Territory distribution and habitat selection of the Serra Finch (Embernagra Longicauda) in Serra do Cipo, Brazil.
|Abstract:||The near-threatened Serra Finch (Embernagra longicauda) is restricted to the main mountain ranges in eastern Brazil inhabiting campos rupestres (rocky fields). We mapped 17 mated pairs in a 138-ha area within Serra do Cipo National Park; a density of 0.25 adults/ha. Estimated average territory size varied from 2.52 [+ or -] 0.77 ha (95% kernel) to 3.35 [+ or -] 0.90 ha (100% minimum convex polygon). The distance between territory centers of neighboring pairs was 162.38 [+ or -] 28.93 m. The overlap between neighboring territories was 15.3 [+ or -] 5.9% (95% kernel) and 2.0 [+ or -] 2.3% (polygon method). Pairs remained together throughout the year in the same territories and defended these against intruding neighbors. Analyses of habitat selection indicated preference for woodland and scrubland habitats associated with humid valleys, while grasslands were avoided. The Serra Finch used the available habitats more than expected from random at different spatial scales. Our data identified habitats that should be priority for conservation of the Serra Finch.|
Computer software industry
(Protection and preservation)
Grasslands (Protection and preservation)
National parks and reserves (Protection and preservation)
Freitas, Guilherme H.S.
|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|
|Product:||Product Code: 7949121 National Parks NAICS Code: 71219 Nature Parks and Other Similar Institutions SIC Code: 7372 Prepackaged software|
|Organization:||Company Name: Environmental Systems Research Institute Inc. Organization: BirdLife International|
|Geographic:||Geographic Scope: Brazil; United States; California Geographic Code: 3BRAZ Brazil; 1U9CA California|
The Serra Finch (Embernagra longicauda) is a poorly understood
species, known for more than a century based on only two specimens from
an unspecified locality in South America (O'Brien 1968, Mattos and
Sick 1985). Recent studies have found it to be restricted to
mountaintops in eastern Brazil, mainly in the states of Minas Gerais and
Bahia, and particularly the Espinhaco Range, Serra da Mantiqueira and
Serra do Caparao (Vasconcelos 2008). This species may be easily observed
on mountains above 900 m, frequenting habitats on quartzite, gneiss, and
iron rich soils. This species typically places its nest on rocky
outcrops (Hoffmann et al. 2009, Rodrigues et al. 2009), tends to forage
in pairs for arthropods and fleshy fruits among herbs, grasses and
bushes (Hoffmann et al. 2009), and pairs are known to sing a duet
(Freitas and Rodrigues 2007), a typical behavior of territory defense
(Catchpole and Slater 1995).
The Serra Finch has been categorized as nearthreatened due to a perceived population decline possibly resulting from significant habitat loss, primarily due to cattle ranching, land conversion, and intensive mining activity (BirdLife International 2011). Thus, there is a need of focused ecological studies of the species to identify its habitat requirements (Stotz et al. 1996, BirdLife International 2011). Accurate assessment of habitat requirements, population size, and density are paramount to undertaking any successful conservation measures (Bibby et al. 2000). The objectives of our study were to: (1) quantify and map all territories of Serra Finches in a study site within Serra do Cipo National Park in southeastern Brazil, and (2) identify specific habitats favored by this species.
Study Area.--Our study was conducted in a 138-ha area near Indaia Stream, in the 'Alto do Palacio' region (19[degrees] 15' S, 43[degrees] 31' W) in the northern part of Serra do Cipo National Park in the southern portion of the greater Espinhaco Range of southeastern Brazil (Fig. 1). Alto do Palacio is near the ridge of the eastern slope of the Serra do Cipo Mountains at an altitude of 1,280 to 1,380 m. This region is humid throughout the year, even in the dry season, and is characterized by frequently misty weather conditions (Ribeiro et al. 2009); the bird community is closely associated with the Atlantic Forest (Rodrigues et al. 2011).
The study area is a mosaic of habitats traversed by numerous small valleys. Seven habitats were identified. (1) Rocky outcrops, referring to areas with soils derived from quartzite, that are dominated by several species of herbs, shrubs (such as Bromeliaceae, Orchidaceae, Velloziaceae, and Cactaceae), and small trees up to 3 m in height, including Eremanthus erythropappus and E. crotonoides (Asteraceae). (2) Dry grasslands, dominated by Lagenocarpus tenuifolius (Cyperaceae), Panicum loreum (Poaceae), and Paepalanthus spp. (Eriocaulaceae). (3) Wet grasslands, dominated by Lagenocarpus rigidus in marshy areas close to watercourses. (4) Dry scrubland, which has some plants that also occur in the rocky outcrop habitat, but which is dominated by Coccoloba acrostichoides (Poligonaceae) and Aulonemia effusa (Poaceae) in sandy soil with substantial gravel in some hilltop areas. (5) Wet scrubland, dominated by small trees up to 2 m in height, such as Tibouchina spp., Lavoisiera imbricata (Melastomataceae), and Baccharis itatiaiae (Asteraceae) in marshy areas. (6) Riparian woodlands, characterized by dense vegetation and dominated by 4 to 10-m tall trees including Miconia chartacea (Melastomataceae) and Hololepis pedunculata (Asteraceae). (7) Candeial, a specific formation of dense groups of 3 to 5-m tall Eremanthus spp. (Asteraceae) trees, known as 'candeia'. The term 'dry' is not associative of arid conditions but differentiates from saturated soil conditions of the 'wet' habitats.
[FIGURE 1 OMITTED]
Data Collection.--Data were collected from February 2007 to May 2008 during monthly surveys of 3 days in duration. The total sampling effort was 245 hrs, divided between capturing and processing individuals (~80 hrs), and locating birds.
We captured Serra Finches using mist nets and banded each with unique combinations of colored leg bands to allow individual identification and numbered metal leg bands supplied by Centro Nacional de Pesquisa para Conservacao de Aves Silvestres (CEMAVE/IBAMA, license # 1161/3 and 1161/5). Surveys were conducted throughout the study area. Locations of Serra Finches were recorded using a hand-held Garmin (Olatbe, KS, USA) global positioning system (GPS) unit to within 9 m accuracy. All GPS locations correspond to pairs because finches were always detected in pairs. Locations were recorded every 10 m of a pair's movements (not caused by observer presence) until lost from view. Habitat type was recorded where birds were observed.
Territory Distribution.--We took GPS locations for all marked and unmarked pairs detected in the study area. Pairs were detected mainly while singing a duet from a perch, which was interpreted as evidence of a particular mated pair's territory. Locations were recorded throughout the year (dry and wet seasons), as we did not observe significant seasonal changes in spatial distribution and general territory defense of mated pairs. We performed spatial analyses using ArcView GIS Version 3.2 (ESRI 1998). The distance between nearest neighboring pairs was calculated from the arithmetic mean of the locations of each mapped pair. We calculated the density of individuals in the study area by mapping territories (Bibby et al. 2000).
Territory size was calculated using the Home Range Extension Version 1.1 (Rodgers and Carr 1998). We used the following estimators: 95% fixed kernel using the least-squares cross-validation smoothing parameter (Wotton 1989, Seaman and Powell 1996); 50% fixed kernel for the core areas (Powell 2000); and 100% minimum convex polygon (MCP; Mohr 1947). We chose the kernel method because it is the most recommended (Laver and Kelly 2008) and MCP for comparison because it is the most used and emphasized the boundaries of space used (Kernohan et al. 2001). This is important for territory characterization based on the concept of use of an exclusive area (Pitelka 1959). We measured territory size only for pairs with [greater than or equal to] 30 locations to avoid imprecision and bias in the size estimates (Seaman et al. 1999, Kernohan et al. 2001).
Habitat Selection.--We analyzed habitat selection by comparing habitat use and that expected based on availability. We used the described habitat classifications for availability and delineated habitats based on satellite images from Google Earth (2010) in combination with direct observations in the field and measured with X-tools extension for ArcView Version 3.2 (ESRI 1998).
We calculated habitat selection using three approaches. First, we compared the number of all Serra Finch locations within each habitat with the proportion of available habitats in the study area. Chi-square tests were used to evaluate the null hypothesis that actual use of different habitat types is directly proportional to their availability (Neu et al. 1974). Bonferroni confidence intervals ([alpha] = 0.05) were calculated from the observed proportions of habitat use to identify which habitat types were selected (Neu et al. 1974, Byers and Steinhorst 1984). Second, we compared the proportions of habitat within the 95% kernel territory boundaries to proportions of available habitats in the study area (second-order habitat selection sensu Johnson 1980). Third, we compared the proportion of locations in each territory to the proportion of available habitats within the 95% kernel territory boundaries (third-order habitat selection sensu Johnson 1980).
We used compositional analysis (Aebischer et al. 1993) for the second and third-order approach. This analysis used the Wilks' lambda statistic test for overall differences in habitat use. Comparisons of particular habitat types were made with paired t-tests (Aebischer et al. 1993) if the Wilks' test suggested differential habitat use. Significance was set at P < 0.05. Habitat types were ranked from most to least selected using a matrix of mean and standard deviation of log ratio differences for all habitat types if selection was significantly non random. Missing values (zero) in the data matrix were replaced by 0.001. The minimum number of individuals needed for compositional analysis is six (Aebischer et al. 1993). Thus, all Serra Finches with >10 locations were used and those with <10 locations were excluded from the analyses. Leban et al. (2001) noted that compositional analysis is primarily affected by the number of animals sampled and did not vary much with number of locations (as few as 10 observations). We conducted the habitat selection analyses using Resource Selection for Windows, Version 1.0 (Leban 1999).
Territory Distribution.--Twenty-one individual birds were marked with colored leg bands and 17 were relocated distributed among 11 mated pairs. The four individuals not relocated included a juvenile and three possibly adult floaters. Six additional mated pairs of unmarked individuals were also detected (during at least 2 different days) occupying areas between some of the marked pair's territories. Those pairs were reliably identified because the adjacent pairs often sang duets at the same time. Thus, at those moments it was possible to hear up to three different pairs. Territories of 17 total pairs were mapped by 318 GPS locations in the entire 138-ha area during the study period (Fig. 1). This was a density of adults with established territories of 0.25 birds/ha or 0.12 pairs/ha. Distance between territory centers was 162.38 [+ or -] 28.93 (range = 119.95-233.85 m, n = 17). Territory size estimates (mean [+ or -] SD) of five pairs that were located most often ranged from 2.52 [+ or -] 0.77 ha (MCP) to 3.35 [+ or -] 0.90 ha (95% kernel) (Table 1).
The area of common use between territories (MCP) was 991.27 [m.sup.2] for pairs A (4.68% of territory) and B (3.00%), and 19.34 [m.sup.2] for pairs C (0.07%) and D (0.06%) (Fig. 1). The estimated overlap in area between territories (95% fixed kernel) was 7,204.59 [m.sup.2] for pairs A (20.72%) and B (19.35%), and 3,673.51 [m.sup.2] for pairs C (13.09%) and D (7.94%) (Fig. 2). The core areas overlapped for 1,155.62 [m.sup.2] for pairs A (12.03%) and B (10.99%), and did not overlap between pairs C and D.
We did not observe any changes in the composition of mated pairs or in their territories, indicating these birds have annual site fidelity and a socially monogamous mating system. All pairs were observed singing duets on perches, apparently defending their territories throughout the year, even outside the breeding season. We observed agonistic events (n = 4) on territory boundaries, when two pairs stayed in close proximity performing a unison duet, and ending in a physical confrontation on two occasions. One of those, between pairs A and B, occurred within the MCP overlap area.
Habitat Selection.--The 318 locations recorded in the habitat types were not distributed as expected from availability in the study area (P < 0.001, [X.sup.2] = 686.82, df = 6; Table 2). The candeial, rocky outcrops, wet and dry scrubland, and riparian woodland habitats were used more than expected by chance as the proportions of expected use were below the Bonferroni confidence intervals for observed use. Use of dry and wet grassland habitats was lower than what could be expected by chance (Table 2).
We used the location data of nine Serra Finches which had > 10 locations (Tables 1-3) to examine habitat selection by the second and third-order approaches. Dry and wet grassland were essentially avoided by all individuals (Table 3), and we excluded them from the compositional analysis. The proportion of habitat use based on 95% kernel territories in the second-order approach was non random relative to that available in the study area ([lambda] = 0.13, df = 4, [X.sup.2] = 18.30, P < 0.05). A ranking matrix ordered the habitat types in the sequence: wet scrubland > riparian woodland > rocky outcrop > candeial > dry scrubland (Table 4). Serra Finch territories had significantly more wet scrubland and riparian woodland than dry scrubland.
Habitat use of territory locations versus 95% kernels in the third-order approach significantly differed from random ([lambda] = 0.20, df = 4, [X.sup.2] = 14.41, P < 0.05). The matrix ranked the habitat in the order: candeial > riparian woodland > rocky outcrop > dry scrubland > wet scrubland. Candeial was significantly more used than rocky outcrop, dry scrubland, and wet scrubland, the last was significantly less used than riparian woodland (Table 4).
Territory Distribution.--The number of adult Serra Finches that we found is high compared with other species that inhabit open areas in central Brazil. Our results revealed higher densities than 11 species studied by Braz (2008), including the closely related Black-masked Finch (Coryphaspiza melanotis) (0.23 individuals/ha) and Wedge-tailed Grass Finch (Emberizoides herbicola) (0.15 individuals/ha). Silva (2008) reported 11 individuals/ha for Serra Finch. Hoffman (2011) estimated a density of 0.44 [+ or -] 0.08 individuals/ha for Gray-backed Tachuri (Polystictus superciliaris), a tyrant-flycatcher with a range distribution similar to the Serra Finch (Vasconcelos 2008). Population density is highly dependent on the quality of a particular ecosystem (Makarieva et al. 2005, Johnson 2007). Our density data are for a population of the Serra Finch occurring in an undisturbed area, which may serve as a baseline for comparison with future studies of this species.
Mattos and Sick (1985) reported 400 m as the distance between pairs of Serra Finch, which is higher than the average distance estimated for the birds at our study site. Our results were derived from small sample sizes, but it is the most accurate estimate available, particularly given that Mattos and Sick (1985) did not specify how their values were calculated.
[FIGURE 2 OMITTED]
The mean MCP territory size of Serra Finch was relatively small compared to published territory estimates for other species that inhabit the Cerrado Biome including the Rufous-fronted Thornbird (Phacellodomus rufifrons) (3.43 [+ or -] 0.62 ha; Rodrigues and Carrara 2004), Suiriri Flycatcher (Suiriri suiriri) (14 [+ or -] 1.9 ha; Lopes and Marini 2006), Chapada Flycatcher (S. islerorum) (11.2 [+ or -] 0.6 ha; Lopes and Marini 2006), Crested Black Tyrant (Knipolegus lophotes) (7.3 [+ or -] 0.57 ha; Ribeiro et al. 2002), Cinnamon Tanager (Schistochlamys ruficapillus) (8.4 [+ or -] 1.9 ha; Domingues and Rodrigues 2007), Shrike-like Tanager (Neothraupis fasciata) (4.3 ha; Alves 1990), Henna-capped Foliage-gleaner (Hylocryptus rectirostris) (2.9 [+ or -] 1.4 ha; Faria et al. 2007), Graybacked Tachuri (4.3 [+ or -] 1.2 ha; Hoffmann 2006), and Cipo Canastero (Asthenes luizae) (4.0 [+ or -] 2.6 ha; Freitas 2011). These studies ali used similar methods, without radiotelemetry (except Freitas 2011). The home range area of those species could be larger than reported (Anich et al. 2009).
The Serra Finch appears to be similar to most tropical bird species, maintaining a long-term pair bond and a territory throughout the year (Stuchbury and Morton 2001), at times making it difficult to discern the territory space of the total home range area (Lopes and Marini 2006). The territorial behavior was illustrated by the annual site fidelity associated with duet song performance, and minimum overlap of the used area visualized by the MCP.
Habitat Selection.--Johnson (1980) proposed a natural order to the process of habitat selection according to the scale of observation. First-order selection identifies the physical or geographical distribution of a species which, for the Serra Finch, is the mountaintop habitat complex that occurs throughout eastern Brazil (Vasconcelos 2008).
Second-order selection delineates the homerange area of an individual or social group. Serra Finches did not establish territories at random and second-order selection was visually demonstrated by the localization of the 17 Serra Finch territories closely associated with watercourses (Fig. 1). We also found that wet scrubland and riparian woodland habitats delineated establishment of the boundaries of the pair's territory. This preference may be the result of higher concentration of fruits, mostly Malastomataceae fruits, which are an important resource for the Serra Finch (Hoffmann et al. 2009; GHSF, pers. obs.). Species of Melastomataceae are highly diverse in the campos rupestres (Giullieti et al. 1997) and produce fleshy ornithocoric fruits, which are abundant in riparian woodlands, rocky outcrops, and scrublands, while almost no fleshy fruits are available in grasslands. The distribution of these particular plants closely matches the pattern of habitat use by Serra Finches in our study area.
Third-order selection examines the specific use of habitat components within the home-range area (Johnson 1980). Our ranking at this spatial scale selection indicates a preference for candeial. This vegetation is typically on the western slopes of the Serra do Cipo (Melo-Junior et al. 2001), and is patchily distributed in the study area. This habitat does not supply fruits but does supply arthropods and perches. We also detected more preference for riparian woodland than wet scrubland at this scale suggesting preference for habitats with woody elements. This contrasts with previous information about Serra Finch preference for open vegetation habitats (Mattos and Sick 1985, Vasconcelos 2001, Hoffmann et al. 2009).
We identified the most important habitat types in the campos rupestres complex for the Serra Finch and described the spatial distribution that should influence development of better long-term conservation strategies. Actions that contribute to reduce habitat quality for the species should be avoided (BirdLife International 2011). The conservation priority should be maintenance of the most preferred habitats of the Serra Finch that possibly contribute most to species fitness (Garshelis 2000). Preservation of valleys, woodlands and scrnblands, and water in the mountains within the species range must be prioritized; especially important is protection and maintenance of riparian woodlands and candeial. Those habitats have been intensely exploited in the region for production of wood and also for oil extraction in the candeial (Ribeiro et al. 2009).
The mountain range where the Serra Finch occurs is critically threatened due to increasing land conversion and no effective conservation plans (Jacobi et al. 2007, Martinelli 2007). The actual habitat protection law is under review which may be detrimental to the currently protected areas (Ribeiro and Freitas 2010).
One of the most common threats for the campos rupestres complex are annual fires, mainly promoted to benefit livestock. Annual fires tend to reduce the extent of scrublands and woody elements while increasing the dominance of grassland habitats (Coutinho 1990, Moreira 2000). The Alto do Palacio is a protected area and there have been no recent reports of fires, unlike in neighboring areas (Franca and Ribeiro 2008). We would expect to find a more grassland-dominated landscape, less woody vegetation, and fewer shrubs in areas with annual burning regimes and, consequently, a lower density of Serra Finches. Thus, areas with annual burning need a fire protection program for habitat restoration and conservation.
We thank the student volunteers who assisted us in the field. We are grateful to the ICMBio Serra do Cipo staff for lodging and support, and to Bruno Crepaldi and G. G. Teixeira for help with the figures. We appreciate the important comments and suggestions on this manuscript hy C. E. Braun and L. M. Costa, and the efforts of J. A. Mobley in reviewing our work and helping with the English language. We also thank Fundacao O Boticario de Protecao Natureza, CNPq (473428/2004-0 and 473809/2008-7) and FAPEMIG (CRA APQ-0434-5.03/07) for supporting this study. M. R. is grateful to CNPq for fellowship support during this study.
Received 8 September 2010. Accepted 24 September 2011.
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GUILHERME H. S. FREITAS (1, 2) AND MARCOS RODRIGUES (1)
(1) Laboratorio de Ornitologia, Departamento de Zoologia, Instituto de Ciencias Biologicas, Universidade Federal de Minas Gerais, GP 486, 31270-901, Belo Horizonte, Minas Gerais, Brazil.
(2) Corresponding author; e-mail: email@example.com
TABLE 1. Data for nine best sampled pairs of the Serra Finch in Serra do Cipo National Park, Brazil. Values of 100% minimum convex polygon (MCP); and by 95% and 50% kernel in ha. Bird Number of Sampling Sampling 100% 95% 50% pair locations days period (a) MCP Kernel Kernel A 43 6 435 2.11 3.29 0.85 B 57 14 412 3.30 3.62 1.02 C 52 9 273 2.68 2.84 0.63 D 44 6 168 3.09 4.69 1.16 E 14 3 32 0.78 2.66 0.75 F 13 3 113 1.46 4.87 1.23 G 30 4 169 1.42 2.32 0.55 H 12 3 112 1.24 4.78 1.15 K 16 4 169 0.88 1.85 0.40 (a) Number of days elapsed between the first and the last locations. TABLE 2. Habitat selection of Serra Finches in Serra do Cipo National Park, Brazil. The proportion expected was calculated from the relative proportion of each habitat available within the study area, and the proportion observed in relation to 318 locations obtained for 17 pairs. Habitat selection is indicated for differences <0.05 significance; positive (+) for expected proportions below the Bonferroni confidence interval, and negative (-) for expected proportions above the interval. Proportion Proportion observed Habitat expected [P.sub.i] Candeial 0.003 0.053 Dry grassland 0.793 0.019 Wet grassland 0.079 0.025 Rocky outcrop 0.047 0.366 Dry scrubland 0.016 0.072 Wet scrubland 0.038 0.234 Riparian 0.022 0.231 woodland Selec Habitat Confidence interval -tion Candeial 0.039 [less than or equal to] [P.sub.1] 0.068 + Dry grassland 0.010 [less than or equal to] [P.sub.2] 0.028 - Wet grassland 0.015 [less than or equal to] [P.sub.3] 0.352 - Rocky outcrop 0.334 [less than or equal to] [P.sub.4] 0.397 + Dry scrubland 0.055 [less than or equal to] [P.sub.5] 0.089 + Wet scrubland 0.207 [less than or equal to] [P.sub.6] 0.262 + Riparian 0.204 [less than or equal to] [P.sub.7] 0.259 + woodland TABLE 3. Habitat distribution (%) in relation to locations of the most sampled pairs of Serra Finches in Serra do Cipo National Park, Brazil, and the habitat composition within each 95% kernel territory area. Habitat A B C D E F G % locations Candeial 16.28 0.00 5.77 0.00 14.29 0.00 6.67 Dry grassland 6.98 0.00 3.85 0.00 0.00 0.00 0.00 Wet grassland 0.00 3.51 3.85 0.00 7.14 0.00 6.67 Rocky outcrop 0.00 8.77 78.85 75.00 28.57 69.23 33.33 Dry scrubland 25.58 0.00 0.00 0.00 14.29 0.00 3.33 Wet scrubland 18.60 50.88 7.69 9.09 21.43 15.38 23.33 Riparian 32.56 36.84 0.00 15.91 14.29 15.38 26.67 woodland % territory Candeial 3.34 0.00 1.83 0.00 1.03 0.00 0.72 Dry grassland 38.77 42.32 30.73 26.46 62.93 67.31 56.73 Wet grassland 1.30 10.54 6.41 1.41 10.98 5.47 8.24 Rocky outcrop 0.00 5.30 53.94 62.55 3.46 22.78 14.56 Dry scrubland 27.24 0.00 0.00 0.00 4.44 0.00 0.61 Wet scrubland 19.56 35.54 7.09 7.09 13.57 2.67 7.54 Riparian 9.80 6.30 0.00 2.49 3.60 1.78 11.60 woodland Habitat H K % locations Candeial 16.67 0.00 Dry grassland 0.00 0.00 Wet grassland 0.00 6.25 Rocky outcrop 16.67 25.00 Dry scrubland 25.00 0.00 Wet scrubland 16.67 37.50 Riparian 25.00 31.25 woodland % territory Candeial 1.27 0.00 Dry grassland 73.22 51.84 Wet grassland 4.74 18.95 Rocky outcrop 1.77 7.69 Dry scrubland 4.88 0.00 Wet scrubland 4.21 16.90 Riparian 9.92 4.62 woodland TABLE 4. Habitat ranking matrices for nine Serra Finch pairs in Serra do Cipo National Park, derived from log-ratio differences based on compositional analysis. Triple sign represents significant deviation from random at P < 0.05. Second-order habitat selection compared proportion of habitat used within 95% kernel territories with proportion of total available habitat in the study area, and third-order compared the proportion of locations for each animal in each habitat type with the proportion of each habitat type within the finches's 95% kernel territories. Rank from the least to the most preferred habitat. Rocky Dry Wet Candeial outcrop scrubland scrubland Second-order Candeial - + - Rocky outcrop + + - Dry scrubland - - --- Wet scrubland + + +++ Woodland + + +++ - Third-order Candeial +++ +++ +++ Rocky outcrop --- + + Dry scrubland --- - + Wet scrubland --- - - Woodland - + + +++ Woodland Rank Second-order Candeial - 1 Rocky outcrop - 2 Dry scrubland --- 0 Wet scrubland + 4 Woodland 2 Third-order Candeial + 4 Rocky outcrop - 2 Dry scrubland - 1 Wet scrubland --- 0 Woodland 3
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