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Observations on the breeding biology of the Pygmy Nightjar Nyctipolus hirundinaceus in the Caatinga of Bahia and Ceará, Brazil

Observations on the breeding biology of the Pygmy Nightjar Nyctipolus hirundinaceus in the... Revista Brasileira de Ornitologia, 22(2), 201-209 ARTICLE June 2014 Observations on the breeding biology of the Pygmy Nightjar Nyctipolus hirundinaceus in the Caatinga of Bahia and Ceará, Brazil 1 2 3 4 Juan Mazar Barnett , Johan Ingels , Andrei Langeloh Roos , Jefferson Luiz Gonçalves de Lima 5,6 and Luciano Nicolas Naka Deceased. Galgenberglaan 9, BE-9070 Destelbergen, Belgium. Instituto Chico Mendes da Biodiversidade. Reserva Extrativista Marinha Pirajubaé, CEP 88047-011, Florianópolis, Santa Catarina, Brazil. Sítio Pau Preto,10, Zona Rural, CEP 63160-000, Potengi, Ceará, Brazil. Laboratório de Ornitologia, Departamento de Zoologia, Universidade Federal de Pernambuco, Av. Prof. Moraes Rego, 1235, CEP 50670-001, Recife, Pernambuco, Brazil. Corresponding author: lnaka1@lsu.edu Received on 24 December 2013. Accepted on 21 April 2014. ABSTRACT: We present the first details of nesting sites, eggs, and chicks of the Pygmy Nightjar (Nyctipolus hirundinaceus), a small nocturnal bird endemic to northeastern Brazil. We conducted behavioral observations near Curaçá in northern Bahia, and at Potengí, southern Ceará, both located in the heart of the Brazilian Caatinga. We found four ‘nests’ in Bahia and another five in Ceará. In all cases, a single egg was laid, and only the females took care of the chick during the day. Pygmy Nightjars in both places bred mostly during the rainy season, as do most of the bird species in the region. By gathering breeding data from throughout the species distribution, we observed that although most populations (c. 75 %) breed during the rainy season, some populations of the race cearae also seem to breed during the dry season. KEY WORDS: Brazil, Caprimulgidae, eggs, nesting biology, reproduction. INTRODUCTION inselbergs in the Atlantic Forest (veilliardi; Cleere 1999). As the name suggests, the Pygmy Nightjar is among the The family Caprimulgidae, which contains smallest Neotropical members of the family (16 – 20 nighthawks and nightjars, includes cryptically colored, cm), and seems to prefer to some extent the vicinity of flat rocky outcrops (lajeiros) both in the Caatinga and the mainly nocturnal species that are often difficult to find and observe. More commonly heard than seen, several Atlantic Forest (Sick 1997; Vasconcelos & Lins 1999; basic aspects of their biology remain poorly known, Ingels et al. 2014). especially for the Neotropical members of the family The evolutionary relationships of this nightjar are (Cleere 1998, 1999; Holyoak 2001). Their breeding only now starting to be unveiled. Formerly located in the genus Caprimulgus, the Pygmy Nightjar was recently biology is particularly under-studied; the most recent monographs on the family have pointed out that ~70% placed in an expanded genus Hydropsalis by the Comitê of all species had either ‘no breeding information’ or their Brasileiro de Registros Ornitológicos (2014), following breeding biology was ‘poorly known’ (Cleere 1998, 1999; the taxonomic recommendations of Han et al. (2010). Holyoak 2001). More recently, the South American Checklist Committee (Remsen et al. 2014) decided to resurrect the genus Among the least known species in the family is the Pygmy Nightjar (Nyctipolus hirundinaceus), endemic to Nyctipolus, including both N. hirundinaceus and the the Brazilian northeast, where three described allopatric Blackish Nightjar (Nyctipolus nigrescens), following the subspecies (hirundinaceus, cearae, and vielliardi) occupy recent findings of the first available molecular sequences open areas in light woodland, scrubland, and areas with of Pygmy Nightjar (Sigurdsson & Cracraft 2014). During January and February 1997, JMB, LNN, xeric vegetation locally known as Caatinga (hirundinaceus and cearae), as well as areas with xeric vegetation on and ALR spent several weeks in the Caatinga of Curaçá, Observations on the breeding biology of the Pygmy Nightjar Nyctipolus hirundinaceus in the Caatinga of Bahia and Ceará, Brazil Juan Mazar Barnett, Johan Ingels, Andrei Langeloh Roos, Jefferson Luiz Gonçalves de Lima and Luciano Nicolas Naka located in the interior of the Brazilian state of Bahia (see surviving wild Spix’s Macaw (Cyanopsitta spixii) (da Ré Mazar Barnett et al. 2014 for a more detailed description 1995, Mazar Barnett et al. 2014). of this site). During that time, while working for the Spix’s Observations around Fazenda Concórdia near Macaw Project (Projeto Ararinha-azul), they observed Curaçá were made between 2 January and 3 February and documented the breeding behavior of 29 different 1997 by JMB, LNN, and ALR, whereas observations at species of birds (Mazar Barnett et al. 2014), including Sítio Pau Preto near Potengi were made by JLGL between several with undescribed nests at the time, such as the November 2013 and May 2014. To evaluate whether Cactus Parakeet (Aratinga cactorum; Naka 1997), the there is a seasonal pattern in the breeding behavior of Lesser Nighthawk (Chordeiles pusillus; Leite et al. 1997), the Pygmy Nightjar, we gathered additional data from the Scarlet-throated Tanager (Compsothraupis loricata; an online source, namely WikiAves (www.wikiaves.org), Mazar Barnett et al. 2014), and the Pygmy Nightjar, of where dated and georeferenced photographs of both eggs which four ‘nests’ were found around the Spix’s Macaw and chicks were available (Table 1). To determine the Project headquarters, at Fazenda Concordia. Although breeding time of the species from available photographs, some basic information from these observations has been we used the date when the egg was photographed. For the reported elsewhere (Cleere 1998, 1999; Holyoak 2001), chicks, we estimated their age in days, and then calculated detailed information on the breeding biology of the the date the eggs were likely laid using a 16-day incubation Pygmy Nightjar remains lacking. More recently, JLGL period, as has been previously reported for similar-sized found, and reports here on, five eggs in southern Ceará, nightjars (Cleere 1999). Although the accuracy of this some 250 km north of Curaçá. method is not ideal, we believe it is accurate enough to Here, we present the first detailed description of the shed light on the general breeding patterns of the species. ‘nest,’ eggs, and chicks of the Pygmy Nightjar, including Because there is considerable geographic variation in observations on the general breeding behavior of this the patterns of rainfall throughout the distribution Brazilian endemic nightjar. We discuss the timing of range of the Pygmy Nightjar, we obtained rainfall data breeding in relation to rainfall, and compare our data for each locality, when available. When these data were with newly available breeding information obtained from not available for the exact location where the eggs and throughout the species’ distributional range. chicks were photographed, we used rainfall data from the closest locality available (Table 1). Rainfall data (1911- 2009) for Bahia, Paraíba, Rio Grande do Norte, and METHODS Ceará were obtained from the Departamento de Ciências Atmosféricas (DCA), Universidade Federal de Campina We studied the breeding behavior of Pygmy Nightjars Grande (http://www.dca.ufcg.edu.br/clima); data for at Fazenda Concórdia (09°09'S, 39°45'W), situated Minas Gerais were obtained from the Instituto Nacional c.25 km south of Curaçá in northern Bahia (Mazar de Pesquisas Espaciais (INPE), Centro de Previsão de Barnett et al. 2014) and at Sítio Pau Preto (07º04'S, Tempo e Estudos Climáticos (CPTEC) (http://www. 40º05'W), located c.5 km west of Potengí in southern cptec.inpe.br/cidades/). We considered records prior to Ceará. Both regions are covered by scrubland and dry the two rainiest months as early rainy season, during the woodland. Fazenda Concórdia is covered by open xeric two rainiest months as rainy season, and after the two vegetation, dominated by bushes and small trees such rainiest months as late rainy season. We considered the as, Cnidoscolus phyllacanthus (Euphorbiaceae), Jatropha first month of drought as early dry season, the last month mollissima (Euphorbiaceae), Caesalpinia pyramidalis of drought as late dry season, and all months in between (Caesalpinoideae), and several species of cacti. Along as the dry season. the seasonal streams filled during the rainy season grows riparian forest dominated by tall Tabebuia caraiba RESULTS (Bignoniaceae) trees. Sítio Pau Preto is known for having an artificial pond that provides water to the town of Potengí. Both areas present clayish soil with abundant General observations gravel, pebbles, and rocks, although Sítio Pau Preto seems to have more rocky outcrops. The natural vegetation Although no quantitative estimates of density or cover of both areas has been severely modified. Fazenda abundance of the Pygmy Nightjar were made either at Concórdia has been subjected to heavy grazing by goats Fazenda Concórdia or Sítio Pau Preto, we noted that and cattle, resulting in a heavily eroded land, whereas corn several individuals or pairs lived in close proximity, and and bean plantations were the main economic activities we believe the species was fairly common in both areas, at Sítio Pau Preto. Although both sites are located in the as individuals were seen on a daily basis. At Fazenda heart of the semi-arid interior of the Caatinga, Curaçá is Concórdia, feeding bouts at night were observed as better known for the presence, until recently, of the last described by Vasconcelos and Figueiredo (1996; Figure Revista Brasileira de Ornitologia, 22(2), 2014 Observations on the breeding biology of the Pygmy Nightjar Nyctipolus hirundinaceus in the Caatinga of Bahia and Ceará, Brazil Juan Mazar Barnett, Johan Ingels, Andrei Langeloh Roos, Jefferson Luiz Gonçalves de Lima and Luciano Nicolas Naka 1), with birds sallying c. 0.5 to 1.0-m upwards from the in an area of open Caatinga with sparse low vegetation ground and landing back on the same spot. We also noted and large areas of bare soil and stony ground. The two individuals that appeared to be feeding while flying very roadside nests B and C, and nest D were within 15 to 20 low over large areas of open ground, mostly in circles or m of each other, whereas nest A was c. 350 m from the loose figures. Once, a female was observed sitting in the other ones. Most observations were made at nest A. We middle of the road, making sharp sideward movements presume that the four eggs belonged to three different with her head, apparently capturing tiny coleopterans pairs: nest A belonged to one pair, nests B and C probably that swarmed in myriads around her. At Sítio Pau to a second pair, and nest D to a third one. The egg of Preto, where rocky outcrops are more abundant, Pygmy nest C is presumed to be a replacement clutch for the Nightjars were seen exclusively in these locations. egg of nest B, which was abandoned and likely eaten by No body mass data are available for this species a predator. (Holyoak 2001). An adult male captured at Fazenda All nests contained a single egg, laid directly on the Concórdia weighed 32 g. The length of his wings and sandy to stony ground. The area immediately around the tail, 133 and 92 mm respectively, seems to place it near eggs was cleared of small gravel, probably as a result of the nominate race, for which Cleere (1998) gives ranges comfort movements when an adult installs itself on the of 119-130 mm and 77-94 mm. egg. All eggs had a pale buffy-cream ground color, with irregular rufous reddish spots and dark blotches, similar to Description of the nest, egg, and chick other eggs of this species found elsewhere in the Caatinga (Figure 1). Three eggs measured 24.1 x 18.3 mm, 24.3 x As is the case for the rest of the family, no actual nest 18.0 mm, and 24.7 x 17.4 mm. is built by Pygmy Nighjars. Four eggs of at least three The egg of nest A was found on 6 January 1997 breeding pairs were found in January 1997 at Fazenda and was probably laid a few days earlier. A second egg Concórdia. Three nests (nests A, B, and C) were found on (nest B) was found in early January, but it already seemed the side of an unpaved road used occasionally by vehicles abandoned and remained unattended for about two and pedestrians, and one nest (nest D) was found c. 15 weeks, when it finally disappeared. The egg of nest C m-away from the same road and within meters of a rocky was found on 24 January, whereas the egg of nest D was outcrop. Nest A was situated near a house surrounded found broken, with an almost fully-grown chick inside, by lush woody vegetation. Nests B and C were found on 3 February. FIGURE 1. Egg of Pygmy Nightjar (Nyctipolus hirundinaceus) found in a depression filled with vegetal litter, gravel, and rock debris on rocky outcrop (lajeiro) Morada Nova (Ceará, 05°07’S, 38°23’W). Photograph by Arthur Grosset. Revista Brasileira de Ornitologia, 22(2), 2014 Observations on the breeding biology of the Pygmy Nightjar Nyctipolus hirundinaceus in the Caatinga of Bahia and Ceará, Brazil Juan Mazar Barnett, Johan Ingels, Andrei Langeloh Roos, Jefferson Luiz Gonçalves de Lima and Luciano Nicolas Naka Two of the eggs found eventually hatched. A chick, a similar looking chick). A day later it weighed 9 g. The estimated to be 2-4 days old, weighing 4 g was found at egg of nest C apparently hatched on 3 February, but we nest A on 24 January. It was covered with grayish white could only find the two halves of the eggshell, with no down with a pattern of rufous-brown lines. On 28 January, sign of a hatchling or any of the adults. Given that an egg when we estimated the chick was 6–8 days old, it weighed of a few days old was found on 6 January at nest A and 7.5 g, and the first feather shafts appeared, rufous on the a 2-4 days old chick was found there 18 days later, we scapulars, and dark in the wings and tail (see Figure 2 for believe that the incubation period lasts at least 16 days. FIGURE 2. Temporal series of an egg, chick, and young of Pygmy Nightjar (Nyctipolus hirundinaceus) at Sítio Pau Preto near Potengí. A) First egg found at site (18 November 2013); B) chick of 1-2 days old (1 February 2014); C) chick of 15-16 days old (16 February 2014); D) same chick with adult female (16 February 2014); E) young bird of 31-32 days old (4 March 2014). Revista Brasileira de Ornitologia, 22(2), 2014 Observations on the breeding biology of the Pygmy Nightjar Nyctipolus hirundinaceus in the Caatinga of Bahia and Ceará, Brazil Juan Mazar Barnett, Johan Ingels, Andrei Langeloh Roos, Jefferson Luiz Gonçalves de Lima and Luciano Nicolas Naka Five eggs were found at Sítio Pau Preto between outcrop. A new visit to rock 1 on 23 March resulted in November and May, 2014. All eggs were found on three the finding of a third egg, laid at exactly the same place rocky outcrops; 3 on rock 1, and 1 on rock 2 and 3, where the chick was found on 16 February. Although it is respectively, located some 300 m apart. All rocky outcrops likely that this third egg was laid by the same pair, several were relatively small: rock 1 measured 40 x 40 m (~1600 adult birds were seen on rock 1, and the use of the same 2 2 m ), rock 2 measured 26 x 15 m (~340 m ), and rock 3 place for breeding by different pairs cannot be discarded. measured 40 x 20 m (~800 m ). A first egg (Figure 2a) This egg could not be relocated on subsequent visits to was found in a crack on rock 1 on 18 November 2013, a the rock. On 26 March 2014 another egg was found at couple of weeks before the beginning of the rainy season, rock 2, and on 31 May 2014 a fifth egg was found on which arrived on 16 December in that particular year. The rock 3 (Figure 3). In all cases, the eggs were laid directly fate of that egg was not followed, but on 1 February 2014 on the bare rock (Figs. 2 and 3). a recently hatched chick (Figure 2b) was found on the same rock, 5 m from the site where the above mentioned Behavior of the chick egg was found, but obviously from a different egg, although likely from the same breeding pair. That same Our observations suggest that a chick remains around chick, with an estimated age of 16 days old (Figure 2c), its nesting site for at least four weeks after hatching. At was observed on 16 February being protected by an adult Curaçá, during the first days after hatching, the chick female (Figure 2d). Presumably the same young bird was of nest A remained in the immediate vicinity of the nest observed again on 4 March at an estimated age of 32-33 site, never moving more than 0.5 m away from that spot. days (Figure 2e). All observations were done on the some These movements were apparently not related to the rocky outcrop and an adult bird was always found in the availability of shade. Six to eight days after hatching, the area. On 4 March, four adult birds were observed at the chick had moved 2.5 m away from the original nesting FIGURE 3. Rocky outcrop (lajeiro) at Sítio Pau Preto (rock 3) near Potengí and egg of Pygmy Nightjar (Nyctipolus hirundinaceus) found on 31 May 2014. Revista Brasileira de Ornitologia, 22(2), 2014 Observations on the breeding biology of the Pygmy Nightjar Nyctipolus hirundinaceus in the Caatinga of Bahia and Ceará, Brazil Juan Mazar Barnett, Johan Ingels, Andrei Langeloh Roos, Jefferson Luiz Gonçalves de Lima and Luciano Nicolas Naka site. At Potengí, the chick found in February remained on Nightjar. Surprisingly, we found reports of nesting birds the rocky outcrop until at least 32 days old, only moving from throughout the year. The only months lacking 1 m away from the egg shells after two weeks, and c.10 m nesting records are August and December. Thus, breeding after 4 weeks. records include all seasons, from the early dry to the late Detailed observations at Curaçá revealed that at dusk rainy season (Table 1). when the adults became active, the chick stretched and walked around. On occasions, it also made short jumps DISCUSSION like the adults do when alarmed. It always remained in the open, but quickly ran to the edge of dense grass to seek cover, when the breeding adult was flushed by The clutch of the Pygmy Nightjar consists of a single egg. approaching humans. When the adult returned and Our own observations and 18 additional records report while still approaching, the chick sometimes ran up to a single egg or chick (Table 1). Although it has been claimed that the clutch size of the Pygmy Nightjar is one it, seeking cover under its raised breast feathers. Food begging was by means of tapping the bill of the adult. or two eggs (Cleere 2010; WikiAves 2013), we have not When handled, the chick uttered soft alarm calls. found any evidence of more than one egg. On the other hand, although we did not mark adult breeding birds, our Behavior of the adults at the nest observations from Potengí suggest that a single pair laid three eggs during five months (November, January, and During daytime, only the female was seen incubating the March) at two individual spots located 5 m apart. egg or brooding the chick, both at nest A (at Curaçá) and Comparing available pictures (Table 1), egg rock 1 (at Potengí). We never saw the male attending the coloration seems very similar throughout the range of the nest before dusk. At Curaçá, throughout the incubation species, which is in contrast to the large color variation found in the adults. Although the Pygmy Nightjar is period, the egg remained almost at the same spot, moving only a few cm sometimes, probably as a result of an adult considered a rocky outcrop specialist (Ingels et al. 2014) turning it. When the female was flushed during the day, our data from Curaçá demonstrate that this species can it took her only c.1 to 2 min to return to the nest site, breed away from these substrates, using bare ground. On and even less, once the chick had hatched. To return, the the other hand, it seems that when rocky outcrops are available as in Potengí, Pygmy Nightjars will lay their adult usually landed 1 to 2 m away from the egg or chick, slowly approaching it by walking in a semi-circle. eggs exclusively on these outcrops. A detailed study of The male that we assumed to be associated with rocky outcrop availability and nesting site selection by nest A roosted within c.5 to 15 m from the nest, in the this species may shed a light on this issue. cover of light leguminous shrubs or trees, as noted by A striking feature of our observations at Fazenda Concórdia was the close proximity of three of the Pygmy Vasconcelos and Figueiredo (1996). Normally, the male was observed taking over the nest at dusk, and was more Nightjar nests. Sick (1997) mentions that Little Nightjars shy and nervous than the female, as he was probably less (Setopagis parvulus) and Sand-colored Nighthawks used to our presence. The male also gave alarm calls more (Chordeiles rupestris) occasionally breed in groups, while often. On occasions, he performed a distraction display Cleere (1999) noted that some caprimulgids breed in a semi-colonial or even colonial way. The Blackish Nightjar, by walking short distances with fanned tail and spread wings raised slightly in V-shape. The female performed a the closest relatives of the Pygmy Nightjar, is known to different distraction display by flattening her body against breed in an almost colonial way on inselbergs in primary the ground and flapping her wings while crawling away rainforest in the Guianas (Ingels et al. 1984, 2009; Cleere from the nest site, a display resembling a ‘broken wing & Ingels 2002). Unfortunately, the small number of nests with detailed observations does not allow a conclusion display.’ The male was seen to do a similar display only once. Both adults responded vigorously to the playback about communal nesting by Pygmy Nightjars. of the chick’s calls, with the female once flying very close, The breeding of nightjars is often linked to moon producing dry sounds by clapping her wings. phases and annual rainfall (Cleere 1999). In most cases, this is likely related to the abundance of food resources, particularly insect abundance. The Caatinga is an Annual rainfall and breeding season extremely seasonal environment, and most breeding seems Five of the nine eggs found during our studies were laid to take place during the beginning of the rainy season, during the early rainy season before the heavier rains when insect abundance peaks in the region (Vasconcelos arrived, three during the rainy season, and only one egg et al. 2010). The rainy season in Curaçá, which lasts a few months, seems to be the preferred season to breed for was laid before the arrival of the rainy season (Table 1). We obtained data for 7 additional eggs and 11 chicks most species (Mazar Barnett et al. 2014), including the found throughout the distribution range of the Pygmy Pygmy Nightjar. From the 27 breeding events we report Revista Brasileira de Ornitologia, 22(2), 2014 Observations on the breeding biology of the Pygmy Nightjar Nyctipolus hirundinaceus in the Caatinga of Bahia and Ceará, Brazil Juan Mazar Barnett, Johan Ingels, Andrei Langeloh Roos, Jefferson Luiz Gonçalves de Lima and Luciano Nicolas Naka Revista Brasileira de Ornitologia, 22(2), 2014 TABLE 1. Breeding records of the Pygmy Nightjar (Nyctipolus hirundinaceus) currently available, including estimated date, coordinates, season when it was laid, and references. Date of eggs is represented by the actual date when they were found and photographed, whereas date of chicks represents our estimate of the date when the egg was laid, considering a 16-day incubation period. Race Locality State Coordinates Date Year Season Author Reference Eggs hirundinaceus Curaçá Bahia 09°09'S, 39°45'W 6 January 1997 Early rainy This study ’’ 8 January 1997 Early rainy This study ’’ 24 January 1997 Early rainy This study ’’ 3 February 1997 Early rainy This study cearae Morada Nova Ceará 05°07'S, 38°22'W 20 October 2008 Dry A. Grosset Ingels et al. 2014 “ 20 October 2008 Dry A. Grosset Ingels et al. 2014 cearae Ipueira Rio Grande do Norte 06°28'S, 37°23'W 1 September 2009 Dry T. Silva WA59203 cearae São Mamede Paraíba 06°56'S, 37°06'W 1 July 2009 Early dry J. Medkraft WA32531 cearae Massapê Ceará 03°31'S, 40°20'W 1 November 2010 Late dry A. Netto WA232763 hirundinaceus Manoel Vitorino Bahia 14°08'S, 40°14'W 14 January 2012 Late rainy O. Borges WA548127 cearae Jaguaribara Ceará 05°40'S, 38°37'W 28 May 2013 Late rainy K. Serra WA973865 cearae Potengi Ceará 07°04'S, 40°05'W 18 November 2013 Late dry This study WA1156756 ’’ 23 March 2014 Rainy This study ’’ 26 March 2014 Rainy This study 31 May 2014 Late rainy This study Fig. 3 Chicks hirundinaceus Mamonas Minas Gerais 16°08'S, 43°32'W 6 March 2005 Rainy E. Luiz WA172359 hirundinaceus Boa Nova Bahia 14°21'S, 40°12'W 20 February 2007 Late rainy E. Luiz WA167354 ’’ 24 February 2007 Late rainy E. Luiz WA167358 cearae Massapê Ceará 03°31'S, 40°20'W 12 October 2009 Dry A. Netto WA75163 cearae Quixadá Ceará 04°58'S, 39°00'W 29 June 2009 Late rainy R. Bessa WA182099 cearae Equador Rio Grande do Norte 06°56'S, 36°43'W 15 February 2009 Early rainy L. Gonzaga WA343446 cearae São Mamede Paraíba 06°56'S, 37°06'W 21 April 2009 Rainy J. Medkraft WA22688 ’’ 17 May 2009 Late rainy J. Medkraft WA26653 ’’ 13 February 2010 Rainy J. Medkraft WA118078 cearae Quixadá Ceará 04°58'S, 39°00'W 9 June 2011 Late rainy C. Albano WA390021 cearae Parelhas Rio Grande do Norte 06°41'S, 36°39'W 21 April 2011 Rainy L. Gonzaga WA355490 cearae Potengi Ceará 07°04'S, 40°05'W 1 February 2014 Early rainy This study WA1285132 Observations on the breeding biology of the Pygmy Nightjar Nyctipolus hirundinaceus in the Caatinga of Bahia and Ceará, Brazil Juan Mazar Barnett, Johan Ingels, Andrei Langeloh Roos, Jefferson Luiz Gonçalves de Lima and Luciano Nicolas Naka in this paper (Table 1), 20 (c.75%) occurred during the REFERENCES early rainy, rainy, or late rainy seasons. One case occurred during the early dry, four during the dry, and two during Ab’Saber, A. N. 1977. Espaços ocupados pela expansão dos climas the late dry season (Table 1). Interestingly, all of the seven secos na América do Sul, por ocasião dos períodos glaciais quaternários. Paleoclimas, 3: 1-19. dry-season breeding records were observed in Ceará, Albano, C. 2011. [WA390021, Hydropsalis hirundinacea (Spix, Paraíba, and Rio Grande do Norte, within the range of 1825)]. www.wikiaves.com/ 390021 (access on 27 April 2014). the race cearae. None of the nesting records further south Borges, O. B. 2012. [WA548127, Hydropsalis hirundinacea (Spix, (presumably within the range of the nominate race) were 1825)]. www.wikiaves.com/548127 (access on 27 June 2014). Bessa, R. 2009. [WA182099, Hydropsalis hirundinacea (Spix, 1825)]. found during the dry season (Table 1). But this does not www.wikiaves.com/182099 (access on 27 June 2014). seem to be the rule, as rainy-season breeders have also Comitê Brasileiro de Registros Ornitológicos. 2014. Listas das been reported in all regions (Table 1). On the other hand, th aves do Brasil. 11 Edition, http://www.cbro.org.br (access on 15 none of the breeding records of the nominate race were January 2014). found during the dry season. Cleere, N. 1998. Nightjars. A Guide to the Nightjars and related Nightbirds. Robertsbridge: Pica Press. Rainfall in the Caatinga, however, can be dramatically Cleere, N. 1999. Family Caprimulgidae (Nightjars), p. 302–386. In: erratic, and can sometimes fail to arrive in any given year del Hoyo, J.; Elliott, A. & Sargatal, J. (eds.). Handbook of the Birds (Ab’Saber 1977). This region is well known for long and of the World. Vol. 5. Barn-owls to Hummingbirds. Barcelona: Lynx fierce draughts that can extend for up to two years. It Edicions. Cleere, N. 2010. Nightjars of the World. Potoos, Frogmouths, Oilbird is possible that local changes in the seasonal patterns of and Owlet-nightjars. Hampshire: WILDGuides Ltd. rainfall may explain these differences, and it would be Cleere, N. & Ingels, J. 2002. Notes on the breeding biology of necessary to collect rainfall and breeding data in the same the Blackish Nightjar Caprimulgus nigrescens in French Guiana. years to make warranted correlations. On the other hand, Alauda, 70: 253–259. ground-nesting nightjars must deal with the danger of Gonzaga, L. (2009). [WA343446, Hydropsalis hirundinacea (Spix, 1825)]. www.wikiaves.com/343446 (access on 27 June 2014). heavy rainfall, which may drown the chick or cool the Gonzaga, L. (2011). [WA355490, Hydropsalis hirundinacea (Spix, eggs. Blackish Nightjars living on inselbergs in French 1825)]. www.wikiaves.com/355490 (access on 27 June 2014). Guiana and Suriname only breed during the dry season, Han, K-L.; Robbins, M. B. & Braun, M. J. 2010. A multi- refraining form breeding when monthly precipitation gene estimate of phylogeny in the nightjars and nighthawks exceeds c. 300 mm of rain (Ingels et al. 1984). In the (Caprimulgidae). Molecular phylogenetics and evolution 55(2): 443-453. case of the Pygmy Nightjar, it seems that the total annual Holyoak, D. T. 2001. Nightjars and their Allies: the Caprimulgiformes. rainfall in the Caatinga is low enough and it rarely rains Oxford: Oxford University Press. more than 200 mm/month, so that avoiding the rainy Ingels, J., Ribot, J. H. & de Jong, B. H. J. 1984. Vulnerability of season may not be necessary. eggs and young of the Blackish Nightjar (Caprimulgus nigrescens) in Suriname. Auk, 101: 388–391. Data on the breeding of the Atlantic Forest race Ingels, J.; Pelletier, V. & Cleere, N. 2009. Nighthawks and nightjars (vielliardi) are not yet available, but given the isolated of the region of Saül, French Guiana. Alauda, 77: 303-308. nature of the three allopatric populations of Pygmy Ingels, J.; Mazar Barnett, J.; Vasconcelos, M. F. & Jackson, H. D. Nightjar, the seasonal timing of reproduction may 2014. The habitat preference of the endemic Pygmy Nightjar Nyctipolus hirundinaceus of Brazil. Revista Brasileira de Ornitologia, represent an important pre-zygotic isolating mechanism 22: (in press). worth exploring in future studies. Leite, L. O., Naka, L. N., Vasconcelos, M. F. & Coelho, M. M. 1997. Aspectos da nidificação do bacurauzinho, Chordeiles pusillus (Caprimulgiformes: Caprimulgidae) nos estados da Bahia e Minas ACKNOWLEDGEMENTS Gerais. Ararajuba, 5: 237–240. Luiz, E. R. 2005. [WA172359, Hydropsalis hirundinacea (Spix, 1825)]. www.wikiaves.com/172359 (access on 27 June 2014). The summer of 1997 was particularly fruitful for the Luiz, E. R. 2005. [WA167354, Hydropsalis hirundinacea (Spix, little town of Curaçá, in the interior of the state of Bahia, 1825)]. www.wikiaves.com/167354 (access on 27 June 2014). mostly because a group of three friends (JMB, LNN and Luiz, E. R. 2005. [WA167358, Hydropsalis hirundinacea (Spix, 1825)]. www.wikiaves.com/167358 (access on 27 June 2014). ALR) were invited by Marcos da Ré to spend the summer Mazar Barnett, J., Silva, C. L. G., Araujo, H. F. P., Roos, A. L., as interns at the Spix’s Macaw Project. We are grateful to Machado, C. G., Uejima, A. M. K. & Naka, L. N. 2014. The Marcos da Ré and Yara de Melo Barros for the wonderful Avifauna of Curaçá (Bahia), the last stronghold of Spix’s Macaw. time spent in the Caatinga. It was a time that will remain in Revista Brasileira de Ornitologia, 22: xxx- our hearts, together with the memories of Juan as a young Medcraft, J. P. 2009. [WA22688, Hydropsalis hirundinacea (Spix, 1825)]. www.wikiaves.com/22688 (access on 27 April 2014). boy running and jumping in the Caatinga after having Medcraft, J. P. 2009. [WA26653, Hydropsalis hirundinacea (Spix, glimpsed the last Spix’s Macaw. We are grateful to Ciro 1825)]. www.wikiaves.com/26653 (access on 27 April 2014). Albano and Arthur Grosset for sending their unpublished Medcraft, J. P. 2009. [WA32531, Hydropsalis hirundinacea (Spix, photographs of eggs and chicks. We are also grateful to 1825)]. www.wikiaves.com/32531 (access on 27 April 2014). Medcraft, J. P. 2009. [WA118178, Hydropsalis hirundinacea (Spix, Marcos Rodrigues, Alexandre Aleixo, and Des Jackson for 1825)]. www.wikiaves.com/118178 (access on 27 April 2014). reading and commenting on earlier drafts of this paper. Revista Brasileira de Ornitologia, 22(2), 2014 Observations on the breeding biology of the Pygmy Nightjar Nyctipolus hirundinaceus in the Caatinga of Bahia and Ceará, Brazil Juan Mazar Barnett, Johan Ingels, Andrei Langeloh Roos, Jefferson Luiz Gonçalves de Lima and Luciano Nicolas Naka Naka, L. N. 1997. Nest and egg description of an endemism of the Sigurdsson, S. & Cracraft, J. 2014. Deciphering the diversity and Brazilian north-east: the Cactus Parakeet (Aratinga cactorum). history of New World nightjars (Aves: Caprimulgidae) using Revista Brasileira de Ornitologia 5: 182-185. molecular phylogenetics. Zoological Journal of the Linnean Society, Netto, A. 2010. [WA75163, Hydropsalis hirundinacea (Spix, 1825)]. 170: 506–545. www.wikiaves.com/75163 (access on 1 June 2014). Vasconcelos, A.; Andreazze, R.; Almeida, A. M.; Araujo, H. F. P.; Netto, A. 2010. [WA232763, Hydropsalis hirundinacea (Spix, 1825)]. Oliveira, E. S. & Oliveira, U. 2010. Seasonality of insects in the www.wikiaves.com/232763 (access on 1 June 2014). semi-arid Caatinga of northeastern Brazil. Revista Brasileira de Silva, T. N. 2009. [WA59203, Hydropsalis hirundinacea (Spix, 1825)]. Entomologia, 54: 471-476. www.wikiaves.com/59203 (access on 1 June, 2014). Vasconcelos, M. F. & Figuereido, C. C. 1996. Observações da Ré, M. 1995. Edging back from the edge of extinction. World preliminares sobre o comportamento do bacurauzinho-da- Birdwatch 17: 17–19. caatinga (Caprimulgus hirundinaceus) na Estação Ecológica de Remsen, J. V.; Jr., Cadena, C. D.; Jaramillo, A.; Nores, M.; Pacheco, Aiuaba-CE. Atualidades Ornitológicas, 73: 13. J. F.; Pérez-Emán, J.; Robbins, M. B.; Stiles, F. G.; Stotz, D. Vasconcelos, M. F. & Lins, L. V. 1999. Photo Spot: Pygmy Nightjar F. & Zimmer, K. J. Version 2014. A classification of the birds Caprimulgus hirundinaceus vielliardi. Cotinga, 11: 74. species of South America. American Ornithologists’Union. http:// WikiAves. 2013. Hydropsalis hirundinacea www.wikiaves.com.br www.museum.lsu.edu/~Remsen/SACCBaseline.htm (accessed on (Access on 22 November 2013). 30 May 2013). Serra, K. S. 2013. [WA973865, Hydropsalis hirundinacea (Spix, 1825)]. www.wikiaves.com/973865 (access on 27 June 2014). Sick, H. 1997. Ornitologia brasileira. Rio de Janeiro: Edições Nova Associate Editor: Alexandre P. Aleixo Fronteira. Revista Brasileira de Ornitologia, 22(2), 2014 http://www.deepdyve.com/assets/images/DeepDyve-Logo-lg.png Ornithology Research Springer Journals

Observations on the breeding biology of the Pygmy Nightjar Nyctipolus hirundinaceus in the Caatinga of Bahia and Ceará, Brazil

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Revista Brasileira de Ornitologia, 22(2), 201-209 ARTICLE June 2014 Observations on the breeding biology of the Pygmy Nightjar Nyctipolus hirundinaceus in the Caatinga of Bahia and Ceará, Brazil 1 2 3 4 Juan Mazar Barnett , Johan Ingels , Andrei Langeloh Roos , Jefferson Luiz Gonçalves de Lima 5,6 and Luciano Nicolas Naka Deceased. Galgenberglaan 9, BE-9070 Destelbergen, Belgium. Instituto Chico Mendes da Biodiversidade. Reserva Extrativista Marinha Pirajubaé, CEP 88047-011, Florianópolis, Santa Catarina, Brazil. Sítio Pau Preto,10, Zona Rural, CEP 63160-000, Potengi, Ceará, Brazil. Laboratório de Ornitologia, Departamento de Zoologia, Universidade Federal de Pernambuco, Av. Prof. Moraes Rego, 1235, CEP 50670-001, Recife, Pernambuco, Brazil. Corresponding author: lnaka1@lsu.edu Received on 24 December 2013. Accepted on 21 April 2014. ABSTRACT: We present the first details of nesting sites, eggs, and chicks of the Pygmy Nightjar (Nyctipolus hirundinaceus), a small nocturnal bird endemic to northeastern Brazil. We conducted behavioral observations near Curaçá in northern Bahia, and at Potengí, southern Ceará, both located in the heart of the Brazilian Caatinga. We found four ‘nests’ in Bahia and another five in Ceará. In all cases, a single egg was laid, and only the females took care of the chick during the day. Pygmy Nightjars in both places bred mostly during the rainy season, as do most of the bird species in the region. By gathering breeding data from throughout the species distribution, we observed that although most populations (c. 75 %) breed during the rainy season, some populations of the race cearae also seem to breed during the dry season. KEY WORDS: Brazil, Caprimulgidae, eggs, nesting biology, reproduction. INTRODUCTION inselbergs in the Atlantic Forest (veilliardi; Cleere 1999). As the name suggests, the Pygmy Nightjar is among the The family Caprimulgidae, which contains smallest Neotropical members of the family (16 – 20 nighthawks and nightjars, includes cryptically colored, cm), and seems to prefer to some extent the vicinity of flat rocky outcrops (lajeiros) both in the Caatinga and the mainly nocturnal species that are often difficult to find and observe. More commonly heard than seen, several Atlantic Forest (Sick 1997; Vasconcelos & Lins 1999; basic aspects of their biology remain poorly known, Ingels et al. 2014). especially for the Neotropical members of the family The evolutionary relationships of this nightjar are (Cleere 1998, 1999; Holyoak 2001). Their breeding only now starting to be unveiled. Formerly located in the genus Caprimulgus, the Pygmy Nightjar was recently biology is particularly under-studied; the most recent monographs on the family have pointed out that ~70% placed in an expanded genus Hydropsalis by the Comitê of all species had either ‘no breeding information’ or their Brasileiro de Registros Ornitológicos (2014), following breeding biology was ‘poorly known’ (Cleere 1998, 1999; the taxonomic recommendations of Han et al. (2010). Holyoak 2001). More recently, the South American Checklist Committee (Remsen et al. 2014) decided to resurrect the genus Among the least known species in the family is the Pygmy Nightjar (Nyctipolus hirundinaceus), endemic to Nyctipolus, including both N. hirundinaceus and the the Brazilian northeast, where three described allopatric Blackish Nightjar (Nyctipolus nigrescens), following the subspecies (hirundinaceus, cearae, and vielliardi) occupy recent findings of the first available molecular sequences open areas in light woodland, scrubland, and areas with of Pygmy Nightjar (Sigurdsson & Cracraft 2014). During January and February 1997, JMB, LNN, xeric vegetation locally known as Caatinga (hirundinaceus and cearae), as well as areas with xeric vegetation on and ALR spent several weeks in the Caatinga of Curaçá, Observations on the breeding biology of the Pygmy Nightjar Nyctipolus hirundinaceus in the Caatinga of Bahia and Ceará, Brazil Juan Mazar Barnett, Johan Ingels, Andrei Langeloh Roos, Jefferson Luiz Gonçalves de Lima and Luciano Nicolas Naka located in the interior of the Brazilian state of Bahia (see surviving wild Spix’s Macaw (Cyanopsitta spixii) (da Ré Mazar Barnett et al. 2014 for a more detailed description 1995, Mazar Barnett et al. 2014). of this site). During that time, while working for the Spix’s Observations around Fazenda Concórdia near Macaw Project (Projeto Ararinha-azul), they observed Curaçá were made between 2 January and 3 February and documented the breeding behavior of 29 different 1997 by JMB, LNN, and ALR, whereas observations at species of birds (Mazar Barnett et al. 2014), including Sítio Pau Preto near Potengi were made by JLGL between several with undescribed nests at the time, such as the November 2013 and May 2014. To evaluate whether Cactus Parakeet (Aratinga cactorum; Naka 1997), the there is a seasonal pattern in the breeding behavior of Lesser Nighthawk (Chordeiles pusillus; Leite et al. 1997), the Pygmy Nightjar, we gathered additional data from the Scarlet-throated Tanager (Compsothraupis loricata; an online source, namely WikiAves (www.wikiaves.org), Mazar Barnett et al. 2014), and the Pygmy Nightjar, of where dated and georeferenced photographs of both eggs which four ‘nests’ were found around the Spix’s Macaw and chicks were available (Table 1). To determine the Project headquarters, at Fazenda Concordia. Although breeding time of the species from available photographs, some basic information from these observations has been we used the date when the egg was photographed. For the reported elsewhere (Cleere 1998, 1999; Holyoak 2001), chicks, we estimated their age in days, and then calculated detailed information on the breeding biology of the the date the eggs were likely laid using a 16-day incubation Pygmy Nightjar remains lacking. More recently, JLGL period, as has been previously reported for similar-sized found, and reports here on, five eggs in southern Ceará, nightjars (Cleere 1999). Although the accuracy of this some 250 km north of Curaçá. method is not ideal, we believe it is accurate enough to Here, we present the first detailed description of the shed light on the general breeding patterns of the species. ‘nest,’ eggs, and chicks of the Pygmy Nightjar, including Because there is considerable geographic variation in observations on the general breeding behavior of this the patterns of rainfall throughout the distribution Brazilian endemic nightjar. We discuss the timing of range of the Pygmy Nightjar, we obtained rainfall data breeding in relation to rainfall, and compare our data for each locality, when available. When these data were with newly available breeding information obtained from not available for the exact location where the eggs and throughout the species’ distributional range. chicks were photographed, we used rainfall data from the closest locality available (Table 1). Rainfall data (1911- 2009) for Bahia, Paraíba, Rio Grande do Norte, and METHODS Ceará were obtained from the Departamento de Ciências Atmosféricas (DCA), Universidade Federal de Campina We studied the breeding behavior of Pygmy Nightjars Grande (http://www.dca.ufcg.edu.br/clima); data for at Fazenda Concórdia (09°09'S, 39°45'W), situated Minas Gerais were obtained from the Instituto Nacional c.25 km south of Curaçá in northern Bahia (Mazar de Pesquisas Espaciais (INPE), Centro de Previsão de Barnett et al. 2014) and at Sítio Pau Preto (07º04'S, Tempo e Estudos Climáticos (CPTEC) (http://www. 40º05'W), located c.5 km west of Potengí in southern cptec.inpe.br/cidades/). We considered records prior to Ceará. Both regions are covered by scrubland and dry the two rainiest months as early rainy season, during the woodland. Fazenda Concórdia is covered by open xeric two rainiest months as rainy season, and after the two vegetation, dominated by bushes and small trees such rainiest months as late rainy season. We considered the as, Cnidoscolus phyllacanthus (Euphorbiaceae), Jatropha first month of drought as early dry season, the last month mollissima (Euphorbiaceae), Caesalpinia pyramidalis of drought as late dry season, and all months in between (Caesalpinoideae), and several species of cacti. Along as the dry season. the seasonal streams filled during the rainy season grows riparian forest dominated by tall Tabebuia caraiba RESULTS (Bignoniaceae) trees. Sítio Pau Preto is known for having an artificial pond that provides water to the town of Potengí. Both areas present clayish soil with abundant General observations gravel, pebbles, and rocks, although Sítio Pau Preto seems to have more rocky outcrops. The natural vegetation Although no quantitative estimates of density or cover of both areas has been severely modified. Fazenda abundance of the Pygmy Nightjar were made either at Concórdia has been subjected to heavy grazing by goats Fazenda Concórdia or Sítio Pau Preto, we noted that and cattle, resulting in a heavily eroded land, whereas corn several individuals or pairs lived in close proximity, and and bean plantations were the main economic activities we believe the species was fairly common in both areas, at Sítio Pau Preto. Although both sites are located in the as individuals were seen on a daily basis. At Fazenda heart of the semi-arid interior of the Caatinga, Curaçá is Concórdia, feeding bouts at night were observed as better known for the presence, until recently, of the last described by Vasconcelos and Figueiredo (1996; Figure Revista Brasileira de Ornitologia, 22(2), 2014 Observations on the breeding biology of the Pygmy Nightjar Nyctipolus hirundinaceus in the Caatinga of Bahia and Ceará, Brazil Juan Mazar Barnett, Johan Ingels, Andrei Langeloh Roos, Jefferson Luiz Gonçalves de Lima and Luciano Nicolas Naka 1), with birds sallying c. 0.5 to 1.0-m upwards from the in an area of open Caatinga with sparse low vegetation ground and landing back on the same spot. We also noted and large areas of bare soil and stony ground. The two individuals that appeared to be feeding while flying very roadside nests B and C, and nest D were within 15 to 20 low over large areas of open ground, mostly in circles or m of each other, whereas nest A was c. 350 m from the loose figures. Once, a female was observed sitting in the other ones. Most observations were made at nest A. We middle of the road, making sharp sideward movements presume that the four eggs belonged to three different with her head, apparently capturing tiny coleopterans pairs: nest A belonged to one pair, nests B and C probably that swarmed in myriads around her. At Sítio Pau to a second pair, and nest D to a third one. The egg of Preto, where rocky outcrops are more abundant, Pygmy nest C is presumed to be a replacement clutch for the Nightjars were seen exclusively in these locations. egg of nest B, which was abandoned and likely eaten by No body mass data are available for this species a predator. (Holyoak 2001). An adult male captured at Fazenda All nests contained a single egg, laid directly on the Concórdia weighed 32 g. The length of his wings and sandy to stony ground. The area immediately around the tail, 133 and 92 mm respectively, seems to place it near eggs was cleared of small gravel, probably as a result of the nominate race, for which Cleere (1998) gives ranges comfort movements when an adult installs itself on the of 119-130 mm and 77-94 mm. egg. All eggs had a pale buffy-cream ground color, with irregular rufous reddish spots and dark blotches, similar to Description of the nest, egg, and chick other eggs of this species found elsewhere in the Caatinga (Figure 1). Three eggs measured 24.1 x 18.3 mm, 24.3 x As is the case for the rest of the family, no actual nest 18.0 mm, and 24.7 x 17.4 mm. is built by Pygmy Nighjars. Four eggs of at least three The egg of nest A was found on 6 January 1997 breeding pairs were found in January 1997 at Fazenda and was probably laid a few days earlier. A second egg Concórdia. Three nests (nests A, B, and C) were found on (nest B) was found in early January, but it already seemed the side of an unpaved road used occasionally by vehicles abandoned and remained unattended for about two and pedestrians, and one nest (nest D) was found c. 15 weeks, when it finally disappeared. The egg of nest C m-away from the same road and within meters of a rocky was found on 24 January, whereas the egg of nest D was outcrop. Nest A was situated near a house surrounded found broken, with an almost fully-grown chick inside, by lush woody vegetation. Nests B and C were found on 3 February. FIGURE 1. Egg of Pygmy Nightjar (Nyctipolus hirundinaceus) found in a depression filled with vegetal litter, gravel, and rock debris on rocky outcrop (lajeiro) Morada Nova (Ceará, 05°07’S, 38°23’W). Photograph by Arthur Grosset. Revista Brasileira de Ornitologia, 22(2), 2014 Observations on the breeding biology of the Pygmy Nightjar Nyctipolus hirundinaceus in the Caatinga of Bahia and Ceará, Brazil Juan Mazar Barnett, Johan Ingels, Andrei Langeloh Roos, Jefferson Luiz Gonçalves de Lima and Luciano Nicolas Naka Two of the eggs found eventually hatched. A chick, a similar looking chick). A day later it weighed 9 g. The estimated to be 2-4 days old, weighing 4 g was found at egg of nest C apparently hatched on 3 February, but we nest A on 24 January. It was covered with grayish white could only find the two halves of the eggshell, with no down with a pattern of rufous-brown lines. On 28 January, sign of a hatchling or any of the adults. Given that an egg when we estimated the chick was 6–8 days old, it weighed of a few days old was found on 6 January at nest A and 7.5 g, and the first feather shafts appeared, rufous on the a 2-4 days old chick was found there 18 days later, we scapulars, and dark in the wings and tail (see Figure 2 for believe that the incubation period lasts at least 16 days. FIGURE 2. Temporal series of an egg, chick, and young of Pygmy Nightjar (Nyctipolus hirundinaceus) at Sítio Pau Preto near Potengí. A) First egg found at site (18 November 2013); B) chick of 1-2 days old (1 February 2014); C) chick of 15-16 days old (16 February 2014); D) same chick with adult female (16 February 2014); E) young bird of 31-32 days old (4 March 2014). Revista Brasileira de Ornitologia, 22(2), 2014 Observations on the breeding biology of the Pygmy Nightjar Nyctipolus hirundinaceus in the Caatinga of Bahia and Ceará, Brazil Juan Mazar Barnett, Johan Ingels, Andrei Langeloh Roos, Jefferson Luiz Gonçalves de Lima and Luciano Nicolas Naka Five eggs were found at Sítio Pau Preto between outcrop. A new visit to rock 1 on 23 March resulted in November and May, 2014. All eggs were found on three the finding of a third egg, laid at exactly the same place rocky outcrops; 3 on rock 1, and 1 on rock 2 and 3, where the chick was found on 16 February. Although it is respectively, located some 300 m apart. All rocky outcrops likely that this third egg was laid by the same pair, several were relatively small: rock 1 measured 40 x 40 m (~1600 adult birds were seen on rock 1, and the use of the same 2 2 m ), rock 2 measured 26 x 15 m (~340 m ), and rock 3 place for breeding by different pairs cannot be discarded. measured 40 x 20 m (~800 m ). A first egg (Figure 2a) This egg could not be relocated on subsequent visits to was found in a crack on rock 1 on 18 November 2013, a the rock. On 26 March 2014 another egg was found at couple of weeks before the beginning of the rainy season, rock 2, and on 31 May 2014 a fifth egg was found on which arrived on 16 December in that particular year. The rock 3 (Figure 3). In all cases, the eggs were laid directly fate of that egg was not followed, but on 1 February 2014 on the bare rock (Figs. 2 and 3). a recently hatched chick (Figure 2b) was found on the same rock, 5 m from the site where the above mentioned Behavior of the chick egg was found, but obviously from a different egg, although likely from the same breeding pair. That same Our observations suggest that a chick remains around chick, with an estimated age of 16 days old (Figure 2c), its nesting site for at least four weeks after hatching. At was observed on 16 February being protected by an adult Curaçá, during the first days after hatching, the chick female (Figure 2d). Presumably the same young bird was of nest A remained in the immediate vicinity of the nest observed again on 4 March at an estimated age of 32-33 site, never moving more than 0.5 m away from that spot. days (Figure 2e). All observations were done on the some These movements were apparently not related to the rocky outcrop and an adult bird was always found in the availability of shade. Six to eight days after hatching, the area. On 4 March, four adult birds were observed at the chick had moved 2.5 m away from the original nesting FIGURE 3. Rocky outcrop (lajeiro) at Sítio Pau Preto (rock 3) near Potengí and egg of Pygmy Nightjar (Nyctipolus hirundinaceus) found on 31 May 2014. Revista Brasileira de Ornitologia, 22(2), 2014 Observations on the breeding biology of the Pygmy Nightjar Nyctipolus hirundinaceus in the Caatinga of Bahia and Ceará, Brazil Juan Mazar Barnett, Johan Ingels, Andrei Langeloh Roos, Jefferson Luiz Gonçalves de Lima and Luciano Nicolas Naka site. At Potengí, the chick found in February remained on Nightjar. Surprisingly, we found reports of nesting birds the rocky outcrop until at least 32 days old, only moving from throughout the year. The only months lacking 1 m away from the egg shells after two weeks, and c.10 m nesting records are August and December. Thus, breeding after 4 weeks. records include all seasons, from the early dry to the late Detailed observations at Curaçá revealed that at dusk rainy season (Table 1). when the adults became active, the chick stretched and walked around. On occasions, it also made short jumps DISCUSSION like the adults do when alarmed. It always remained in the open, but quickly ran to the edge of dense grass to seek cover, when the breeding adult was flushed by The clutch of the Pygmy Nightjar consists of a single egg. approaching humans. When the adult returned and Our own observations and 18 additional records report while still approaching, the chick sometimes ran up to a single egg or chick (Table 1). Although it has been claimed that the clutch size of the Pygmy Nightjar is one it, seeking cover under its raised breast feathers. Food begging was by means of tapping the bill of the adult. or two eggs (Cleere 2010; WikiAves 2013), we have not When handled, the chick uttered soft alarm calls. found any evidence of more than one egg. On the other hand, although we did not mark adult breeding birds, our Behavior of the adults at the nest observations from Potengí suggest that a single pair laid three eggs during five months (November, January, and During daytime, only the female was seen incubating the March) at two individual spots located 5 m apart. egg or brooding the chick, both at nest A (at Curaçá) and Comparing available pictures (Table 1), egg rock 1 (at Potengí). We never saw the male attending the coloration seems very similar throughout the range of the nest before dusk. At Curaçá, throughout the incubation species, which is in contrast to the large color variation found in the adults. Although the Pygmy Nightjar is period, the egg remained almost at the same spot, moving only a few cm sometimes, probably as a result of an adult considered a rocky outcrop specialist (Ingels et al. 2014) turning it. When the female was flushed during the day, our data from Curaçá demonstrate that this species can it took her only c.1 to 2 min to return to the nest site, breed away from these substrates, using bare ground. On and even less, once the chick had hatched. To return, the the other hand, it seems that when rocky outcrops are available as in Potengí, Pygmy Nightjars will lay their adult usually landed 1 to 2 m away from the egg or chick, slowly approaching it by walking in a semi-circle. eggs exclusively on these outcrops. A detailed study of The male that we assumed to be associated with rocky outcrop availability and nesting site selection by nest A roosted within c.5 to 15 m from the nest, in the this species may shed a light on this issue. cover of light leguminous shrubs or trees, as noted by A striking feature of our observations at Fazenda Concórdia was the close proximity of three of the Pygmy Vasconcelos and Figueiredo (1996). Normally, the male was observed taking over the nest at dusk, and was more Nightjar nests. Sick (1997) mentions that Little Nightjars shy and nervous than the female, as he was probably less (Setopagis parvulus) and Sand-colored Nighthawks used to our presence. The male also gave alarm calls more (Chordeiles rupestris) occasionally breed in groups, while often. On occasions, he performed a distraction display Cleere (1999) noted that some caprimulgids breed in a semi-colonial or even colonial way. The Blackish Nightjar, by walking short distances with fanned tail and spread wings raised slightly in V-shape. The female performed a the closest relatives of the Pygmy Nightjar, is known to different distraction display by flattening her body against breed in an almost colonial way on inselbergs in primary the ground and flapping her wings while crawling away rainforest in the Guianas (Ingels et al. 1984, 2009; Cleere from the nest site, a display resembling a ‘broken wing & Ingels 2002). Unfortunately, the small number of nests with detailed observations does not allow a conclusion display.’ The male was seen to do a similar display only once. Both adults responded vigorously to the playback about communal nesting by Pygmy Nightjars. of the chick’s calls, with the female once flying very close, The breeding of nightjars is often linked to moon producing dry sounds by clapping her wings. phases and annual rainfall (Cleere 1999). In most cases, this is likely related to the abundance of food resources, particularly insect abundance. The Caatinga is an Annual rainfall and breeding season extremely seasonal environment, and most breeding seems Five of the nine eggs found during our studies were laid to take place during the beginning of the rainy season, during the early rainy season before the heavier rains when insect abundance peaks in the region (Vasconcelos arrived, three during the rainy season, and only one egg et al. 2010). The rainy season in Curaçá, which lasts a few months, seems to be the preferred season to breed for was laid before the arrival of the rainy season (Table 1). We obtained data for 7 additional eggs and 11 chicks most species (Mazar Barnett et al. 2014), including the found throughout the distribution range of the Pygmy Pygmy Nightjar. From the 27 breeding events we report Revista Brasileira de Ornitologia, 22(2), 2014 Observations on the breeding biology of the Pygmy Nightjar Nyctipolus hirundinaceus in the Caatinga of Bahia and Ceará, Brazil Juan Mazar Barnett, Johan Ingels, Andrei Langeloh Roos, Jefferson Luiz Gonçalves de Lima and Luciano Nicolas Naka Revista Brasileira de Ornitologia, 22(2), 2014 TABLE 1. Breeding records of the Pygmy Nightjar (Nyctipolus hirundinaceus) currently available, including estimated date, coordinates, season when it was laid, and references. Date of eggs is represented by the actual date when they were found and photographed, whereas date of chicks represents our estimate of the date when the egg was laid, considering a 16-day incubation period. Race Locality State Coordinates Date Year Season Author Reference Eggs hirundinaceus Curaçá Bahia 09°09'S, 39°45'W 6 January 1997 Early rainy This study ’’ 8 January 1997 Early rainy This study ’’ 24 January 1997 Early rainy This study ’’ 3 February 1997 Early rainy This study cearae Morada Nova Ceará 05°07'S, 38°22'W 20 October 2008 Dry A. Grosset Ingels et al. 2014 “ 20 October 2008 Dry A. Grosset Ingels et al. 2014 cearae Ipueira Rio Grande do Norte 06°28'S, 37°23'W 1 September 2009 Dry T. Silva WA59203 cearae São Mamede Paraíba 06°56'S, 37°06'W 1 July 2009 Early dry J. Medkraft WA32531 cearae Massapê Ceará 03°31'S, 40°20'W 1 November 2010 Late dry A. Netto WA232763 hirundinaceus Manoel Vitorino Bahia 14°08'S, 40°14'W 14 January 2012 Late rainy O. Borges WA548127 cearae Jaguaribara Ceará 05°40'S, 38°37'W 28 May 2013 Late rainy K. Serra WA973865 cearae Potengi Ceará 07°04'S, 40°05'W 18 November 2013 Late dry This study WA1156756 ’’ 23 March 2014 Rainy This study ’’ 26 March 2014 Rainy This study 31 May 2014 Late rainy This study Fig. 3 Chicks hirundinaceus Mamonas Minas Gerais 16°08'S, 43°32'W 6 March 2005 Rainy E. Luiz WA172359 hirundinaceus Boa Nova Bahia 14°21'S, 40°12'W 20 February 2007 Late rainy E. Luiz WA167354 ’’ 24 February 2007 Late rainy E. Luiz WA167358 cearae Massapê Ceará 03°31'S, 40°20'W 12 October 2009 Dry A. Netto WA75163 cearae Quixadá Ceará 04°58'S, 39°00'W 29 June 2009 Late rainy R. Bessa WA182099 cearae Equador Rio Grande do Norte 06°56'S, 36°43'W 15 February 2009 Early rainy L. Gonzaga WA343446 cearae São Mamede Paraíba 06°56'S, 37°06'W 21 April 2009 Rainy J. Medkraft WA22688 ’’ 17 May 2009 Late rainy J. Medkraft WA26653 ’’ 13 February 2010 Rainy J. Medkraft WA118078 cearae Quixadá Ceará 04°58'S, 39°00'W 9 June 2011 Late rainy C. Albano WA390021 cearae Parelhas Rio Grande do Norte 06°41'S, 36°39'W 21 April 2011 Rainy L. Gonzaga WA355490 cearae Potengi Ceará 07°04'S, 40°05'W 1 February 2014 Early rainy This study WA1285132 Observations on the breeding biology of the Pygmy Nightjar Nyctipolus hirundinaceus in the Caatinga of Bahia and Ceará, Brazil Juan Mazar Barnett, Johan Ingels, Andrei Langeloh Roos, Jefferson Luiz Gonçalves de Lima and Luciano Nicolas Naka in this paper (Table 1), 20 (c.75%) occurred during the REFERENCES early rainy, rainy, or late rainy seasons. One case occurred during the early dry, four during the dry, and two during Ab’Saber, A. N. 1977. Espaços ocupados pela expansão dos climas the late dry season (Table 1). Interestingly, all of the seven secos na América do Sul, por ocasião dos períodos glaciais quaternários. Paleoclimas, 3: 1-19. dry-season breeding records were observed in Ceará, Albano, C. 2011. [WA390021, Hydropsalis hirundinacea (Spix, Paraíba, and Rio Grande do Norte, within the range of 1825)]. www.wikiaves.com/ 390021 (access on 27 April 2014). the race cearae. None of the nesting records further south Borges, O. B. 2012. [WA548127, Hydropsalis hirundinacea (Spix, (presumably within the range of the nominate race) were 1825)]. www.wikiaves.com/548127 (access on 27 June 2014). Bessa, R. 2009. [WA182099, Hydropsalis hirundinacea (Spix, 1825)]. found during the dry season (Table 1). But this does not www.wikiaves.com/182099 (access on 27 June 2014). seem to be the rule, as rainy-season breeders have also Comitê Brasileiro de Registros Ornitológicos. 2014. Listas das been reported in all regions (Table 1). On the other hand, th aves do Brasil. 11 Edition, http://www.cbro.org.br (access on 15 none of the breeding records of the nominate race were January 2014). found during the dry season. Cleere, N. 1998. Nightjars. A Guide to the Nightjars and related Nightbirds. Robertsbridge: Pica Press. Rainfall in the Caatinga, however, can be dramatically Cleere, N. 1999. Family Caprimulgidae (Nightjars), p. 302–386. In: erratic, and can sometimes fail to arrive in any given year del Hoyo, J.; Elliott, A. & Sargatal, J. (eds.). Handbook of the Birds (Ab’Saber 1977). This region is well known for long and of the World. Vol. 5. Barn-owls to Hummingbirds. Barcelona: Lynx fierce draughts that can extend for up to two years. It Edicions. Cleere, N. 2010. Nightjars of the World. Potoos, Frogmouths, Oilbird is possible that local changes in the seasonal patterns of and Owlet-nightjars. Hampshire: WILDGuides Ltd. rainfall may explain these differences, and it would be Cleere, N. & Ingels, J. 2002. Notes on the breeding biology of necessary to collect rainfall and breeding data in the same the Blackish Nightjar Caprimulgus nigrescens in French Guiana. years to make warranted correlations. On the other hand, Alauda, 70: 253–259. ground-nesting nightjars must deal with the danger of Gonzaga, L. (2009). [WA343446, Hydropsalis hirundinacea (Spix, 1825)]. www.wikiaves.com/343446 (access on 27 June 2014). heavy rainfall, which may drown the chick or cool the Gonzaga, L. (2011). [WA355490, Hydropsalis hirundinacea (Spix, eggs. Blackish Nightjars living on inselbergs in French 1825)]. www.wikiaves.com/355490 (access on 27 June 2014). Guiana and Suriname only breed during the dry season, Han, K-L.; Robbins, M. B. & Braun, M. J. 2010. A multi- refraining form breeding when monthly precipitation gene estimate of phylogeny in the nightjars and nighthawks exceeds c. 300 mm of rain (Ingels et al. 1984). In the (Caprimulgidae). Molecular phylogenetics and evolution 55(2): 443-453. case of the Pygmy Nightjar, it seems that the total annual Holyoak, D. T. 2001. Nightjars and their Allies: the Caprimulgiformes. rainfall in the Caatinga is low enough and it rarely rains Oxford: Oxford University Press. more than 200 mm/month, so that avoiding the rainy Ingels, J., Ribot, J. H. & de Jong, B. H. J. 1984. Vulnerability of season may not be necessary. eggs and young of the Blackish Nightjar (Caprimulgus nigrescens) in Suriname. Auk, 101: 388–391. Data on the breeding of the Atlantic Forest race Ingels, J.; Pelletier, V. & Cleere, N. 2009. Nighthawks and nightjars (vielliardi) are not yet available, but given the isolated of the region of Saül, French Guiana. Alauda, 77: 303-308. nature of the three allopatric populations of Pygmy Ingels, J.; Mazar Barnett, J.; Vasconcelos, M. F. & Jackson, H. D. Nightjar, the seasonal timing of reproduction may 2014. The habitat preference of the endemic Pygmy Nightjar Nyctipolus hirundinaceus of Brazil. Revista Brasileira de Ornitologia, represent an important pre-zygotic isolating mechanism 22: (in press). worth exploring in future studies. Leite, L. O., Naka, L. N., Vasconcelos, M. F. & Coelho, M. M. 1997. Aspectos da nidificação do bacurauzinho, Chordeiles pusillus (Caprimulgiformes: Caprimulgidae) nos estados da Bahia e Minas ACKNOWLEDGEMENTS Gerais. Ararajuba, 5: 237–240. Luiz, E. R. 2005. [WA172359, Hydropsalis hirundinacea (Spix, 1825)]. www.wikiaves.com/172359 (access on 27 June 2014). The summer of 1997 was particularly fruitful for the Luiz, E. R. 2005. [WA167354, Hydropsalis hirundinacea (Spix, little town of Curaçá, in the interior of the state of Bahia, 1825)]. www.wikiaves.com/167354 (access on 27 June 2014). mostly because a group of three friends (JMB, LNN and Luiz, E. R. 2005. [WA167358, Hydropsalis hirundinacea (Spix, 1825)]. www.wikiaves.com/167358 (access on 27 June 2014). ALR) were invited by Marcos da Ré to spend the summer Mazar Barnett, J., Silva, C. L. G., Araujo, H. F. P., Roos, A. L., as interns at the Spix’s Macaw Project. We are grateful to Machado, C. G., Uejima, A. M. K. & Naka, L. N. 2014. The Marcos da Ré and Yara de Melo Barros for the wonderful Avifauna of Curaçá (Bahia), the last stronghold of Spix’s Macaw. time spent in the Caatinga. It was a time that will remain in Revista Brasileira de Ornitologia, 22: xxx- our hearts, together with the memories of Juan as a young Medcraft, J. P. 2009. [WA22688, Hydropsalis hirundinacea (Spix, 1825)]. www.wikiaves.com/22688 (access on 27 April 2014). boy running and jumping in the Caatinga after having Medcraft, J. P. 2009. [WA26653, Hydropsalis hirundinacea (Spix, glimpsed the last Spix’s Macaw. We are grateful to Ciro 1825)]. www.wikiaves.com/26653 (access on 27 April 2014). Albano and Arthur Grosset for sending their unpublished Medcraft, J. P. 2009. [WA32531, Hydropsalis hirundinacea (Spix, photographs of eggs and chicks. We are also grateful to 1825)]. www.wikiaves.com/32531 (access on 27 April 2014). Medcraft, J. P. 2009. [WA118178, Hydropsalis hirundinacea (Spix, Marcos Rodrigues, Alexandre Aleixo, and Des Jackson for 1825)]. www.wikiaves.com/118178 (access on 27 April 2014). reading and commenting on earlier drafts of this paper. Revista Brasileira de Ornitologia, 22(2), 2014 Observations on the breeding biology of the Pygmy Nightjar Nyctipolus hirundinaceus in the Caatinga of Bahia and Ceará, Brazil Juan Mazar Barnett, Johan Ingels, Andrei Langeloh Roos, Jefferson Luiz Gonçalves de Lima and Luciano Nicolas Naka Naka, L. N. 1997. Nest and egg description of an endemism of the Sigurdsson, S. & Cracraft, J. 2014. Deciphering the diversity and Brazilian north-east: the Cactus Parakeet (Aratinga cactorum). history of New World nightjars (Aves: Caprimulgidae) using Revista Brasileira de Ornitologia 5: 182-185. molecular phylogenetics. Zoological Journal of the Linnean Society, Netto, A. 2010. [WA75163, Hydropsalis hirundinacea (Spix, 1825)]. 170: 506–545. www.wikiaves.com/75163 (access on 1 June 2014). Vasconcelos, A.; Andreazze, R.; Almeida, A. M.; Araujo, H. F. P.; Netto, A. 2010. [WA232763, Hydropsalis hirundinacea (Spix, 1825)]. Oliveira, E. S. & Oliveira, U. 2010. Seasonality of insects in the www.wikiaves.com/232763 (access on 1 June 2014). semi-arid Caatinga of northeastern Brazil. Revista Brasileira de Silva, T. N. 2009. [WA59203, Hydropsalis hirundinacea (Spix, 1825)]. Entomologia, 54: 471-476. www.wikiaves.com/59203 (access on 1 June, 2014). Vasconcelos, M. F. & Figuereido, C. C. 1996. Observações da Ré, M. 1995. Edging back from the edge of extinction. World preliminares sobre o comportamento do bacurauzinho-da- Birdwatch 17: 17–19. caatinga (Caprimulgus hirundinaceus) na Estação Ecológica de Remsen, J. V.; Jr., Cadena, C. D.; Jaramillo, A.; Nores, M.; Pacheco, Aiuaba-CE. Atualidades Ornitológicas, 73: 13. J. F.; Pérez-Emán, J.; Robbins, M. B.; Stiles, F. G.; Stotz, D. Vasconcelos, M. F. & Lins, L. V. 1999. Photo Spot: Pygmy Nightjar F. & Zimmer, K. J. Version 2014. A classification of the birds Caprimulgus hirundinaceus vielliardi. Cotinga, 11: 74. species of South America. American Ornithologists’Union. http:// WikiAves. 2013. Hydropsalis hirundinacea www.wikiaves.com.br www.museum.lsu.edu/~Remsen/SACCBaseline.htm (accessed on (Access on 22 November 2013). 30 May 2013). Serra, K. S. 2013. [WA973865, Hydropsalis hirundinacea (Spix, 1825)]. www.wikiaves.com/973865 (access on 27 June 2014). Sick, H. 1997. Ornitologia brasileira. Rio de Janeiro: Edições Nova Associate Editor: Alexandre P. Aleixo Fronteira. Revista Brasileira de Ornitologia, 22(2), 2014

Journal

Ornithology ResearchSpringer Journals

Published: Jun 1, 2014

Keywords: Brazil; Caprimulgidae; eggs; nesting biology; reproduction

References