Breeding and foraging variation of the plush-crested jay (Cyanocorax chrysops) in the Brazilian Atlantic Forest.
|Abstract:||We monitored six flocks and five active nests of the Plush-crested Jay (Cyanocorax chrysops) at three sites in the Atlantic Forest in southern Brazil. The sites had different vegetation compositions and spanned different levels of anthropogenic disturbance. Home range size in fall/winter was 20-30 ha and the breeding territory size in spring/summer was 5-10 ha in size. Territories were smaller across sites with higher anthropogenic food supplementation. Flock sizes were 5-11 individuals during spring/summer and 8-15 individuals during fall/winter. The Plush-crested Jay is a cooperative breeder, nests were 4-7 m above ground level, the incubation period was 18-20 days, brood size ([bar.x] [+ or -] SD) was 3.4 [+ or -] 0.80 eggs per nest, and nestlings fledged 23 [+ or -] 1.26 days after hatching. This species occupies all forest strata but tends to use the understory and middle levels most (G = 178.2; P < 0.01). Invertebrates were the most frequently consumed item in all areas, but percent consumption varied among sites (G = 105.06; P < 0.01). We observed 110 food caching events throughout the year, primarily seeds of Araucaria angustifolia, maize, and coconuts. Caches were on the ground (n = 40), in epiphytes (n = 47), and on branches (n = 23). Levels of anthropogenic food supplementation resulted in variation in territory and home-range size, nestling survival rates, strata occupation, and diet composition of the Plush-crested Jay.|
Jays (Physiological aspects)
Jays (Environmental aspects)
Uejima, Angelica Maria Kazue
Boesing, Andrea Larissa
Anjos, Luiz Dos
|Publication:||Name: The Wilson Journal of Ornithology Publisher: Wilson Ornithological Society Audience: Academic Format: Magazine/Journal Subject: Biological sciences Copyright: COPYRIGHT 2012 Wilson Ornithological Society ISSN: 1559-4491|
|Issue:||Date: March, 2012 Source Volume: 124 Source Issue: 1|
|Topic:||Event Code: 310 Science & research|
|Geographic:||Geographic Scope: Brazil Geographic Code: 3BRAZ Brazil|
The family Corvidae is represented in South America by the genera
Cyanocorax and Cyanolyca (Madge and Burn 1994). These genera occur
across the neotropical region and include 17 species of Cyanocorax and
nine species of Cyanolyca (Gill and Donsker 2011). The Plush-crested Jay
(Cyanocorax chrysops) is widely distributed throughout South America
from Amazonia to northeastern Argentina occupying a variety of habitats
that include several types of forests: mixed rainforest, semi-deciduous
forest, Cerrado (one of the largest tropical savannas of the world)
(Silva and Bates 2002), scrub, and also semi-urban areas. It is not
usually found in the interior of primary forest, but tends to be more
frequent at the borders and in clearings (Goodwin 1976, Anjos 2009). It
is poorly studied, although relatively common and conspicuous in its
range. Only one species-specific study has been published on habitat
occupancy and vocal repertoire (Brunetta and Anjos 2010).
The objectives of our study were to describe the breeding and foraging behaviors of the Plush-crested Jay at three sites in the Atlantic Forest in southern Brazil. The sites studied each have different vegetation structure and anthropogenic food supplementation.
Study Areas.--Field observations were conducted at three sites in Parana State in southern Brazil (Fig. 1). Locations and natural features of the three areas varied (Table 1). The first site, Vila Velha State Park (VVP), was mostly natural open fields including grassland/wetland and undisturbed mixed rainforest (MF) beside an administrative area of the Park. The sample area included forest merged with a rocky outcrop and grassland. There was one small restaurant open for visitors to the VVP within this sample area. The second area was Klabin Ecological Park (KEP), which was mostly covered by MF with the remaining area grassland, tree plantations for timber production, and administrative areas of the Park. The area sampled was represented by forest with a few patches of grassland beside the administrative area, which included an animal breeding center (7 ha). The third area was the Ribeirao do Tigre Farm (RTF) where well-preserved seasonal semi-deciduous forest (SF) merged with disturbed areas used for agriculture and livestock grazing.
General Observations.--Field observations were made 6 days per month in each area from October 1995 to December 1996. We used 8 x 35 binoculars for observations that began at sunrise and lasted all day. Field observations were conducted during 1,800 hrs. The daily procedure was to walk along previously established transects to find one flock, and to follow this flock throughout the day. Particular methods were used to document breeding and foraging behaviors. The birds were not banded but we were able to distinguish different flocks (at least 2) at each site as American jays maintain an area of dominance around active nests (Anjos et al. 2009). We followed Gill and Donsker (2011) for nomenclature of birds and the Missouri Botanical Garden (2010) for nomenclature of vegetation.
[FIGURE 1 OMITTED]
Flock Size, Home Range, and Territory.--We recorded the number of individuals in the flocks in the different sites and periods. We separated the year into two periods: the first when aggressive territorial behavior was observed between different flocks (the territorial period), and the second when aggressive behavior was not observed (the home-range period). We plotted each observation of Plush-crested Jay flocks on each transect onto a map (Luginbuhl et al. 2001) for both territorial and home-range estimates. We used these observations to calculate the size of the overall territory and home range of the jays using the minimum convex polygon method (MCP) (Mohr 1947, Odum and Kuenzler 1955). This consists of joining the outermost observation points for each flock with a straight line. The largest polygon obtained was taken as the size of the territory of a flock. This procedure was chosen due to its simplicity and wide use in ornithology (e.g., Jullien and Thiollay 1998, Wiktander et al. 2001, Ribeiro et al. 2002, Duca et al. 2006).
Reproduction.--Reproductive activities were observed during ~720 hrs of field observations (240 hrs/area). We recorded the number of eggs laid and hatching data, and monitored the development of nestlings. Nestlings were measured the second week post-hatching. A nest was considered depredated when the entire brood disappeared or when there was no evidence inside or underneath the nest of any other kind of loss. Activities related to incubation, provisioning of the brooding bird or nestlings, and cleaning of the nest were also recorded. Two fledglings were color banded. The nests were measured (height, diameter, brood chamber diameter and diameter of twigs used in its construction) after nesting activities had ceased, collected, and deposited at the Capao da Imbuia Museum of Natural History (Curitiba, Brazil) and the Klabin Ecological Park Museum (Telemaco Borba, Brazil).
Foraging.--We made field observations of foraging behavior during 1,080 hrs (360 hrs/area). We used focal animal sampling following Altmann (1974) with 5-min breaks between observation of different individuals or flocks (scan-sampling). This procedure should have increased data independence. The food item, capture substrate, and stratum were recorded for each foraging event of an individual jay. We considered three levels of forest strata, besides the ground: undergrowth (up to 2 m above the ground), middle level (from 2 to 7 m), and subcanopy (from 7 m above the ground to 1 m below the canopy). The substrates were: ground, branches, leaves or epiphytes, and air for aerial foraging.
Statistical Analyses.--The R X C test for independence (G-test) was used to evaluate the significance (P = 0.01) of proportions of events related to foraging behaviors among the three sites (Fowler and Cohen 1986).
Flock Sizes, Home Range, and Territory.--Territorial behavior was first observed in early October (spring). The size ([bar.x] [+ or -] SD) of flocks during this period was smaller at VVP (6 [+ or -] 0.8 individuals) and at RTF (5 [+ or -] 0.2 individuals) than at KEP (11 [+ or -] 0.6 individuals). Territories were smaller at VVP (5 [+ or -] 0.3 ha) and KEP (5 [+ or -] 0.7 ha) than at RTF (10 [+ or -] 0.4 ha; Fig. 2). Territories at KEP and RFT did not overlap and were separated by 350 and 500 m, respectively. However, overlapping (~ 20%) of territories was detected at VVP. The area of greatest anthropogenic food supplementation had the greatest overlap at VVP, and aggressive territorial behavior was frequently observed at this site. Aggressive territorial behavior between individuals started with "bobbing", (described by Hardy 1961), followed by alarm calls. Direct contact between individuals was recorded when the dominant individual touched the back of an opponent after sweeping flights. At times, the opponent persisted and the two individuals attacked each other with their feet while the remainder of the individuals in the flock stayed close, emitting alarm calls. Territorial behavior was relaxed throughout the end of summer (Mar). Flock sizes were largest in the home-range period (Apr-Sep, mostly fall/winter) at KEP (15 [+ or -] 1.92 individuals), followed by RTF (11 [+ or -] 1.48 individuals), and VVP (8 [+ or -] 1.63 individuals). Home ranges were smaller at VVP (20 [+ or -] 1.34 ha) than at KEP (30 [+ or -] 0.89 ha) and RTF (30 [+ or -] 1.04 ha; Fig. 2). Overlapping of home ranges (where aggressive behavior was not recorded among individuals of different flocks) was ~25% in all areas.
Reproduction.--Five active nests were found, three at VVP and two at KEP. The first active nest was found in mid-October at VVP. Nests were placed closer to the animal breeding center at KEP and to the restaurant at VVP. Only one nest was successful at VVP. The other two nests found in this area were depredated several times (n =4) after hatching. Nests were reconstructed (each 18 to 21 days after depredation) into February, but none was successful. Two new nests were 50 m from the previous nests and the other two were in trees that were closer, but no new nests were found in the same tree in which a nest was previously depredated.
Nests (n = 5) were placed in a fork between 4 and 7 m above the ground in the middle stratum of the forest, and were rounded and consisted of a platform with a brood chamber in the center. They were built with sticks up to 3 mm in diameter, and the brood chamber was lined with twigs < 1 mm in diameter. Nests measured 151 [+ or -] 31.27 mm in height and 295.4 [+ or -] 24.33 mm in diameter. The brood chamber measured 162 [+ or -] 12.07 mm in internal diameter and 99.4 [+ or -] 13.88 mm in depth (Table 2).
[FIGURE 2 OMITTED]
The eggs (3.4 [+ or -] 0.80 eggs/nest) were spotted or blotched with dark brown concentrated on the rhombic pole and less scattered in the center on a cream-colored background (Fig. 3). Three, four, and two nestlings were recorded in the three non-depredated nests. The newly hatched chicks had red-orange skin, and were almost black around the eyes, wings, and spine. The bill was completely yellow, especially the mandible. Measurements of nestlings (n = 9) 2 weeks post-hatching were 115 [+ or -] 12.47 g, tarsus = 37.9 [+ or -] 1.46 mm, culmen = 18.4 [+ or -] 1.53 mm, nostrils = 11.9 [+ or -] 0.99 mm, and wing length = 90.8 [+ or -] 10.90 mm (Table 3).
Plush-crested Jays are cooperative breeders with auxiliaries at the nest (helpers). The incubation period was between 18 and 20 days. Individuals took turns in the nest during incubation and offspring care. Food was brought to the brooding bird by helpers in the flock after constant calling for food. The helpers either directly delivered food to the nestlings or to the adult in the nest; this individual received the food and redistributed it to the nestlings. The average number of visits to provision nestlings varied throughout their development: the average was five visits/hr in the first week; seven visits/hr in the second week, and nine visits/hr in the third week. The average visitation rate was the same regardless of brood size. Helpers were recorded removing fecal sacs from the nest and providing defense against predators in addition to helping provision the nestlings. Up to six helpers were clearly observed defending a nest against tufted capuchins (Cebus appela), which were kept far from the nest.
[FIGURE 3 OMITTED]
Nestlings left the nest ~23 [+ or -] 1.26 days after hatching and, for the next 20 days, they solely depended on food provided by the parents and helpers. Fledglings began feeding by themselves after 25 days and were completely independent at 90 days. The two color banded fledglings marked in 2005 were observed helping in 2006.
Foraging.--We observed 3,508 foraging events (1,209 at VVP; 1,700 at KEP; and 599 at RTF; Table 4). The capture technique used by Plush-crested Jays was 'gleaning' (described by Robinson and Holmes 1982). The diet (recorded in all 3,508 events) included invertebrates (spiders, millipedes, beetles, caterpillars, grasshoppers, and mollusks), fruit, many kinds of foodstuffs given or discarded by humans (garbage, bread, meat, cookies, grain plants, and maize), and vertebrates (frogs, eggs, and nestlings, and small reptiles). Invertebrates were the most commonly used food resource at all three areas, but the percentages of each item varied (G = 105.06; df = 6; P < 0.01; Table 4). Food resulting from human activities was used more at VVP and KEP than at RTF where the most commonly consumed items were insects and invertebrates.
Plush-crested Jays fed on fruit from 24 plant species, but only the fruits of Syagrus romanzoffiana were commonly eaten in all three areas (Table 5). A greater variety of fruit was observed to be consumed at KEP (13 species), while 10 and five fruit species were recorded at VVP and RTF, respectively. The fruits most frequently recorded were: Eriobotrya japonica, Diospyros kaki, Philodendron spp., Casearia sylvestris, Rapanea ferruginea, Syagrus romanzoffianum, and Ficus enormis.
The foraging forest stratum and capture substrate were recorded at 2,980 foraging events (964 VVP, 1,457 KEP, 559 RTF; Table 6). The proportion of observations related to foraging stratum differed among the areas studied (G = 178.2; df = 6; P < 0.01). The understory and the middle level were the most frequently used strata in all areas, but the ground and the subcanopy were proportionally used more at VVP and RTF, respectively.
The proportions of use of the capture substrates also differed among the three areas (G = 80.84; df = 8; P < 0.01). Branches and leaves were generally the most frequently used substrates in the understory and the middle level in all areas (G = 24.18; df = 6; P < 0.01); epiphytes were less used, particularly in the subcanopy, and the air was the least used.
Food caching behavior was also observed and recorded (n = 110). Each individual had its own cache up to 100 m from where the food was captured. Each cache contained only one unit of a given item (except for maize, for which each cache could contain up to 12 seeds). Leaves or small sticks were laid over the food cache. We did not observe individuals defending their caches. Caching locations were in the ground (n = 40), in epiphytes (n = 47), and on branches (n = 23). The caches mainly contained seeds of A. angustifolia, bones, maize, and seeds of S. romanzoffianum). The number of foods cached did not differ between seasons (G = 2.06; df = 3; P > 0.01).
The larger size of the territory at RTF than at VVP and KEP was probably related to the amount of supplementary food at the two latter sites. Larger territory sizes at RTF may compensate for areas with lower food availability. Marzluff and Neatherlin (2006) observed that crows and ravens (Corvus spp.) had smaller home ranges and higher numbers of fledglings near settlements. However, we found larger home ranges at KEP than at VVP, although flock size was similar. This may be due to higher predation rates at VVP (lower fledgling production) than at KEP, allowing a smaller home range. Territory size expanded at all sites during the summer (Fig. 2), probably due to addition of fledglings to flocks or possible variation in food abundance.
Most of the available data published on Cyanocorax species indicate communal breeding with an active nest in each group and helpers in the nest during the nesting period and while the young are dependent (Anjos et al. 2009). We documented this system in our study of the Plush-crested Jay. Alternation among individuals was observed during the incubation period, unlike for most Cyanocorax jays, where only one individual incubates (e.g., Inca Jay [C. yncas]; Alvarez 1975). This behavior appears similar to that of the Curl-crested Jay (C. cristatellus), where short substitutions were observed (Amaral and Macedo 2003). Two individuals were observed incubating for Azure Jays (C. caeruleus), at times with relief (Boesing and Anjos, In Press). Brooding Plush-crested Jays were observed being fed by several individuals of the flock, as reported for most Cyanocorax jays (e.g., Moore 1938, Crossin 1967). The type of vocalization (pair call) before feeding has also been recorded for other Cyanocorax species: Curl-crested Jay (Amaral and Macedo 2003), Tufted Jay (C. dickeyi) (Moore 1938, Skutch 1987), and Azure Jay (Anjos and Vielliard 1993; Boesing and Anjos, In Press).
Feeding of both nestlings and the brooding bird by helpers is common in several species of Cyanocorax jays (e.g., Bosque and Molina 2002, Amaral and Macedo 2003). Breeders assisted by helpers were observed to fledge more young in the case of the Florida Scrub Jay (Aphelocoma coerulescens) (Woolfenden and Fitzpatrick 1984), a cooperative jay in North America. We observed possible predation of nestlings and fledglings by hawks (Roadside Hawk [Buteo magnirostris] and Yellow-headed Caracara [Milvago chimachima]), marsupials (e.g., Didelphis spp.), and reptiles (e.g., Tupinambis spp.). One South American coati (Nasua nasua) was recorded preying on a fledgling on the ground. The most common cause of the lack of reproductive success in nests of corvids is predation (Marzluff 1985).
Neotropical jays occupy different forest strata, but preferences vary among species (Anjos et al. 2009). The Plush-crested Jay frequents all forest strata, but seems to prefer the understory and middle level of the forest, despite frequently using the ground, where food is available (VVP and KEP). All available layers, including the ground, are used by Inca Jays (Alvarez 1975) and Purplish-backed Jays (C. beecheii) (Raitt and Hardy 1979). Azure Jays often only frequent the canopy of the forest (Anjos 1991). The availability of food at VVP favors use of resources on the ground, whereas the density of vegetation promotes the use of higher strata at RTF and KEP.
Caching behaviors are widespread throughout the Corvidae. Food storage is commonly observed when birds have more food available than they are able or willing to eat at once, especially if the food is of a kind that will remain edible for some time (Goodwin 1976). This behavior is poorly documented in Cyanocorax jays. Caching behavior was observed in captivity by the Tufted Jay, but it does not appear to have been observed in the wild (Crossin 1967). Anjos (1991) observed this behavior in Azure Jays in both captivity and in the wild; this species was observed storing seeds of Araucaria angustifolia in the canopy of the MF (Anjos 1991). The Plush-crested Jay caches the seed of A. angustifolia on the ground, and it should be considered as an incidental disperser of this tree species. Some seedlings of A. angustifolia were recorded in places, around the animal breeding center at KEP, where individual Plush-crested Jays were observed caching seeds of this tree species.
The Plush-crested Jay exhibited variations in: (1) forest strata occupancy (preference for the understory and middle level of the forest, frequently using the ground where there was human food supplementation); (2) territory and home range sizes (both smaller when food supplementation was available); and (3) foraging behaviors (more diverse in less disturbed habitats). These characteristics demonstrate the plasticity of this species in habitat use, similar to most species of Corvidae.
We are grateful for logistical support in Vila Velha State Park, Klabin Ecological Park, and Ribeirao do Tigre Farm. Personal of the Environmental Police of Parana State also assisted us during field work. Plants were identified by M. C. Dias (State University of Londrina). Financial support for field work was provided by the Federal University of Parana (UFPR). Pedro Sherer-Neto, L. C. Milleo, and F. C. Straube made important suggestions during the field work and revised an earlier version of the manuscript, which was presented as a Master Thesis at UFPR. We appreciate the valuable suggestions by J. M. Marzluff, Tom Webber, and C. E. Braun, which improved the last version of this manuscript. The authors received research grants from CNPq (Brazilian Council for Development of Science and Technology) to the first and third authors, and from CAPES (Coordination for the Improvement of Higher Level Personnel, DS), to the second author.
Received 6 February 2011. Accepted 29 August 2011.
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ANGELICA MARIA KAZUE UEJIMA, (1,3) ANDREA LARISSA BOESING, (2,4) AND LUIZ DOS ANJOS (2)
(1) Programa de Pos-Graduacao em Zoologia, Universidade Federal do Parana (UFPR), Postal Code 81531-990, Curitiba, Parana, Brazil.
(2) Departamento de Biologia Animal e Vegetal-Universidade Estadual de Londrina, Rodovia Celso Garcia Cid, Campus Universitfirio, Postal Code 86050-980, Londrina, Parana, Brazil.
(3) Current adress: Universidade Federal de Pernambuco, Centro Academico de Vitoria, Rua Alto do Reservatorio, Postal Code 55600-000, Vitoria de Santo Antao, Pernambuco, Brazil.
(4) Corresponding author; e-mail: firstname.lastname@example.org
TABLE 1. Characteristics of areas where Plush-crested Jays were monitored in the Atlantic Forest, southern Brazil: Vila Velha State Park (VVP), Klabin Ecological Park (KEP), and Ribeirao do Tigre Farm (RTF). VVP KEP Vegetation type * Mixed rainforest (MF) No. tree species ** 350 species; 13% endemic Dominant tree species Araucaria angustifolia, Ilex paraguariensis, Camponesia xanthocarpa, Ocotea porosa, O. odorifera Climate * Cfb Average temperature * 18 [degrees]C 19.5[degrees]C Average rainfall * 1,550 mm 1,700 mm Coordinates 25[degrees] 15' 24[degrees]17'S, S, 50[degrees] 50[degrees]35' W 05'W Total area 3,790 ha 11,196 ha Sample area 150 ha 140 ha Forest cover 23% 60% Grassland 60% 30% Human built 17% 10% Agriculture RTF Vegetation type * Seasonal semi-deciduous forest (SF) No. tree species ** 220 species; 10% endemic Dominant tree species Aspidosperma polyneurum, Cedrella fissilis, Balfourodendron riedelianum Climate * Cfa Average temperature * 21[degrees]C Average rainfall * 1,600 mm Coordinates 23[degrees]27' S, 51[degrees] 15'W Total area 1,285 ha Sample area 150 ha Forest cover 20% Grassland 20% Human built Agriculture 60% * Mendonca and Danni-Oliveira 2002; ** Oliveira and Rotta 1982; Cbf = subtropical humid with warm summer to mildly warm; Cfa = subtropical humid with warm summer. TABLE 2. Plush-crested Jay nest measurements (mm) in the Atlantic Forest, southern Brazil: Klabin Ecological Park (KEP) and Vila Velha State Park (VVP). Nest # Nest Nest Brood Brood height diameter chamber chamber height diameter 1 KEP 152 287 99 180 2 VVP 180 282 120 170 3 VVP 190 275 108 162 4 KEP 125 290 90 150 5 KEP 108 343 80 148 Mean 152 295.4 99.4 162 [+ or -] [+ or -] [+ or -] [+ or -] [+ or -] SD 31.27 24.33 13.88 12.07 TABLE 3. Measurements of 2-week-old Plush-crested Jay offspring (mm, g) in southern Brazil: Klabin Ecological Park (KEP) and Vila Velha State Park (VVP). Offspring Tarsus Culmen Nostril/Bill Wing Mass 1 KEP 38.8 20.2 12.2 92 140 2 KEP 36.4 18.2 11 80 115 3 KEP 36.5 19.4 12.7 89.3 120 4 KEP 39 20.6 12.1 89 130 5 VVP 38 15.2 12 110 100 6 VVP 36 19 14.1 110 100 7 KEP 39.1 17.6 10.7 82.4 8 KEP 36.8 17.5 11.3 82.4 9 KEP 40.5 18.2 11.1 82.5 Mean 37.9 18.4 11.9 90.8 115 [+ or -] [+ or -] [+ or -] [+ or -] [+ or -] [+ or -] SD 1.46 1.53 0.99 10.90 12.47 TABLE 4. Observed frequency (percentage) of foods consumed by adult Plush-crested Jays in southern Brazil: Vila Velha State Park (VVP), Klabin Ecological Park (KEP), and Ribeirao do Tigre Farm (RTF). Food item VVP KEP RTF Total Insects/ invertebrates 964 (80) 1,457 (86) 559 (93) 2,980 (84.9) External resources * 134 (11) 103 (6) 21 (4) 258 (7.4) Fruit 31 (3) 89 (5) 11 (2) 131 (3.7) Vertebrates 80 (7) 51 (3) 8 (1) 139 (4) Totals 1,209 (100) 1,700 (100) 599 (100) 3,508 (100) * Garbage, maize seeds. TABLE 5. Plant species used by Plush-crested Jays and relative frequencies of fruits consumed in southern Brazil: Vila Velha State Park (VVP), Klabin Ecological Park (KEP), and Ribeirao do Tigre Farm (RTF). Plant species VVP KEP RTF Urera baccifera 3 0 0 Psidium cattleianum 4 0 0 Paullinia carpopoda 2 0 0 Lepismium eruciforme 1 0 0 Ficus enormis 3 0 0 F. insipida 0 0 1 F. monckii 2 0 0 Melia azedarach 1 0 1 Syagrus romanzoffiana 4 2 7 Casearia svlvestris 7 12 0 Rapanea ferruginea 4 15 0 R. umbellata 0 6 0 Philodendron spp. 0 14 0 Cestrum calvcinum 0 4 0 Miconia spp. 0 1 0 St yrax leprosus 0 1 0 Citronella gongonha 0 1 0 Morus nigra 0 6 0 Trema micrantha 0 1 0 Diospyrus kaki 0 9 0 Citrus spp. 0 0 1 Eriobotrya japonica 0 13 0 Peschiera australis 0 0 1 Celtis iguanaea 0 0 1 Totals 31 85 12 TABLE 6. Number of feeding events by strata and foraging substrate of Plush-crested Jays at each of the st sites in southern Brazil: Vila Velha State Park (VVP), Klabin Ecological Park (KEP), and Ribeirao do Tigre Farm (RTF). Strata VVP KEP RTF Totals Understory Branches 164 204 56 424 Leaves 132 187 70 389 Epiphytes 81 106 30 217 Air 8 17 13 38 Subtotals 385 514 169 1,068 Middle level Branches 122 168 89 379 Leaves 138 224 90 452 Epiphytes 82 137 39 258 Air 6 17 8 31 Subtotals 348 546 226 1,120 Subcanopy Branches 11 118 42 171 Leaves 17 93 56 166 Epiphytes 17 32 18 67 Air 0 12 6 18 Subtotals 45 255 122 422 Ground 186 142 42 370 Totals 964 1,457 559 2,980
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