Do Bird Perching Structures Elevate Seed Rain and Seedling Establishment in Abandoned Tropical Pasture?

Karen D. Holl1
Abstract

Lack of seed dispersal has been shown to be a major factor limiting tropical forest recovery in abandoned pasture land. The goal of this work was to determine whether bird perching structures serve to enhance seed dispersal and seedling establishment in an abandoned pasture in Costa Rica. Two types of perching structures (crossbar and branch) were tested. Bird visitation rates were significantly higher on branch than on crossbar perches. The number of animal-dispersed seeds was significantly higher below branch perches than below crossbar perches or in open pasture. Despite differences in seed rain, percent cover of animaldispersed plants and the number of seedlings of animal-dispersed plant species were similar below both perch types and in open pasture. Baiting perches with bananas did not increase either bird visitation rates or seed rain. These results suggest that, although bird perching structures increase seed dispersal, they do not overcome other barriers to tropical forest recovery such as seed predation and low seed germination.
Introduction

lthough global estimates vary widely, it is unquestionable that tropical forests are being cleared at a staggering rate (Myers 1989; Food and Agriculture Or-

1Center

for Conservation Biology, Stanford University, Stanford, CA 94305, U.S.A. Current address: Environmental Studies Department, Natural Sciences II, Room 349, University of California, Santa Cruz, CA 95064, U.S.A.
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ganization 1993). Nowhere is this trend more obvious than Costa Rica, where land covered by forest has dropped from 80% to 25% in the past 50 years (Ramrez & Maldonado 1988; Sader & Joyce 1988). The vast majority of the forest clearing in Costa Rica and Latin America is for agricultural purposes, in particular to create pasture land for cattle grazing (Amelung & Diehl 1992; Fearnside 1993). Extensive deforestation has caused excessive erosion and associated water-quality degradation (Hartshorn 1982; Ramrez & Maldonado 1988). Tropical deforestation also has profound effects on carbon cycling (Houghton 1995), the hydrological cycle (Lean & Warrilow 1989; Shukla et al. 1990), and the conservation of biodiversity (Wilson 1988). In Latin America, agricultural lands are commonly abandoned due to declining productivity and changing economic incentives (Uhl et al. 1988; Aide et al. 1995). This process of clearing and abandonment has led to large areas of highly degraded lands that are often slow to recover, particularly in areas intensely disturbed with agricultural machinery (Buschbacher et al. 1988; Uhl et al. 1988). A number of factors may impede tropical forest recovery in areas of pasture land, including lack of soil nutrients, soil compaction, competition with aggressive, nonnative pasture grasses, seasonal drought, low rates of seed colonization, as well as seed and seedling predation (Uhl 1987; Buschbacher et al. 1988; Nepstad et al. 1991; Aide & Cavelier 1994; Fernandes & Sanford 1995; Nepstad et al. 1996; Holl in press). In many cases the overriding factor impeding forest recovery appears to be lack of dispersal of forest seeds (Nepstad et al. 1991; Aide & Cavelier 1994; Holl in press). Indeed, if seeds of forest species are not dispersed into the pasture, questions of seed predation and competition with pasture grasses become secondary. At the site of the current study, seed rain of animal-dispersed seeds dropped dramatically beyond 5 m of the forest edge (Holl in press). This lack of seed dispersal is likely due to the fact that few birds venture into open pasture areas (Cardoso da Silva et al. 1996). At the current study site, for example, forest bird abundance in the pasture was about one-fifth that in the forest (Sisk 1991). One method for increasing seed dispersal, and thereby facilitating recovery, is the use of artificial bird perching structures. Increased vertical structure may attract birds farther into the pasture, thereby enhancing seed dispersal. Artificial perching structures have successfully encouraged seed dispersal into disturbed temperate ecosystems (McDonnell & Stiles 1983; McDonnell 1986; McClanahan & Wolfe 1987, 1993) but have received little study in tropical rain forest ecosystems. The few studies of bird perching structures to facilitate recovery of tropical rain forest have mostly been less than 1 year in duration (Aide & Cavelier 1994; Ferguson 1995) and have looked only at post and crossbar bird perching struc253

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tures. Determining the utility of bird perching structures for increasing seed rain is of particular interest in tropical forests, where the seeds of 5090% of canopy trees and nearly 100% of shrubs and subcanopy trees have adaptations for animal dispersal (Howe & Smallwood 1982). In addition, passage of seeds through the digestive tracts of birds enhances the rate of germination for many tropical plant species (Ellison et al. 1993). The overall goal of this research was to test whether bird perching structures increased seed dispersal and seedling establishment of forest plants in abandoned pasture land in southern Costa Rica. Specifically, I compared the effect of perch typepost and crossbar versus branchand distance of the perch to the forest edge on bird visitation, seed dispersal, and seedling establishment.
Methods
Study Site

open pasture. Common forest frugivores include a number of species of tanagers, manakins, and pigeons. Many of the common birds in the pasture, such as Crotophaga ani (smooth-billed ani), Tiaris olivaceae (yellow-faced grassquit), and Zonotrichia capensis (rufous-collared sparrow), are predominantly insectivores or granivores. Some frugivores, however, such as Dacnis venusta (scarlet-thighed dacnis), Ramphocelus passerinii (scarlet-rumped tanager), and Thraupis episcopus (blue gray tanager) are regularly observed in the pasture.
Experimental Design

In May 1995, 12 perches (six each of two types) were installed in the pasture. Crossbar perches consisted of a post 6 m tall and 10 10 cm wide; two crossbars 2.2 m long were attached perpendicularly at the top (Fig. 1). Branch perches consisted of a branch of Inga spp., which ranged in height from 5.8 to 7 m (mean 6.2 m) (Fig. 2). The wood for the crossbar perches was ob-

This study was conducted in abandoned pasture adjacent to the Las Alturas Biological Station in southern Costa Rica (8 57 N, 82 50 W, 1500 m elevation). Average annual rainfall is approximately 3000 mm, more than 95% of which normally falls between April and December (Instituto Costarricense de Electricidad, unpublished data). Average annual maximum and minimum temperatures are 24.6 C and 13.2 C, respectively (Instituto Costarricense de Electricidad, unpublished data). The soils are volcanic in origin; previous research at this site (Holl in press) shows that the soils have many of the characteristics of Andisols including a pH over 5.5, high organic matter content, low bulk density, and low available phosphorus, due to high fixation rates. The pasture directly abuts primary forest, forming an abrupt, linear forest-pasture edge. The primary forest is seasonal montane wet forest (Holdridge et al. 1971) and is part of the Las Tablas Protected Area (19,602 ha), which is contiguous with La Amistad Biosphere Reserve. The 5 ha pasture where the study was conducted is part of a mosaic of agricultural land uses covering approximately 2500 ha. The pasture was cleared 25 years ago with heavy machinery. The land was used for 15 years for the cultivation of coffee and for the subsequent 10 years for cattle grazing. Cattle were removed from the pasture in February 1995. At that time, the pasture vegetation consisted predominantly of nonnative grasses such as Axonopus scoparius (Flgge) Kuhlm. (Poaceae), Digitaria decumbens Stent. (Poaceae), and Melinus minutiflora Beauv. (Poaceae). Isolated trees, primarily Inga edulis Mart. (Fabaceae) and Inga punctata (Fabaceae) Willd., are scattered throughout the pasture. A total of approximately 325 bird species has been observed at the site (list available from author), but only a small fraction of these are commonly observed in the
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Figure 1. Crossbar perch with seed traps below. Restoration Ecology

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Figure 3. Schematic drawing of perches and nearby trees. Crossbar perch, square; branch perch, triangle; tree, circle.

Figure 2. Crotophaga ani perching on a branch perch.

and 30 hours during baiting trials (see below). Each pair of perches was observed for a total of 13 hours. Observation periods were distributed between June 1995 and March 1996 to allow for variation in the bird communities during different seasons. All observations were made between 530 and 900 hours. Observation periods lasted between 30 and 60 minutes, and the order in which perches were observed was randomized. Observers recorded the species, time of arrival and departure, direction of arrival and departure, and behaviors for each bird that visited a perch.
Seed Rain

tained from a nearby sawmill. The branch perches were obtained by removing branches from remnant trees in the pasture with a chain saw. Perches were transported and manually installed to a depth of 1 m, so the final height of all perches was approximately 5 m. Three perches of each type were located at approximately 25 m and 250 m from the forest-pasture edge (Fig. 3). Perches of each type were installed in pairs, separated by 10 m. At each distance from the edge, pairs of perches were separated by approximately 100 m. All perches were located a minimum of 10 m from the edge of the canopy of remnant pasture trees. At the time of installation the distance to the nearest tree was measured. Shrub cover in the pasture at the time of perch installation was minimal, as shrubs were regularly cleared by the landowner when the pasture was actively used.
Bird Visitation

Bird visits to perches were recorded for a total of 78 hours: 48 hours when there were no bananas on perches
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Seed rain was measured below perches and in open pasture between 1 June 1995 and 31 May 1996. Two seed traps were placed below each perch; one was placed adjacent to the base of the perch and the second was placed 1 m from the base of the perch below a branch or crossbar. Seed rain for the two traps below each perch was combined for analysis. In addition, seed rain was monitored in six traps in open pasture that were a minimum of 10 m from the perches and remnant trees. Seed traps consisted of fine mesh hardware cloth suspended in an inverted pyramid below 0.5 0.5 m wood frames raised above the ground by 0.5 m legs. Traps were emptied twice monthly. All visible, apparently healthy seeds were separated from leaves and other debris, identified to morphospecies, and counted. Only the number of animal-dispersed seeds is reported here, as the focus of this study was on increasing the number of animal-dispersed seeds. Seeds were identified to the low255

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est possible taxonomic group by collecting flowering and fruiting specimens of plants and comparing them with the collections at the California Academy of Sciences, the Costa Rican National Herbarium, and the La Selva Biological Station. Previous research at this site suggests that seed predation in traps is negligible (Holl in press).
Seedling Establishment

Vegetation measurements were made at the initiation of the study (May 1995), after 1 year (June 1996), and at the end of the study (February 1997). Because the majority of animal-dispersed seeds at this site are dispersed at the beginning of the rainy season (May-July; Holl in press) and germinate soon after dispersal, the final sampling period included 2 years of seedling establishment. Total percent cover and percent cover of individual species were recorded in 1 m2 quadrats adjacent to each seed trap. In addition, the number of seedlings of broadleaved species was counted in a 0.25 m2 quadrat within the larger quadrat. Voucher specimens were deposited at the University of Costa Rica Herbarium.
Baiting Trials

complete block within a split-plot design. The split was distance from the forest edge. At each distance, each replicate was considered a block because each replicate was certainly influenced by minor differences in the surrounding landscape. The effect of distance from the forest edge was tested with the block-within-distance error term. The other main effects and interactions were tested with the residual error term. Because short-term banana hanging trials appeared to have no influence on bird visitation rates and seed rain, the data for the entire year were combined. Relationships between visitation rates, seed rain, and seedling establishment were tested by linear regression. When necessary, variables were log-transformed in order to meet assumptions of homogeneity of variance. Values of p 0.05 are reported as significant throughout.
Results
Bird Visitation

Because few birds were observed visiting perches during the first 2 months of the study, I decided to experiment with baiting perches with fruits to attract frugivorous birds, as in two previous short-term studies (Aide & Cavelier 1994; Press 1996). I used bananas, which are commonly interplanted with coffee in this region to provide shade. A number of tropical bird species have been observed to visit banana feeders (Stiles & Skutch 1989). On 1 August 1995, 10 bananas were hung directly from the top of each perch. Bananas were contained in 2.4 cm square nylon netting to allow birds to eat the bananas but to prevent them from dropping when they ripened. Bananas that had been removed (presumably by mammals) were replaced after 1 week. The experiment was terminated after 2 weeks because no birds were ever observed eating bananas, and most bananas were rapidly removed by mammals. This experiment was repeated between 14 and 28 February 1996, because previous research has suggested that fruit tends to be a more limiting resource during the dry season (Denslow & Moermond 1985). Because treatment differences were confounded temporally, the results of these experiments cannot be tested statistically. Therefore, results are discussed qualitatively.
Statistical Analyses

Ten bird species were observed at branch perches, whereas only three of those species were observed at crossbar perches (Table 1). Most of the species observed at perches are omnivorous, utilizing fruit as part of their diet, and are common in pastures and second-growth vegetation (Table 1). The most common visitor was Crotophaga ani (smooth-billed ani), a predominantly insectivorous bird that also consumes fruit. Visitation rates were very low: 0.41 and 0.12 visits per hour on branch and crossbar perches, respectively. Bird visitation rates were significantly higher for branch than for crossbar perches (F 24.1, p 0.01) and significantly higher at perches farther from the forest edge (F 52.9, p 0.01). Bird visitation rates were not significantly correlated with distance to the nearest tree for either perch type. For 35 birds, the location from which they flew to the perch was observed; their origination points were distributed as follows: 12, remnant pasture trees; 8, shrubs or snags; 7, ground; 4, forest; and 4, fence posts. The average visit length was 3.2 3.8 min. In two cases, birds were observed to defecate while sitting on the perches.
Seed Rain

The effects of distance from the forest edge and perch type on bird visitation rates, seed rain, and seedling establishment were analyzed by means of a randomized
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The number of animal-dispersed seeds was significantly higher below branch perches than in open pasture or below crossbar perches (Fig. 4; F 4.52, p 0.05), although the variance was quite high. There was no significant difference in seed rain at different distances from the forest edge (F 0.21, p 0.05). The number of animal-dispersed seeds was significantly dependent upon the number of bird visits to perches (n 12, F 7.8, r2 0.44, p 0.05).
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Table 1. Number of bird visits to perches during 72 hours of observation.* No. of Visits Scientific Name Common Name Diet Habitat Crossbar Branch

Chlorospingus ophthalmicus Crotophaga ani Geothlypis poliocephala Myiozetes similis Ramphocelus passerinii Thraupis episcopus Tiaris olivacea Tyrannus melancholicus Zimmerius vilissimus Zonotrichia capensis Unidentified Total

common bush-tanager smooth-billed ani gray-crowned yellowthroat social flycatcher scarlet-rumped tanager blue-gray tanager yellow-faced grassquit tropical kingbird mistletoe tyrannulet rufous-collared sparrow

I, F, N I, F I, F I, F F, I F, N, I G, F, I I, F F, I G, I

F, P P P P P, F P P P F, P P

0 1 0 0 0 0 4 1 0 0 3 9

1 12 2 2 1 3 3 2 1 2 3 32

*Dietary and habitat preferences are listed in order of importance as discussed in Stiles and Kutch (1989). Diet: F, frugivore; G, granivore; I, insectivore; N, nectarivore. Habitat: P, pasture; F, forest.

A total of 2, 6, and 13 seed types was identified in open pasture and below crossbar and branch perches, respectively (Table 2). Of the 14 species found below perches, only 6 were from trees, shrubs, or vines commonly observed in the forest. The remainder were from second-growth species or remnant trees already present in the pasture. It is impossible to determine the source of most seeds, however, because a number of species are present in both the forest and pasture. The three most commonly observed seed types included Dendropanax arboreus (L.) Decne & Planch. (Araliaceae), Ficus spp. (fig), and Solanum spp. (nightshade). D. arboreus is a small tree common in the forest. Ficus spp. is a common tree and vine in the forest but also grows on

remnant trees in the pasture. A number of Solanum species are common, early-colonizing shrubs in the pasture.
Seedling Establishment

Figure 4. Total number of seeds dispersed in open pasture and below crossbar and branch perches between 1 June 1995 and 31 May 1996. Error bars represent 1 SD. Means with the same letter are not significantly different (p 0.05), based on Tukeys LSD. Restoration Ecology

Total herbaceous cover, cover of herbaceous plants with animal-dispersed seeds, and the number of seedlings of plants with animal-dispersed seeds did not differ significantly in open pasture and below either perch type at any of the sampling periods (Fig. 5). Cover of plants with animal-dispersed seeds was significantly higher (F 11.0, p 0.05) at sites farther from the forest edge in February 1997, which was due primarily to a single replicate (13% for the one replicate versus a mean of 1% for the other replicates). At the end of the study (2 years after pasture abandonment), the vast majority of cover was still comprised of grasses and other wind-dispersed species such as Elephantopus mollis H. B. K. (Asteraceae) and Heterocondylus vitalbae (D. C.) King & H. Robins (Asteraceae). The number of seedlings of animal-dispersed species per square meter was extremely low and did not differ significantly between habitat types (open pasture, 1.0 1.7; crossbar perch, 2.0 4.0; branch perch, 2.3 1.5; values are means 1 SD). Most of the seedlings of animal-dispersed species were Rubus spp. (blackberry) or Solanum spp., two second-growth shrubs common in the pasture. In the final sampling period, only two of the seedlings of animal-dispersed species recorded were of a forest species, Ocotea whitei Woodson (Lauraceae). The number of animal-dispersed seeds explained a significant amount of the variation in both percent cover of plants with animal-dispersed seeds (n 18, r2 0.23, F 4.8, p 0.05) and the number of seedlings of animaldispersed species (n 18, r2 0.39, F 10.1, p 0.01). Percent grass cover was negatively correlated with cover of plants with animal-dispersed seeds (Spearman corre0.56, p 0.05). lation coefficient, n 18, r
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Table 2. Animal-dispersed seeds recorded in seed traps in open pasture and under crossbar and branch perches.* Number of Seeds/m2 Species Type Location Open Pasture Crossbar Perch Branch Perch

Cecropia polyphlebia Donn. Sm. Dendropanax arboreus (L.) Decne. & Planch. Ficus spp. Heliconia tortuosa Grigg Hyeronima oblonga (Tul.) Mll. Arg. Inga punctata Willd. Ocotea whitei Woodson Paullinia sp. Psychotria goldmanii Standl. Rubus urticaefolius Poir. Sapium sp. Solanum acerifolium Dunal Solanum spp. Strutanthus costaricensis Standl. Unknown

T T T H T T T V S S T S S V

P, F F F, P P, F F P, F F F, P F P P, F P P P

0.8

1.6 (1) 0 0 0 0 0 0 0 0 0 0 0 6.4 (3) 0 0

1.2 0.4

0.4 1.2 8.0 3.2 0.4

2.0 (3) 0 0.8 (1) 0 0 0 0 0.8 (1) 0 2.4 (1) 0 16.3 (1) 5.6 (2) 0 0.8 (1)

3.6 25.6 20.8 0.4 16.0 0.4 0.4 0.8 12.8 2.0 25.2 24.0 14.4 2.4

13.0 (4) 18.3 (6) 40.5 (2) 0.8 (1) 16.7 (4) 0.8 (1) 0.8 (1) 0 1.6 (1) 25.1 (2) 2.7 (2) 53.0 (1) 30.1 (4) 29.4 (1) 4 (2)

*Values are mean number of seeds/m2 1 SD. Values in parentheses are the number of perches out of six for each habitat type below which seeds of each species were recorded. Growth form: H, herb; S, shrub; T, tree; V, vine. Location where a species is found: P, pasture; F, forest. When a plant is found in both habitats, the area where it is most commonly found is listed first.

Baiting Trials

Discussion
Bird Visitation

Overall, visitation rates of birds were nearly identical during observation periods with and without bananas (0.25 visits/hour with bananas; 0.27 visits/hour without bananas). Similarly, the number of seeds dispersed during the 2 weeks preceding banana hanging trials and when bananas were hung were nearly identical (Table 3). An average of only 12.3% of the bananas were left after 1 week during the 4 weeks when bananas were hung. But bananas with holes typical of bird consumption were never observed. Generally, entire bananas were removed. In two cases, Nasua narica (coatis) were observed on perches with bananas.

Whereas a number of bird species were observed on branch perches, visitation rates were relatively low, and few of the species observed were predominantly frugivorous. Few previous studies have quantified the visitation rates of birds to perches. Aide and Cavelier (1994) observed six species of birds at perches baited with plantains in a tropical pasture in Colombia. In the one other study in which visitation rates were quantified, Press (1996) recorded 13.6 visits/hour and 0.2 visits/ hour on crossbar perches baited with bananas and not baited, respectively, in southern Costa Rica. Contrary to the results of Press (1996), bird visitation rates did not increase with baiting in this study, even though most of the same bird species were observed in the two studies. The lack of visitation of birds to bananas is somewhat surprising because a number of tropical birds are commonly observed at banana feeders (Stiles & Skutch 1989). The results of my study and those of Press

Table 3. Total number of animal-dispersed seeds falling below perches with and without bananas during 2-week periods in July and August 1995 and February 1996. July/August 1995 Perch Type No Bananas Bananas February 1996 No Bananas Bananas

Figure 5. Total percent cover (crosshatch) and percent cover of animal-dispersed species (solid) in open pasture and below crossbar and branch perches. Error bars represent 1 SD.

Crossbar Branch

0 8

0 10

3 20

1 19

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(1996) must be interpreted cautiously because both studies were short-term and were confounded by the fact that baiting trials were performed on the same perches that were used for nonbaited measurements, at different times. One possible explanation for these differences may be differences in the surrounding landscape. The pasture where the study of Press (1996) was conducted is located in an extremely fragmented habitat dominated by agricultural land uses, with only small patches of secondary forest nearby. In contrast, the site of the current study is located adjacent to extensive tracts of primary forest. It may be that this forest provides sufficient resources so that birds are not attracted to fruits in the pasture, or that birds are less accustomed to entering the more open pasture habitat. These ideas need further experimentation before conclusions can be drawn. Bird visitation rates were not higher near the forest edge, as might be expected given that birds had to cross less open area to reach nearer perches from the forest; in fact, more birds were observed at perches farther from the forest edge. McClanahan and Wolfe (1987), working on reclaimed phosphate mines in Florida, did not report a difference in bird visitation with distance from the forest edge. In the current study, most of the bird species observed on perches, including the common visitor Crotophaga ani (smooth-billed Ani), are common in the pasture, and few of these species regularly forage in the forest. Only a few birds were observed flying from the forest to the perches, which explains why visitation was higher near the forest edge. There is no obvious explanation for the higher visitation rates farther from the forest edge. There may have been other variables that were not quantified in this study, such as nesting sites, that favored visitation to the farther perches. Interestingly, bird visitation was not correlated with distance to the nearest tree. Remnant trees serve as focal points for bird activity in open pastures (Guevara et al. 1986; K. D. Holl, personal observation). The lack of a significant correlation may have been a result of the small sample size or the small range in distances from the perches to the nearest tree (1434 m). Also, a number of the birds flew to perches from either shrubs, the ground, or fence posts, suggesting that the birds that readily use artificial perches are adapted to open areas and may perch on other anthropogenic structures.
Seed Rain and Seedling Establishment

Results of this and previous studies in both tropical and temperate zones clearly demonstrate that perches increase the number of seeds dispersed (McDonnell & Stiles 1983; McDonnell 1986; McClanahan & Wolfe 1987, 1993; Press 1996; Miriti 1997). The results of the current study also agree with those of McDonnell and Stiles (1983), which demonstrated that branch perches increased
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seed rain more than simpler perching structures. This result would be expected given that bird diversity has been shown to be closely correlated with the structural diversity of vegetation (MacArthur & MacArthur 1961; Karr 1968). If seeds falling in the pasture do not become established, however, bird perching structures will not help to accelerate forest recovery. Few previous studies have investigated the effects of perches on both seed rain and seedling establishment. As in the current study, Miriti (1997) reported that establishment of seedlings with animal-dispersed seeds was not higher below perches in pastures in Brazil. McClanahan and Wolfe (1993) showed a small increase in seedlings of animal-dispersed species below perches on areas that were previously mined for phosphate in Florida. This lack of increase in seedlings despite elevated seed dispersal is likely due to a number of other factors that may inhibit seedling establishment in the pasture. First, seeds in tropical pastures commonly show low rates of germination (Aide & Cavelier 1994; Holl in press) and are subjected to high rates of predation (Janzen 1971; Uhl 1987; Nepstad et al. 1990; Aide & Cavelier 1994; Holl & Lulow 1997). Second, pasture grasses have been suggested as a major factor limiting growth of certain plant species in tropical pastures (Nepstad et al. 1991; Guariguata et al. 1995; Sun & Dickinson 1996). Ferguson (1995) and Miriti (1997) found that seedling establishment of nongrass species below perches was elevated when grasses near perches were killed with herbicides, supporting this idea. Third, seedling survival may be limited by stressful microclimatic conditions, herbivory, and nutrient limitation (Uhl 1987; Nepstad et al. 1991; Aide & Cavelier 1994; Miriti 1997; Holl in press). Although perching structures increase seed dispersal, the number and species richness of seeds falling in the current study was still well below that in the forest (1395 seeds/m2 and 63 seed types; Holl in press). It appears that the increase in seed rain as a result of the provision of perching structures is insufficient to overcome the other numerous barriers to seedling establishment and growth. An additional consideration in evaluating the utility of perching structures in facilitating forest recovery is the composition of seeds that are dispersed and become established. In this and previous studies, most of the seeds dispersed (Aide & Cavelier 1994) and seedlings recorded below perches (Ferguson 1995) were from second-growth species that were already present in the pasture in occasional shrub patches below remnant trees; few of the seeds and almost none of the seedlings were from forest species, which is not surprising given that few birds were observed flying between the forest and pasture. But these second-growth shrubs and trees may play an important role in facilitating recovery (Vieira et al. 1994).
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Existing studies report remarkably similar results of providing bird perching structures in temperate and tropical regions, despite large differences in the avifaunal communities. A higher percentage of plants have fleshy fruits in the tropics, and there are a greater number of frugivores in the region (Howe & Smallwood 1982; Fleming 1991). But plants with animal-dispersed seeds also play an important and often underestimated role in succession in temperate forests (Wheelwright 1988). In both regions, seeds dispersed into open old fields and pastures have to compete with aggressive existing vegetation, which explains their limited growth and survival. Clearly, more studies of bird perching structures in both tropical and temperate zones are necessary before rigorous comparisons can be made.
Management Implications

M. Loik, F. Rein, I. Samuels, and two anonymous reviewers provided helpful comments on earlier drafts of the manuscript.
LITERATURE CITED

Bird perching structures appear to be an ideal means of increasing seed dispersal in disturbed areas given that they are relatively inexpensive to install and require little maintenance. But they do not appear to accelerate early recovery of tropical forest at the site studied. Bird perching structures address only one of the obstacles to recovery; although they serve to increase seed dispersal, they do not overcome other barriers such as nutrient limitation and competition with pasture grasses. Whereas planting seedlings of woody species is a much more expensive undertaking, this practice has been shown to have a substantial effect on forest recovery in areas of abandoned pastures (Montagnini & Sancho 1990; Prinsley 1991; Parrotta 1992; Montagnini et al. 1995). After a few years of growth, young trees not only provide the structure to attract birds and encourage seed dispersal but they also serve to shade out pasture grasses, increase soil nutrients, and modify microclimatic conditions. Research by Vieira et al. (1994) suggests that early successional shrubs may play a similar role. By the end of this study, isolated shrub patches were beginning to become established in the pasture, predominantly from resprouting shrubs that had been cut. Planting or seeding shrubs or trees may provide a more integrated solution to accelerating forest recovery in abandoned pastures.

Acknowledgments

This research was supported in part by a Global Change Distinguished Postdoctoral Fellowship sponsored by the Office of Health and Environmental Research of the U.S. Department of Energy. Additional funding was provided by a grant from the American Philosophical Society and grant DEB-9508683 from the National Science Foundation. I am grateful to M. Lulow, D. Press, and E. Quiros Nietzen for assistance with field work. B. Lindh,
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