Influence of Plant Abundance on Disease Incidence in a Mexican Tropical Forest Author(S): Graciela García-Guzmán and Rodolfo Dirzo Source: Ecoscience, 13(4):523-530
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Influence of plant abundance on disease incidence in a Mexican tropical forest Author(s): Graciela García-Guzmán and Rodolfo Dirzo Source: Ecoscience, 13(4):523-530. Published By: Centre d'études nordiques, Université Laval DOI: http://dx.doi.org/10.2980/1195-6860(2006)13[523:IOPAOD]2.0.CO;2 URL: http://www.bioone.org/doi/full/10.2980/1195-6860%282006%2913%5B523%3AIOPAOD %5D2.0.CO%3B2 BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. 13 (4): 523-530 (2006) Influence of plant abundance on disease incidence in a Mexican tropical forest1 Graciela GARCÍA-GUZMÁN2 & Rodolfo DIRZO3, Departamento de Ecología Evolutiva, Instituto de Ecología, Universidad Nacional Autónoma de México, A.P. 70-275, C. P. 04510, México, D. F. México, e-mail: [email protected] Abstract: In this paper we investigate to what extent the occurrence of foliar diseases is affected by plant relative abundance in the understory community of the Los Tuxtlas tropical rain forest, how this changes with season of the year, and how plant species with different life histories present at the understory are affected by disease. To provide a context to these analyses we also include a general description of the floristic composition of the understory community. Using the eight most common species, we found that their relative abundance in each sampling location significantly explained the proportion of diseased plants. Accordingly, using the relative abundance of all plant species, we found that the probability of a host plant species being free of infection showed a significant decrement with abundance. At the plant level, we found that relative abundance had the greatest effect on the variation in leaf area/plant affected by pathogens, although the proportion of explained deviance was only 12%. Seasonality did not affect disease incidence and disease levels per plant. Throughout the year, plant relative abundance was very much lower in lianas, tree seedlings, and palms than in perennial herbs and ferns, and disease incidence was very much higher in the latter two, the most abundant life forms. These results collectively suggest that both intraspecific and interspecific variation in plant relative abundance explain variation in leaf damage by pathogenic fungi in tropical forest understories. Keywords: leaf pathogens, plant abundance, Los Tuxtlas, tropical rain forest. Résumé : Dans cette étude, nous avons examiné dans quelle mesure l’abondance relative des plantes en sous-étage dans la forêt tropicale de Los Tuxtlas influence l’incidence de maladies des feuilles. Nous avons aussi examiné si l’incidence de maladies change avec les saisons et comment les espèces de plantes du sous-étage forestier ayant différentes histoires de vie sont affectées par les maladies. Afin de mettre ces analyses en contexte, nous incluons également une description générale de la composition floristique de la communauté du sous-étage forestier. En utilisant les huit espèces les plus communes, nous avons déterminé que leur abondance relative à chaque site d’échantillonnage expliquait de façon significative la proportion de plantes malades. Ainsi, en utilisant l’abondance relative de toutes les espèces de plantes présentes, nous avons observé que la probabilité qu’une plante hôte soit non infectée diminuait significativement avec son abondance. Au niveau de la plante individuelle, nous avons trouvé que c’est l’abondance relative qui avait le plus d’effet sur la variation de la superficie des feuilles par plante infectée par des pathogènes même si la proportion de la variation expliquée était seulement de 12%. La saison n’avait pas d’effet sur l’incidence de maladies et le niveau d’infection par plante. Tout au long de l’année, l’abondance relative des lianes, des semis d’arbres et des palmiers était beaucoup plus faible que celle des herbes pérennes et des fougères et l’incidence de maladies était beaucoup plus élevée chez ces deux dernières formes de vie les plus abondantes. L’ensemble de ces résultats suggèrent qu’à la fois la variabilité intraspécifique et interspécifique dans l’abondance relative des plantes expliquent la variation du niveau de dommage des feuilles par des champignons pathogènes dans les communautés du sous-étage forestier tropical. Mots-clés : abondance des plantes, forêt tropicale, Los Tuxtlas, pathogènes de feuilles. Nomenclature: Ibarra-Manríquez & Sinaca-Colín, 1987. Introduction Several aspects of the abiotic and biotic environment, Guzmán & Wennström, 2001) have been shown to influence acting singly or in combination, have a profound effect the incidence of plant diseases in a wide range of natural on the spatial occurrence and severity of plant diseases systems. Surprisingly, the influence of plant abundance (Burdon, Jarosz & Kirby, 1989). For example, soil fungal and species composition on pathogenic incidence has been pathogens kill fewer tree seedlings in gaps than in the shad- poorly explored in natural systems, although a few limited ed understory of a tropical forest (Wenny, 2000). Likewise, studies indicate that differences in both the number of host biotic factors such as plant density (Burdon & Chilvers, plants available for pathogens and host plant spatial distri- 1982), competition between pathogens (Daamen & Stol, bution affect the establishment and spread of disease among 1990), and the presence of particular plant species (García- plants in temperate ecosystems (Burdon & Chilvers, 1982). In species-rich ecosystems, such as tropical rain forests, dispersal of pathogens with some degree of specificity for a 1Rec. 2005-09-06; acc. 2006-06-05. host-plant species may be restricted by the low abundance Associate Editor: John Klironomos. 2Author for correspondence. of a potential host-plant species (Burdon, 1978; May, 1991; 3Present address: Department of Biological Sciences, Stanford University, Stanford, Gilbert & Hubbell, 1996). However, observations in a spe- California 94305, USA. cies-rich tropical rain forest at Los Tuxtlas, Mexico suggest GARCíA-GUZMÀN & DIRZO: PLANT ABUNDANCE AND DISEASE INCIDENCE that ca 60% of plant species in the understory commu- Veracruz, Mexico (18° 34' to 18° 36' N, 95° 04' to 95° 09' W) nity exhibit leaf spots caused by fungal pathogens (García- (see details in Dirzo, González & Vogt, 1997). Mean monthly Guzmán & Dirzo, 2001). The floor of this forest is densely temperature is 27 °C, ranging from mean temperatures of covered by extensive carpets of high-density tree seedlings, 17 °C (January) to 29 °C (June). Total mean annual rain- frequently near parent trees of several species (see Dirzo & fall is 4750 mm, with a relatively dry season from April Miranda, 1990), or by high-density patches of herbaceous to the end of May. The rainy season extends from June to plants (García-Guzmán & Dirzo, 2001). Such high plant mid February (Soto & Gama, 1997). The vegetation of the seedling density can alter local environmental factors that area, tropical rain forest (Ibarra-Manriquez et al., 1997; favour disease development (e.g., increased humidity and Pennington & Sarukhán, 1998), is characterized by the pres- temperature) as well as increase disease spread through ence of evergreen vegetation with trees exceeding 35 m. greater host–pathogen encounter rates (Burdon & Chilvers, The forest understory is composed of herbaceous plants and 1982). In addition, given that environmental factors such as seedlings and saplings of trees, shrubs, lianas, and palms temperature and humidity change with time of the year, it (Dirzo et al., 1992; Ibarra-Manriquez et al., 1997). is possible that the effects of plant density will covary with season of the year. Environmental conditions are known to PLANT RELATIVE ABUNDANCE be particularly favourable for the attack of crop plants by We assessed the effect of plant relative abundance on different types of pathogens in the wet season of the year the incidence and levels of damage by leaf fungal patho- (Waller, 1976; Agrios, 1997), and they may also influence gens in four independent sampling sites in mature forest at disease incidence in the case of leaf pathogens, but this has Los Tuxtlas. In each site a sampling plot was established not been studied in tropical natural systems. On the other using a 50-m north–south straight line. Along the line, hand, plant life history could be another important source of 10 points were selected to establish 5-m-long transects, variation in pathogenic incidence. For example, attack could perpendicular to the central line. Their position (right or be lower in short-lived, ephemeral species (e.g., annual left of the 50-m line) and distance from the origin were herbs) than in long-lived,