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«Brooks, D.M. 2003. The role of size assortment in structuring Neotropical bird communities. Tx. J. Sci. 55: 59-74. THE ROLE OF SIZE ASSORTMENT IN ...»

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Brooks, D.M. 2003. The role of size assortment in structuring Neotropical bird communities. Tx. J. Sci. 55: 59-74.

THE ROLE OF SIZE ASSORTMENT

IN STRUCTURING NEOTROPICAL BIRD COMMUNITIES

Daniel M. Brooks

Houston Museum of Natural Science; Department of Vertebrate Zoology;

One Hermann Circle Dr.; Houston, Texas 77030-1799, USA

Abstract

- I tested confamilial size assortment at three different latitudes, representing a gradient of productivity and stability: the northern subtropics (Rio Grande Valley), the equatorial zone (Amazonian Peru) and the austral subtropics (Paraguayan Chaco). Size assortment is the likely diminished persistence of a species by presence of morphologically similar species; temporally synchronous and spatially sympatric species competing for similar resources should exhibit distinct characters in ecomorphological space, molded over time to reduce the chance of competition. Despite least intensive sampling effort at the Amazon site, it is the most speciose (238 species, 78 common) compared to the Chaco (147, 76) and Rio Grande (61, 24) sites. Size assortment was tested by comparing mean mandibular measurements of confamilials in a real pool against those in a null pool. The pattern of size assortment was pervasive in 68% of the 22 families tested, with most being animal consumers or omnivores, represented by a high percentage of insectivores.

EL PAPEL DE LA VARIEDAD DE TAMAÑO EN LA ESTRUCTURACIÓN DE LAS

COMUNIDADES DE AVES NEOTROPICALES - La variedad del tamaño confamiliar (miembros de la misma familia) fue probada en tres latitudes diferentes representando un gradiente de productividad y estabilidad: el subtrópico septentrional (Valle del Río Grande), la zona ecuatorial (Amazonas peruano) y el subtrópico austral (Chaco paraguayo). La variedad de tamaño es la persistencia disminuida por la presencia de especies morfológicamente similares; especies sincrónicas temporalmente y simpátricas espacialmente que se encuentran compitiendo por recursos similares deben exhibir caracteres distintos en espacio ecomorfológicos, los cuales son moldeados a través del tiempo con el fin de reducir la competición. A pesar del mínimo esfuerzo de muestreo en el sitio amazónico, este es el más especioso (238 especies, 78 comunes) comparado con los sitios del Chaco (147, 76) y Río Grande (61, 24). La variedad de tamaño fue probada al comparar las medidas de la media mandibular de confamiliares en un “real pool” o grupo verdad (grupo actual de especies) contra un “null pool” o grupo posibilidad (especies que potencialmente podrían estar presentes). El patrón en la variedad de tamaño fue penetrante en 68% de las 22 familias estudiadas, siendo la mayoría animales consumidores u omnívoros, representados por un alto porcentaje de insectívoros.

Elton (1927) first proposed the question of whether communities have ‘limited membership’. In his classic work on species assembly, Diamond (1975) suggested that species in a community are selected, and their niches and abundances coadjusted, so that the community possesses “stability”. By stability, Diamond implied that species composition would change only if the physical environment did. Such patterns of coexistence in species assemblages with respect to interspecific competition have been the subject of intensive debate by evolutionary ecologists during the latter half of the 1900’s. Perhaps one of the reasons for the lengthy and intensive debate is that these hypotheses are so difficult to test in tropical, volant bird communities compared to situations involving terrestrial small mammals (Vassallo 1993) for example, where one can more easily measure direct competition by manipulating study subjects (Brooks 1997a).

The role of competition in determining species composition can be tested against null models of comparative regional assemblages. Additionally, evolutionary ecology has a long history of testing competition for resources by examining morphologies of closely related species (Brown & Bowers 1985). Consider the classic case of Darwin’s Galapagos finches, in which a series of empty niches led to rapid evolution of divergent morphologies in colonizing species, permitting coexistence by apportioning empty niche spaces (Darwin 1859, Lack 1947, Grant 1982). Avian competition studies are extrememly important to test hypotheses of species composition processes in volant tropical assemblages (Weiher & Keddy 1999).

The shapes of latitudinal productivity gradients in taxonomic diversity are known for many living organisms, but quantitative data on morphological patterns remain scarce (Roy & Foote 1997). Although some studies have addressed whether communities in similar environments will converge ecomorphologically (Ricklefs & Miles 1994), these studies took place in vegetative communities at opposite ends of the globe, such as California and Chile (Cody 1974, Kelt et al. 1996). Moreover, such studies compare temperate environments or temperate versus tropical environments, but have not addressed how such patterns may vary among avian communities across various sites within the Neotropics. My objective herein is to examine the role of size assortment in structuring Neotropical bird assemblages at three latitudes in the Neotropics, representing a gradient of productivity and seasonality.





Grant (1969) recognized that character displacement could arise as a result of: 1) divergent natural selection of coexisting species, or 2) selective survival of immigrating species as a function of distinctness from species already present. It appears that the first pattern evolved into size adjustment, whereas the latter became size assortment (Case & Sidell 1983). One way to investigate fine-scale tuning among interacting species is to look at variation in related taxa because ecological similarity could result in morphological divergence resulting from dietary differences (Karr & James 1975). Size assortment is the likely diminished persistence of a species by presence of morphologically similar species (Case & Sidell 1983). In other words, from a random pool of species that could potentially colonize an area, the species actually present are those that could coexist because they were different sizes and therefore did not compete for resources. The reason for species not persisting within the community has been debated. The two main theories are that similar-sized species: 1) became locally extinct (Willig & Moulton 1989) or 2) could not colonize a region due to intense competition from a similar-sized species that shared the same resources (Diamond 1975).

I will test size assortment by comparing bill morphologies of confamilials that utilize similar resources (same habitat, similar diet, spatially and temporally sympatric), examining whether ecomorphological distances in a real community are overdispersed compared to a null community. Such phylogenetically related confamilials have similar environmentally constrained phenotypes and therefore are good candidates for testing hypotheses of interspecific competition (Ricklefs & Miles 1994). All taxa that may coexist through other mechanisms (habitat allotopy, temporal asynchrony, dietary differences, asynchronous activity patterns, social mimicry or species packing) will be filtered from analyses, insuring that complete competitiors are used in analyses. I will compare size assortment in the northern subtropics (Rio Grande River Valley), the equatorial tropics (northern Peruvian Amazon), and the austral subtropics (central Paraguayan Chaco). Analysis of pervasiveness of size assortment in the Neotropics will help provide insight of underlying processes that govern these patterns, and therefore mold the communities themselves. My specific hypothesis is: size assortment is equally prevalent among different latitudinal gradients.

METHODS

Methods are provided in detail in Brooks (1998), but are summarized below. I collected data along transects in the field to determine community composition at each of the three sites (Rio Grande - 26015’N; 98030’W, Amazonia ~ 2045’S; 72055’W, and Chaco S; 60030’W), and also to study the ecology of species within those communities;

this was important to designate which species could be used in analyses. From the comprehensive species lists, I designated confamilials for size assortment analyses.

Although competition could occur among more distantly related taxa, my objective is to examine competition in species with similar morphologies; hence comparisons at the confamilial level were used.

Accounting for other factors that may enhance coexistence a-priori will increase the chance that species selected to test hypotheses are (or were) competitors. Therefore, upon determination of confamilial groups, I excluded species from analyses that might coexist through mechanisms other than size differences. Ecological factors controlled for include species used in analyses being common (at least 2 individuals/sampling duration, for at least one-third of all durations), temporally synchronous seasonally, spatially sympatric (in terms of habitat use), utilizing similar dietary resources, and temporally synchronous daily (sharing activity periods). Additionally, evolutionary factors to be controlled for include: species used in analyses must not be constrained by social mimicry (two more distantly related species appearing more chromatically similar than other, more closely related species), phylogeny (restricted to familial level, see above), or be candidates for species packing (reverse pattern of size assortment - real pool pool underdispersed when compared to null pool). While species filtered from analyses may have been morphologically structured within their communities, I could not test my hypotheses without analyzing patterns in fully competing species.

I considered competing species remaining after filtering as the real pool, and I constructed regional null pools of potentially colonizing species to compare against real pools for analyses. A null pool is a pooled assemblage occurring within a designated geographic region that shares characteristics (residents or semi-residents, sharing similar habitat/diet/activity patterns, and lacking potential for social mimicry) with actual species from the study site (real pool) (slightly modified from Ricklefs 1987). Null pools are used to test for structured patterns in bird communities, by comparing the real pool against the null pool. I selected the geographic region for each null pool by drawing a concentric circle from the study site using a radius of approximately 333 km. I selected this radius based upon its relative capacity to encircle biomes at country-wide scales for the South American sites (the Peruvian Napo-intersect region, the Paraguayan Chaco), whereas the Rio Grande site encompassed the Texas Rio Grande Valley (south of Corpus Christi) and portions of two northeastern states in Mexico (central and eastern Nuevo Leon, and north and central Tamaulipas). I obtained the source pools and data on habitat association, migrant status, and other ecological attributes by sampling bird communities at nearby localities, supplemented with available regional avifauna references. These data from other sites, complemented with references, provide complete and accurate species lists for comparison, insuring availability of an adequate pool for comparison (Willig & Moulton 1989). I did not analyze communities where null and real pools contained the same species, because lack of species selection in the real pool is a rare occurrence, resulting in uncomparable species pools (Willig & Moulton 1989).

I obtained measurements from museum specimens for species comprising the null and real pools. I analyzed the data using graphic analyses, by plotting mean bill height against mean bill width of species in two-dimensional space, and comparing the segment lengths between species comprising real and null pools.

I used shortest spanning trees (hereafter referred to as SSTs), as used by Ricklefs & Travis (1980) following the idea of a Prim Network (Prim 1957). An SST represents the shortest composite line that connects all congeners in a guild and contains one fewer line segments than the number of species in the community (if N-species in a guild, the SST has N - 1 line segments). Each individual line segment between species measures nearest neighbor distance (hereafter referred to as NND) between species. Measurement of segment lengths (NNDs) between related taxa provides an index of ecomorphological distance between species (Brooks 1997a).

I plotted the ecomorpholocial means between species in each community on twodimensional graphs using the computer program SPSS (SPSS 1996). Using a statitical package to create graphs permits standardization of axes for each SST across sites, insuring that measurements among separate taxa and sites would not be biased. Using the two-dimensional graph plots, I measured NNDs from real pool and null pool SSTs.

I tested size assortment by comparing mean NNDs of the real and null pool assemblages in cases where the real pool was hyperdispersed to the null (Strong et al.

1979, Willig & Moulton 1989). If mean NNDs of the real species are hyperdispersed in ecomorphological space compared to the null pool this indicates that size assortment governs these assemblages (Willig & Moulton 1989). A graphic depiction is provided in Figure 1, where mean NND length of the null pool (n = 8 NNDs) = 18.7 (21 + 29.5 + 25.5 + 19 + 20 + 12 + 5.5 + 17 / 8) and mean NND length of the actual species assemblage (n = 2 NNDs) = 36.8 (56.5 + 17 / 2). In this example the assemblage is significantly overdispersed from the null pool (36.8 versus 18.7, respectively); therefore size assortment structures this icterid assemblage at the Amazon site.

I applied the goodness-of-fit chi-square test (Sokal & Rohlf 1969, Ricklefs & Travis 1980, Case & Sidell 1983) to test for significance (P 0.05) between NND means of real and null confamial groups.

RESULTS



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