ORNITOLOGIA NEOTROPICAL 15: 71–85, 2004 © The Neotropical Ornithological Society

SPATIAL DISTRIBUTION OF ON THREE ISLANDS IN THE UPPER RIVER PARANÁ, SOUTHERN

Márcio Rodrigo Gimenes1 & Luiz dos Anjos2

1Universidade Estadual de Maringá, Pós-Graduação em Ecologia de Ambientes Aquáticos Continentais (PEA), Av. Colombo 5790, Maringá, CEP 87020-900, Paraná, Brasil. E-mail: [email protected] 2Universidade Estadual de Londrina, Depto Biologia e Vegetal, CX 6001, Londrina, CEP 86051-970, Paraná, Brasil. E-mail: [email protected]

Resumo. – Distribuição espacial de aves em três ilhas no alto rio Paraná, Sul do Brasil. – O número e a composição das espécies de aves em três ilhas no alto rio Paraná foram comparados. As ilhas Porto Rico (103 ha) e Mutum (976 ha) sofreram forte desflorestamento, enquanto a ilha Bandeira (14 ha) foi leve- mente desflorestada. Cinco ambientes foram reconhecidos nas ilhas: florestas, zonas arbustivas, campos abertos, zonas aquáticas e bancos de areia. Um transecto incluindo todos estes ambientes sobre as ilhas foi percorrido mensalmente de Dezembro de 1999 a Setembro de 2000. Foram registradas 113 espécies de aves nas três ilhas: 99 na ilha Mutum, 86 na ilha Bandeira e 82 na ilha Porto Rico. A melhor explicação para o fato do número de espécies da ilha Bandeira ser similar ao das outras ilhas, apesar da sua pequena área, possivelmente é a presença de uma floresta contínua e pouco perturbada sobre a ilha Bandeira, ao contrá- rio das ilhas Mutum e Porto Rico, onde as florestas são fragmentadas. Ilhas bem preservadas, mesmo que pequenas, devem ter prioridade em programas conservacionistas. Abstract. – The number and composition of species on three islands in the upper Paraná river, Brazil, were compared. Porto Rico island, 103 ha, and Mutum island, 976 ha, are highly deforested; Bandeira island, 14 ha, is slightly deforested. Five habitats have been recognized: forests, shrubs, open fields, aquatic areas and sand bars. A transect passing through all the above habitats on the islands was sampled monthly, from December 1999 to September 2000. Birds on the three islands totaled 113 species: 99 on Mutum, 86 on Bandeira, and 82 on Porto Rico islands. The high number of species on Bandeira island, taken its small size, is probably due to the presence on it of a continuous and little disturbed forest. On the contrary, for- ests on Mutum and Porto Rico islands are fragmented. Well preserved islands, albeit small in size, should have priority in conservation programs. Accepted 26 May 2003. Key words: Paraná river; islands; birds; conservation; southern Brazil.

INTRODUCTION discharge in the world, and second in basin drainage in (2,800,000 km2), is The semideciduous forests of southern Bra- approximately 3800 km long (Maack 1981). zil, as part of the Atlantic forest, present a The Paraná river basin has suffered strong high number of Neotropical endemic bird human impact (dams building and deforesta- species, particularly forest species, but have tion). The 250-km river stretch between the undergone such intense fragmentation as to mouth of the Paranapanema river (its main make them among the most endangered zoo- affluent) and the town of Guaíra PR, Brazil, geographical regions in South America (Stotz in the upper Paraná river is the only segment et al. 1996). The Paraná river, ranking tenth in within Brazilian territory which is free from

71 GIMENES & DOS ANJOS

53º 20’ 53º 10’

’

’

0

South America 0

4

4

º

º

2

2 2 r e 2 iv R Mato Grosso do Sul ía a State B

r e iv R r ía r e a ve iv B Ri R á land á aran Is an P ar a Chann P ub el m rut utu Cu M Araçatuba Channel Bandeira Island Porto Rico Island

N

Paraná State

,

’

0

0 0 1 2 3 4 5km

5

5

º

º

2

2

Scale 2

2

53º 20’ 53º 10’ FIG. 1. Paraná river stretch with Mutum, Porto Rico and Bandeira islands. dams. On the right bank (Mato Grosso do Sul Martin 1981, Wright et al. 1985, Wiens 1989, State) of this stretch of the river, there is a Bierregaard 1990, Anjos & Boçon 1999) have fairly well preserved floodplain. On the left shown that larger islands or larger forest frag- bank (Paraná State), with a higher land eleva- ments support higher bird species richness tion, semideciduous forests have almost all than smaller ones. However, other studies been transformed into pasture lands. have shown that habitat diversity (Williams More than one hundred islands, ranging 1964, Boecklen 1986, Martin et al. 1995) and from less than 10 ha to approximately 2000 vegetation disturbance (Anjos & Seger 1988, ha, are distributed over this river stretch Aleixo & Vielliard 1995, Restrepo et al. 1997) (Stevaux et al. 1997). On many islands, where also have a strong influence on bird species forests once covered the larger part of the richness. Therefore, the islands of the Paraná land, and other natural habitats covered very river are appropriate for the study of the small areas, intensive deforestation has cre- influence of area, habitat diversity and vegeta- ated fragmented landscapes (Souza et al. tion disturbance on bird species richness. 1997). However, a few small islands still main- Few ornithological studies have been con- tain continuous non-fragmented forest cover. ducted in the region. Anjos & Seger (1988) The island biogeography theory (MacArthur recorded 133 species in about 100 ha of main- & Wilson 1967) and posterior studies (Moore land and islands. Straube et al. (1996) and & Hooper 1975, Forman et al. 1976, Galli et al. Straube & Urben-Filho (2002) provided a 1976, Connor & McCoy 1979, Willis 1979, checklist of birds on the islands and associ-

72 BIRDS OF ISLANDS OF UPPER PARANÁ RIVER, BRAZIL

FIG. 2. Habitats on the studied islands, Paraná River, south Brazil. The square indicates sampled area of Mutum island. ated floodplain between the mouth of the STUDY AREA AND METHODS Paranapanema river and Guaíra with a total of some 298 species. However, there is no long Mutum (976 ha), Porto Rico (103 ha) and term study encompassing the bird richness of Bandeira (14 ha) islands lie in the upper the islands and the floodplain of the region. Paraná river (between 22°44’S and 22°48’S This study compares the number and compo- and 53°21’W and 53°15’W), at an altitude of sition of bird species on three islands on the 230 m a.s.l. (Fig. 1). According to the Köep- upper Paraná river. pen’s system, the region’s climate is classified

73 GIMENES & DOS ANJOS

TABLE 1. Area (ha) and percentage of habitats on the islands.

Forests Shrubs Open fields Aquatic habitats Sand bars Total (ha) Mutum (total) 205 625 127 19 — 976 (21%) (64%) (13%) (2%) Mutum (sampling) 40 115 30 15 — 200 (20%) (57.5%) (15%) (7.5%) Porto Rico 8 79 13 3 — 103 (7.7%) (76.7%) (12.6%) (3%) Bandeira 9 1 0.4 0.3 3.3 14 (64.3%) (7.1%) (2.8%) (2.1%) (23.7%) as Cfa (tropical-subtropical) with an average sis (Mimosaceae), Peschiera australis (Apocy- annual temperature of 22°C (summer average naceae) and Ficus obtosiuscula (Moraceae); and 26°C, and winter average 17°C), and an aver- (5) sand bars, present only on Bandeira island age annual rainfall of 1500 mm (Eletrosul (Table 1). 1986). The islands are made of clayey sedi- Samplings were conducted from Decem- ments, clayey-sandy sediments and sand ber 1999 to September 2000, and each island deposited by the river channel (Stevaux et al. was visited monthly. One sampling is equal to 1997). The area lies within the phytoecologi- one day of field work in which the transect cal region of semideciduous seasonal forest was ran on one island (total of 10 samplings with the local vegetation classified as alluvial on each island). Once a month, a standard semideciduous seasonal forest (Souza et al. transect was ran on each island in the same 1997). Mutum and Porto Rico islands have direction throughout all pre-established habi- been heavily deforested, leaving only small tats. Samplings always began one hour after forest fragments. Bandeira island is slightly sunrise, with an 8-h duration on each island. disturbed, but most of its area still has contin- The transects sampled all areas of the Ban- uous forest (Fig. 2). deira and Porto Rico islands. On Mutum Five habitat categories are found on the island, the transect comprised approximately islands (Souza et al. 1997): (1) open fields, 200 ha and all habitats were covered. Two without any or with low vegetation cover of other visits were made (in 2 days, totaling plant, such as Panicum, Setaria, Paspalum eight hours) to sections of the island not cov- (Poaceae), Cyperus (Cyperaceae) species, Lippia ered by the transect. Since no bird species was alba (Lamiaceae) and Ageratum conyzoides registered which had not already been (Asteraceae); (2) shrubs, with the predomi- detected along the transect, we believe the nance of Palicourea crocea (Rubiaceae), Cordia sampled area is representative of the Mutum monosperma (Boraginaceae) and Solanum evony- island. The observer (MRG) identified each moides (Solanaceae); (3) aquatic areas (lakes, bird encountered along the transect using bin- backwaters and bogs), with Eichhornia crassipes, oculars, and recorded the habitat in which Eichhornia azurea (Pontederiaceae), Salvinia each bird was sighted or heard. Each species auriculata (Salviniaceae) and Polygonum acumina- was registered only once in each habitat on a tum (Polygonaceae); (4) forests, whose most single sampling day, thus, abundance informa- common tree species are Cecropia pachystachya tion is not available. (Cecropiaceae), Croton urucurana (Euphorbi- The frequency of occurrence in percent- aceae), Celtis iguanaea (Ulmaceae), Inga uruguen- age was calculated for of each species 1) at the

74 BIRDS OF ISLANDS OF UPPER PARANÁ RIVER, BRAZIL

A 110 100 90 80 70 Mutum 60 Bandeira 50 40 Porto Rico 30

Number ofNumber species 20 10 0 DJFMAMJJAS Months

B

100 More than 75% 80 25.1% to 75% Up to 25% 60

40

20 Numberof species (%) 0 Mutum Porto Rico Bandeira

FIG. 3. Monthly cumulative number of bird species on each island (A), and number of species recorded during up to 25%, between 25.1 and 75%, and more than 75% of samplings for each island (B). island level, as the number of samplings (or was recorded on that island, and multiplied by days) in which each species was recorded on a 100. Analysis of variance (one way ANOVA, P particular island, divided by 10 (total number < 0.05) was used to evaluate whether there of samplings or days of field work for each was any significant difference between aver- island) and multiplied by 100, and 2) at the age numbers of species on the islands. Chi- habitat level, as the number of samplings (or square and contingency table analyses (α = days) in which each species was recorded in 0.05) determined whether there was any sig- one particular habitat on one island, divided nificant difference in the numbers of species by number of samplings (or days) in which it recorded during up to 25%, between 25.1% to

75 GIMENES & DOS ANJOS

75%, or more than 75% of samplings on each species (χ2 = 6.89, df = 3, P > 0.05), contrary island or habitat.. Similarity regarding the bird to Porto Rico χ2 = 19.66, df = 3, P < 0.05) species composition of the islands and habi- and Bandeira χ2 = 47.98, df = 4, P < 0.05) tats was estimated by use of the qualitative islands (see Appendix 1). Forests also had the Sørensen’s index: 2j/(a + b), where “j” is the highest number of exclusive species (21), fol- number of species common to both sites, “a” lowed by aquatic areas (11), shrubs (3), and is the number of species in site “A”, and “b” is sand bars (2). Highest similarity values were the number of species in site “B”. verified between the forests and shrubs, and between the shrubs and open fields (Table 2). RESULTS Highest similarity values among the habitats of a same island was obtained between forests Number of species on the islands. Birds recorded and shrubs of Mutum and Porto Rico while, on the three islands totaled 113 species: 99 on on Bandeira, the highest value was between Mutum, 86 on Bandeira and 82 on Porto Rico shrubs and open fields. The forests of Ban- (Appendix 1). These values are statistically deira were less similar to shrubs and open similar (F = 2.43, df = 2, P > 0.05), even fields than on the other two islands (Table 2). excluding the 12 species which occurred only The highest species richness of forests on the sand bars of Bandeira island (F = 3.22, was recorded on Bandeira island (Appendix df = 2, P > 0.05). Therefore, although the 1). Bandeira was the only island for which sampling area of each island was different, species richness of forests was significantly species richness was similar. On all islands, greater than that of the habitat with the sec- species-accumulation curves reached an ond highest species number (χ2 = 4.45, df = asymptotic plateau (Fig. 3A). The frequency 1, P < 0.05). The same island had the highest of occurrence of species was statistically proportion of species recorded in more than similar (χ2 = 2.11, df = 4, P > 0.05) on the 75% of samplings, and the least proportion three islands (Fig. 3B). Highest similarity val- occurring in no more than 25% of forest sam- ues occurred between Mutum and Porto Rico plings (Fig. 4). Based on contingency table, (0.84), followed by Mutum and Bandeira the species richness of Bandeira and Porto (0.81) and Porto Rico and Bandeira (0.76). Rico forests appeared as high as that of Excluding the 12 species which occurred only Mutum forests, in spite of the fact that forests on the sand bars of Bandeira island, the simi- are smaller on Bandeira and Porto Rico (χ2 = larity value between Mutum and Bandeira 14.08, df = 1, P < 0.05 and χ2 = 12.71, df = 1, decreased to 0.75, while it increased to 0.78 P < 0.05, respectively). When the forests of between Porto Rico and Bandeira. the Bandeira and Porto Rico islands are taken into account, species richness appears related Species richness according to habitats. The fre- to the size of the forest (χ2 = 0.0001, df = 1, quency of occurrence of species differed sig- P > 0.05). nificantly between habitats, on Mutum (χ2 = Highest species richness in shrubs and 35.8, df = 6, P < 0.05), Porto Rico (χ2 = 16.6, open fields were recorded on Mutum and df = 6, P < 0.05) and Bandeira islands (χ2 = Porto Rico islands (Appendix 1). However, 42.6, df = 6, P < 0.05) (Fig. 4). On all three considering the size of the area occupied by islands, the forests presented the highest shrubs on each island, species richness of number of bird species (Appendix 1). How- Bandeira shrubs resulted higher when com- ever, the four habitat categories of Mutum pared to Mutum (χ2 = 57.22, df = 1, P < tended to have roughly the same number of 0.05) and Porto Rico (χ2 = 40.61, df = 1, P <

76 BIRDS OF ISLANDS OF UPPER PARANÁ RIVER, BRAZIL

M utum island

100 90 More than 75% 80 25 .1 % to 75 % 70 Up to 25% 60 50 40 30 20

Number of species (%) species of Number 10 0 FO SH OF AQ

Porto Rico island

100 90 More than 75% 80 25.1% to 75% 70 Up to 25% 60 50 40 30 20

Number of species (%) species of Number 10 0 FO SH OF A Q

Bande ira island

100 90 More than 75% 80 25.1% to 75% 70 U p to 2 5% 60 50 40 30 20

Number of species (%) species of Number 10 0 FO SH OF AQ SB

FIG. 4. Number of species recorded during up to 25%, between 25.1 and 75%, and more than 75% of samplings of each habitat for Mutum, Porto Rico and Bandeira islands. FO: forests, SH: shrubs, OF: open fields, AQ: aquatic habitats, and SB: sand bars.

0.05) ones. The same occurred with the small Nevertheless, the use of shrubs and open open fields of Bandeira. In spite of the fact fields on Bandeira island, based on the fre- that these habitats are much smaller on Ban- quency of occurrence of species during the deira island, the number of species did not samplings, was significantly less than that of decrease significantly. forests of the same island (χ2 = 29.52, df = 2

77 GIMENES & DOS ANJOS

TABLE 2. Similarity index among different habitats.

Forests Shrubs Open fields Aquatic habitats Sand bars All three islands Forests 0.72 0.56 0.21 0.21 Shrubs 0.72 0.22 0.26 Open fields 0.20 0.28 Aquatic habitats 0.54 Bandeira island Forests 0.62 0.40 0.05 0.22 Shrubs 0.64 0.12 0.26 Open fields 0.06 0.27 Aquatic habitats 0.27 Mutum island Forests 0.73 0.56 0.08 Shrubs 0.71 0.16 Open fields 0.11 Porto Rico island Forests 0.69 0.52 0.13 Shrubs 0.66 0.14 Open fields 0.20

P < 0.05). The proportion of species which The total number of species on the three have been recorded during more than 75% of islands (total of 317 ha) is close to that samplings in shrubs and open fields was recorded by Anjos & Seger (1988) from a somewhat less on Bandeira than on the other much smaller area (133 species; 100 ha) in the two islands (Fig. 4). same geographic region. However, Anjos & The species number in aquatic areas seem Seger (1988) sampled mainly the mainland, to be related to the size of the areas, as the χ2 which commonly supports higher bird rich- test (Contingency table) showed when ness than islands in a same geographical Mutum was compared to Porto Rico (χ2 = region (MacArthur & Wilson 1967, Blondel 1.83, df = 1, P > 0.05), and as the small num- 1991). ber of species on Bandeira island demon- The three islands, despite their very differ- strated with its extremely restricted area of ent sizes, presented a similar number of spe- this habitat (Appendix 1). Higher proportion cies. When similar habitats were compared of species occurring in more than 75% of between the islands, the contingency table test samplings in this habitat was recorded on showed that the number of species did not Mutum island (Fig. 4). increase in accordance with the size of the habitat, except in aquatic environments. Vari- DISCUSSION ations in the number of species parallel to the size of an habitat were found particularly The richness of 113 bird species on the three when highly homogeneous forest fragments islands is much lower than the 298 species or islands were compared (Moore & Hooper recorded for the 250 km of river and flood- 1975, Forman et al. 1976, Galli et al. 1976, plains of the Upper Paraná river (Straube et al. Connor & McCoy 1979, Martin 1981, Anjos 1996, Straube & Urben-Filho 2002), possibly & Boçon 1999). However, when islands with because of the intense human disturbance different habitat diversity and vegetation dis- and lower natural habitat diversity on the turbance are compared, the influence of the islands than on the associated floodplains. size of an habitat area on the number of spe-

78 BIRDS OF ISLANDS OF UPPER PARANÁ RIVER, BRAZIL cies appears to decrease (Boecklen 1986, island, the forest is continuous and well pre- Martin et al. 1995, Warburton 1997). The fact served while, on the other two islands, it has that the three islands had similar frequency suffered intense fragmentation. Several stud- of occurrence of species is an indication ies have reported a decrease in bird species that the proportion of species either sporadi- richness following the fragmentation of con- cally and commonly recorded is about the tinuous forests due to low dispersion between same on the three islands. Mutum and Porto fragments (Simberloff & Abelle 1982, Bierre- Rico islands are very similar in terms of land- gaard 1990, Terborgh et al. 1997), microhabi- scape, presenting a mosaic of habitats. Ban- tat loss (Aleixo & Vielliard 1995, Restrepo et deira island, on the contrary, still has a al. 1997), food decrease (Willis 1979), increase continuous forest, and this difference in the in predation and competition (Wilcove & landscape possibly explains its lower similarity Robinson 1990), and edge effect (Lovejoy et in bird species composition with the other al. 1986, Wilcove et al. 1986, Bierregaard 1990, islands. Sisk et al. 1997). Forest specialist species are The most remarkable result of this study more susceptible to habitat fragmentation is the number of species on the very small (Blondel 1991) and, in sites where there is a Bandeira island being similar to that of the mosaic of small forest fragments and open other two islands. Williams (1964) stated that areas, generalists species predominate, as the increase in the number of species accord- commonly verified in southern Brazil ing to the size of an area depends on habitat (Gimenes & Anjos 2000, Krugel & Anjos diversity. Bandeira island has sand bards, an 2000, Anjos 2001). In summary, the forest on habitat type lacking on Mutum and Porto Bandeira island presented a richer and more Rico. Sand bars certainly contributed to the differentiated avifauna than the forests on the species richness of Bandeira. Indeed, on Ban- other islands, and thus resulted in higher over- deira, twelve species were recorded exclusively all richness of bird community on this island. on sand bars. However, 10 of these species As stated by Martin et al. (1995), habitat were recorded in other habitats of the other heterogeneity and forest integrity might be islands. Only Collared Plovers (Charadrius col- more important than the size of an area in laris) and Black Skimmers (Rynchops niger) predicting the number of species found on occurred exclusively on the sand bars. Exclud- islands. Small islands with almost unaltered ing these 12 species, richness remained statis- forests should have priority in conservation tically similar on the three islands. Therefore, programs over larger but extremely altered the species richness of Bandeira island might ones. be due to factors other than the presence of an extra habitat. ACKNOWLEDGMENTS Bird species richness increased in accor- dance with habitat succession, from sand bars, This study had the financial support of the to open fields, to shrubs, with a peak in the CNPq (National Research Council, Brasília) forests on all the islands, as verified by Will- and the Curso de Pós-Graduação em Ecolo- son (1974) in Champaign-Urbana, USA, and gia de Ambientes Aquáticos Continentais by Anjos et al. (1997) in the Tibagi River basin, (Universidade Estadual de Maringá). MRG Brazil. Forests also presented the highest received M.Sc. scholarship from CAPES and number of exclusive species. However, the LdA received a Research Productivity grant highest species richness in forests was from CNPq (350054/95-9). A. da Silva and S. encountered on Bandeira island. On this Rodrigues helped in field work. Comments of

79 GIMENES & DOS ANJOS two anonymous reviewers improved substan- Avian distribution patterns in forest islands of tially the manuscript. different sizes in central New Jersey. Auk 93: 356–365. Gimenes, M. R., & L. dos Anjos. 2000. Distri- REFERENCES buição espacial de aves em um fragmento flor- estal do campus da Universidade Estadual de Aleixo, A., & J. M. E. Vielliard. 1995. Composição Londrina, norte do Paraná, Brasil. Rev. Bras. e dinâmica da avifauna da mata de Santa Gene- Zool. 17: 263–271. bra, Campinas, São Paulo, Brasil. Rev. Bras. Krugel, M. M., & L. dos Anjos. 2000. Bird commu- Zool. 12: 493–511. nities in forest remnants in the city of Maringá, Anjos, L. dos. 2001. Bird communities in five Paraná State, southern Brazil. Ornitol. Atlantic forest fragments in southern Brazil. Neotrop. 11: 315–330. Ornitol. Neotrop. 12: 11–27. Lovejoy, T. E., R. O. Bierregaard, Jr., A. B. Rylands, Anjos, L. dos, & R. Boçon. 1999. Bird communi- J. R. Malcolm, C. E. Quintela, L. H. Harper, K. ties in natural forest patches in southern Brazil. S. Brown, A. H. Powell, G. V. N. Powell, H. O. Wilson Bull. 111: 397–414. Schubart, & M. B. Hays. 1986. Edge and other Anjos, L. dos, K.-L. Schuchmann, & R. Berndt. effects of isolation on Amazon forest frag- 1997. Avifaunal composition, species richness, ments. Pp. 257–285 in Soulé, M. E. (ed). Con- and status in the Tibagi river basin, Paraná servation biology: the science of scarcity and State, southern Brazil. Ornitol. Neotrop. 8: diversity. Sinauer Associates, Sunderland, Mas- 145–173. sachusetts. Anjos, L. dos, & C. Seger. 1988. Análise da distri- Maack, R. 1981. Geografia física do Estado do buição das aves em um trecho do rio Paraná, Paraná. Livraria José Olympio, Rio de Janeiro, divisa entre os Estados do Paraná e Mato Brasil. Grosso do Sul. Arq. Biol. Tecnol. (Curitiba) 31: MacArthur, R., & E. O. Wilson. 1967. The theory 603–612. of island biogeography. Princeton Univ. Press, Bierregaard, Jr., R. O. 1990. Avian communities in Princeton, New Jersey. the understory of Amazonian forest fragments. Martin, J.-L., A. J. Gaston, & S. Hitier. 1995. The Pp. 333–343 in Keast, A. (ed). Biogeography effect of island size and isolation and old and ecology of forest bird communities. SPB growth forest habitat and bird diversity in Academic Publishing, The Hague, The Nether- Gwaii Haanas (Queen Charlotte Islands, Can- lands. ada). Oikos 72: 115–131. Blondel, J. 1991. Birds in biological isolates. Pp. Martin, T. E. 1981. Limitation in small habitat 45–72 in Perrins, C. M., J. D. Lebreton, & G. J. islands: chance or competition. Auk 98: 715– M. Hirons (eds.). Bird population studies. 734. Oxford Univ. Press, Oxford, UK. Moore, N. W., & M. D. Hooper. 1975. On the Boecklen, W. J. 1986. Effects of habitat heteroge- number of bird species in British woods. Biol. neity on the species-area relationships of forest Conserv. 8: 239–250. birds. J. Biogeogr. 13: 59–68. Narosky, T., & D. Izurieta. 1987. Guia para la iden- Connor, E. F., & E. D. McCoy. 1979. The statistics tificación de las aves de y . and biology of the species-area relationship. Vazques Mazzini Editores, Buenos Aires, Am. Nat. 113: 791–833. Argentina. Eletrosul. 1986. Ilha Grande. Centrais Elétricas do Restrepo, C., L. M. Renjifo, & P. Marples. 1997. Sul do Brasil, Florianópolis, Brasil. Frugivorous birds in fragmented Neotropical Forman, R. T. T., A. E. Galli, & C. F. LECK. 1976. montane forests: landscape pattern and body Forest size and avian diversity in New Jersey mass distribution. Pp. 171–189 in: Laurance, W. woodlots with some land use implications. F., & R. O. Bierregaard, Jr. (eds.). Tropical for- Oecologia 26: 1–8. est remnants: ecology, management and con- Galli, A. E., C. F. Leck, & R. T. T. Forman. 1976. servation of fragmented communities. Univ.of

80 BIRDS OF ISLANDS OF UPPER PARANÁ RIVER, BRAZIL

Chicago Press, Chicago, Illinois. Tropical forest remnants: ecology, management Sick, H. 1997. Ornitologia brasileira. Editora Nova and conservation of fragmented communities. fronteira, Rio de Janeiro, Brasil. Univ. of Chicago Press, Chicago, Illinois. Simberloff, D., & L. G. Abelle. 1982. Refuge design Warburton, N. H. 1997. Structure and conservation and island biogeographic theory: effects of of forest avifauna in isolated rainforest rem- fragmentation. Am. Nat. 120: 41–50. nants in tropical Australia. Pp. 190–206 in Lau- Sisk, T. D., N. M. Haddad, & P. R. Ehrlich. 1997. rance, W. F., & R. O. Bierregaard, Jr. (eds.). Bird assemblages in patchy woodlands: model- Tropical forest remnants: ecology, management ing the effects of edge and matrix habitats. and conservation of fragmented communities. Ecol. Appl. 7: 1170–1180. Univ. of Chicago Press, Chicago, Illinois. Souza, M. C., J. Cislinski, & M. B. Romagnolo. Wiens, J. A. 1989. The ecology of bird communi- 1997. Levantamento florístico. Pp. 343–368 in ties. Cambridge Univ. Press, Cambridge, UK. Vazzoler, A. E. A. M., A. A. Agostinho, & N. S. WiIcove, D. S., C. H. McLellan, & A. P. Dobson. Hahn (eds.). A planície de inundação do alto rio 1986. Habitat fragmentation in the temperate Paraná: aspectos físicos, biológicos e socio- zone. Pp. 237–256 in Soulé, M. E. (ed). Conser- econômicos. Editora da Univ. Estadual de Mar- vation biology: the science of scarcity and ingá/Nupélia, Maringá, Brasil. diversity. Sinauer Associates, Sunderland, Mas- Stevaux, J. C., E. E. Souza-Filho, & I. C. Jabur. sachusetts. 1997. A história quaternária do rio Paraná em Wilcove, D. S., & S. K. Robinson. 1990. The impact seu alto curso. Pp. 47–72 in Vazzoler, A. E. A. of forest fragmentation on bird communities in M., A. A. Agostinho, & N. S. Hahn (eds.). A eastern North America. Pp. 319–331 in Keast, planície de inundação do alto rio Paraná: aspec- A. (ed). Biogeography and ecology of forest tos físicos, biológicos e socioeconômicos. Edi- bird communities. SPB Academic Publishing, tora da Univ. Estadual de Maringá/Nupélia, The Hague, The Netherlands. Maringá, Brasil. Williams, C. B. 1964. Patterns in the balance of Stotz, D. F., J. W. Fitzpatrick, T. A. Parker III, & D. nature. New York Academic Press, New York, K. Moskovits. 1996. Neotropical birds: ecology New York. and conservation. Univ. of Chicago Press, Chi- Willis, E. O. 1979. The composition of avian com- cago, Illinois. munities in remanescent woodlots in southern Straube, F. C., M. R. Bornschein, & P. Scherer- Brazil. Pap. Avulsos Zool. (São Paulo) 33: 1–25. Neto. 1996. Coletânea da avifauna da região Willson, M. F. 1974. Avian community organiza- noroeste do Estado do Paraná e áreas limítro- tion and habitat structure. Ecology 55: 1017– fes. Arq. Biol. Tecnol. (Curitiba) 39: 193–214. 1029. Straube, F. C., & A. Urben-Filho. 2002. Análise do Wright, S. J., J. Faaborg, & C. J. Campbell. 1985. conhecimento ornitológico da região noroeste Birds form tightly structured communities in do Paraná e áreas adjacentes. Cad. Biodivers. 3: the Pearl Archipelago, Panama. Pp. 798–812 in 4–11. Buckley, P. A., M. S. Foster, E. S. Morton, R. S. Terborgh, J., L. Lopez, J. Tello, D. Yu, & A. R. Ridgely, & F. G. Buckley (eds.). Neotropical Bruni. 1997. Transitory states in relaxing eco- ornithology. American Ornithologists Union, systems of land bridge islands. Pp. 256–274 in Washington, DC. Laurance, W. F., & R. O. Bierregaard, Jr. (eds.).

81 GIMENES & DOS ANJOS 70 20 20 20 10 10 40 30 20 20 10 20 30 20 90 FQ 100 50 50 SB 100 100 100 100 100 100 100 100 33.3 50 50 100 100 100 AQ 33.3 ng to Narosky & Yzu- & Narosky ng to : aquatic, and SB: sand and : aquatic, 50 i OF 11.1 Q Bandeira (14 ha) (14 Bandeira 50 50 SH 100 33.3 66.6

50 FO 100 100 66.6 66.6 60 60 10 30 50 10 40 10 20 30 30 40 10 30 20 FQ 100 100 50 100 100 100 100 100 100 100 100 100 100 100 AQ 66.6 OF 100 25 10 SH 66.6 Porto Rico (103 ha) (103 Rico Porto FO 100 100 100 33.3 33.3 50 10 60 80 70 30 20 40 10 10 40 50 20 60 50 40 30 50 40 60 10 10 30 70 90 FQ 100 50 80 80 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 AQ 33.3 33.3 20 90 OF 16.6 66.6 20 20 10 d nomenclature according to Sick (1997) and common names accord names (1997) and common to Sick according d nomenclature SH 100 Mutum (200 ha) 16.6 66.6 66.6 50 FO 100 100 66.6 66.6 ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) Nycticorax nycticorax Nycticorax ) ) ) ) ) ) ) Milvago chimachimaMilvago ) Dendrocygna viduata Tigrisoma lineatum Tigrisoma Phalacrocorax brasilianus Ardea cocoi ) Vanellus chilensis Vanellus Buteogallus urubitinga Buteogallus Platalea ajaja Polyborus plancus Polyborus Rupornis magnirostris Cairina moschata Cathartes aura Rostrhamus sociabilis Rostrhamus Amazonetta brasiliensis Butorides striatus Butorides Jacana jacana Ciconia maguari Coragyps atratus Egretta thula Geranospiza caerulescens Mycteria americana Mycteria Casmerodius albus Mesembrinibis cayennensis Mesembrinibis Anhinga anhinga Anhinga Aramus guarauna Aramus Jabiru mycteria Neotropic CormorantNeotropic ( Anhinga ( White-necked Heron ( EgretGreat ( EgretSnowy ( Heron Striated ( ( Night-Heron Black-crowned Rufescent Tiger-Heron ( Ibis ( Green Spoonbill( Roseate Wood-Stork ( ( Maguari Stork Jabiru ( ( Vulture Blake ( Vulture Turkey ( White-faced Tree-Duck ( Duck Muscovy ( Brazilian Duck Kite ( Everglade ( Hawk Roadside ( Hawk Black Great ( Crane Hawk ( Caracara Yellow-headed Crested Caracara ( Limpkin ( ( Jacana Wattled Southern( Lapwing APPENDIX 1. Frequency of occurrence percentages of bird species at the habitat level (FO: forest, SH: shrub, OF: open fields, A fields, open OF: shrub, SH: forest, (FO: level habitat at the species bird of percentages occurrence of Frequency 1. APPENDIX an on the (FQ) threeisland islands. and the bar) level rieta (1987).

82 BIRDS OF ISLANDS OF UPPER PARANÁ RIVER, BRAZIL 50 40 70 70 20 20 80 40 70 10 80 30 30 60 20 30 10 90 10 20 50 30 20 20 40 100 100 100 100 100 100 12.5 12.5 66.6 33.3 40 66.6 50 50 50 100 88.8 14.2 12.5 33.3 25 75 50 50 50 100 100 100 12.5 28.5 33.3 16.6 20 50 100 100 100 71.4 62.5 66.6 33.3 100 100 100 100 100 80 10 20 40 90 20 90 60 20 40 20 20 30 40 10 10 30 100 100 100 100 100 11.1 30 40 25 50 50 37.5 33.3 44.4 30 50 50 50 100 100 100 100 100 44.4 77.7 33.3 33.3 60 50 100 62.5 66.6 33.3 66.6 100 100 100 100 100 10 60 30 10 70 80 90 30 10 40 70 40 30 10 10 30 10 60 20 20 20 40 100 100 100 100 100 100 100 100 100 50 50 50 10 75 100 14.2 87.5 77.7 14.2 33.3 50 50 25 50 100 100 100 Mutum (200 ha) (200 Mutum ha) (103 Rico Porto ha) (14 Bandeira 28.5 55.5 33.3 42.8 66.6 90 50 20 50 50 FO SH OF AQ FQ FO SH OF AQ FQ FO SH OF AQ SB FQ 100 100 100 100 100 37.5 44.4 66.6 57.1 ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) Forpus xanthopterygius Galbula ruficauda Columbina talpacoti Columbina Picumnus cirratus Himantopus himantopus Himantopus Aratinga leucophthalmus Aratinga Columbina picui Columbina Pionus maximiliani Chloroceryle amazona Leptotila verreauxi Columba cayennensis Columba Crotophaga ani Sterna superciliaris Ceryle torquata Chloroceryle americana Tringa flavipes Phaetusa simplex Phaetusa Columba picazuro Hylocharis chrysuraHylocharis Phaethornis pretrei Speotyto cunicularia Tapera naevia Tapera Charadrius collaris Charadrius Rynchops niger Rynchops Caprimulgus parvulusCaprimulgus Guira guira Scardafella squammata Zenaida auriculata Nyctidromus albicollis White-barred Piculet ( Green Kingfisher ( Kingfisher Green ( Jacamar Rufous-tailed Collared Plover ( Collared Plover Lesser Yellowlegs ( Lesser Yellowlegs Eared Dove ( Eared Dove ( Scaled Dove ( White-tipped Dove ( White-eyed Parakeet Pale-vented Pigeon Pale-vented ( Black Skimmer ( Black Blue-winged ( Parrotlet Guira Cuckoo ( Cuckoo Guira ( Cuckoo Striped Burrowing( Owl ( Pauraque Scaly-headed Parrot ( Parrot Scaly-headed Smooth-billed Ani ( ( Little Planalto Hermit( ( Sapphire Gilded Ringed Kingfisher ( Amazon Kingfisher ( South American ( Stilt American South Large-billed Tern ( Large-billed Tern ( Tern Yellow-billed ( Ground-Dove Ruddy Picazuro Pigeon ( Picui ( Ground-Dove APPENDIX 1. 1. Continuation. APPENDIX

83 GIMENES & DOS ANJOS 40 40 20 50 20 10 60 10 10 70 10 20 50 40 80 60 40 50 40 70 20 60 100 100 100 40 25 60 40 25 37.5 16.6 57.1 50 20 40 70 25 40 40 25 7.1 100 33.3 16.6 70 90 75 75 50 85.7 71.4 66.6 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 10 10 10 20 50 80 70 30 40 90 20 40 60 50 10 80 20 70 10 20 100 100 50 70 20 75 100 44.4 16.6 57.1 40 70 20 60 100 100 100 100 100 37.5 85.7 55.5 16.6 62.5 85.7 25 50 90 50 6.6 60 50 60 71.4 83.3 85.7 100 100 100 100 100 100 100 30 10 10 10 10 70 40 30 20 90 10 10 50 70 40 10 20 30 40 30 60 20 40 100 100 25 100 100 90 20 25 50 100 100 100 28.5 33.3 66.6 40 20 25 10 50 50 25 100 100 Mutum (200 ha) (200 Mutum ha) (103 Rico Porto ha) (14 Bandeira 14.2 55.5 42.8 66.6 83.3 25 40 80 75 50 FO SH OF AQ FQ FO SH OF AQ FQ FO SH OF AQ SB FQ 100 100 100 100 100 100 100 100 100 100 100 100 100 77.7 66.6 33.3 ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) Camptostoma obsoletum Camptostoma ) ) ) ) ) Melanerpes flavifrons Arundinicola leucocephala Automolus leucophthalmus Automolus Colaptes melanochlorosColaptes Todirostrum cinereum Todirostrum Thamnophilus ruficapillusThamnophilus Serpophaga subcristata Conopias trivirgata Synallaxis frontalisSynallaxis Tyrannus savana Tyrannus Cranioleuca vulpinaCranioleuca Elaenia flavogaster Elaenia Picumnus albosquamatus Empidonomus varius Empidonomus Herpsilochmus longirostris Campylorhamphus trochilirostrisCampylorhamphus Melanerpes candidus Phacellodomus ruber Phacellodomus Veniliornis passerinus Veniliornis Fluvicola pica Fluvicola Tyrannus melancholicus Tyrannus Thamnophilus doliatus Thamnophilus Furnarius rufus Hirundinea ferruginea Dysithamnus mentalis Elaenia spectabilis Machetornis rixosus Colaptes campestris Colaptes White Woodpecker ( Woodpecker White ( Woodpecker Yellow-fronted White-wedged Piculet ( ( Little Woodpecker ( Woodpecker Green-barred ( Field Flicker White-crested Tyrannulet ( White-crested Tyrannulet Barred ( Antshrike ( Tyrannulet Beardless Southern Three-striped ( Flycatcher Rufous-capped Antshrike ( Antshrike Rufous-capped Plain Antvireo( Large-billed Antwren ( Scythebill ( Red-billed Elaenia ( Yellow-bellied ( Flycatcher Variegated Sooty-fronted Spinetail ( Spinetail Sooty-fronted Rufous Hornero( Spinetail ( Rusty-backed Greater Thornbird ( Large Elaenia ( ( Common Tody-Flycatcher Cliff Flycatcher ( ( Kingbird Tropical White-eyed( Foliage-Gleaner ( Cattle Tyrant ( Flycatcher Fork-Tailed White-headed Marsh-Tyrant ( White-headed Marsh-Tyrant Pied Water-Tyrant ( Pied Water-Tyrant APPENDIX 1. 1. Continuation. APPENDIX

84 BIRDS OF ISLANDS OF UPPER PARANÁ RIVER, BRAZIL 20 20 90 50 40 30 30 60 90 20 30 40 10 80 50 50 10 60 30 80 100 28 100 22.2 33.3 11.1 50 50 40 50 25 23 08 33.3 22.2 25 20 25 60 39 100 100 33.3 12.5 33.3 37.5 50 50 60 66.6 87.5 66.6 44.4 100 100 100 100 100 100 100 100 100 100 100 100 30 20 80 60 10 30 80 90 10 40 30 10 60 10 10 10 30 20 40 20 40 20 50 100 21 40 100 33.3 16.6 70 50 25 80 27 100 100 33.3 12.5 66.6 33.3 50 60 50 50 50 75 40 48 100 100 100 33.3 33.3 33.3 62.5 77.7 66.6 16.6 50 50 50 25 53 6.6 6.6 80 40 16.6 62.5 22.2 66.6 100 100 100 100 100 100 30 10 90 10 30 20 20 30 10 90 50 30 80 40 70 10 10 20 50 40 30 80 10 70 35 100 66.6 33.3 20 60 25 50 38 100 100 100 55.5 66.6 44.4 37.5 28.5 71.4 75 25 50 25 50 49 100 100 100 100 100 100 Mutum (200 ha) (200 Mutum ha) (103 Rico Porto ha) (14 Bandeira 33.3 66.6 33.3 28.5 57.1 20 75 57 40 75 25 FO SH OF AQ FQ FO SH OF AQ FQ FO SH OF AQ SB FQ 100 100 100 100 100 100 100 33.3 66.6 37.5 85.7 71.4 ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) Myiozetetes similis Myiozetetes Donacobius atricapillus ) ) Mimus saturninusMimus ) ) Sporophila caerulescens Euphonia chlorotica Conirostrum speciosum Zonotrichia capensis Thlypopsis sordida Tangara cayana Tangara Stelgidopteryx ruficollis Ramphocelus carbo Ramphocelus Tachycineta albiventer Tachycineta Turdus rufiventris Turdus Phaeoprogne tapera Paroaria capitata Paroaria Euphonia violacea Turdus leucomelas Turdus Volatinia jacarina Volatinia Myiodynastes maculatus Myiodynastes Nemosia pileata Nemosia Sporophila lineola Pitangus sulphuratus hraupis palmarum Thraupis sayaca Molothrus bonariensis Icterus cayanensis Sicalis flaveola Sicalis Troglodytes aedon Troglodytes Purple-throated Euphonia ( Violaceous Euphonia ( Silver-beacked Tanager ( Tanager Silver-beacked Palm Tanager (T Burnished-buff ( Tanager Conebill ( Chestnut-vented Sparrow ( Rufous-collared ( Finch Saffron GrassquitBlue-black ( Lined Seedeater ( Double-collared Seedeater ( Yellow-billed Cardinal ( Yellow-billed Epaulet Oriole ( ( Shiny Cowbird Total of species Total Sayaca Tanager ( Tanager Sayaca Hooded Tanager ( Hooded Tanager Pale-breasted ThrushPale-breasted ( Ingbird ( Mock Chalk-browed ( Orange-headed Tanager Rufous-bellied Thrush( Great Kiskadee ( Kiskadee Great White-winged ( Swallow Martin ( Brown-chested Rough-winged Swallow ( ( Wren House Black-capped Mock Ingthrush Mock Black-capped ( Streaked Flycatcher ( Flycatcher Streaked Flycatcher ( Vermilion-crowned APPENDIX 1. 1. Continuation. APPENDIX

85