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Clark, David B. Clark, Rosa Sandoval M., Marco Vinicio Castro C. Source: Ecology, Vol. 76, No. 8 (Dec., 1995), pp. 2581-2594 Published by: Ecological Society of America Stable URL: http://www.jstor.org/stable/2265829 Accessed: 23/09/2009 17:03
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The most abundant species. accepted 4 March 1995. I Present address: Instituto Nacional de Biodiversidad.Ecology. Socratea. and ecosystems. CLARK. AND MARCO VINICIO CASTRO C. current ecological studI Manuscript received 12 September 1994. Of the seven species.2 ROSA SANDOVAL M.5-103 ha). All others showed significant edaphic variation in their distribution and/or estimated density. Basnet 1992. These findings and evidence from other sites suggest that marked spatial heterogeneity in community structure.O. To develop predictive power further requires understanding the processes. We studied the landscape-level spatial variation in distribution and abundance of seven species of canopy and subcanopy palms in a neotropical rain forest. at small to large scales (0. while present at all sample points. This substructuring of the arborescent palm guild results from the interplay of edaphic variation and past human activity. Within 568 ha of nonswamp old-growth lowland forest at the La Selva Biological Station. 1985. tropical rain forest. edaphic variation. Two other closely related species. Louis. Total stem density of this guild varied with both edaphic factors. but not for larger individuals. communities. Euterpe macrospadix was biased toward steep topography and less fertile sites. Another potential source of small-to-medium scale spatial heterogeneity within tropical wet . palms. while virtually omnipresent across soil types and topographic positions. Much of the Amazon Basin is a highly interdigitated mosaic of forest types produced by the erosion/deposition cycles of the major rivers (Salo and Rasanen 1989. confertum) were rare. Welfia. Welfia georgii. a first need is to determine how the distributions and abundances of the organisms vary across the landscape.. spatial heterogeneity. Two closely related species had strong and opposite edaphic associations. Study of the spatial scales and causes of this variability will produce a more robust understanding of these complex ecosystems. Johnston 1992. 8001 Natural Bridge Road. Chauvel et al. Lescure and Boulet 1985. showed marked reciprocal variation in density between related soils. Louis. Prestoea decurrens was nearly ommipresent on soil types with gentle topography while absent from half the points on soils with steep slopes. INTERLINK-341. We found mosaics of community structure related to marked within-forest variability in both soil and topography. Missouri 63121 USA (mailing address remains as above). Key words: Astrocaryum. Iriartea's spatial distribution further indicates local removal of this species from one sector of the old growth by human harvesting (with subsequent apparent "release" of Socratea in this site). Local (point) species richness of the subcanopy and canopy palms varied among soil types. Miami. INTRODUCTION To understand many aspects of populations. Euterpe. P. Box 02-5635. ies often lack the underpinnings of a clear understanding of the important scales of spatial variation within study sites. University of Missouri-St. In the wet tropics a growing number of studies at small-to-large scales within old-growth forest have demonstrated strong effects of edaphic heterogeneity on floristic composition (Ashton 1969. Prestoea. Costa Rica. landscape ecology. Ruokolainen and Tuomisto 1993. Guillaumet 1987). we sampled at 516 intersections of a reserve-wide grid to systematically assess the species' distributions across the major edaphic gradients. 2 Present institutional address: Department of Biology. Iriartea deltoidea and Socratea exorrhiza. Steep sites had twice as many large palms (> 10 m tall) per hectare than those on gentler topography or at lower slope positions. pp. The combined density of subcanopy and canopy palms also varied significantly among soil types in the smallest size class (1-5 m tall). Oliveira-Filho et al. 1994). 1995. Costa Rica. 2581-2594 ? 1995 by the Ecological Society of America EDAPHIC AND HUMAN EFFECTS ON LANDSCAPE-SCALE DISTRIBUTIONS OF TROPICAL RAIN FOREST PALMS1 DEBORAH A.2 DAVID B. Newbery and Proctor 1984 and cited references. 1987. Florida 33152 USA Abstract. revised 21 February 1995. two (Astrocaryum alatum and A. Foster 1990) and marked variation in terra firme parent material and geochemistry (Jordan 1985. Although these ideas are generally recognized by most ecologists. 76(8). may be general among tropical wet forests. Iriartea. Costa Rica. final version received 27 March 1995. St. Gentry and Ortiz 1993. past and present. La Selva Biological Station. CLARK. Lieberman et al. showed significant among-soil variation in density. Santo Domingo de Heredia.3 Organization for Tropical Studies. underlying the spatial heterogeneity thus identified (Levin 1992).
2582 TABLE DEBORAH A. Weimann & P. Four species have been locally harvested (for heart-of-palm. On the nonalluvial soils we sampled at the in- . Understanding the patterns and causes of spatial variability in the community structure of tropical wet forests can contribute to the resolution of important questions about these ecosystems. METHODS The La Selva Biological Station of the Organization for Tropical Studies (OTS) is a 1550-ha reserve in the To evaluate landscape-scale distributions of the large palms in upland old-growth forest. as is the frequency of gap formation (Hartshorn 1978. Stevens 24559] Prestoea decurrens (Wendl. Our study species are the seven subcanopy to canopy palms found in old-growth at La Selva (Table 1). Chazdon 1985. size distribution. H. Moore [M. soil map. McDade and Hartshorn 1994). construction wood. The seven species of canopy and subcanopy palms (Arecaceae. construction wood construction wood (less used than Iriartea). The flora includes 323 tree species (Hartshorn and Hammel 1994) and is rich in palms (31 native species. Although not subject to large-scale disturbances such as hurricanes. Stem turnover is high (2.) H. Chac6n. 1) we sampled at intersections of La Selva's reserve-wide grid (posts every 50 m along lines at 100m intervals. Tribe designations are from Uhl and Dransfield (1987). at the scale of 500 ha of contiguous forest: (1) How do the abundance. which they identified from line drawings. D. 1985). Tribe Iriarteeae Species (voucher no. Voucher specimens are from the Costa Rican National Herbarium. Mora 1968] Socratea exorrhiza (Martius) Wendl. 1990). Stevens 24625] Welfia georgii Wendl. de Nevers. 1994).4 ha of restricted access plots. Jermy 6783] Euterpe macrospadix Oersted [M. Is small-to-medium scale edaphic heterogeneity more characteristic of these forests than of temperate stands? Could this be a factor in the maintenance of high species richness in tropical rain forests? Are wet tropical forests characterized by major internal spatial heterogeneity in ecosystem-level processes such as primary productivity? Is variability in site conditions and forest communities likely to have major impacts for efforts in tropical forest conservation and restoration? Our goal in this study was to assess landscape-scale spatial heterogeneity within an old-growth neotropical rain forest. Bush and Colinvaux 1994). An unprecedented set of research tools for landscape-level studies of tropical rain forest (a reserve-wide grid system. 8 1. Mean annual rainfall is 3962 mm. 84?00' W. H. ex Burret [M. Lieberman et al. Lieberman et al. In the limited areas of alluvial soils. even within stands considered to be old-growth (Gordon 1982. La Selva's soils range from relatively fertile entisols and inceptisols to infertile ultisols (Sollins et al. Grayum. from silviculture to selective harvesting. C. as reflected in the distribution and abundance patterns of the large palms. and local species diversity of the guild of subcanopy and canopy palms vary with respect to the landscape-scale variation in soil and topography? (2) Do the distribution and abundance of the individual species vary with edaphic conditions. D. rarely heart-of-palm none heart-of-palm (less used than Iriartea) none none thatching Areceae Cocoeae Geonomeae forests is the historical or current impact of local human activity. It is classified in the Holdridge life zone system as tropical wet forest (Hartshorn and Hammel 1994). Grayum 7813] Astrocaryum confertum Wendl. accuracy ?20 cm). Vol. secondary forest. Gomez-Pompa and Kaus 1990. 1994). E. Fig. Ecology.3%/ yr for trees ?10 cm dbh. CLARK ET AL. [W.0-2. 76. and the 12. with every month averaging at least 100 mm of rain (Sanford et al. 25% of all woody stems ?10 cm in diameter. ex Burret [G. the forest is very dynamic. No. we used the original La Selva reserve ("Original La Selva". Sanford et al. elevation 37-150 m). E. Henceforth all except the Astrocaryum species are designated by genus name only. M. Anderson 1990. We used them to ask the following questions. B. after excluding swamp. Hammel 7820] Astrocaryum alatum Loomis [W. Within the resulting study area (568 ha. Local uses were determined by interviewing eight long-term residents of the surrounding region (Cant6n de Sarapiquf) about recent historical uses of each palm species. we sampled at all grid intersections (N = 78). and do these patterns differ among species? (3) Is there evidence of human impact on the distributions of large palms in the oldgrowth forest? STUDY SITE AND SPECIES Atlantic lowlands of Costa Rica (10?26' N. The 1:10 000 soils map of La Selva (Sancho and Mata 1987) demarcates 23 consociations (mapping units within which ?75% of the area is the described soil type) and one complex. disturbed habitats. 1986). Rich 137] Local uses heart-of-palm. thatching) in recent history (Table 1). and Geographical Information System [GIS]) is newly available at the study site. Subfamily Arecoideae) in old-growth forest at La Selva and their local uses.) Iriartea deltoidea Ruiz & Pav6n [Chac6n.
We excluded points that fell within streams or floodable microsites m as well as those 30() from secondary forest or altered habitat.. they have higher pH. For each size class. S. J. Denslow and G. > 10(-15. The is distributionof Iri(artea dleltoidleat shown by: present (0). soil Residualsoil. El Surd-Saltito (Typic Humitropept). Streamis [swampsare indicatedby the figuredhatchingpattern]). S. 1994). and topography to refine this map. N = 398).ipublishled daita). unpublished data). The resulting GIS coverage was used to classify each sample point by major soil type. The Jaguar and Maitabuievconsociations. While low in pH and exchangeable bases. The Stream consociations occur in the valley bottoms of principal streams: El Scalto (Lithic Humitropept). N = 438 grid points). and/or exchangeable bases than the other three soil types (Sollins et al.1. the grid survey data (OTS records) show these sectors to be at equivalent elevations (Experimental: mode = 42-44 m. Steep Slope. At each point we used this technique tor each of four size classes: 1-5. J. and with poor-to-moderate drainage (Sollins et al. Although divided into the Experinental and Holdridge consociations.0. Gentle Slope (all the slope positions between Crest and Base.-In the field we categorized each sampling point (grid post) by topographic position: Slope Crest (just above to just below the upper breakpoint of a slope). by the proportion of that species among the Closest Individuals sampled in the area. N = 132. 2. This "Closest Individual Method" (Cottam and Curtis 1956) enables estimation of stem density [Densitv (N/ha) = 10 000/(4D2)J.. and El Taconazo (Typic Tropaquept). classified as Ultisols (Typic Tropohumults). absent (0).December1995 TROPICALPALM COMMUNITYSTRUCTURE 2583 tersections along every other grid line (thus along lines separated by 200 m. Trailsare shown as dashedlines. cover 45% of La Selva (Sollins et al. 1994). Alluvium is an Andic Humitropept (Inceptisol) on gently undulating terrain. Chaverri. we measured the distance from each sample point to the nearest individual of any of the species and noted the species of that palm. 1994). but equivalent in pH to the Residual soils (J.-For each of four size classes. 1. 4 '4 1~~~~~0p 1 0 100 300 800 metres MVCC-JJP/Jan95 Ftc. The study area in old-growth(nonswamnp) forest (the thick blackborderindicatesthe boundaryof the sampled area). A telescoping 15--mmeasuring pole was used to evaluate palm heights. Pailnm samnplinig. in a given soil type or at a given topographic position) was calculated by multiplying the total stem density of canopy and subcanopy palm species (see formula above). Estimnated densitv. Chaverri. 1994. Matabttey. with slopes classed by relative steep- . We used our field notes regarding streams. where D is the mean distance in metres to the closest individual (the mean of the distances of Closest Individuals from all sample points). which were previously considered distinct terraces (Sollins et al. They are acidic and low in exchangeable bases and extractable P (Sollins et al. >5-10. Moderate Slope. Edap)hic factors: To)pograiphvy. on Sun Sparc II workstations). 44 m. Edaiphic factors: Soils. Arboleda soil. unlpublished data) The Residual consociations (Jagutar. This soil appears intermediate between the Alluvium (Holdridge consociation) and Residual soils in extractable P.-From data at each sample point (grid post) we evaluated distributions of the seven palm species in two ways: in terms of estimated density and presence/absence. 1. extractable phosphorus (P). Denslow and G. and then grouped related soil map units into four categories. Arboleda is a consociation thought to represent old.-The La Selva soils map is in the station's GIS (Arc/Info 6. and Esquina) are highly weathered soils derived from old lava flows and overlie broken ridge-valley topography. All are strongly acidic. 1994. S. un. Differentgrey tones indicate the four soil types (from lightest to darkest grey tone: Alluvium [Holdridge and ExperimentalConsociations of the Alluvium are labeled as "Ho" and "Ex' l.e. swamps. and > 15 m tall. Denslow and G.-For each of the seven study species we noted whether any individual > 1 m tall was visible (at any distance) in any direction from the grid post or while measuring the Closest Individuals. highly weathered alluvial deposits that overlie very broken terrain (Sollins et al. Holdridge: mode = 42- 4 S @0 N 4. Circles denote the 516 points sampledalong the 50 x 200 m grid covering La Selva. density of each species in a given area (i. very low in exchangeable bases. Chaverri. 1994). Preseele/Absence.
from left to right (with N.2584 550 500cu 450o4001350 300z ii DEBORAH A. The relation of total palm density (all seven species combined) to topographic position (all soils combined). gentle slope-91. (A) Individuals in the two smaller size classes. mean slope angles were (N. 3A). 24. the probability used for significance is 0. 8 sZZ l 15 IEI RESIDUAL 111ARBOLEDA STREAMS ALLUVIUM 250- 200C< 15010050 1m > 5-lOin >10-15m >15m HEIGHTSIZE CLASS 2.60 (150). total estimated density across topographic and soil gradients produce marked spatial heterogeneity in their local abundance. Probabilities are for a Kruskal-Wallis test of the distances to the Closest Individual (see Methods). the presence-absence data were tested with chi-square (number of sample points with a given species present and absent vs. 6.001. (B) The two larger size classes (note different y axis). 76. 0 LU 200 -J z Statistical analysis.-By far the highest estimated densities (203-509 palms/ha. Probabilities are for a Kruskal-Wallis test of the distances to the Closest Individual (see Methods). In the two larger size classes (>10-15 m tall.70 (124). CLARK ET AL.60 (60). Changes in .. -: 400 630- ness). FIG. B. *** P ' 0. Vol. steep moderate slope-42. and Flat (flat terrain. Topographic effects. In both size classes palm density varied twofold among topographic positions. however. Significant among-soil variation in a species' representation among Closest Individuals indicates that the species' density varies significantly among soils. Clark et al. the seven species combined) occurred in the smallest size class (1-5 m tall. To test whether total palm density in a given size class differed among soil types (or topographic positions). Ecology. Moderate Slope.01. did not vary significantly with topography (Fig. by * P ' 0. Gentle Slope. >5-lOinm * 50o 40 CO 30 z 20-J- 0 >10-15 m >15 m HEIGHTSIZE CLASS 3. 3B). RESULTS 0- iS5m 60 B. Total palm density The abundance of subcanopy and canopy palms varies greatly across the La Selva landscape. A chi-square test was used to compare the soil types in terms of the Closest Individuals that were and were not the species in question. a Kruskal-Wallis test was performed on the Closest Individual distances for that size class in each soil type (or topographic position). Throughout. Base of Slope (just above to just below the lower slope breakpoint). Variation in total palm density (all seven species combined) by size class and soil type. >15 m tall). 15. Abundance dropped sharply at larger sizes. flat-83.by topographic position. total density decreased continuously from slope crests. soil types or topographic positions). Fig. slope base-57. to slopes of decreasing steepness. Fill patterns. points): Steep Slope. FIG. whether on broad ridges or terraces or in swales). and Flat. slope-105. ** P ' 0. When this field index was combined with slope angle measurement at the same point in another La Selva study (D. unpublished data).70 (91). species could be individually tested for an association between their density and soil. No. The number of smaller palms per hectare.05.-Estimated stem density varied significantly in relation to topography. When total palm densities did not differ across soils. significantly different distances to Closest Individuals indicate significant density differences among the edaphic conditions (greater distances indicate lower density). 1. soil type.-To test for species associations with edaphic conditions. to slope bases and flat terrain (Fig. Density and palm size. points sampled): slope crest-138.05. Sample sizes are given in Table 2. 2).
The combined data on presence/absence and estimated density reveal: (1) general rarity of the two Astrocaryum species. Species distributions The seven palm species have different distribution patterns across La Selva. 21%. 1%. however.-The relation between estimated palm density (all seven species combined) and soil type (Fig. P < 0. EU-Euterpe.0-3. Sample sizes as in Table 2. this species occurred at only 10% of the points in Alluvium. 38%. we carried out two additional analyses of Euterpe's distribution patterns. A single individual occurred in the density sampling on the Alluvium. palm densities were equivalent across the four soil types. 2) contrasted with the topographic patterns. confertum (from Closest Individual data) was 0. Euterpe's abundance also significantly varied with soil type at all sizes (Appendix).December 1995 TROPICAL PALM COMMUNITY STRUCTURE 2585 100RESIDUAL 900 H l_ARBOLEDA w E - 5 _ V/A STREAMS ALLUVIUM w W z 80 C/) CE W 70 6050 0~~~~~~~~~~~~~~~~~~~~~~~~~~R 1 w 00 20 10 . Species abbreviations: WE-Welfia. did vary significantly among soils (Kruskal-Wallis. A parallel pattern is seen with the Closest Individual data (percent of sample points with Euterpe as ?1 of the four Closest Individuals: Residual. A.'30l l WE SO IR EU SPECIES PR AL CO The distribution of the seven canopy and subcanopy palm species with respect to the four major soil classes. Comparing presence/ . IR-Iriartea. with intermediate frequencies in the other two soil types. Its distribution was also strongly associated with soil type per se. Euterpe was absent from half of the Flat sites. Moderate Slope). (3) evidence of local removal of one species (Iriartea) through human harvesting. P < 0. Abundance of the smallest palms. SO-Socratea. df = 3. P < 0. Steep Slope. Arboleda. AL-Astrocaryum alatum. Similarly. FIG. alatum estimated density was 0. Data for each species are the percent of sample points in each soil type at which any individual >1 m tall was observed. Euterpe's presence/absence patterns within the Residual soils (Fig. SA) strongly varied with topographic position. Although present at 91% of sample points in the Residual soils. (4) reciprocal density differences between Iriartea and Socratea. Stem density at this size was much lower on the Streams soils than on the other three soil types (Tukey a posteriori pairwise comparison. they occurred at few sample points (3-5%. a species' association with soil type could reflect responses either to topography and/or to substrate per se.5 stems/ha (?10 cm diameter) in a 2-ha plot in swamp. Alluvium. The many sample points on the Residual soils allow assessment of species' topographic associations within a single soil type.0 stems/ha across sizes and soils. and (5) significant intersoil variation in the density of one ubiquitous species (Welfia). CO-Astrocaryum confertum. is abundant in swamps at La Selva.0 stems/ ha in all size classes on all soils. (2) strong and opposite associations with soil type and topography for the species Euterpe and Prestoea. confertum. Soil effects. In all four soil types. While present at >90% of points associated with steeper topography (Slope Crest. The estimated density of A. 4). 4. alatum).-The presence/absence data (Fig. To separate these factors. This species.2 stems/ha).0001). PR-Prestoea. A. except for the 1-5 m height class on Residual soils (1. Astrocaryum. 25%. df = 3. however. A. Euterpe. Estimated densities were markedly higher on the Residual soils than on the other three soils. 4) revealed strong soil-related heterogeneity in Euterpe's distribution (chi-square.-Both Astrocaryum species were rare in the sampled area (Fig.03). 2-i 1%.05). chisquare. P < 0. Because topography varies with soil type (Table 2). Hartshorn (1983) reported 30. For the three larger size classes. Streams.0001).
27 0. 20. Together these patterns suggest balanced avoidance of gentle topography (Table 2) and more fertile conditions (based on the limited nutrient data for La Selva soils.15 0. 14.2586 TABLE DEBORAH A.9. 52.28 Total 1.00 N 44 329 65 78 Area sampled (ha)* 340 43 65 64 * Additional area was sampled at upland sites embedded within the 55 ha of swamp in the total study area. for each.3%.8 (1-5 m tall palms) and 3.0001). and its density on the Residual soils (Density ratios.00 0. Socratea. Prestoea density was lowest on the Residual soils. While present at only 29 and 47% of points associated with steeper topography (Slope Crest and Steep Slopes. and nearly absent from Alluvial soils (gentle topography. density on the Streams soils and Arboleda soils.30 0. Prestoea's occurrence within the Residual soils (Fig.00 0.69 0. 6A).00 1.00 1. CLARK ET AL.31 Steep slope 0. but in directions opposite to those found with Euterpe. Stronger shifts with increasing size class occurred in the ratios between Prestoea's z 0 30- U_ 20-'0 100 EUTERPE 100 B w Ul) w PRESTOEA 80- cE cE o~7060- 50- ft u) Z 4030 q 0 u_ 0 20 10S} 0 S IRIARTEASOCRATEA WELFIA FIG. see Methods). and Welfia. Its distribution 100 A. Euterpe's significant density differences among soils (Appendix) paralleled these presence/absence patterns. 3.0001) is from a chi-square test for association of species presence/absence with topographic position. These patterns suggest that in both the 1-5 m and >5-10 m size classes. steep slope (70).09 0.11 0. respectively). from 3. Prestoea suffers higher mortality on the Residual soils than on the Streams or Arboleda soils. Prestoea's estimated density (stems per hectare.0 (10-15 m tall).06 Flat 0.05 0. from left to right (with N. P < 0. df = 3. gentle slope (30). 4) varied markedly among soil types (chi-square. 8 2.3. points): slope crest (98).00 0. Alluvium.1. Arboleda/Residual-0. Euterpe's distribution was: nearly omnipresent on Residual soils (steep topography. two pairs of adjacent topographic positions were combined to eliminate expected values <5). it occurred at 91-92% of points in the Streams and Alluvial soils.15 0. within each soil type.19 Base of slope 0.7.02 0. Vol. Prestoea only occurred once as a Closest Neighbor at >15 m height).00 1. infertile).3%. flat (9).0001). .36 0. within the Residual soils. P < 0.21 0. controlling for soil type. and >10-15 m height classes.10 0. Prestoea. A similar pattern exists in the percent of sample points with Prestoea as any of the four Closest Individuals (Residual. Prestoea's presence/absence patterns (Fig. intermediate occurrence on both the Arboleda soils (steep topography. 2.03 Moderate slope 0. infertile). z w w w Tul) H 90 705040- absence at the same topographic position (the four po- sitions with adequate samples) showed much more frequent occurrence on the Residual soils than on the Alluvium (Fig. While at only 48-53% of points in the Residual and Arboleda soils. (B) Iriartea. Ecology. No. moderate slope (90). Prestoea was at 78 and 81% of the Slope Base and Flat sites. such withinswamp points were classified to soil type based on the local topography and proximity to soil map units. Proportion within each soil type Residual Arboleda Stream Alluvial Topographic position Slope crest 0.8.03 0.25 0. Fill patterns. (A) Euterpe and Prestoea (df = 3 for both chi-square tests. intermediate fertility) and the Streams soils (base of slopes/flat terrain. but in the opposite direction to that of Euterpe.5%.-This close relative of Euterpe was also strongly associated with both topographic and soil variation. Data are the percent of sample points at which each species was present at each topographic position. 76. Probability (**** P ' 0. 5.0). Streams. The ratio between its density on the Alluvium and on the Residual soils increased with size class. >5-10 m.5 (5-10 m tall) to 5. Topographic variation on the four major soil types (see Methodsfor descriptions of soil classes and topographic positions).13 Gentle slope 0. 1-5 m. fertile). Overall. As with Euterpe. slope base (32). 5. SA) was strongly associated with topographic position. chi-square.21 0. 7. Arboleda. Appendix) varied significantly across soil types in all three size classes (smaller than the other six species.2%. Data are the proportion of sample points in each topographic position. df = 3. respectively: Streams/Residual-2. 46. Topographic affinities of the abundant canopy/ subcanopy palm species.
Its presence/absence patterns (Fig. df = 3. suggest a stronger association with topography than with soil. In contrast. flat. 6B) and by the size-related increase in relative density on the Arboleda soil. Taken together. 4).December 1995 TROPICAL PALM COMMUNITY STRUCTURE A. and have equivalent distributions of topographic positions (chi-square.001. Iriartea. 2587 was also significantly associated with soil type per se. sample points (Residual soils and Alluvium. in spite of these soils' apparent fertility difference. respectively): slope crest. P < 0. N. A nonedaphic factor distinguishing the two soil map units. are relatively remote from modern human habitation. In the largest size class.-Unlike Euterpe and Prestoea. with this process occurring over a protracted period. however. these patterns indicate that Prestoea's distribution is associated with both soil and topography. Iriartea did not show an association with topography (Fig. had equal estimated densities on the two Alluvium consociations (Appendix). 5B).01. A bias toward more fertile soil is suggested by the Alluvium/Residual comparison at given topographic positions (Fig. Mapping (Fig. An edaphic explanation for Iriartea's distribution within the Alluvium is unlikely given: (1) the species' lack of variation with topography. Each probabilityis for a chisquare test for association of the species' presence/absence with soil type. black bars). df = 3. df = 1.0001 and P = 0. 30 and 15. although the pattern is less clear. 90 and 10. Prestoea's density is lowest on the steep Residual soils and highest on the flat Streams soils. these lines of evidence indicate that Iriartea's absence from the Experimental consociation is due to intense past harvesting of the species from this sector of the La Selva old-growth forest. at a fixed topographicposition (** P < 0. SLOPE GENTLE SLOPE FLAT FIG. while contrasting in fertility). 6. *** P < 0. Su Ul) 80- 70w LU 60I50- U) Z 403020- 0 aIL O 10 0 SLOPE CREST MOD.89) and elevations (see Methods). (2) its virtual omnipresence in all four soil types (within the Alluvium. however. The Experimental consociation lies between a former commercial cacao plantation (operative until the 1980s) and the former housing for La Selva farm workers (abandoned in the early 1980s). is location 10080- *** *** Ul) 70m UJ 60 I50- cl Z 4030- aO 20 10C -0 SLOPE CREST MOD SLOPE GENTLE SLOPE FLAT 100F . where it occurred at 50% of the points. Iriartea's absence from this consociation in all size classes >1 m tall suggests complete local removal of large stems by harvesting. The presence/absence data. did differ significantly among soils (chi-square. omnipresent in the Holdridge consociation). but topographically equivalent) Residual soils. the species was virtually absent from the Experimental consociation (present at only 2 of 42 points). Similarly. P = 0.0000). 4). Prestoea was nearly omnipresent on the Streams and Alluvium soils (both soils of low slope positions/gentle topography.0001). in analyses controlling for topography (Fig. moderateslope. Iriartea was present at 91-98% of sample points on all soils but Alluvium. compared to the (less fertile. 6B). P = 0. (3) that both strongly edaphically varying species. 9 and 22. The significant among-soil differences in Prestoea's stem density in the two smaller size classes (Appendix) follow the same general trend (markedly lower densities in the Residual and Arboleda soils than in the Streams and Alluvium soils). Euterpe and Prestoea. 6A). 1) revealed a strong spatial bias in Iriartea's distribution within the Alluvium. In contrast to Euterpe (Fig. with respect to human activity and access. Such ac- . and (4) the similarity of the two consociations. **** P < 0. however. Although found at all 36 sample points in the Holdridge consociation of the Alluvium. 98 and 24. The two consociations are both classified as Andic Humitropepts (Sollins et al. 1994). Fig.01 for two of four comparisons). Soil affinities of (A) Euterpe and (B) Prestoea. Prestoea occurred more frequently on the Alluvium than on Residual soils at all four topographic positions that could be compared (chi-square. Taken together. the species occurred at only about half the sample points on the two soils on highly dissected terrain (Residual. Arboleda. the old-growth areas of the Holdridge consociation. controlling for topographicposition (Residual soils: white bars. gentle slope. Alluvium. while not far from a neighboring cattle ranch.
did vary significantly among soil types (two of four size classes. the summed density of the two palm species in each of these size classes is remarkably similar in the two consociations. To compare all soils we excluded Iriartea because of the confounding effect of human harvesting of this species from the Experimental consociation.01 (two-tailed). which have similar topography (Table 2. Appendix). 4.001. Its estimated densities were overall the highest among the seven palm species. these tests effectively compare densities (see Methods) of a given species on the two soils of each pair (or of the two species combined across the soil pair): *** P < 0.51. Between these two soil types. Welfia was least abundant on the Streams soils. Residual.58. Nevertheless. Probabilities are for chi-square tests of Closest Individual data. however. 4). The Binomial probability of such an outcome is <0. SB). In spite of each species' large density differences between consociations. and Welfia) showed density differences between these two Alluvial consociations (Appendix). however. Socratea and Iriartea have reciprocal density patterns on similar soils (Table 3). but is unlikely to have occurred after strict site protection was established in the early 1980s. The mean numbers of palm species present at a sample point were: Streams. In two of the four size classes.-The ubiquitousness of this palm was shown by the presence/absence survey (Fig. . We examined this relationship two ways. number of species/sample point on five soil types. 8 3.05. Holdridge. Socratea's lowest density occurred on the Streams soils. The intersoil differences. there is a reciprocal relationship between the densities of Iriartea and Socratea (one species' relatively higher density in the first soil compared to the second soil is complemented by the op- posite pattern for the other species). Experimental vs.0001). Within the Residual soils. it is not surprising that its occurrence within La Selva showed no association with topography (Fig. are opposite for the two species. Socratea. Socratea's estimated density did vary among soils (Appendix). Arboleda vs.3% [1-5 m tall] and 50.0000). Iriartea and Socratea also showed reciprocal density patterns between the Residual and Arboleda soils (Table 3). Between the Experimental and Holdridge consociations of the Alluvium. chisquare. Alluvium. both species showed significant density differences in three of the four size classes. CLARK ET AL. Socratea and Iriartea had large significant differences in density between these soils. although the sum of the two species' densities in this size class was remarkably similar between soils. No. In the 1-5 m size class. 63% [1- 5 m]). Its density.0% [10-15 m]. but excluding the Experimental Consociation to remove the effect of harvesting of Iriartea. Arboleda. for example. 52. In all four size classes. Density (stems/ha) Height class (m): Soil I Alluvial consociation Experimental 0 NS 1-5 S 58 NS >5-10 (S + I) (58) NS >10-15 (S + I) (11) NS * >15 (S + I) (17) NS * I 0 ** S 11 ** I 1 13 5 NS S 16 * I 0 10 8 * S 23 * (S + I) (23) NS Holdridge Arboleda Residual 47 17 ** 35 121 *** (82) (138) NS 13 11 NS 0 19 NS (13) (30) NS 4 8 NS (17) (13) NS 5 17 NS (15) (25) NS 84 51 (135) 17 15 (32) 9 7 (16) 16 14 (30) tivity could have been relatively recent.07). P < 0. Local species richness and soil The local species richness of subcanopy/canopy palms varied significantly among soil types. * P < 0. a much smaller proportion of points on the other soils had this many palm species present (Arboleda25%. four size classes. While at least four of the six palm species were present at 68% of sample points on the Streams soils and at 50% of points on the Residual soils. Kruskal-Wallis. Prestoea.01. all seven palm species had similar frequencies of occurrence (Fig. however. 4). 76. number of species per sample point on four soils. df = 4. 5B). P = 0. The relationship between the densities of Socratea exorrhiza (S) and Iriartea deltoidea (I) on related soil types. ** P < 0.23. 4.05. We repeated this analysis with all seven species. 4). 4. In all eight comparisons for these two soil pairs (four size classes.-As for Iriartea. the distribution of this closely related species was not associated with edaphic conditions. Ecology. In all four size classes. The patterns were unchanged (Kruskal-Wallis. None of the other abundant palm species (Euterpe. Welfia was one or more of the four Closest Individuals at 79-90% of the sample points in all four soil types.2588 TABLE DEBORAH A. Socratea's presence/absence patterns did not vary with topographic position (Fig. and Experimental Consociation-17%. NS. Residual). Welfia. Given Welfia's widespread distribution. Alluvium: Holdridge Consociation-19%. particularly in the two smallest size classes (Appendix). the among-soil density differences were significant. P > 0. Welfia was also the only species accounting for 50% or more of the canopy and subcanopy palm stems per hectare in any size class on any soil (percent of stems that were Welfia: Arboleda. Vol. The species was present at 9 1-100% of the sample points in each of the four soil types (Fig. P = 0.
and edaphic variation in density for many species size classes. While very suggestive.-Over the landscape we found highly significant variation in the local (point) species richness of subcanopy and canopy palms. in spite of major intersoil variation in the density of individual species. Kahn and de Castro (1985) found substantially higher palm densities on slope crests than at midslope. All five common species showed marked distributional and/or density changes over the edaphic gradients within La Selva. The density-topography variation of the larger size classes of palms may be due to the favoring of growth into these sizes by more frequent gaps on steeper terrain. The limited additional data from La Selva support these patterns. Topographic variation in tree stem density may be frequent in tropical wet forests. 1994. the total density of stems in this species group was remarkably similar in the four soil types (Fig. S. they speculated that the higher densities at slope crests reflected greater gap formation on these more wind-prone sites. 4. Within three 4-ha forest inventory plots (one each on the Experimental and Holdridge consociations of the Alluvium and one on the Arboleda soil). The limited available soil data suggest that this diversity trend is inversely related to soil fertility. Only for the 1-5 m height class did total density vary significantly among soils. Total palm density. 2].g. For all other size classes. control for several factors that can vary among geographically separated sites (e. Conversely. all five abundant species varied edaphically in density.2-4. In a floristic study at La Selva (J.-The abundance patterns of canopy and subcanopy palms within the La Selva oldgrowth revealed marked topographic variation in this tree guild. unpublished data) Euterpe occurred in plots on both the Arboleda . elevation. Gentry and Ortiz 1993). with intermediate local palm diversity. B. Palm species richness. S. J. definitive analysis of their relation to soil fertility will await more systematic soil sampling. species pool). The lowest local richness of palm species occurs in the Alluvium. Although we as yet can propose no mechanisms that could produce such patterning. unpublished data). the Arboleda soil.-These closely related species showed marked associations with both soils and topography (Euterpe with steep topography and less fertile soils. The soils with highest local palm diversity. unpublished data). DISCUSSION Edaphic factors and forest heterogeneity The variation in soil and topography within this oldgrowth forest has major impacts on the large palms: a mesoscale (hundreds of hectares) gradient in local species diversity. in the 1-5 m size class. Chaverri. Denslow and G. In a Central Amazon forest. the most fertile soil type (Sollins et al. which could produce spurious differences in species richness (palm densities were equivalent across soils in three of the four size classes. Nevertheless. They thus potentially provide interesting support for Huston's hypothesis (contingent on confirmation by more extensive soil nutrient data from La Selva). Chaverri. 1985). This constancy in total abundance of an internally varying species group suggests the operation of some sort of floristic "assembly rules" with respect to the trees in this forest (cf. appears intermediate in fertility between the Alluvium and Residual soils (see Methods).. Similarly.19 [Holdridge]. Denslow and G. however. in a Puerto Rican montane forest. In contrast. On no soil type were there any sample points with fewer than two of the palm species present. Basnet (1992) found that the stem density of trees -10 cm dbh and of saplings 2. stem density was lowest in the Streams soil [Fig. This trend is not confounded by stem density differences. On steep slopes in La Selva old-growth forest height is lower.. his findings involved possibly confounding effects from wide differences in elevation (which affects the species pool. A negative correlation between tree species richness and soil nutrients was found by Huston (1980) in a data set from 46 forest plots distributed around Costa Rica. however. Estimated stem density of the two larger size classes varied twofold among topographic positions. the mean number of large palm species at each sample point was high (4. this potentially important feature of tropical forests merits study. where local species richness was highest). see Methods). Although the intersoil differences in local species diversity are clear. appear to be the two least fertile (Sollins et al.6 of seven species). with lowest densities on the plateaus. On all soils. The patterns we found for the large palms at La Selva. Euterpe occurred at all the higher elevations (3872 m) but was absent from the lowest elevations (3236 m) (Lieberman et al. Euterpe and Prestoea. with highest densities on the slope crests and steeper slopes. He attributed this pattern to lowered dominance by superior competitors in less productive sites. at no sample points in any soil did all the species co-occur. Finally. Streams and Residual. Gentry 1982) and rainfall patterns. and recent gaps (with canopy ?5 m high) are three times more frequent than at other topographic positions (D. topographic changes in the group's overall abundance. one species' absence from a major soil type. 2). 1994. climate.December 1995 TROPICAL PALM COMMUNITY STRUCTURE 2589 Alluvium. Prestoea with gentle topography and more fertile soils). the soil variation at La Selva had little impact on the estimated density of canopy and subcanopy palms.5-10 cm dbh decreased significantly from ridges to slopes to valley positions. Clark et al. Species responses to edaphic variation The patterns of local species richness of large palms demonstrated a high level of species mixing in this guild.
from numerous small plots over hundreds of square kilometres (cf." Human activity is likely to have affected the composition of most neotropical forests (e. Johnston 1992). 0. Given the increasing evidence of anthropogenic impacts within neotropical old growth. Ashton 1969. these species showed reciprocal density variation at all sizes (significant at 1-5 m tall). and Welfia. the distribution of Iriartea revealed a footprint of human activities (Hamburg and Sanford 1986) in forest previously considered "virgin.2590 DEBORAH A. In all size classes. Basnet 1992. Other sites. CLARK ET AL. researchers studying the structure and function of these forests should explicitly seek indications of human activities in their study sites (Hamburg and Sanford 1986). among the other soils all three showed marked density differences (Appendix). In the present study. Hubbell and Foster (1986) found half of the 303 tree and shrub species in a 50-ha plot to be significantly associated with one of four broad topographic site types.-These palm species showed no distributional variation with either topography or soil. edaphic variation in forest composition has been found at all scales: from very local effects (within a 60 x 120 m plot. 1993). Newbery and Proctor 1984. No. and L exorrhiza showed no edaphic associations within a Brazilian Amazonian forest (Kahn and de Castro 1985). Comparative field observations and experiments to assess the factor(s) underlying these species' reciprocal distributions would be very interesting. Phillips et al. Human impact in old-growth tropical rain forest In addition to the now well-documented impacts of edaphic variation. 1994). their densities varied greatly within La Selva. and their summed density was remarkably similar between soils. is likely to have resulted from more recent harvesting by local residents (otherwise. 1985). If Euterpe's distribution reflects lowered competitive ability on more productive sites. and 75% of the stems -10 cm dbh in this forest are nonpalms (Lieberman et al. Kahn and de Castro 1985. 1969). No other species varied in density between these consociations. The distributional anomaly found for Iriartea. Puerto Rico. Socratea and Iriartea would seem to have a low probability of interspecific encounter. Socratea's current abundance is higher on the Experimental soil (Iriartea absent) than on the Holdridge (Iriartea present) (significant in the three larger size classes).g. Such a study would be especially valuable given the widespread human use of Iriartea deltoidea in neotropical forests (Pinard 1993. On the Residual and Arboleda soils. French Guiana. and Welfia were nearly omnipresent in the four soil types. Euterpe was virtually absent from the Alluvium. Garcia-Montiel and Scatena 1994). but not on the two consociations of the Alluvium. As in Asia. these pre-Columbians could have affected the composition of La Selva's current old growth. (1994) in all seven forest types in a lowland Amazonian reserve in Peru. Peres 1994. On the Alluvium." Archaeological work at La Selva (I.. revealed unrecognized spatial heterogeneity in the guild. Similar evidence has come from neotropical wet forests. subtropical Queensland. Lescure and Boulet 1985. Anderson 1990. 1987). When clearing patches of forest they protect Iriartea and Socratea. Williams et al. to small-scale within-forest variation (four 1-ha plots. Iriartea. ter Steege et al. Determining the proximal causes of these edaphic associations will require experimental work and intensive soil analysis. into their "tree gardens. given these developmental differences and their relatively low densities. Although their relation to the Alluvium is affected by human harvesting of Iriartea. Quintanilla. and they often plant these species.BC 300 to AD 300. Ecology. Sarawak. Many cases are known from Southeast Asia. however. to catenas or watersheds on scales of 0. Brunei. the Guaymi Indians practice selective silviculture in forests similar to La Selva (Gordon 1982). Similarly. Guyana. 76. the size-related increases in Prestoea's among-soil density differences suggest continuing mortality effects through regeneration. In Panama. unpublished data cited by Horn and Sanford 1992) revealed habitation sites and pottery shards from . the harvesting of Iriartea from the Experimental Consociation created a natural experiment. Socratea.4 ha. 8 and Residual soils. Interspecific interactions: the case of Iriartea and Socratea The local abundances of these palms (Table 3) suggest some type of interspecific interaction.-There is increasing evidence of the importance of edaphically related spatial heterogeneity within old-growth tropical wet forests. In Panama. Puerto Rico. as well as Welfia. Gomez-Pompa and Kaus 1990. Baillie et al. anthropogenic effects are a relatively little-studied but potentially important source of heterogeneity within old-growth tropical forests. recolonization from neighboring populations should have occurred). Socratea. Even in the smallest (1-5 m) size class. Prestoea showed the opposite pattern. Iriartea deltoidea was found by Phillips et al.5-500 ha (Central Amazon. It is unlikely that direct competition between Euterpe and Prestoea causes their opposite edaphic associations. for example. and led . Neither accounts for >40-46% of the large palms per hectare in any size class on any soil (Appendix). In contrast. Methods for landscape-scale plant community studies The techniques in this study enabled a survey of palm distributions and densities over a large area. such competitive interactions must occur early in development. Possible forms of interaction between the two include competition for pollinators or indirect effects mediated by specific vesicular-arbuscular mycorrhizal fungi or natural enemies. Although Iriartea. Vol. Sarawak.
Tropical forest disturbance: paleoecological records from Darien. and The La Selva GIS and reserve-widegrid were madepossible for by donationsto the Organization TropicalStudies (OTS) from the U. L. Lucas. We also thank OTS for on-going logistic support. New York. we recommend the following approach for studies of plant community variation over large landscapes. however. 1990.and Fred Scatena for their constructivereviews of the manuscript. Turner. Based on this experience. at little additional field effort. and J. Fitzpatrick. and topography) within small-to-large landscapes in these forests. and P. M. 1987. Although the Closest Individual technique produces quantitative density data. 1994. B. to directly link the plant distributions to local site conditions. Panama. J. Anderson. Such unbiased inference is difficult to attain from few samples or plots. This small-plot method would provide more quantitative and more site-controlled data than the techniques we used in the present study.. Grubb. Court. Principes 29:74-78. 1) Distributing many sample points over the major environmental gradients in the study area enables generalization to the landscape. JennyJuarezfor help with GIS mapping. A. Ashton... On the genesis of the soil mantle of the region of Manaus. M. ACKNOWLEDGMENTS We gratefullyacknowledgethe financialsupportprovided (BSR89-18185) and by the U. B. Alternatives to deforestation. K. A. 1994. P. Conclusions The findings from this study and from many other sites indicate that tropical wet forests are characterized by considerable internal heterogeneity in community structure. Boulet. Site characteristics and the distribution of tree species in Mixed Dipterocarp Forest on Tertiary sediments in central Sarawak. New York. USA. and Sun Microsystems. Chauvel. B. 1994). we recommend complete stem inventory and identification within a small. The palm flora of Finca La Selva. J. and L.December 1995 TROPICAL PALM COMMUNITY STRUCTURE 2591 to inferences about underlying factors. This would provide data at each point for each species of interest and would enable analysis of presence/absence patterns and local density. A. Both sampling methods. what factors lead to the opposite distributions of Euterpe and Prestoea? Possibilities include differential impacts of drainage conditions or nutrients on their ability to grow or survive in low light. Simpson. Mellon Foundation. LITERATURE CITED Anderson. Biomass of the North American boreal forest. N.Phil Sollins introduced us to La Selva's soil variability. I. editor. The Presence/ Absence sampling provided data on each species at all sample points. D. Ashton. the Andrew W. Ashton 1969)? Do these forests have relatively more plant species with strong edaphic associations and/or more pronounced edaphic variability than less speciesrich communities? Comparing these properties in temperate and tropical forests could be very revealing. (2) Does high internal spatial heterogeneity promote the exceptional species richness of tropical wet forests (cf. Mellon Foundation. and R. Baillie. and soil characteristics should be taken in each plot. (1) What processes underly the nonuniform distributions of the large palms at La Selva? For example. What processes could produce the reciprocal densities of Socratea and Iriartea? Resolution of these issues will require field and shadehouse experiments (outplantings along soil/topographic gradients. standard-radius circular plot (large enough to contain several stems of the target group) at each sample point. Y. Pages 65-85 in A. however. intersite density comparisons for individual species require equivalent overall density of the total guild between sites. Biological Journal of the Linnean Society 1:155-196. We thank David Ackerly. I. slope. Speciation among tropical forest trees: some deductions in the light of recent evidence. P. Anderson. Tinsley. Ecology 75:1761-1768.Leonel Camposand William Brenes carriedout the field work and data input with skill and quality control. 1969. 1985. G. Additional heterogeneity can derive from local human impacts. R. Carol Augspurger. population structure could be quantified for more abundant species and for the species group as a whole. If not provided by a GIS or other means.Environmental Systems Research Institute. Much of this spatial variation is due to the frequent occurrence of edaphic mosaics (varying soil chemistry and texture. Columbia University Press. B. 1990. P. Biogeochemistry 9:161174. With Closest Individuals evaluated on a species group basis.Inc. . Burslem et al.S. A. especially those not located with strict random sampling (see Botkin and Simpson 1990). 1987. Experientia 43:234-241. Malaysia. S. S. C. Extraction and forest management by rural inhabitants in the Amazon estuary. Goldberg 1982. one abundant species could mask variation in the others (taking data on each species at each sample point would have required seven times the field effort and would have involved data truncation at points where a species is absent or rare). E. Journal of Tropical Ecology 10:579-599. Mineral nutrient status of coastal hill dipterocarp forest and adinandra belukar in Singapore: bioassays of nutrient limitation.S.. M. Combining methods helped resolve these issues and strengthened conclusions when the techniques produced equivalent patterns. Chazdon. 2) To assess community or guild structure. National Science Foundation. have drawbacks. cf. With stem measurement. F R. Journal of Tropical Ecology 3:201-220. Brazil. but within a variable "plot" size (due to greater visibility in more open forest or steeper terrain). NationalScience Foundation the Andrew W. Two challenging questions arise from this study. Burslem. Additional noise in the data could result from edaphic variability within the search area. Effect of topography on the pattern of trees in Tabonuco (Dacryodes excelsa) dominated rain forest in Puerto Rico. Central Amazonia. Basnet. Biotropica 24:31-42. drainage. More complex edaphic effects may produce the among-soil density changes in widespread species such as Welfia. A. Colinvaux. even in old-growth stands. Botkin. and P. R. field data on the topographic position. soil trials with fertilization and watering.. D. We also thank those interviewed about local uses of palms. Bush. 1992.
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Socratea exorrhiza 50.4 13.0001 14.003 0.4 0..0 2.0 0.01 23..9 0.9 0.8 6.6 5..2 0.2 16.9 0.002 0.3 20.7 0.0001 1.0 0.0003 2.7 45.0 .9 NS 11.0 18..2 0.5 6.0 0.6 110.2 4.0 17.5 25..December 1995 TROPICAL PALM COMMUNITY STRUCTURE APPENDIX 2593 Density of the five abundant palm species by size and soil type..0 Prestoea decurrens 5.0000 ..0001 Iriartea deltoidea Soil type Residual Arboleda Stream Alluvium P Consociation Experimental Holdridge P 83.0 4.0 <0..3 14.0002* 0.5 <0. Density (stems/ha) Height class (m): Soil type Residual Arboleda Stream Alluvium P Consociation Experimental Holdridge P Soil type Residual Arboleda Stream Alluvium P Consociation Experimental Holdridge P Soil type Residual Arboleda Stream Alluvium P 1-5 >5-10 Euterpe macrospadix >10-15 >15 45.4 0.3 8..5 NS 15.9 25.0000 14.3 4.2 3.0 10. 29.4 0.6 0..3 25.4 11.6 8.0 12.4 0..02 0.1 0.3 1.0006* 129..9 11.001* 14.9 68.4 0.9 14.6 11..0 NS 6.0 .9 NS .5 0..0 0. across soils.0 0. Probabilities are for chi-square tests of the numbers of Closest Individuals that were and were not a given species. this tests for variation in a given species' density across soils (see Methods).9 121. and Alluvium). .0 NS 0.2 5. 6.0 0.7 34.1 1.8 0.1 31.01 9. .1 28. . Only in the 1-5 m height class did total density vary among soils (see Results). for this size..8 0.0 6.5 NS 5.8 7.7 3.5 2.0 18.4 5..001 0.8 0.2 7.4 18.6 0.7 NS 16. Arboleda.0 <0.9 93..5 2. When total palm density does not differ among soils. .9 17.0 46.5 5.002 16.0001 0. Holdridge).0 0. For the Alluvium. .0 NS Consociation Experimental Holdridge P 57.0 NS 6.8 6. data are presented both for the overall soil type and separately by consociation (Experimental. P* is for a chi-square test (as above) of a species' representation among Closest Individuals on the three soil types of equivalent palm density (Residual. .001* 15.3 1.
Continued.6 14.3 15.9 15.8 16.3 0.3 23.0 199.6 NS .8 13.0 NS 8.0 53.8 245.5 35.2594 APPENDIX.2 319. Ecology.2 NS 15. DEBORAH A.8 21.3 10.8 8.7 25.6 10. 76.0000 21. No.1 NS 14. CLARK ET AL. 8 Density (stems/ha) Height class (m): Soil type Residual Arboleda Stream Alluvium P Consociation Experimental Holdridge P 1-5 >5-10 Welfia georgii 177.02* 404.5 NS >10-15 >15 29.7 NS 8. Vol.8 0.
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