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Small-Scale Spatial Patterns of Copernicia Alba Morong Near Bahia Negra, Paraguay

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Small-Scale Spatial Patterns of Copernicia Alba Morong Near Bahia Negra, Paraguay Natural Resources, 2013, 4, 369-377 369 http://dx.doi.org/10.4236/nr.2013.45045 Published Online September 2013 (http://www.scirp.org/journal/nr) Small-Scale Spatial Patterns of Copernicia alba Morong near Bahia Negra, Paraguay Michelle E. Cisz, Michael J. Falkowski, Blair Orr* School of Forest Resources and Environmental Science, Michigan Technological University, Houghton, USA. Email: *[email protected] Received June 23rd, 2013; revised July 20th, 2013; accepted July 31st, 2013 Copyright © 2013 Michelle E. Cisz et al. This is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. ABSTRACT We assessed the spatial distribution of Copernicia alba Morong. In the study area, a lowland palm savanna floodplain, C. alba is the only overstory species. We hypothesized C. alba would be randomly distributed within natural stands. Palms were tallied in six randomly located 0.25 ha plots and analyzed using a first-order, Ripley’s K function to assess the distribution of juvenile, adult, and total palm populations. While the total population had either aggregated or ran- dom distributions, when analyzing juvenile and adult population separately, we found juveniles were consistently more aggregated than the adults. Keywords: Ripley’s K-Function; First Order Analysis; Floodplain; Fire; Chaco; Monotypic Stands; Disturbance Ecology; Population Dynamics 1. Introduction ture have also been explored as variables [22,26,27] spe- cific to the plant rather than its environment and can be The spatial distribution of plant species is considered an scale dependent [11]. important ecological subject as distribution results from Physical patterns within the landscape such as soil, and mediates complex ecological processes [1]. Studies moisture or topography may generate patterns that are of the spatial distribution of palms and trees may be dissimilar from the pattern that would develop in a more purely descriptive, (this was especially true for the early uniform landscape [28]. For example, seedlings and ju- theoretical development of spatial distribution models) veniles of Borassus aethiopum, an African savanna palm, [2,3]; they are designed to elicit the underlying ecologi- are more aggregated than adults and the aggregated clus- cal mechanisms for the patterns [4,5], or observe or infer ters tend to occur on sites with better nutrient status [1]. changes in and the dynamics of spatial distribution over In general, spatial heterogeneity of resources [29,30] and time [1,4,6]. Frequently studies contain elements from all species diversity [31] may increase aggregation. A single three purposes; for example, it is the underlying ecologi- plant species may aggregate when growing in heteroge- cal processes that may drive temporal changes in species nous landscapes and in dense enough stands to generate distribution [1,7,8]. competition between plants even when the pattern for all Ecologists have hypothesized that plant distribution species combined is not aggregated [32]. may be based on seed dispersal patterns [4,9], underlying Populations may have different distributions at differ- heterogeneity of local environment [1,10], competition or ent scales [2,5,11,13]. Plants may be aggregated at larger facilitation between plants—both within and between scales but plant competition generates a more regular species [5-7,11], climate [12], and relative abundance of pattern at smaller scales [3]. In the early stages of plant a species within the plant community [13]. Ecological establishment species may benefit from the protection or factors influencing plant distributions may include mois- the amelioration of soil properties provided by a more ture availability [14], soil heterogeneity [1,15], topogra- mature plant, fostering aggregation of plants. This is es- phy [16,17], inter-specific competition [18-22], and seed pecially true in arid and semi-arid environments [6]. dispersal [11,23-25]. The plant’s role in temporal succes- Skellam’s [3] paper on theoretical random distribu- sion, intraspecific competition, along with forest struc- tions of plants assumed an open environment, one re- *Corresponding author. cently disturbed so that initially there is no inter-specific Copyright © 2013 SciRes. NR 370 Small-Scale Spatial Patterns of Copernicia alba Morong near Bahia Negra, Paraguay competition and one where the plants’ environment is area and were located within 0.5 km of two focal points. uniform. Most studies of spatial distribution include Each plot was a 50 × 50 m quadrat. The plots are not more than one species and some form of spatial hetero- replicates and statistical analysis was performed on each geneity. For example, in a study of different tropical for- plot separately. ests almost all species were aggregated and none were C. alba dominates both the overstory and the midstory dispersed; however, the number of species in the forest throughout the study area. The understory is codomi- ranged from 70 to over 800 [33]. Pillay and Ward’s study nated by members of Poaceae and Fabaceae family [34] of the small woody tree Acacia karroo is an excep- (Acacia and Albizia genera). Members of the Verbena- tion; all trees in their study plots were Acacia karroo. ceae, Asteraceae, and Apocynaceae families can also be The purpose of our study was to examine the spatial dis- found in the understory. tribution of Copernicia alba (Arecaceae) without the confounding influences of inter-specific overstory com- 2.3. Copernicia alba petition or a heterogenous environment. Our study loca- C. alba is 10 to 13 meters tall, with a maximum height of tion contained only one overstory species, Copernicia 25 m [48-51]. It is monoecious and generally does not alba, and the study plots were located on uniform flat produce flowers or seeds until it reaches at least 3 m in section of the Chaco of Paraguay near Bahía Negra, but height [52,53]. C. alba blooms between the months of distant from the Rio Paraguay. We used Ripley’s K func- August to October [46,53,54]. Fruits are 1.0 to 1.5 cm in tion as it allows for independent testing for random, ag- diameter [50,55]. The rachises that carry seeds grow gregated or dispersed distribution at different scales. Our from the center of the crown and can range from 0.5 m to null hypothesis was that species would be randomly dis- 2 m in length. Fruit are produced irregularly throughout tributed at all measured scales. the seasons and the majority of individuals drops their seed fruits in the late dry season (winter) with the onset 2. Methods of sporadic rainfalls, strong northern winds, and fire sea- 2.1. The Gran Chaco Basin son [46,54]. In the open canopied savanna C. alba is ex- posed to strong winds that could aid in dislodging the The Chaco is part of a Pleistocene Epoch lake that de- light fruits [56]. C. alba’s role with many of the endan- posited a thick layer of fine-grained sediment and poorly gered fauna and migratory birds has received limited drained soils along the Chaco’s northeastern edge [35,36]. study. Birds, small rodents, and native herbivores such as Fire is a historically common disturbance agent in the dry the greater rhea (Rhea Americana) and white-lipped and humid Chaco and throughout the neighboring Pan- peccary (Tayassu pecari) could play a role in dispersing tanal wetland [37,38]. The Paraguay River system sup- seed [4,53,56,57]. Other studies have shown that herbi- ports a landscape of monotypic stands of naturally re- vory may increase aggregation [4] or lead to a more re- generated palm forest intermixed with riparian edges rich gular dispersion, especially in more diverse plant com- in species characteristic of the Brazilian Amazon, Cer- munities [58]. The corky vascular meristems of palms rado, and Humid Chaco [35,39-43]. make them resistant to most savanna fires while second- dary adventitious roots sustain the palm in times of 2.2. The Study Area flooding [11]. The study area is located between the Pantanal wetlands Copernicia alba Morong has been documented from and the Humid Chaco forest, approximately 13 km west latitudes 14˚S to 27˚S and longitudes 56˚03'W to of the Paraguay River, W20˚04'5", S58˚17'16", with ele- 66˚00'W, along the Pilcomayo and Paraguay Rivers. vations of 82 - 85 m. Temperature ranges from 12˚C - Mature palms occupy the overstory in both forest and grassland habitats. Within forests, C. alba can codomi- 40˚C with an average rainfall of 995 mm/year [44,45]. nate the overstory and midstories. Throughout the Chaco The rainy season is from September to April, with De- C. alba is associated with planar alluvial flood zones that cember and January having the greatest rainfall [45,46]. cycle between floods and long periods of drought [46, The dry season is from August to October when hot 53,59-61]. northerly winds and wild fires are more common [47]. The region has a return fire interval of one to five years. 2.4. Study Design Key informants described the study area as being grazed in the 1970s by approximately 3000 cattle and palms A preliminary trip to the savanna was made in April of were harvested until a major flood had forced owners to 2010 and the first dataset was collected from Plots 1 - 3 move to higher lands. The land is currently operated as a during the middle of the dry season (winter) 2010. Field biological reserve maintained by Guyra Paraguay, a non- measurements for Plots 4 - 6 were taken during the end profit organization. of a light drought in November, 2010. Plot locations were chosen randomly within the study Within each plot all palms with a stem height of at Copyright © 2013 SciRes.
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