, 1981). It receives different names according to the ®ve including different mediterranean-climate regions (Na- different regions of the world where it grows, including veh, 1967; Mooney and Dunn, 1972; Parsons, 1976; Pig- ªchaparralº in California, ªfynbosº in the Cape Province natti and Pignatti, 1985; Cowling and Witkowski, 1994, of South Africa, ªmatorralº in Chile, ªmaquiaº in the among others). Mediterranean basin, and ªmalleeº in the south and Although the convergent viewpoint has played an im- south-western Australia (Cody and Mooney, 1978). Al- portant role in promoting comparative studies among though these regions occupy ,5% of the earth's surface, mediterranean-type ecosystems focused on plant physi- they harbor almost 20% of the known vascular plants of ology (Mabry and Difeo, 1973), plant anatomy (Kum- the world (Cowling et al., 1996). These plant communi- merow, 1973), phenology (Kummerow, 1983), ¯oristics ties, except the Chilean matorral, have also been tradi- (Pignatti and Pignatti, 1985; Arroyo et al., 1995), life tionally characterized by ®res. Resprouting ability by history (Armesto, Vidiella, and JimeÂnez, 1995; Zedler, means of lignotubers, burls, etc., has been considered an 1995), seed dispersal (Milewski and Bond, 1982: Hoff- adaptation to these disturbances (Naveh, 1974). mann and Armesto, 1995), and vegetation (Parsons and Under the view of the ªsuperorganismº or Clementsian Moldenke, 1975; Parsons, 1976; Naveh and Whittaker, paradigm, this ecological system has been analyzed ex- 1979; Cowling and Campbell, 1980), this conceptual 1 Manuscript received 24 April 1997; revision accepted 27 January framework has limited the approach in which the scle- 1998. rophyllous vegetation can be integrally viewed and un- The authors thank D. Axelrod, J. Rzedowski, P. GarcõÂa-Fayos, F.Llor- derstood. For example, paleobotanical evidence has been et, and one anonymous reviewer for comments on the manuscript; Leo- underestimated, although many studies of the paleobo- poldo Valiente and Ignacio MeÂndez for their help in statistical analysis tanical development of the chaparral have been described at the IIMAS. This study was supported by a grant provided by the DireccioÂn General de Asuntos del Personal AcadeÂmico de la UNAM in different publications by Axelrod (1950, 1958, 1973, (DGAPA-IN208195). 1975, 1977, 1989), emphasizing that a signi®cant number 4 Author for correspondence. of chaparral genera are represented in Tertiary ¯oras and 1398 October 1998] VALIENTE-BANUET ET AL.ÐCHAPARRAL VEGETATION UNDER NONMEDITERRANEAN CLIMATE 1399 in many cases the living species are very similar to the 1978; Pons, 1981; and Palamarev, 1989) than by similar fossils. These records provide reliable evidence that scle- (convergent) selection pressures under a Mediterranean rophyllous vegetation, which is distributed all over the climate. Keeley (1995) supports this explanation due to world, constitutes a reminiscence of the Madrean-Tethyan the remarkable similarity in seed germination, dispersal, sclerophyllous vegetation that occupied a subhumid belt and seedling recruitment patterns observed in California across much of North America-Eurasia region by the and Mediterranean species of Quercus, Rhamnus, and middle Eocene. This vegetation had originated from al- Prunus. liances in older laurophyllous forests that adapted to a The presence of the evergreen sclerophyllous vegeta- spreading dry climate (Axelrod, 1977). In North Ameri- tion in Mexico under a tropical climate of summer rains ca, the Madro-Tertiary Geo¯ora appeared on the drier has been reported by different authors (Muller, 1939, borders of the North American tropics by the Middle Eo- 1947; Miranda and HernaÂndez, 1963; Axelrod, 1975, cene and probably occupied much of the southwestern 1989; Rzedowski, 1978). Taking into consideration the United States and adjacent Mexico by the end of the Oli- information from these studies, it is possible to assume gocene, and it expanded its range north and southward, that the sclerophyllous vegetation in Mexico might be a as well as east and westward in response to expanding relict of the Madro-Tertiary Geo¯ora that constitutes the dry climate conditions during the Miocene epoch (Ax- principal element of the Mexican chaparral (named here elrod, 1958). Similar processes occurred in the Northern mexical) in nonmediterranean climates and that its cur- Hemisphere and also explain many of the broad ¯oristic rent patchy distribution probably responds to a gradual changes in the actual mediterranean-type ecosystems of trend toward increased dryness during the Tertiary and the Southern Hemisphere (Rundel [1981] for Chilean ma- Quaternary (Axelrod, 1958). If the sclerophyllous vege- torral, but see Arroyo et al. [1995] for a different hy- tation in Mexico exhibits the same plant characters as- pothesis on the origin of Chilean sclerophyllous vegeta- sociated with the chaparral vegetation under mediterra- tion; Specht [1981] for Australian Mallee and Linder, nean climates, this would allow analysis of what factors Meadows, and Cowling [1992] for South African Fyn- have contributed to similar plant traits dominating under bos). different climates. At the same time, and considering that Since the seminal model of Mooney and Dunn (1970), ®re has not played an important ecological factor in the proposing that in the mediterranean-type climates, ®re, TehuacaÂn mexical (only a 1-ha ®re occurring in 1995 has drought, high temperatures, rainfall unpredictability, and been reported in the study zone for .50 yr), the resprout- mineral de®ciencies have selected resprouting evergreen ing ability of plants and characters in plants that have sclerophyllous shrubs as the predominant growth form, been referred as adaptations to ®re need to be reconsid- most of the studies were designed to test for this evolu- ered. tionary convergence. Therefore, the central idea of the This paper is the ®rst attempt to assess a comparison model relating to the existence of an evergreen sclero- of the evergreen sclerophyllous vegetation located in phyllous vegetation with mediterranean climate has re- south-central Mexico (TehuacaÂn Valley), with other med- mained untouched, even though much evidence of its iterranean regions of the world. The aim of this study is presence in nonmediterranean climates all over the world to test whether the characters at the community and pop- should also be considered to expand the model. For ex- ulation level that have been claimed to converge under ample, Muller (1939) described the western montane mediterranean climates all
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