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Rapid southward and upward range expansion of a tropical songbird, the Thrush-like Wren (Campylorhynchus turdinus), in South America: a consequence of habitat or climate change?

Rapid southward and upward range expansion of a tropical songbird, the Thrush-like Wren... Revista Brasileira de Ornitologia 26(1): 57–64. ARTICLE March 2018 Rapid southward and upward range expansion of a tropical songbird, the Thrush-like Wren (Campylorhynchus turdinus), in South America: a consequence of habitat or climate change? 1,3 1 2 Floyd E. Hayes , Peter Lecourt & Hugo del Castillo Department of Biology, Pacific Union College, 1 Angwin Ave., Angwin, CA 94508, USA. Asociación Guyra Paraguay, Parque Ecológico Capital Verde. Avda. Carlos Bóveda CC 1719. Asunción, Paraguay. Corresponding author: floy d_hayes@yahoo.com Received on 05 July 2017. Accepted on 28 March 2018. ABSTRACT: The Thrush-like Wren (Campylorhynchus turdinus) is a polytypic, non-migratory, cooperatively breeding species of bird widely distributed in central South America. In recent decades it has expanded its range by approximately 24%, based on a published map of its distribution in the mid-1980s and recent reports submitted to eBird up through March 2017. The northwestern subspecies, C. t. hypostictus, dramatically expanded its elevation range upward from approximately 1200 m to 4200 m a.s.l. in the Peruvian Andes. During 1977–2015 the southwestern subspecies, C. t. unicolor, dramatically expanded the southern border of its range from central Brazil and northern Paraguay (approximately 22 06'S) southward into northeastern Argentina and throughout Paraguay to 29 40'S in southeastern Brazil, extending its range east-southeast approximately 934 km at a rate of 24.6 km/yr, and latitudinally southward 838 km at a rate of 22.1 km/yr. It also expanded its range westward into the relatively dry Chaco of western Paraguay and north-central Argentina. Midwinter records near the southern border of its range in northeastern Argentina suggest it is non-migratory. Because of its affinity for disturbed habitats, its range expansion has been attributed to deforestation, but its range expansion is also consistent with the prediction that organisms are extending their geographic distribution toward higher latitudes and elevations as a consequence of climate change. KEY-WORDS: Argentina, Brazil, Colombia, distribution, elevation, global warming, Paraguay, Peru. INTRODUCTION Parmesan 2006, Chen et al. 2011). For example, analyses of the breeding ranges of birds in the temperate northern Because birds are abundant and conspicuous organisms hemisphere reveal that they are shifting northward at an that can be easily monitored, they represent ideal average rate of 2.4 km/year in North America (Hitch & bellwethers for the impacts of anthropogenic habitat Leberg 2007), 1.6 km/year in Finland (Brommer 2004), change and climate change on biodiversity (Morrison and 1.0 km/year in Britain (Thomas & Lennon 1999). The winter ranges of temperate North American birds are 1986, Hutto 1998). Short-term changes in the distributional ranges of birds are usually caused by also shifting northward at an average rate of about 1.5 stochastic population fluctuations or habitat change, but km/year (La Sorte & Thompson-III 2007). may also be attributed to climate change (Parmesan & Yohe Distributional range shifts by birds in the southern 2003, Root et al. 2003, Hickling et al. 2006, Parmesan hemisphere are not as well documented, but many species of birds in South Africa and some in Australia 2006, Chen et al. 2011). Birds may respond to recent increases in temperature and changes in precipitation appear to be rapidly expanding their ranges southward by either attempting to adapt to changes in situ or by (Chambers et al. 2005, Olsen 2007, Hockey et al. 2011). spatially shifting their distribution to areas with more In South America, several tropical bird species appear optimal conditions (Huntley et al. 2006). Distributional to be rapidly expanding their ranges southward into subtropical Argentina, Paraguay, and Brazil (e.g., Straube shifts toward higher latitudes and elevations, in concert with phenological shifts toward earlier breeding, occur et al. 2006, 2007, Piacentini et al. 2009, del Castillo et globally across a wide array of taxonomic groups, al. 2012, Pagano & Bodrati 2017). And several species providing a fingerprint of climate change (Parmesan of birds appear to be rapidly expanding their ranges & Yohe 2003, Root et al. 2003, Hickling et al. 2006, upward in elevation in the Andes (e.g., Henry 2005, Range expansion of the Thrush-like Wren (Campylorhynchus turdinus) Hayes et al. 2012, Solano-Ugalde & Real-Jibaja 2010, Gibbons et al. the southernmost record published by Vargas-Peixoto 2011, Avendaño et al. 2013). The driver of such rapid & Bosholn (2016). Google Earth was used to measure changes in bird distribution in South America is usually the rate of range expansion as a straight line between attributed to habitat change, even though such changes two points and in a southward line between degrees of are predicted by a warming climate (Parmesan & Yohe latitude. 2003, Root et al. 2003, Hickling et al. 2006, Parmesan 2006, Jetz et al. 2007, Chen et al. 2011). More detailed analyses of individual species are needed to assess the RESULTS validity of claims that distributional ranges are shifting toward higher latitudes and elevations, and to document A comparison of the past and present ranges of C. turdinus the rate of change. In this paper we document unusually revealed that its range has expanded by about 24% (Fig. rapid range expansion, both southward and upward in 1), from approximately 4,329,243 km in the mid-1980s elevation, in a tropical songbird, providing evidence that (Ridgely & Tudor 1989) to approximately 5,671,727 climate change in addition to (or instead of) habitat km by March 2017 (eBird). In the following accounts change may be accelerating its range expansion. we describe its range expansion by country. Colombia METHODS Although the northwestern subspecies, C. t. hypostictus, Study subject has not expanded its range northward, it has expanded its range northeastward along the Inírida River and Guaviare The Thrush-like Wren (Campylorhynchus turdinus) is River in Guaviare (eBird; Fig. 1). The northeasternmost o o a polytypic species of the avian family Troglodytidae record is from Guainía-Puerto Inírida (04 01'S; 67 45'W), with three distinct subspecies inhabiting tropical South only about 5 km west of the Venezuelan border, on 24 America (Hellmayr 1934, Ridgely & Tudor 1989, August 2016 (Valerie La May, Marie Lister & Warren Kroodsma & Brewer 2005). The northwestern subspecies, Stevens in eBird). C. t. hypostictus, occurs from the east slope of the Andes of Colombia, Ecuador, Peru, and northern Bolivia eastward Peru through Amazonian Brazil. The southwestern subspecies, C. t. unicolor, occurs in the lowlands of Bolivia, Paraguay, The north western subspecies, C. t. hypostictus, has and southwestern Brazil (Ridgely & Tudor 1989, expanded its range westward up into the Peruvian Andes. Kroodsma & Brewer 2005), but has recently expanded Fjeldså & Krabbe (1990) did not include it in their book its range southward into northern Argentina (Contreras on birds of the Andes because it had not been recorded in & Contreras 1986) and southeastern Brazil (Bencke et al. the temperate zone above 2300 m a.s.l. Ridgely & Tudor 2008). The eastern subspecies, nominate C. t. tur dinus, (1989) reported its maximum elevation range along the east occurs in a disjunct population along the coast of eastern slopes of the Andes as 1200 m a.s.l., which was repeated by Brazil. Clements & Shany (2001). In 1998, Hornbuckle (1999) reported it at a site with an elevation range of 1300–1600 Data gathering m a.s.l. east of Abra Patricia Pass, San Martín (05 41'S; 77 41'W), which may account for Kroodsma & Brewer To document the southward and upward range expansion (2005) reporting its maximum elevation as 1300 m a.s.l. of C. turdinus, we reviewed published distributional Schulenberg et al. (2007) reported it occurring up to 1500 records and unpublished reports submitted to eBird m a.s.l. In 2009, Robbins et al. (2011) observed it at an (ebird.org) through mid-March 2017. The latitude and elevation of about 1700 m a.s.l. at Alto Materiato, Cusco o o longitude of each location were obtained from the original (12 42'S; 72 53'W). It has now expanded its distribution source or, if absent, estimated by consulting either an dramatically upward to well over 3000 m a.s.l. in Cusco ornithological gazetteer for each country (Paynter-Jr. (several 2016 records in eBird), with a maximum elevation & Traylor 1981, 1991, Stephens & Traylor-Jr. 1983, of about 4200 m a.s.l. at ACP Ábra Málaga Thastayoc , o o Paynter-Jr. 1985, 1989) or Google Earth (www.google. Cusco (13 09'S; 72 18'W), on 18 October 2014 (Celeste com/earth). The elevation of each location was obtained Morien in eBird). from the original source. Using ArcGIS software (www. arcgis.com), we constructed maps and calculated the Paraguay area (km ) of its range in the 1980s, based on Ridgely & Tudor (1989), and the area of its current range, based The first recor d of C. turdinus in Paraguay was a specimen on reports submitted to eBird through March 2017 plus collected in 1928 on the west bank of the upper Paraguay Revista Brasileira de Ornitologia 26(1): 2018 Range expansion of the Thrush-like Wren (Campylorhynchus turdinus) Hayes et al. Figure 1. The known distribution of the Thrush-like Wren (Campylorhynchus turdinus) in the mid-1980s (A), based on Ridgely & Tudor (1989), and in February 2017 (B), based on observations reported to eBird (ebird.org) and Vargas-Peixoto & Bosholn (2016). River at Puerto Guaraní in southeastern Alto Paraguay thousand bird specimens, but none of C. turdinus, had o o (21 17'S; 57 55'W; Zotta 1940). Given the paucity of been collected prior to 1975 (Hayes 1995). Numerous specimen records at the time from the upper Paraguay individuals were seen in the area, indicating a population River region, it remains uncertain whether the specimen had recently become established. The first recor d for was within the ancestral range of the species or represented southern Presidente Hayes occurred at Estancia La o o an actively expanding population. Golondrina (24 59'S; 57 43'W) during 6–9 November In July 1977, Robert Ridgely observed several along 1995 (James Lowen). It is now widespread throughout the Apa River in northern Concepción (Ridgely & Tudor the eastern Paraguayan Chaco (eBird). 1989). The specific locality was not reported, so we On 16 July 1993, one was observed on the east side of assume it was in the vicinity of San Lázaro in northwestern the Paraguay River 5 km east of Villeta in western Central o o o o Concepción (22 06'S; 57 58'W), which was relatively (25 30'S; 57 31'W; Contreras et al. 1993), providing the accessible at the time. By 1988 and 1989 C. turdinus had first recor d for southern Paraguay. The first for Bahía de become common along both banks of the Paraguay River Asunción, Central, which had been frequently visited by as far south as Puerto Militar (formerly Villa Militar) in ornithologists since 1987, was noted in October 1998 o o northeastern Presidente Hayes (23 24'S; 57 28'W) and (H.C., unpub. data). o o Concepción in western Concepción (23 24'S; 57 26'W; In subsequent years C. turdinus spread rapidly Hayes et al. 1990). Nesting was subsequently confirmed throughout eastern Paraguay (eBird), although the first at Puerto Militar in 1991 (Contreras et al. 1993). Because department records occurred sporadically rather than no C. turdinus had been collected during extensive field progressively southward. During 3–4 October 1989, six o o work resulting in the collection of perhaps more than a were observed at Puerto Olivares (25 09'S; 57 15'W), thousand bird specimens in southeastern Alto Paraguay providing the first recor d for Cordillera (Paul Scharf). (e.g., Puerto La Victoria, formerly Puerto Casado; 22 17'S; One was observed during 4–5 May 1990 somewhere o o o 57 57'W), northeastern Presidente Hayes (e.g., Puerto between Curuguaty (24 35'S; 55 25'W) and Colonia o o o o Pinasco; 22 43'S; 57 57'W), and northern Concepción Nueva Durango (24 15'S; 55 50'W), representing the (e.g., Apa River; 22 05'S) up through 1945 (Paynter- first recor d for Canendiyú (Paul Scharf). On 8 June o o o Jr. 1989, Hayes 1995), all records south of 22 S almost 1999 it was recorded at Ybytyruzú (25 50'S; 56 13'W), certainly represented a subsequent range expansion rather providing the first recor d for Guairá (Guyra Paraguay, than a previously undetected population. in eBird). The first recor d for San Pedro occurred on o o In 1986 a specimen was collected 140 km west of 23 January 2000 at Laguna Blanca (23 49'S; 56 17'W; the Paraguay River at Pozo Colorado in central Presidente Guyra Paraguay in eBird). By 2000 it appeared in o o Hayes (23 25'S; 58 50'W; Contreras & Contreras several areas of Alto Paraná, including Presidente Franco o o 1986), in a region of the Paraguayan Chaco where a few (27 03'S; 58 37'W; Nelson Pérez in Savigny 2010). On Revista Brasileira de Ornitologia 26(1): 2018 Range expansion of the Thrush-like Wren (Campylorhynchus turdinus) Hayes et al. 22 January 2005 it was observed at Mamoreí (26 23'S; to disperse southward in the Argentinian Chaco. In 57 02'W), representing the first recor d for Paraguarí September 2009, two vocalizing C. turdinus were observed o o (Guyra Paraguay in eBird). The first recor d for Amambay at Puerto Las Palmas in eastern Chaco (27 03'S; 58 37'W; occurred on 24 March 2005, when it was noted south Bodrati et al. 2012). On 6 October 2016 a group of eight o o of Bella Vista Norte (22 10'S; 56 30'W; Guyra Paraguay was building a nest at Isla del Cerrito in eastern Chaco o o in eBird). In 2005 it was observed at Estancia Tapytá (27 17'S; 58 37'W; Pagano & Bodrati 2016). And on 10 o o (26 14'S; 55 58'W), providing the first recor d for Caazapá October 2012, two were observed at Puerto Antequeras o o (Velázquez et al. 2016). The first recor ds for Itapúa in eastern Chaco (27 27'S; 58 51'W; Fabricio Gorleri in occurred on 18 February 2006 in the vicinity of Estación eBird), providing the southernmost record for Argentina. o o Biológica Kanguery (26 26'S; 55 48'W; Guyra Paraguay Since Ridgely's observations in 1977 along the Apa in eBird). During 2–4 February 2010 it was recorded River of Paraguay (Ridgely & Tudor 1989), C. turdinus o o at Estancia La Graciela (26 35'S; 56 49'W), providing extended its range slightly west of south about 595 km the first recor d for Misiones (Silvia Centrón & Cristina at a rate of 17.1 km/yr and southward 593 km at a rate Morales). By 2012 multiple C. turdinus had reached the of 17 km/yr. And since its first arrival in the Argentinian southern border of Paraguay at Encarnación in southern Chaco in about 1986 (Contreras & Contreras 1986), C. o o Itapúa (27 21'S; 55 52'W; Smith et al. 2013). It was turdinus extended its range slightly west of south 274 km not recorded in Caaguazú until 9 January 2013, when it at a rate of about 10.5 km/yr and southward 265 km at a o o was reported from Rancho Rosalba (25 15'S; 56 17'W; rate of 10.2 km/yr. A. Lesterhuis in eBird). The first recor d for Ñeembucú The first recor d in northeastern Argentina occurred occurred on 16 September 2014, when one was observed on 23 May 2003, when three C. turdinus were observed o o o o at Humaitá (27 04'S; 58 30'W; Sergio Rios in eBird), in at Puerto Iguazú, Misiones (25 37'S; 54 35'W; Rey southwestern Paraguay. & Zurita 2004). The southernmost recor ds from In addition to expanding its range southward northeastern Argentina are from Posadas, Misiones o o and eastward, C. turdinus has also expanded its range (27 23'S; 55 57'W), where it was reported in four westward, albeit more slowly, into the dry Alto Chaco localities in 2016 (eBird). region of western Paraguay. The first recor d for Boquerón In addition to expanding its range southward and comprised a pair observed at Estancia Teniente Montania eastward, C. turdinus continues to expand its range o o in eastern Boquerón (21 57'S; 60 07'W), representing westward into the dry Argentinian Chaco, with the the westernmost record in Paraguay, on 29 September westernmost record at Bartolomé de las Casas, Formosa o o 2016 (Alberto Esquivel in eBird). (25 24'S; 59 35'W), on 14 December 2016 (Sebastán Dardanelli in eBird). Argentina There are midwinter recor ds from July near the southernmost extent of its range at Posadas (Joel Martínez Dabbene (1910) reported a specimen collected in 1880 in eBird), suggesting that the southernmost populations o o from Córdoba in Córdoba (31 24'S; 64 11'W), but are non-migratory or, possibly, partially migratory. Hellmayr (1934, p. 135) stated that the locality “can hardly be correct” and Zotta (1940) concurred. Thus, Brazil a sight record by Pablo Canevari at Parque Nacional o o Río Pilcomayo (25 04'S; 58 09'W) in eastern Formosa In central Brazil, C. turdinus has expanded its range into the (Contreras & Contreras 1986) provided the first gap between Pará and Mato Grosso do Sul states since the accepted record for Argentina. Unfortunately the date of 1980s (Fig. 1), but it is unclear whether the northwestern Canevari's observation was not published, not even in his subspecies C. t. hypostictus or the southwestern subspecies own book (Canevari et al. 1991), but occurred no later C. t. unicolor inhabits this area. than the publication of Contreras & Contreras (1986). In the isolated population along the coast of eastern Canevari's sighting was subsequently verified by sightings Brazil, nominate C. t. turdinus has not expanded its range of multiple birds in 1988 (Finch 1991) and 1993 southward, but it has expanded its range northward (Fig. (Fortabat et al. 1995). Canevari's sighting represented 1). The northernmost recor d is from Reserva Biológico de o o a remarkable southward range extension of at least 330 Saltinho, Pernambuco, Brazil (8 44'S; 35 11'W), on 30 km since Ridgely's observations in 1977 along the Apa October 2013 (Forest Rowland in eBird). River of Paraguay (Ridgely & Tudor 1989). Assuming In 1941, a specimen of C. t. unicolor was collected Canevari's sighting was in the year 1986, C. turdinus had on the east bank of the Paraguay River at Pôrto o o extended its range southward at a rate of 36.7 km/yr from Quebracho (21 50'S; 57 53'W), in southwest Mato 1977–1986. Grosso do Sul (Contreras et al. 1993), representing the In subsequent decades C. turdinus continued southwesternmost record for Brazil (Naumburg 1930, Revista Brasileira de Ornitologia 26(1): 2018 Range expansion of the Thrush-like Wren (Campylorhynchus turdinus) Hayes et al. Hellmayr 1934). This locality is about 60 km south of Castillo et al. 2012, Pagano & Bodrati 2017). Vagrancy, where the nearest specimen had been previously collected the long-distance dispersal of individuals beyond their at Puerto Guaraní (21 17'S), Paraguay. Again, given the normal distribution or migratory path (e.g., Thomson paucity of specimen records at the time from the upper 1964, Veit 2000), occurs less frequently among non- Paraguay River region, it remains uncertain whether the migratory than migratory species of birds (Lees & Gilroy specimen was within the ancestral range of the species or 2009). Data from eBird indicate that C. turdinus occurs represented an actively expanding population. near its southernmost limit in Argentina throughout the Farther south, in southern Rio Grande do Sul, winter, indicating that it is non-migratory or, possibly, Helmut Sick (in Belton 1985) reported hearing a C. partially migratory. Vagrancy was thought to occur less turdinus vocalizing at Fazenda da Invernada (31 05'S; frequently among cooperatively breeding than among 52 52'W), on 25 November 1972. According to Belton non-cooperatively breeding species of birds (Zack 1990), (1985, p. 105), Sick felt “absolutely sure of it, as voice but Rusk et al. (2013) provided evidence that vagrancy is very easy to recognize with nothing similar to it”. occurs just as frequently in cooperatively breeding Nevertheless, the record was considered unreliable and species. Rabenold (1990) reported that C. turdinus breeds rejected by Bencke et al. (2010). In 2008, a singing C. cooperatively, although no data have been published. turdinus was heard at Foz do Iguaçu in central Paraná Despite its apparent non-migratory and cooperatively o o (25 37'S; 54 29'W; Bencke et al. 2008). A remarkable breeding habits, long-distance vagrancy appears to be a life range extension occurred in 2015, when a singing C. history trait in C. turdinus, and is presumably increasing turdinus was photographed at Santa Maria in central Rio as a consequence of rapid demographic population o o Grande do Sul (29 40'S; 53 48'W; Vargas-Peixoto & growth (Veit 2000). Bosholn 2016), providing the southernmost confirmed It is difficult to attribute changes in the spatial record for the species. Although the authors did not assign distribution of any single species to either habitat or climate it to a subspecies, photographs reveal the bird's unspotted change. Because of its affinity for disturbed habitats, C. creamy underparts, which are typical of C. t. unicolor, turdinus's rapid southward range expansion has been contrasting with the spotted underparts of nominate C. t. attributed to deforestation (Rey & Zurita 2004, Bodrati turdinus of coastal eastern Brazil (Ridgely & Tudor 1989, et al. 2012). The subtropical moist broa dleaf forests of Sick 1993). Since Ridgely's observations in 1977 along eastern Paraguay, southern Brazil, and northeastern the Apa River of Paraguay (Ridgely & Tudor 1989), by Argentina have been subjected to extensive deforestation 2015 C. turdinus had extended its range slightly east of in recent decades (Hansen & DeFries 2004, Fleytas 2007, south 934 km at a rate of 24.6 km/yr and southward 841 Huang et al. 2007, Aide et al. 2013, Hansen et al. 2013), km at a rate of 22.1 km/yr (Fig. 2). plausibly facilitating the southward range expansion of C. tudinus. However, the rate of deforestation on the east slopes of the Andes, including the region east of Cusco, DISCUSSION Peru, has been considerably slower (Aide et al. 2013, Zegarra & Gayoso 2015), suggesting that habitat change Campylorhynchus turdinus is just one of several tropical may not adequately account for the rapid upward range bird species in southern South America that is rapidly expansion of C. turdinus in Cusco, Peru. expanding its range southward into subtropical latitudes The rapid range expansion of C. tur dinus both (e.g., Straube et al. 2006, 2007, Piacentini et al. 2009, del southward and upward is consistent with the prediction that organisms are extending their geographic distribution toward higher latitudes and elevations as a consequence of global warming (Parmesan & Yohe 2003, Root et al. 2003, Hickling et al. 2006, Parmesan 2006, Chen et al. 2011). The region in which C. turdinus is expanding its range southward in southern South America is warming relatively rapidly (Easterling et al. 1997, Rosenblüth et al. 1997, Vincent et al. 2005). Because the northern boundaries of North American songbirds appear to be limited by winter nighttime temperatures (Root 1988), the southern boundaries of South American songbirds may also be limited by winter nighttime temperatures. If so, songbirds in southern South America may be able Figure 2. Southward range expansion of the Thrush-like Wren to extend their ranges southward as winter nighttime (Campylorhynchus turdinus) illustrated by the southernmost temperatures increase, which may be the case with C. record vs. year, based on data in the Results section. turdinus and several other species of birds in southern Revista Brasileira de Ornitologia 26(1): 2018 Range expansion of the Thrush-like Wren (Campylorhynchus turdinus) Hayes et al. Belton W. 1985. Birds of Rio Grande do Sul, Brazil. Part 2. South America (e.g., Straube et al. 2006, 2007, Piacentini Formicariidae through Corvidae. Bulletin of the American Museum et al. 2009, del Castillo et al. 2012, Pagano & Bodrati of Natural History 180: 1–242. 2017). However, temperatures are not rising much Bencke G.A., Dias R.A., Bugoni L., Agne C.E., Fontana C.S., in the Peruvian Andes (Easterling et al. 1997, Vincent Maurício G.N. & Machado D.B. 2010. Revisão e atualização da lista das aves do Rio Grande do Sul, Brasil. Iheringia, Série et al. 2005), suggesting that climate change may not Zoologia 100: 519–556. adequately account for the rapid upward range expansion Bencke G.A., Dias R.A. & Fontana C.S. 2008. Observações of C. turdinus in Cusco, Peru. Perhaps habitat change ornitológicas relevantes no Parque Nacional do Iguaçu e arredores, and climate change combined best accounts for its rapid incluindo o primeiro registro de Campylorhynchus turdinus para o Paraná. Atualidades Ornitológicas 145: 6–7. upward range expansion in Cusco, Peru. Bodrati A., Areta J.I. & White E. 2012. La avifauna de La Posada y Because tropical species of birds tend to have Reserva Puerto Bemberg, Misiones, Argentina. Nuestras Aves 57: relatively small distributional ranges resulting in a high 63–79. degree of endemicity, and occur in climatically stable Botero E.U. 2015. El cambio climático y sus efectos en la biodiversidad environments where they tend to be K-selected with en América Latina. Santiago: Comisión Económica para América Latina y el Caribe (CEPAL), Naciones Unidas. greater longevity and smaller clutch sizes, resulting in Brommer J.E. 2004. The range margins of northern birds shift lower demographic flexibility, they are thought to be polewards. Annales Zoologici Fennici 41: 391–397. more vulnerable to anthropogenic changes in habitat Canevari M., Canevari P., Carrizo G.R., Harris G., Mata J.R. & and climate than temperate species of birds (Jetz et al. Straneck R.J. 1991. Nueva guía de las aves argentinas. Buenos Aires: Fundación Acindar. 2007, Şekercioğlu et al. 2012, Reif & Štěpánková 2016). Chambers L.E., Hughes L. & Weston M.A. 2005. Climate change Tropical birds may be adversely affected by climate change and its impact on Australia's avifauna. Emu 105: 1–20. in many different ways, such as susceptibility to extreme Chen I-C., Hill J.K., Ohlemüller R., Roy D.B. & Thomas C.D. 2011. weather events, habitat loss, emerging diseases, invasive Rapid range shifts of species associated with high levels of climate warming. Science 333: 1024–1026. species, and hunting (Jetz et al. 2007, Şekercioğlu et Clements J.F. & Shany N. 2001. A field guide to the bir ds of Peru. al. 2012, Botero 2015). Nevertheless, some tropical Temecula: Ibis Publishing Company. bird species, such as C. turdinus, may benefit from Contreras J.R. & Contreras A.O. 1986. Acerca de Campylorhynchus anthropogenic changes in the environment by expanding turdinus unicolor Wied en Paraguay y en la República Argentina their ranges into disturbed habitats in response to habitat (Aves: Trogloditidae [sic]). Historia Natural 6: 75–76. Contreras J.R., Contreras A.O. & Argaña J.E. 1993. Comentarios change or expanding their ranges toward higher latitudes acerca de algunas especies de aves nuevas o poco conocidas para el and elevations in response to climate change. It remains Paraguay. Nótulas Faunísticas 43: 1–7. uncertain which of these two drivers, or both combined, Dabbene R. 1910. Ornitología argentina. Catálogo sistemático y or if there is another yet to be identified, best explains descriptivo de las aves de la República Argentina. Anales del Museo Nacional de Buenos Aires 18: 1–513. the rapid range expansion of C. turdinus. 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Rapid southward and upward range expansion of a tropical songbird, the Thrush-like Wren (Campylorhynchus turdinus), in South America: a consequence of habitat or climate change?

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Abstract

Revista Brasileira de Ornitologia 26(1): 57–64. ARTICLE March 2018 Rapid southward and upward range expansion of a tropical songbird, the Thrush-like Wren (Campylorhynchus turdinus), in South America: a consequence of habitat or climate change? 1,3 1 2 Floyd E. Hayes , Peter Lecourt & Hugo del Castillo Department of Biology, Pacific Union College, 1 Angwin Ave., Angwin, CA 94508, USA. Asociación Guyra Paraguay, Parque Ecológico Capital Verde. Avda. Carlos Bóveda CC 1719. Asunción, Paraguay. Corresponding author: floy d_hayes@yahoo.com Received on 05 July 2017. Accepted on 28 March 2018. ABSTRACT: The Thrush-like Wren (Campylorhynchus turdinus) is a polytypic, non-migratory, cooperatively breeding species of bird widely distributed in central South America. In recent decades it has expanded its range by approximately 24%, based on a published map of its distribution in the mid-1980s and recent reports submitted to eBird up through March 2017. The northwestern subspecies, C. t. hypostictus, dramatically expanded its elevation range upward from approximately 1200 m to 4200 m a.s.l. in the Peruvian Andes. During 1977–2015 the southwestern subspecies, C. t. unicolor, dramatically expanded the southern border of its range from central Brazil and northern Paraguay (approximately 22 06'S) southward into northeastern Argentina and throughout Paraguay to 29 40'S in southeastern Brazil, extending its range east-southeast approximately 934 km at a rate of 24.6 km/yr, and latitudinally southward 838 km at a rate of 22.1 km/yr. It also expanded its range westward into the relatively dry Chaco of western Paraguay and north-central Argentina. Midwinter records near the southern border of its range in northeastern Argentina suggest it is non-migratory. Because of its affinity for disturbed habitats, its range expansion has been attributed to deforestation, but its range expansion is also consistent with the prediction that organisms are extending their geographic distribution toward higher latitudes and elevations as a consequence of climate change. KEY-WORDS: Argentina, Brazil, Colombia, distribution, elevation, global warming, Paraguay, Peru. INTRODUCTION Parmesan 2006, Chen et al. 2011). For example, analyses of the breeding ranges of birds in the temperate northern Because birds are abundant and conspicuous organisms hemisphere reveal that they are shifting northward at an that can be easily monitored, they represent ideal average rate of 2.4 km/year in North America (Hitch & bellwethers for the impacts of anthropogenic habitat Leberg 2007), 1.6 km/year in Finland (Brommer 2004), change and climate change on biodiversity (Morrison and 1.0 km/year in Britain (Thomas & Lennon 1999). The winter ranges of temperate North American birds are 1986, Hutto 1998). Short-term changes in the distributional ranges of birds are usually caused by also shifting northward at an average rate of about 1.5 stochastic population fluctuations or habitat change, but km/year (La Sorte & Thompson-III 2007). may also be attributed to climate change (Parmesan & Yohe Distributional range shifts by birds in the southern 2003, Root et al. 2003, Hickling et al. 2006, Parmesan hemisphere are not as well documented, but many species of birds in South Africa and some in Australia 2006, Chen et al. 2011). Birds may respond to recent increases in temperature and changes in precipitation appear to be rapidly expanding their ranges southward by either attempting to adapt to changes in situ or by (Chambers et al. 2005, Olsen 2007, Hockey et al. 2011). spatially shifting their distribution to areas with more In South America, several tropical bird species appear optimal conditions (Huntley et al. 2006). Distributional to be rapidly expanding their ranges southward into subtropical Argentina, Paraguay, and Brazil (e.g., Straube shifts toward higher latitudes and elevations, in concert with phenological shifts toward earlier breeding, occur et al. 2006, 2007, Piacentini et al. 2009, del Castillo et globally across a wide array of taxonomic groups, al. 2012, Pagano & Bodrati 2017). And several species providing a fingerprint of climate change (Parmesan of birds appear to be rapidly expanding their ranges & Yohe 2003, Root et al. 2003, Hickling et al. 2006, upward in elevation in the Andes (e.g., Henry 2005, Range expansion of the Thrush-like Wren (Campylorhynchus turdinus) Hayes et al. 2012, Solano-Ugalde & Real-Jibaja 2010, Gibbons et al. the southernmost record published by Vargas-Peixoto 2011, Avendaño et al. 2013). The driver of such rapid & Bosholn (2016). Google Earth was used to measure changes in bird distribution in South America is usually the rate of range expansion as a straight line between attributed to habitat change, even though such changes two points and in a southward line between degrees of are predicted by a warming climate (Parmesan & Yohe latitude. 2003, Root et al. 2003, Hickling et al. 2006, Parmesan 2006, Jetz et al. 2007, Chen et al. 2011). More detailed analyses of individual species are needed to assess the RESULTS validity of claims that distributional ranges are shifting toward higher latitudes and elevations, and to document A comparison of the past and present ranges of C. turdinus the rate of change. In this paper we document unusually revealed that its range has expanded by about 24% (Fig. rapid range expansion, both southward and upward in 1), from approximately 4,329,243 km in the mid-1980s elevation, in a tropical songbird, providing evidence that (Ridgely & Tudor 1989) to approximately 5,671,727 climate change in addition to (or instead of) habitat km by March 2017 (eBird). In the following accounts change may be accelerating its range expansion. we describe its range expansion by country. Colombia METHODS Although the northwestern subspecies, C. t. hypostictus, Study subject has not expanded its range northward, it has expanded its range northeastward along the Inírida River and Guaviare The Thrush-like Wren (Campylorhynchus turdinus) is River in Guaviare (eBird; Fig. 1). The northeasternmost o o a polytypic species of the avian family Troglodytidae record is from Guainía-Puerto Inírida (04 01'S; 67 45'W), with three distinct subspecies inhabiting tropical South only about 5 km west of the Venezuelan border, on 24 America (Hellmayr 1934, Ridgely & Tudor 1989, August 2016 (Valerie La May, Marie Lister & Warren Kroodsma & Brewer 2005). The northwestern subspecies, Stevens in eBird). C. t. hypostictus, occurs from the east slope of the Andes of Colombia, Ecuador, Peru, and northern Bolivia eastward Peru through Amazonian Brazil. The southwestern subspecies, C. t. unicolor, occurs in the lowlands of Bolivia, Paraguay, The north western subspecies, C. t. hypostictus, has and southwestern Brazil (Ridgely & Tudor 1989, expanded its range westward up into the Peruvian Andes. Kroodsma & Brewer 2005), but has recently expanded Fjeldså & Krabbe (1990) did not include it in their book its range southward into northern Argentina (Contreras on birds of the Andes because it had not been recorded in & Contreras 1986) and southeastern Brazil (Bencke et al. the temperate zone above 2300 m a.s.l. Ridgely & Tudor 2008). The eastern subspecies, nominate C. t. tur dinus, (1989) reported its maximum elevation range along the east occurs in a disjunct population along the coast of eastern slopes of the Andes as 1200 m a.s.l., which was repeated by Brazil. Clements & Shany (2001). In 1998, Hornbuckle (1999) reported it at a site with an elevation range of 1300–1600 Data gathering m a.s.l. east of Abra Patricia Pass, San Martín (05 41'S; 77 41'W), which may account for Kroodsma & Brewer To document the southward and upward range expansion (2005) reporting its maximum elevation as 1300 m a.s.l. of C. turdinus, we reviewed published distributional Schulenberg et al. (2007) reported it occurring up to 1500 records and unpublished reports submitted to eBird m a.s.l. In 2009, Robbins et al. (2011) observed it at an (ebird.org) through mid-March 2017. The latitude and elevation of about 1700 m a.s.l. at Alto Materiato, Cusco o o longitude of each location were obtained from the original (12 42'S; 72 53'W). It has now expanded its distribution source or, if absent, estimated by consulting either an dramatically upward to well over 3000 m a.s.l. in Cusco ornithological gazetteer for each country (Paynter-Jr. (several 2016 records in eBird), with a maximum elevation & Traylor 1981, 1991, Stephens & Traylor-Jr. 1983, of about 4200 m a.s.l. at ACP Ábra Málaga Thastayoc , o o Paynter-Jr. 1985, 1989) or Google Earth (www.google. Cusco (13 09'S; 72 18'W), on 18 October 2014 (Celeste com/earth). The elevation of each location was obtained Morien in eBird). from the original source. Using ArcGIS software (www. arcgis.com), we constructed maps and calculated the Paraguay area (km ) of its range in the 1980s, based on Ridgely & Tudor (1989), and the area of its current range, based The first recor d of C. turdinus in Paraguay was a specimen on reports submitted to eBird through March 2017 plus collected in 1928 on the west bank of the upper Paraguay Revista Brasileira de Ornitologia 26(1): 2018 Range expansion of the Thrush-like Wren (Campylorhynchus turdinus) Hayes et al. Figure 1. The known distribution of the Thrush-like Wren (Campylorhynchus turdinus) in the mid-1980s (A), based on Ridgely & Tudor (1989), and in February 2017 (B), based on observations reported to eBird (ebird.org) and Vargas-Peixoto & Bosholn (2016). River at Puerto Guaraní in southeastern Alto Paraguay thousand bird specimens, but none of C. turdinus, had o o (21 17'S; 57 55'W; Zotta 1940). Given the paucity of been collected prior to 1975 (Hayes 1995). Numerous specimen records at the time from the upper Paraguay individuals were seen in the area, indicating a population River region, it remains uncertain whether the specimen had recently become established. The first recor d for was within the ancestral range of the species or represented southern Presidente Hayes occurred at Estancia La o o an actively expanding population. Golondrina (24 59'S; 57 43'W) during 6–9 November In July 1977, Robert Ridgely observed several along 1995 (James Lowen). It is now widespread throughout the Apa River in northern Concepción (Ridgely & Tudor the eastern Paraguayan Chaco (eBird). 1989). The specific locality was not reported, so we On 16 July 1993, one was observed on the east side of assume it was in the vicinity of San Lázaro in northwestern the Paraguay River 5 km east of Villeta in western Central o o o o Concepción (22 06'S; 57 58'W), which was relatively (25 30'S; 57 31'W; Contreras et al. 1993), providing the accessible at the time. By 1988 and 1989 C. turdinus had first recor d for southern Paraguay. The first for Bahía de become common along both banks of the Paraguay River Asunción, Central, which had been frequently visited by as far south as Puerto Militar (formerly Villa Militar) in ornithologists since 1987, was noted in October 1998 o o northeastern Presidente Hayes (23 24'S; 57 28'W) and (H.C., unpub. data). o o Concepción in western Concepción (23 24'S; 57 26'W; In subsequent years C. turdinus spread rapidly Hayes et al. 1990). Nesting was subsequently confirmed throughout eastern Paraguay (eBird), although the first at Puerto Militar in 1991 (Contreras et al. 1993). Because department records occurred sporadically rather than no C. turdinus had been collected during extensive field progressively southward. During 3–4 October 1989, six o o work resulting in the collection of perhaps more than a were observed at Puerto Olivares (25 09'S; 57 15'W), thousand bird specimens in southeastern Alto Paraguay providing the first recor d for Cordillera (Paul Scharf). (e.g., Puerto La Victoria, formerly Puerto Casado; 22 17'S; One was observed during 4–5 May 1990 somewhere o o o 57 57'W), northeastern Presidente Hayes (e.g., Puerto between Curuguaty (24 35'S; 55 25'W) and Colonia o o o o Pinasco; 22 43'S; 57 57'W), and northern Concepción Nueva Durango (24 15'S; 55 50'W), representing the (e.g., Apa River; 22 05'S) up through 1945 (Paynter- first recor d for Canendiyú (Paul Scharf). On 8 June o o o Jr. 1989, Hayes 1995), all records south of 22 S almost 1999 it was recorded at Ybytyruzú (25 50'S; 56 13'W), certainly represented a subsequent range expansion rather providing the first recor d for Guairá (Guyra Paraguay, than a previously undetected population. in eBird). The first recor d for San Pedro occurred on o o In 1986 a specimen was collected 140 km west of 23 January 2000 at Laguna Blanca (23 49'S; 56 17'W; the Paraguay River at Pozo Colorado in central Presidente Guyra Paraguay in eBird). By 2000 it appeared in o o Hayes (23 25'S; 58 50'W; Contreras & Contreras several areas of Alto Paraná, including Presidente Franco o o 1986), in a region of the Paraguayan Chaco where a few (27 03'S; 58 37'W; Nelson Pérez in Savigny 2010). On Revista Brasileira de Ornitologia 26(1): 2018 Range expansion of the Thrush-like Wren (Campylorhynchus turdinus) Hayes et al. 22 January 2005 it was observed at Mamoreí (26 23'S; to disperse southward in the Argentinian Chaco. In 57 02'W), representing the first recor d for Paraguarí September 2009, two vocalizing C. turdinus were observed o o (Guyra Paraguay in eBird). The first recor d for Amambay at Puerto Las Palmas in eastern Chaco (27 03'S; 58 37'W; occurred on 24 March 2005, when it was noted south Bodrati et al. 2012). On 6 October 2016 a group of eight o o of Bella Vista Norte (22 10'S; 56 30'W; Guyra Paraguay was building a nest at Isla del Cerrito in eastern Chaco o o in eBird). In 2005 it was observed at Estancia Tapytá (27 17'S; 58 37'W; Pagano & Bodrati 2016). And on 10 o o (26 14'S; 55 58'W), providing the first recor d for Caazapá October 2012, two were observed at Puerto Antequeras o o (Velázquez et al. 2016). The first recor ds for Itapúa in eastern Chaco (27 27'S; 58 51'W; Fabricio Gorleri in occurred on 18 February 2006 in the vicinity of Estación eBird), providing the southernmost record for Argentina. o o Biológica Kanguery (26 26'S; 55 48'W; Guyra Paraguay Since Ridgely's observations in 1977 along the Apa in eBird). During 2–4 February 2010 it was recorded River of Paraguay (Ridgely & Tudor 1989), C. turdinus o o at Estancia La Graciela (26 35'S; 56 49'W), providing extended its range slightly west of south about 595 km the first recor d for Misiones (Silvia Centrón & Cristina at a rate of 17.1 km/yr and southward 593 km at a rate Morales). By 2012 multiple C. turdinus had reached the of 17 km/yr. And since its first arrival in the Argentinian southern border of Paraguay at Encarnación in southern Chaco in about 1986 (Contreras & Contreras 1986), C. o o Itapúa (27 21'S; 55 52'W; Smith et al. 2013). It was turdinus extended its range slightly west of south 274 km not recorded in Caaguazú until 9 January 2013, when it at a rate of about 10.5 km/yr and southward 265 km at a o o was reported from Rancho Rosalba (25 15'S; 56 17'W; rate of 10.2 km/yr. A. Lesterhuis in eBird). The first recor d for Ñeembucú The first recor d in northeastern Argentina occurred occurred on 16 September 2014, when one was observed on 23 May 2003, when three C. turdinus were observed o o o o at Humaitá (27 04'S; 58 30'W; Sergio Rios in eBird), in at Puerto Iguazú, Misiones (25 37'S; 54 35'W; Rey southwestern Paraguay. & Zurita 2004). The southernmost recor ds from In addition to expanding its range southward northeastern Argentina are from Posadas, Misiones o o and eastward, C. turdinus has also expanded its range (27 23'S; 55 57'W), where it was reported in four westward, albeit more slowly, into the dry Alto Chaco localities in 2016 (eBird). region of western Paraguay. The first recor d for Boquerón In addition to expanding its range southward and comprised a pair observed at Estancia Teniente Montania eastward, C. turdinus continues to expand its range o o in eastern Boquerón (21 57'S; 60 07'W), representing westward into the dry Argentinian Chaco, with the the westernmost record in Paraguay, on 29 September westernmost record at Bartolomé de las Casas, Formosa o o 2016 (Alberto Esquivel in eBird). (25 24'S; 59 35'W), on 14 December 2016 (Sebastán Dardanelli in eBird). Argentina There are midwinter recor ds from July near the southernmost extent of its range at Posadas (Joel Martínez Dabbene (1910) reported a specimen collected in 1880 in eBird), suggesting that the southernmost populations o o from Córdoba in Córdoba (31 24'S; 64 11'W), but are non-migratory or, possibly, partially migratory. Hellmayr (1934, p. 135) stated that the locality “can hardly be correct” and Zotta (1940) concurred. Thus, Brazil a sight record by Pablo Canevari at Parque Nacional o o Río Pilcomayo (25 04'S; 58 09'W) in eastern Formosa In central Brazil, C. turdinus has expanded its range into the (Contreras & Contreras 1986) provided the first gap between Pará and Mato Grosso do Sul states since the accepted record for Argentina. Unfortunately the date of 1980s (Fig. 1), but it is unclear whether the northwestern Canevari's observation was not published, not even in his subspecies C. t. hypostictus or the southwestern subspecies own book (Canevari et al. 1991), but occurred no later C. t. unicolor inhabits this area. than the publication of Contreras & Contreras (1986). In the isolated population along the coast of eastern Canevari's sighting was subsequently verified by sightings Brazil, nominate C. t. turdinus has not expanded its range of multiple birds in 1988 (Finch 1991) and 1993 southward, but it has expanded its range northward (Fig. (Fortabat et al. 1995). Canevari's sighting represented 1). The northernmost recor d is from Reserva Biológico de o o a remarkable southward range extension of at least 330 Saltinho, Pernambuco, Brazil (8 44'S; 35 11'W), on 30 km since Ridgely's observations in 1977 along the Apa October 2013 (Forest Rowland in eBird). River of Paraguay (Ridgely & Tudor 1989). Assuming In 1941, a specimen of C. t. unicolor was collected Canevari's sighting was in the year 1986, C. turdinus had on the east bank of the Paraguay River at Pôrto o o extended its range southward at a rate of 36.7 km/yr from Quebracho (21 50'S; 57 53'W), in southwest Mato 1977–1986. Grosso do Sul (Contreras et al. 1993), representing the In subsequent decades C. turdinus continued southwesternmost record for Brazil (Naumburg 1930, Revista Brasileira de Ornitologia 26(1): 2018 Range expansion of the Thrush-like Wren (Campylorhynchus turdinus) Hayes et al. Hellmayr 1934). This locality is about 60 km south of Castillo et al. 2012, Pagano & Bodrati 2017). Vagrancy, where the nearest specimen had been previously collected the long-distance dispersal of individuals beyond their at Puerto Guaraní (21 17'S), Paraguay. Again, given the normal distribution or migratory path (e.g., Thomson paucity of specimen records at the time from the upper 1964, Veit 2000), occurs less frequently among non- Paraguay River region, it remains uncertain whether the migratory than migratory species of birds (Lees & Gilroy specimen was within the ancestral range of the species or 2009). Data from eBird indicate that C. turdinus occurs represented an actively expanding population. near its southernmost limit in Argentina throughout the Farther south, in southern Rio Grande do Sul, winter, indicating that it is non-migratory or, possibly, Helmut Sick (in Belton 1985) reported hearing a C. partially migratory. Vagrancy was thought to occur less turdinus vocalizing at Fazenda da Invernada (31 05'S; frequently among cooperatively breeding than among 52 52'W), on 25 November 1972. According to Belton non-cooperatively breeding species of birds (Zack 1990), (1985, p. 105), Sick felt “absolutely sure of it, as voice but Rusk et al. (2013) provided evidence that vagrancy is very easy to recognize with nothing similar to it”. occurs just as frequently in cooperatively breeding Nevertheless, the record was considered unreliable and species. Rabenold (1990) reported that C. turdinus breeds rejected by Bencke et al. (2010). In 2008, a singing C. cooperatively, although no data have been published. turdinus was heard at Foz do Iguaçu in central Paraná Despite its apparent non-migratory and cooperatively o o (25 37'S; 54 29'W; Bencke et al. 2008). A remarkable breeding habits, long-distance vagrancy appears to be a life range extension occurred in 2015, when a singing C. history trait in C. turdinus, and is presumably increasing turdinus was photographed at Santa Maria in central Rio as a consequence of rapid demographic population o o Grande do Sul (29 40'S; 53 48'W; Vargas-Peixoto & growth (Veit 2000). Bosholn 2016), providing the southernmost confirmed It is difficult to attribute changes in the spatial record for the species. Although the authors did not assign distribution of any single species to either habitat or climate it to a subspecies, photographs reveal the bird's unspotted change. Because of its affinity for disturbed habitats, C. creamy underparts, which are typical of C. t. unicolor, turdinus's rapid southward range expansion has been contrasting with the spotted underparts of nominate C. t. attributed to deforestation (Rey & Zurita 2004, Bodrati turdinus of coastal eastern Brazil (Ridgely & Tudor 1989, et al. 2012). The subtropical moist broa dleaf forests of Sick 1993). Since Ridgely's observations in 1977 along eastern Paraguay, southern Brazil, and northeastern the Apa River of Paraguay (Ridgely & Tudor 1989), by Argentina have been subjected to extensive deforestation 2015 C. turdinus had extended its range slightly east of in recent decades (Hansen & DeFries 2004, Fleytas 2007, south 934 km at a rate of 24.6 km/yr and southward 841 Huang et al. 2007, Aide et al. 2013, Hansen et al. 2013), km at a rate of 22.1 km/yr (Fig. 2). plausibly facilitating the southward range expansion of C. tudinus. However, the rate of deforestation on the east slopes of the Andes, including the region east of Cusco, DISCUSSION Peru, has been considerably slower (Aide et al. 2013, Zegarra & Gayoso 2015), suggesting that habitat change Campylorhynchus turdinus is just one of several tropical may not adequately account for the rapid upward range bird species in southern South America that is rapidly expansion of C. turdinus in Cusco, Peru. expanding its range southward into subtropical latitudes The rapid range expansion of C. tur dinus both (e.g., Straube et al. 2006, 2007, Piacentini et al. 2009, del southward and upward is consistent with the prediction that organisms are extending their geographic distribution toward higher latitudes and elevations as a consequence of global warming (Parmesan & Yohe 2003, Root et al. 2003, Hickling et al. 2006, Parmesan 2006, Chen et al. 2011). The region in which C. turdinus is expanding its range southward in southern South America is warming relatively rapidly (Easterling et al. 1997, Rosenblüth et al. 1997, Vincent et al. 2005). Because the northern boundaries of North American songbirds appear to be limited by winter nighttime temperatures (Root 1988), the southern boundaries of South American songbirds may also be limited by winter nighttime temperatures. If so, songbirds in southern South America may be able Figure 2. Southward range expansion of the Thrush-like Wren to extend their ranges southward as winter nighttime (Campylorhynchus turdinus) illustrated by the southernmost temperatures increase, which may be the case with C. record vs. year, based on data in the Results section. turdinus and several other species of birds in southern Revista Brasileira de Ornitologia 26(1): 2018 Range expansion of the Thrush-like Wren (Campylorhynchus turdinus) Hayes et al. Belton W. 1985. Birds of Rio Grande do Sul, Brazil. Part 2. South America (e.g., Straube et al. 2006, 2007, Piacentini Formicariidae through Corvidae. Bulletin of the American Museum et al. 2009, del Castillo et al. 2012, Pagano & Bodrati of Natural History 180: 1–242. 2017). However, temperatures are not rising much Bencke G.A., Dias R.A., Bugoni L., Agne C.E., Fontana C.S., in the Peruvian Andes (Easterling et al. 1997, Vincent Maurício G.N. & Machado D.B. 2010. Revisão e atualização da lista das aves do Rio Grande do Sul, Brasil. Iheringia, Série et al. 2005), suggesting that climate change may not Zoologia 100: 519–556. adequately account for the rapid upward range expansion Bencke G.A., Dias R.A. & Fontana C.S. 2008. Observações of C. turdinus in Cusco, Peru. Perhaps habitat change ornitológicas relevantes no Parque Nacional do Iguaçu e arredores, and climate change combined best accounts for its rapid incluindo o primeiro registro de Campylorhynchus turdinus para o Paraná. Atualidades Ornitológicas 145: 6–7. upward range expansion in Cusco, Peru. Bodrati A., Areta J.I. & White E. 2012. La avifauna de La Posada y Because tropical species of birds tend to have Reserva Puerto Bemberg, Misiones, Argentina. Nuestras Aves 57: relatively small distributional ranges resulting in a high 63–79. degree of endemicity, and occur in climatically stable Botero E.U. 2015. El cambio climático y sus efectos en la biodiversidad environments where they tend to be K-selected with en América Latina. Santiago: Comisión Económica para América Latina y el Caribe (CEPAL), Naciones Unidas. greater longevity and smaller clutch sizes, resulting in Brommer J.E. 2004. The range margins of northern birds shift lower demographic flexibility, they are thought to be polewards. Annales Zoologici Fennici 41: 391–397. more vulnerable to anthropogenic changes in habitat Canevari M., Canevari P., Carrizo G.R., Harris G., Mata J.R. & and climate than temperate species of birds (Jetz et al. Straneck R.J. 1991. Nueva guía de las aves argentinas. Buenos Aires: Fundación Acindar. 2007, Şekercioğlu et al. 2012, Reif & Štěpánková 2016). Chambers L.E., Hughes L. & Weston M.A. 2005. Climate change Tropical birds may be adversely affected by climate change and its impact on Australia's avifauna. Emu 105: 1–20. in many different ways, such as susceptibility to extreme Chen I-C., Hill J.K., Ohlemüller R., Roy D.B. & Thomas C.D. 2011. weather events, habitat loss, emerging diseases, invasive Rapid range shifts of species associated with high levels of climate warming. Science 333: 1024–1026. species, and hunting (Jetz et al. 2007, Şekercioğlu et Clements J.F. & Shany N. 2001. A field guide to the bir ds of Peru. al. 2012, Botero 2015). Nevertheless, some tropical Temecula: Ibis Publishing Company. bird species, such as C. turdinus, may benefit from Contreras J.R. & Contreras A.O. 1986. Acerca de Campylorhynchus anthropogenic changes in the environment by expanding turdinus unicolor Wied en Paraguay y en la República Argentina their ranges into disturbed habitats in response to habitat (Aves: Trogloditidae [sic]). Historia Natural 6: 75–76. Contreras J.R., Contreras A.O. & Argaña J.E. 1993. Comentarios change or expanding their ranges toward higher latitudes acerca de algunas especies de aves nuevas o poco conocidas para el and elevations in response to climate change. It remains Paraguay. Nótulas Faunísticas 43: 1–7. uncertain which of these two drivers, or both combined, Dabbene R. 1910. Ornitología argentina. Catálogo sistemático y or if there is another yet to be identified, best explains descriptivo de las aves de la República Argentina. Anales del Museo Nacional de Buenos Aires 18: 1–513. the rapid range expansion of C. turdinus. 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Journal

Ornithology ResearchSpringer Journals

Published: Mar 1, 2018

Keywords: Argentina; Brazil; Colombia; distribution; elevation; global warming; Paraguay; Peru

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