BIRD COMMUNITIES IN FIVE ATLANTIC FOREST ......E-mail: [email protected] Resumo. – Comunidades de aves em cinco fragmentos de floresta Atlântica no sul do Brasil. – Comunidades

ORNITOLOGIA NEOTROPICAL 12: 11–27, 2001© The Neotropical Ornithological Society

BIRD COMMUNITIES IN FIVE ATLANTIC FOREST FRAGMENTS IN SOUTHERN BRAZIL

Luiz dos Anjos

Universidade Estadual de Londrina, Depto Biologia Animal e Vegetal, CX 6001, Londrina, 86051-970, Paraná, Brasil. E-mail: [email protected]

Resumo. – Comunidades de aves em cinco fragmentos de floresta Atlântica no sul do Brasil. –Comunidades de aves em cinco fragmentos florestais da floresta atlântica foram estudados usando con-tagens pontuais de distância ilimitada, de Janeiro a Dezembro de 1996, na região de Londrina (23o17’S,51o15’W), Estado do Paraná, sul do Brasil. Em quatro destes fragmentos florestais, os quais não estão amais de 1100 m entre si, o número de espécies diminuiu e foi altamente correlacionado com o tamanhoda área (r = 0.998, df = 2, P = 0.002): 134, 129, 113 e 106 espécies em fragmentos de 656 ha (em torno de70 ha foram somente amostrados), 56 ha, 25 ha e 11 ha, respectivamente. No quinto fragmento de 60 ha,o qual está isolado por ao menos 2 km de qualquer outra grande área de floresta, somente104 espéciesforam registradas. O número médio de espécies por mês no fragmento de 656 ha (64 ± 11.3 spe-cies) foi estatisticamente similar ao de 56 ha (59.8 ± 8.1 espécies; P > 0.05) e ao de 25 ha (54.8 ± 6.1 espé-cies; P > 0.05) mas não ao de 11 ha (49.5 ± 7.6 espécies: P < 0.05) e ao de 60 ha (45.8 ± 7.3 espécies; P <0.05, teste de Tukey). A estrutura da comunidade (tendo em consideração os hábitos alimentares das espé-cies) do fragmento de 25 ha foi smilar ao de 656 ha (rs = 0.81), provavelmente, em função de um corredorde 100 m de largura que os conectava; os outros fragmentos, os quais eram completamente isolados,tiveram menor correlação (rs < 0.72) em relação ao fragmento de 656 ha. Aumento da abundância relativacom a diminuição do tamanho da área dos fragmentos (densidade compensatória) foi encontrada em 18espécies. Embora algumas tendências gerais serem apresentadas neste estudo para certos grupos de aves,os efeitos da fragmentação florestal são certamente específicos para cada espécie.

Abstract. – Bird communities in five Atlantic forest fragments were studied using point counts of unlim-ited distance monthly, January to December of 1996, in the Londrina region (23o17’S, 51o15’W), ParanáState, southern Brazil. In four of these forest fragments, which are no farther than 1100 m from eachother, the species number decreased and was strongly correlated with area size (r = 0.998, df = 2, P =0.002): 134, 129, 113 and 106 species in fragments of 656 ha (only about 70 ha were sampled), 56 ha, 25 haand 11 ha, respectively. In the fifth fragment of 60 ha, which is isolated by at least 2 km from anyother large forest area, only 104 species were recorded. The mean number of species per month in the656 ha fragment (64 ± 11.3 species) was statistically similar to that in the 56 ha (59.8 ± 8.1 species;P > 0.05) and 25 ha fragments (54.8 ± 6.1 species; P > 0.05) but not to the 11 ha fragment (49.5 ± 7.6species; P < 0.05) or the isolated fragment of 60 ha (45.8 ± 7.3 species; P < 0.05, Tukey test). Thecommunity structure (taking into account the species` feeding habits) of the 25 ha fragment wassimilar to the 656 ha fragment, probably, because of a 100 m wide forest corridor connectingthem (rs = 0.81); the other fragments, which were fully isolated, had lower correlation (rs < 0.72) tothe 656 ha fragment. Increase of relative abundance with decrease in area size of fragments (densitycompensation) was found for 18 species. Although some general tendencies are presented in this studyfor certain bird groups, the effects of forest fragmentation are certainly species-specific. Accepted 20 August2000.

Key words: Forest fragmentation, Paraná, Londrina, conservation, density compensation, Brazil, bird communities.

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INTRODUCTION

Atlantic forest stretches along the Atlanticcoast of Brazil from Ceará state, in the north,to Rio Grande do Sul state, in the south, andhas undergone such intense fragmentation asto make it the most endangered zoogeo-graphical region in South America (Stotz et al.1996). Vegetation in northern Paraná state(Fig. 1), for example, was made up of continu-ous forest until the 1920s. Since then therehas been rapid deforestation in the region foragriculture. The continuous forest wasreduced by this process of land use to smalland sparse forest fragments which currentlytotal less than 1% of the original area in thenorth of Paraná. The largest and the mostrepresentative forest fragment in northernParaná is Parque Estadual Mata dos Godoy(656 ha).

In Brazil, studies on bird fauna in forestfragments have been carried out particularlyin southern and southeastern regions and inAmazonia (Willis 1979; Anjos & Laroca 1989,Bierregaard & Lovejoy 1989, Bierregaard1990, Vielliard & Silva 1990, Melo & Marini1997, Anjos 1992, Aleixo & Vielliard 1995,Stouffer & Bierregaard 1995a, b; Bierregaard& Stouffer 1997, Aleixo 1999, Almeida et al.1999, Anjos & Boçon 1999, Soares & Anjos1999). Most of the studies have detected adecrease in number of species and modifica-tion in bird community structure (e.g., Willis1979, Bierregaard 1990); these effects tend tobe minimized in small fragments linked tolarger fragments by forest corridors, or thatare simply close to them (e.g., Stouffer &Bierregaard 1995b, Anjos & Boçon 1999).Some species benefit from fragmentation andincrease their numbers because of an expan-sion of the ecological niche in the smaller for-est fragments, a phenomena which has beencalled density compensation (MacArthur et al.1972, Wright 1980).

The goal of this study was to analyze what

differences in the composition of the birdcommunity are found among five Atlanticforest fragments in the northern region ofParaná state.

STUDY AREA AND METHODS

Five forest fragments were studied, includingthe Parque Estadual Mata dos Godoy (PG).All are close to the town of Londrina, Paranástate, southern Brazil (Fig. 1). PG (23°17’S,51°15’W) 656 ha, is located 15 km south ofLondrina. Three different sized forest frag-ments called A, B and C fragments are southof PG and close to it; fragment A (FA, 56 ha)is 500 m from PG; fragment B (FB, 25 ha) is400 m from PG and linked to it by a 100 mwide forest corridor (a continuous forest witha similar vegetation structure to the frag-ments), and fragment C (FC, 11 ha), which istotally isolated and 1100 m from PG, 500 mfrom FA and 250 m from FB (Fig. 1). Thespace between these four forest fragments,the matrix habitat (Sisk et al. 1997), is used forranching and has some bushes and scatteredtrees. The forest fragments are isolated forthis matrix habitat for at least 40 years. PGhas been a permanent preservation area since1989; before that, hunting and cutting werenot permitted by the owner of the area. Vege-tation is well preserved in PG and in the otherthree fragments. The fifth fragment studied isthe Horto Municipal de Ibiporã (HI, 60 ha),35 km northeast of PG. This fragment alsohas well preserved vegetation, but it is moreisolated (at least 2 km from any other largeforest fragment). The matrix habitat aroundHI is 60% agriculture and 40% urban. Theforest fragments are seasonal semi-deciduousforest. The dominant trees are: Aspidospermapolyneuron (Apocynaceae), Euterpe edulis(Areceae), Galesia intergrifolia (Phytolaccaceae)and Ficus glabra (Moraceae). The mean annualrainfall is 1,200 mm in the region, and is high-est from October to March (130 to 200 mm

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BIRD COMMUNITIES OF FOREST REMMANTS IN SOUTHERN BRAZIL

per month). The annual temperature varia-tion shows that the means from December toFebruary are higher (around 23°C) and theJune and July means are lower (around 17°C)(Maack 1981). PG is 650 m a.s.l.

Forest fragments were censused bymonthly point counts of unlimited distance(Blondel et al. 1970, Vielliard & Silva 1990)from January to December 1996. A total of45 points were used to estimate relative abun-dance: 15 in PG, 10 in HI, 10 in FA, 6 in FBand 4 in FC. These points covered the totalareas of HI, FA, FB and FC but just part ofthe total area of PG (the area sampled in PGwas around 70 ha). Each point was 100 mfrom another and at least 50 m from the edgeof forest. Each month five points in eachfragment were chosen by random selectionfor sampling; in FC the fourth point wasalways repeated once each month. So, 60

counts were performed in each fragment dur-ing the year. The time for sampling at eachpoint was 20 min. The Index of Point Abun-dance (I.P.A.) of each species (Blondel et al.1970; Vielliard & Silva 1990), considered inthis work as the relative abundance of eachspecies, was estimated by dividing the totalnumber of species contacts by the total num-ber of points (60) sampled in each fragment.Sampling began at dawn at the first randomlyselected point and finished about 3 h later atthe fifth point. Species were identified prima-rily by sound (99%). Each pair or flock ofeach species was counted once (one contact)while vocalizing. Precaution was taken not tocount the same individual or group morethan once (a form was used in order to locatethe counted individuals), particularly forhighly mobile species. Bird recordings (2,300recordings of 470 species), deposited in the

FIG. 1. Forest fragments (PG, 656 ha; FA, 56 ha; FB, 25 ha; FC, 11 ha) studied in Londrina region (1)north of Paraná State, southern Brazil. PG and FB are linked by a forest corridor. The fifth fragment stud-ied (HI, 60 ha) is located 35 km northeast of PG.

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Bioacoustics Laboratory of the UniversidadeEstadual de Londrina, were used to aid identi-fication.

Taxonomy mostly follows Meyer deSchauensee (1982) and Sick (1997). Feedinghabit was determined for each species basedon field observations and references (Fitz-patrick 1980, Belton 1984, 1985, Ridgely &Tudor 1989, 1994, Cabot 1992, del Hoyo1994, Baptista et al. 1997, Collar 1997, Sick1997) and was categorized as: nectarivore,frugivore (feeding especially on fruit pulp),herbivore (including pulp, seeds, buds and/orleaves), insectivore, omnivore (feeding oninvertebrates and small vertebrates), carnivore(feeding especially on small vertebrates) andherbivore/omnivore (feeding on either plantor animal sources). For insectivores, the areawhere the insect or its larvae was most oftencaptured was also recorded. Three classes ofcapture sites were defined: (1) trunk (andtwigs), (2) leaves, and (3) other sites (calledgeneralized insectivores).

The accumulated number of species wasobtained monthly in each fragment based onthe five point counts performed each month.

Analyses of variance (one way ANOVA) andthe Tukey multiple range test (P < 0.05) wereused to evaluate the average number of spe-cies and of contacts per month in forest frag-ments. Similarities in the bird speciescomposition among forest fragments weredetermined using the Sørenson Index (Cs;Magurran 1988): Cs = 2j/(a + b), where j isthe number of species found in both sites anda is the number of species in site A, and b isthe number of species in site B. Correlation(r) was used to measure the relation betweenthe species number and area size of fragment.Spearman’s rank correlation (rs) was used tomeasure the relation between the communitystructures of the fragments. Differences innumber of contacts of species between thefragments were tested with χ2 analysis at α <0.05.

RESULTS

Number of species and similarity. Birds in the fiveforest fragments totaled 184 species: 134 inPG (656 ha but only 70 ha were sampled),104 in HI (60 ha), 129 in FA (56 ha), 113 in

FIG. 2. Monthly accumulated number of bird species in the studied forest fragments.

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FB (25 ha) and 106 in FC (11 ha; Appendix).The accumulated number of species in thestudy months, show that no species wasadded after September in FB, October in FAand November in PG and FA. One specieswas added to the list in HI in December (Fig.2).

FA and FB (59.8 ± 8.1 and 54.8 ± 6.1 spe-cies respectively) had a statistically similarmean number of species per month (consid-ering the bird lists obtained in the five sam-plings each month) to PG (64 species ± 11.3;Tukey test, P > 0.05); HI and FC (45.8 ± 7.3and 49.5 ± 7.6 species respectively) had lowermeans (Tukey test, P < 0.05).

HI, the most isolated forest fragment, hadthe lowest similarity value (Sørenson) com-pared with PG (0.64); FA, FB and FC hadsimilarities between 0.73 and 0.78 comparedwith PG. The highest similarity values wereamong FA, FB and FC (0.80 to 0.93).

In the four closer fragments (PG, FA, FBand FC), the species number decreased andwas strongly correlated with area size (r =0.998, df = 2, P = 0.002) when consideringonly the sampled area of PG (70 ha); it wasnot correlated, however, when consideringthe total area of PG (656 ha) (r = 0.73, df =2, P = 0.27). After adding the isolated frag-

ment HI to the sampled area of PG, the num-ber of species remained uncorrelated witharea of forest fragments (r = 0.60, df = 3, P =0.29)

Of the 184 species recorded in the fiveforest fragments, 75 were insectivores, 42herbivores/omnivores, 24 herbivores, 18 fru-givores, 10 carnivores, 9 omnivores and 6nectarivores. Among the insectivores, 36 wereleaf insectivores, 20 were trunk insectivores,and 19 were generalized insectivores. Thenumber of species of leaf insectivores wascorrelated with area (Table 1; r = 0.96, df = 3,P = 0.011), even considering HI in the analy-ses; all the other groups of species numbersshowed weak correlations (not statisticallysignificant) with fragment area (r = –0.33 fornectarivores; r = 0.58 for frugivores; r =–0.08 for herbivores; r = 0.81 for generalizedinsectivores; r = 0.45 for trunk insectivores;r = –0.14 for omnivores; r = 0.63 for carni-vores; r = 0.17 for herbivores/omnivores; df= 3; P > 0.05). If HI is not considered in theanalysis, correlation increases substantially forcarnivores (r = 0.96; df = 2; P = 0.042) andfrugivores (r = 0.93; df = 2; P = 0.068).

Relative abundance. The mean numbers of spe-cies contacts per month were not significantly

TABLE 1. Number of bird species in the study forest fragments (including only the sampled area of PG)according to their feeding habits.

Feeding habits Forest fragments/area size (ha)

PG/70 HI/60 FA/56 FB/25 FC/11NectarivoresFrugivoresHerbivoresTrunk insectivoresLeaf insectivoresGeneralized insectivoresOmnivoresCarnivoresHerbivores/omnivoresTotal Species

216161526136733134

48111324115325104

412141623145536129

511151220115430113

3814142086330106

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different between the fragments (PG = 149.6± 63; HI = 123.8 ± 26; FA = 147.6 ± 26; FB =147.9 ± 25; FC = 133.7 ± 19; Tukey test, P >0.05). But the ten most abundant species in

HI correspond to 44% of the total contacts;in PG this percentage was much lower (33%).The percentage of contacts of the ten mostabundant species in the fragments near to PG

FIG. 3. Ratio between total relative abundance of the bird species grouped according to their feeding hab-its in the study forest fragments (considering only the sampled area of PG). The pie graphs were schemat-ically placed according to the area size and isolation of the forest fragment; PG (70 ha) and FB (25 ha)forest fragments are linked by a forest corridor. Correlation (Spearman's correlation rank) among the com-munity structures (pie graphs) are also showed (* indicates significant correlation, P < 0.05).

AR

EA S

IZE

ISOLATION

HO29%

OM12%

GI10%

LI16%

TI9%

HE17%

FR7%

FC (11 ha)

HO27%

OM7%GI5%

LI16%

TI17%

HE20%

FR8%

PG (70 ha)

HO33%

OM12% GI

4%

LI16%

TI12%

HE15%

FR8%

FB (25 ha)

HO27%

OM10%

GI11%

LI21%

TI10%

HE15%

FR6%

FA (56 ha)HO34%

OM4% GI

7%LI28%

TI7%

HE14%

FR6%

HI (60 ha)

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increased with decrease in area (36% in FA,37% in FB and 39% in FC).

The sum of the relative abundance ofspecies grouped according to feeding habitsdenotes the participation of each group in thecommunity structure in each fragment (Table2). Herbivores/omnivores were the mostabundant species group accounting for 25%or more of the total relative abundance ineach studied fragment; nectarivores and car-nivores had the lowest participation (between0.2% to 3%, Table 2). The community struc-ture in FB, considering the relative abundanceof the species groups (but excluding nectari-vores and carnivores because their low partic-ipation in the community), had the highestcorrelation to PG (Spearman’s rank correla-tion: rs = 0.81, n = 7, P > 0.05); HI (rs =0.72), FA (rs = 0.56) and FC (rs = 0.61) hadno significant correlation to PG (P < 0.05).The highest correlation were between HI andFA (rs = 0.92, P > 0.05) and FA and FC (rs =0.90, P > 0.05; Fig 3).

Sixty-three species recorded in three, fouror five forest fragments had relative abun-dance significantly different (χ2, P < 0.05;Appendix). Of these 63 species, 18 increasedrelative abundance with decrease in area (den-sity compensation) while 17 had the highestrelative abundance in HI, the most isolated

forest fragment (Table 3).Twenty species decreased significantly rel-

ative abundance with decrease in area. Cryp-turellus obsoletus (χ2 = 25.2, df = 4), Aratingaleucophtalmus (χ2 = 42.8, df = 3), Pyrrhura fron-talis (χ2 = 58.4, df = 4), Melanerpes flavifrons (χ2

= 70, df = 4), Sittasomus griseicapillus (χ2 =98.7, df = 4), Xiphocolaptes albicollis (χ2 = 34.7,df = 4), Chiroxiphia caudata (χ2 = 28.4, df = 2),Piprites chloris (χ2 = 23.7, df = 4), Sirystes sibila-tor (χ2 = 54.9, df = 3) and Myiopagis caniceps (χ2

= 52.4, df = 4) were the species whichdecreased more in abundance with decreasein area.

DISCUSSION

The curves of the cumulative number of spe-cies suggest that general totals were reason-ably complete in the study forest fragments.The exception is PG; only a sample area wasconsidered in this study due to its large size.A total of 288 bird species were in factrecorded during an exhaustive survey for thetotal PG area (Anjos et al. 1997). The differ-ence in the number of species recorded in PGmay be explained by the heterogeneous distri-bution of the bird species in a more continu-ous forest; small differences in plantcomposition and structure form mosaics of

TABLE 2. Total relative abundances of bird species in the study forest fragments (including only the sam-pled area of PG) according to their feeding habits.

Feeding habits Forest fragments/area size (ha)

PG/70 HI/60 FA/56 FB/25 FC/11NectarivoresFrugivoresHerbivoresTrunk insectivoresLeaf insectivoresGeneralized insectivoresOmnivoresCarnivoresHerbivores/omnivores

0.232.355.714.884.621.502.050.737.29

0.661.393.311.656.801.680.900.058.38

0.201.824.152.876.053.032.920.277.70

0.332.244.433.554.771.283.620.249.49

0.251.774.652.424.292.753.310.167.52

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environmental conditions for the birds(Tomialojc et al. 1984, Blondel 1986). A moreheterogeneous distribution of bird species

had already been detected in a study carriedout in a continuous area of mixed temperaterain forest (also part of the Atlantic forest)when compared with the fragments; a frag-ment may contain a similar number of birdspecies to a similar sampled area of mixedtemperate rain forest (Anjos & Boçon 1999).The same pattern is repeated in forest frag-ments in the Londrina region, for PG (134species) and FA (129 species); the area of FA(56 ha) is rather similar in size to the sampledarea covered by the 15 points of PG (70 ha)and which resulted in a similar number ofspecies. The mean number of species permonth, which was statistically similar betweenPG (64 species) and FA (59.8 species), cor-roborates this result.

Fewer species were recorded in HI (104species) than in FA (129 species) although theareas are similar (60 ha and 56 ha respec-tively). The degree of isolation of an island (orforest fragment) has been considered animportant impediment to colonization (Mac-Arthur & Wilson 1967). In the present studyHI is the most isolated forest fragment, whichwould explain the low number of species (andthe lowest similarity index to PG, Sørensonindex = 0.64). In addition, correlationbetween number of species and area was onlyseen when HI was not considered; the othersfragments are much closer to each other.

Absence of competitors has been consid-ered an important factor in density compen-sation (MacArthur et al. 1972, Wright 1980,Blondel 1991). It was tested using the groupsof species with similar feeding habits. In thepresent study, leaf insectivores decreased inspecies number with decrease in area; the sig-nificant increase in relative abundance of theleaf insectivores Synallaxis ruficapilla and Dys-ithamnus mentalis with decrease in area couldbe related to the reduced competition in thesmall fragments. However, five other leafinsectivores (Thamnophilus doliatus, Thamnophi-lus caerulescens, Tolmomyias sulphurescens, Myiornis

TABLE 3. Species occurring in three, four or fiveforest fragments that increased significantly theirrelative abundance with a decrease in area (A), andspecies that had the highest relative abundance inHI (B). In both cases χ2 tests (P < 0.05) wereused.

Species χ2; df TendencyCrypturellus tataupaCrypturellus parvirostrisColumba cayennensisLeptotila verreauxiPhaetornis eurynomeDryocopus lineatusPicumnus temminckiiSynallaxis ruficapillaXenops rutilansMackenziaena severaThamnophilus doliatusThamnophilus caerulescensDysithamnus mentalisConopophaga lineataPsilorhamphus guttatusScytalopus indigoticusMegarhynchus pitanguaMyiodynastes maculatusPitangus sulphuratusEmpidonax euleriTolmomyias sulphurescensMyiornis auricularisLeptopogon amaurocephalusCyanocorax chrysopsTurdus rufiventrisTurdus leucomelasCyclarhis gujanensisConirostrum speciosumEuphonia chloroticaThraupis sayacaTrichothraupis melanopsCissops leverianaSaltator similisPitylus fuliginosusArremon flavirostris

37.0; 414.5; 418.5; 445.3; 410.7; 495.9; 317.0; 373.8; 49.00; 322.3; 450.8; 240.0; 414.0; 462.2; 462.8; 319.4; 332.4; 415.9; 428.7; 482.8; 316.6; 460.7; 411.7; 412.7; 340.1; 429.2; 440.7; 420.1; 412.0; 322.6; 416.5; 418.1; 417.2; 411.7; 311.6; 2

BBABBABABABBAAAAAAABBBBAAABBBBAAAAB

18


auricularis, and Leptopogon amaurocephalus) hadthe highest relative abundance in HI whichpresented similar number of leaf insectivoresto PG. Eight herbivores/omnivores alsoincreased relative abundance with decrease inarea (Megarhynchus pitangua, Myiodynastes macula-tus, Cyanocorax chrysops, Turdus rufiventris, Turdusleucomelas, Trichothraupis melanops, Saltator similis,and Pitylus fuliginosus) but the species numbersof this group did not seem to decrease witharea size. Ricklefs & Cox (1978) and Blondel(1991) have suggested slight habitat differ-ences and differences in colonization, besidereduced competition, as additional reasonsfor density compensation which have to beconsidered for the results obtained in thepresent study.

Trunk insectivores had similar speciesnumbers in the studied fragments but the rel-ative abundance decreased substantially withthe decrease in area in the closer fragments(PG, FA, FB, and FC), and was the lowest inthe isolated fragment HI. This may suggestlow populations for some species. In threeother fragments of 100 ha, 86 ha, and 12 hain the studied region, which are more isolatedthan HI, only five, seven, and one species oftrunk insectivores were respectively recorded(Soares & Anjos 1999).

Some species of frugivorous birds werenot mist-netted in the small fragments ofAmazonia monitored after isolation duringthe Minimum Critical Size of EcosystemsProject; the capture numbers of some spe-cies, however, increased in isolated fragmentsof 1 ha (Bierregaard & Stouffer 1997). Similarpattern was obtained in this study; frugivoresdecreased in species numbers with decreasein area, and were represented by few speciesin the isolated fragment HI. But the frugivo-rous Euphonia chlorotica and Arremon flavirostrishad the highest relative abundance in the iso-lated fragment HI.

Nectarivores were not considered vulner-able to fragmentation in Amazonia because

the species had equal proportion of capturesor even increased them in fragments after iso-lation (Stouffer & Bierregaard 1995a). Nec-tarivores in the studied fragments in thenorth of Paraná tended to increase in totalrelative abundance with decrease in area, andpresented the highest value in the isolatedfragment HI (Table 2).

Although some general tendencies werepresented for certain bird groups, the effectsof forest fragmentation are certainly differentfor each species. A study in 270 forest frag-ments in Maryland suggested that theimpacts of forest fragmentation on bird com-munities are complex, species specific andnot related only to fragment area or fragmentisolation (Lynch & Whigham 1984).

FB was the forest fragment most similarto PG when the total relative abundance ofthe species grouped according to their feed-ing habits was considered. FA had a lowersimilarity in relation to the community struc-ture of PG in spite of its larger size andgreater species number. The forest corridorbetween PG and FB may have enabled agreater similarity in the community structureand isolation seems to have importance forthe low similarity of FA. However, the impor-tance of forest corridors for movement oforganisms is yet controversial because of theweakness of available data (Simberloff et al.1992). Crome (1997) comparing several for-est fragmentation studies suggested that eachlandscape appears unique in its pattern offragmentation. So, it is too early for broadgeneralizations of the data presented in thisstudy.

ACKNOWLEDGMENTS

This study was funded by CNPq (NationalResearch Council, Brasília) and by FINEP(National Funding for Studies and Projects,Rio de Janeiro). This work is part of a largerenviromental plan (Tibagi Project) and also

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had the financial support of the UniversidadeEstadual de Londrina, the Consórcio Inter-municipal para Proteção Ambiental do rioTibagi and the Klabin Fabricadora de Papel eCelulose. I also thank the Productivity inResearch grant from CNPq (350054/95-9).A. R. J. Ferreira, V. F. Nassaro, M. Rodrigues,R. A. Depieri and E. R. Barbosa helped infield work. R. A. Depieri helped in thegraphic presentation of the data. A. H. F. deToledo prepared an early English version ofthis manuscript. M. C. Dias helped me withthe vegetation description. W. Belton madeimportant suggestions and helped in the last-est English version of this manuscript. Com-ments of two anonymous reviewers improvedsubstantially the manuscript.

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21

ANJOS

APPENDIX. Relative abundance of bird species recorded in the studied forest fragments (consideringonly the sampled area of PG); (*) indicates the 10 species with greatest relative abundance values in eachforest fragment and (s) indicates significant difference (χ2, P < 0.05) between number of contacts whenspecies is registered in 3, 4 or 5 forest fragments. Feeding habits are shown for each species: NE, nectari-vore; FR, frugivore; HE, herbivore; TI, trunk insectivore; LI, leaf insectivore; GI, generalized insectivore;OM, omnivore; CA, carnivore; HO, herbivore/omnivore.

Families and species Forest fragments-ha Feeding habits

PG-70 HI-60 FA-56 FB-25 FC-11TINAMIDAE

Tinamus solitarius Crypturellus obsoletus (s) Crypturellus undulatus Crypturellus parvirostris (s)Crypturellus tataupa (s)

ACCIPITRIDAEHarpagus diodon Ictinia plumbea Buteo magnirostris Buteo leucorrhous Buteo nitidus

FALCONIDAEHerpetotheres cachinnans (s) Micrastur semitorquatus Micrastur ruficollis

CRACIDAEPenelope superciliaris Pipile jacutinga Crax fasciolata

PHASIANIDAEOdontophorus capueira

RALLIDAEAramides cajanea Aramides saracura

COLUMBIDAEColumba speciosa Columba picazuro Columba maculosa Columba cayennensis (s)Columba plumbea Zenaida auriculata Columbina minuta Columbina talpacoti Columbina picui Claravis pretiosa Scardafella squammata Leptotila verreauxi (s)Leptotila rufaxilla

0.220.62 * 0.050.050.07

0.250.150.1

0.070.020.02

1.7 *

0.270.02

0.47 0.32

0.1

0.270.7 *

0.02

0.07

0.02

0.02

0.17

0.10

0.021.7 *

0.75 *

0.140.020.220.090.03

1.15 *0.44

0.4

0.170.48

0.02

0.020.02

0.08

0.05

0.08

0.02

0.050.03

0.071.93 * 0.02

0.73 *0.02

0.40.37

0.4

0.050.32

0.09

0.07

0.02

0.21

0.02

0.05

1.91 *

0.54

0.02

0.04

0.020.530.37

0.26

0.160.68

0.05

0.11

0.02

0.25

0.02

0.18

2.07 *0.050.81*

0.330.37

HOHOHOHOHO

OMGIOMCACA

CACACA

FRFRHE

HE

HOHO

FRHEHEFRFRHEHEHEHEHEHEHOHO

22


APPENDIX. Continuation.


PG-70 HI-60 FA-56 FB-25 FC-11Geotrygon montana

PSITTACIDAEAratinga leucophthalmus (s) Aratinga solstitialis Pyrrhura frontalis (s)Forpus xanthopterygius Brotogeris tirica (s)Pionopsitta pileata Pionus maximiliani Amazona aestiva Triclaria malachitacea (s)

CUCULIDAEPiaya cayana Crotophaga ani

STRIGIDAEPulsatrix perspicillata Pulsatrix koeniswaldiana Glaucidium brasilianum Ciccaba virgata Strix hylophila

NYCTIBIIDAENyctibius griseus

CAPRIMULGIDAELurocalis semitorquatus

TROCHILIDAEPhaethornis eurynome (s)Anthracothorax nigricollis Chlorostilbon aureoventris Leucochloris albicollis Amazilia fimbriata

TROGONIDAETrogon viridis (s)Trogon surrucura (s)

MOMOTIDAEBaryphthengus ruficapillus

BUCCONIDAENotharcus macrorhynchus

RAMPHASTIDAEPteroglossus aracari Selenidera maculirostris (s)Baillonius bailloni Ramphastos dicolorus

PICIDAEPicumnus temminckii (s)

0.3

0.53 0.17

0.87 *0.030.250.13

0.78 * 0.050.17

0.5

0.020.020.18

0.02

0.1

0.20.03

0.30.62

0.6

0.02

0.070.7 *0.050.12

0.15

0.22

0.020.03

0.53

0.02

0.02

0.03

0.390.150.100.02

0.02

0.02

0.54

0.37

0.12

0.30.050.030.030.53 0.050.02

0.60.02

0.1

0.02

0.12

0.15

0.020.020.02

0.150.58

0.4

0.37

0.03

0.5

0.25

0.11

0.420.040.090.12

0.84 *0.05

0.74

0.02

0.14

0.11

0.190.040.050.04

0.090.6

0.81 *

0.070.68

0.07

0.28

0.28

0.07

0.370.040.050.110.670.090.02

0.70

0.04

0.09

0.160.070.02

0.070.3

0.6

0.28

0.02

HO

HEHEHEHEHEHEHEHEHE

GIHO

CACACACACA

GI

GI

NENENENENE

HOHO

HO

HO

FRFRFRFR

TI

23

ANJOS



PG-70 HI-60 FA-56 FB-25 FC-11Picumnus cirratus Picumnus albosquamatusChrysoptilus melanochloros (s) Piculus aurulentus Celeus flavescens (s)Dryocopus lineatus (s) Melanerpes flavifrons (s)Leuconerpes candidus Veniliornis spilogaster Phloeoceastes melanoleucos

DENDROCOLAPTIDAEDendrocincla turdina Sittasomus griseicapillus (s)Xiphocolaptes albicollis (s) Dendrocolaptes platyrostris (s) Lepidocolaptes angustirostris Lepidocolaptes fuscus (s)

FURNARIIDAEClibanornis dendrocolaptoides Furnarius rufus Synallaxis ruficapilla (s)Synallaxis frontalis Synallaxis spixi Synallaxis cinerascens Cranioleuca obsoleta Syndactyla rufosuperciliata Anabacerthia amaurotis Philydor lichtensteini Philydor rufus (s)Automolus leucophthalmus Heliobletus contaminatus Xenops rutilans (s)Sclerurus scansor

FORMICARIIDAEHypoedaleus guttatus Mackenziaena leachii Mackenziaena severa (s)Thamnophilus doliatus (s)Thamnophilus caerulescens (s) Thamnophilus ruficapillus Dysithamnus mentalis (s)Herpsilochmus rufimarginatus Drymophila malura Pyriglena leucoptera

0.22

0.150.070.55

0.30.03

0.11.28 * 0.43

0.85 *

0.4

0.02

0.20.050.020.020.050.08

0.030.070.050.180.120.03

0.93 *

0.18

0.15

0.550.080.020.52

0.170.050.070.030.02

0.14

0.090.030.19

0.02

0.030.050.02

0.02

0.020.030.10.2

0.030.410.64

0.88 *0.100.420.14

0.03

0.080.030.020.030.08

0.18

0.030.480.270.62 0.020.27

1.03 *

0.020.02

0.020.030.17

0.20.020.02

0.98 *

0.7 *

0.68

0.77 * 0.05

0.82 *

0.3

0.18

0.16

0.16

0.070.75 0.251 *0.09

0.65

0.02

0.04

0.190.12

1.32 *

0.650.040.320.02

0.98 *

0.9 *

0.020.050.050.18

0.75 *0.040.020.14

0.210.040.510.070.25

0.79 *0.040.02

0.02

0.11

1.14 *0.020.7 0.110.51

0.75 * 0.02

0.82 *

TITITITITITITITITITI

TITITITITITI

GIGILILILILITILILILILILITITI

OM

OMOMOMLILILILILIGIOM

24




PG-70 HI-60 FA-56 FB-25 FC-11Chamaeza campanisona Grallaria varia Conopophaga lineata (s)

RHINOCRYPTIDAEPsilorhamphus guttatus (s) Scytalopus indigoticus (s)

COTINGIDAEPachyramphus viridis Pachyramphus castaneus Pachyramphus polychopterus Pachyramphus validus Tityra cayana (s)Tityra inquisitor

PIPRIDAEManacus manacus Chiroxiphia caudata (s)Piprites chloris (s)Schiffornis virescens

TYRANNIDAEColonia colonus Sirystes sibilator (s)Tyrannus melancholicus Empidonomus varius Megarhynchus pitangua (s)Myiodynastes maculatus (s)Myiozetetes similis Pitangus sulphuratus (s)Myiarchus swainsoni Empidonax euleri (s)Cnemotriccus fuscatus Myiophobus fasciatus Platyrinchus mystaceus Tolmomyias sulphurescens (s) Todirostrum cinereum Todirostrum plumbeiceps Idioptilon nidipendulum Myiornis auricularis (s)Hemitriccus obsoletus (s)Phylloscartes eximius Phylloscartes ventralis Capsiempis flaveola (s) Serpophaga subcristata Elaenia flavogaster Elaenia mesoleuca

0.420.320.05

0.080.03

0.020.03

0.080.370.03

0.050.420.430.02

0.080.450.02

0.380.070.020.07

0.05

0.02

0.07

0.030.10.23

0.020.150.05

0.02

0.10

0.020.360.07

0.02

0.050.02

0.020.03

0.370.29

0.370.050.8 *0.030.03

0.250.05

0.83 *0.02

0.140.02

0.02

0.65

0.470.25

0.070.050.230.1

0.020.22

0.050.020.020.030.320.250.020.4

0.18

0.080.020.220.050.25

0.230.23

0.170.23

0.02

0.12

0.020.02

0.05

0.160.160.02

0.120.21

0.090.050.041 *0.260.020.67

0.05

0.020.020.02

0.330.02

0.180.11

0.51

0.75 *0.19

0.05

0.160.07

0.14

0.470.460.020.580.16

0.02

0.160.12

0.050.02

0.040.020.020.02

HOOMGI

GIGI

HOHOHOHOHOHO

OMFRFRHO

GIGIGIGIHOHOHOOMGIGIGIGILILILILILILILILILILILI

HOHO

25




PG-70 HI-60 FA-56 FB-25 FC-11Myiopagis caniceps (s)Myiopagis viridicata Camptostoma obsoletum Leptopogon amaurocephalus (s)Pipromorpha rufiventris (s)Corythopis delalandi

OXYRUNCIDAEOxyruncus cristatus

CORVIDAECyanocorax chrysops (s)

TROGLODYTIDAETroglodytes aedon

TURDIDAETurdus nigriceps Turdus rufiventris (s)Turdus leucomelas (s)Turdus amaurochalinus Turdus albicollis (s)

SYLVIIDAEPolioptila lactea

VIREONIDAECyclarhis gujanensis (s)Vireo olivaceus Hylophilus poicilotis

ICTERIDAECacicus haemorrhous (s)

PARULIDAEParula pitiayumi (s)Basileuterus culicivorus (s) Basileuterus leucoblepharus

COEREBIDAECoereba flaveola Conirostrum speciosum (s)

THRAUPIDAEEuphonia musica Euphonia chlorotica (s)Euphonia violacea Tangara cayana Thraupis sayaca (s)Habia rubica (s)Tachyphonus coronatus Trichothraupis melanops (s)Hemithraupis guira (s)Orchesticus abeillei

0.7 *0.020.070.180.3

0.05

0.15

0.050.130.030.12

0.02

0.55

0.22

0.271.12 * 0.08

0.35

0.020.050.02

0.030.380.20.350.48

0.29

0.100.46

0.7

0.02

0.240.59 0.100.03

1.78 *0.03

0.83 *

0.02

0.171.41 *

0.41

0.2

0.250.10

0.090.51

0.13

0.080.250.08

0.02

0.4

0.030.250.120.10.02

1.4 *

0.02

0.03

0.071.25 *0.53

0.15

0.030.03

0.020.120.130.380.330.02

0.12

0.160.260.02

0.54

0.02

0.030.670.440.210.02

1.05 *

0.18

0.07 1.4 * 0.04

0.020.11

0.070.09

0.190.250.070.510.21

0.16

0.040.18

0.32

0.040.230.420.11

1.33 *

0.07

0.041.23 *0.33

0.14

0.020.020.070.180.160.440.3

LILILILILILI

FR

HO

LI

HOHOHOHOHO

LI

HOLI

HO

HO

LILIGI

NEHO

FRFRFRHOHOHOHOHOHOHO

26



PG-70 HI-60 FA-56 FB-25 FC-11Cissopis leveriana (s)

FRINGILLIDAESaltator similis (s)Pitylus fuliginosus (s)Arremon flavirostris (s)

0.02

0.070.1

0.03

0.36

0.15

0.2

0.380.37

0.16

0.390.280.02

0.25

0.190.160.02

OM

HOHOFR

27

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