Reproductive life history traits of the Yellowish Pipit (Anthus lutescens).
Abstract: We describe reproductive traits of the Yellowish Pipit (Anthus lutescens) in the State of Sao Paulo, Brazil. We found 32 active nests during three breeding seasons (2008-2010). Domed nests were built exclusively on the ground where the grass was sufficiently tall to conceal them. Clutch initiation across years occurred from July to October and average [+ or -] SD clutch size was 3.05 [+ or -] 0.4 eggs or young. Yellowish Pipits were predominantly single-brooded. Eggs were pale white with brown spots and blotches that could be more concentrated at the larger end or homogeneously distributed over the entire surface. Eggs were 18.2 [+ or -] 0.8 mm in length, 13.7 [+ or -] 0.3 mm in width, and weighed 1.7 [+ or -] 0.12 g. Incubation and nestling periods lasted 13.03 [+ or -] 0.2 and 14.5 [+ or -] 1.0 days, respectively. Mean time incubating/hr was 38 [+ or -] 7.1 rain, and incubation recesses averaged 9.4 [+ or -] 4 min. Young were provisioned on average 13.3 [+ or -] 7.9 times/hr, by both males and females. Estimated overall nesting success using a null model of constant nest survival rates was 87% (95% CI, 56-97%). Model selection analyses indicated survival was negatively correlated to nest age and time within the breeding season. Comparisons of Yellowish Pipit life history traits with northern temperate congeners provided support for the premises that clutch sizes are smaller and young development is slower in the tropics. The hypothesis that annual fecundity can be similar across latitudes due to a negative correlation between clutch size and number of renesting attempts was not supported. Our data contradicted the commonly claimed, but poorly tested hypothesis, that smaller clutch sizes in the tropics can be explained by a longer breeding season that permit more opportunities to renest within the same breeding season.
Article Type: Report
Subject: Life history theory (Research)
Animal reproduction (Research)
Motacillidae (Physiological aspects)
Authors: Freitas, Maikon S.
Francisco, Mercival R.
Pub Date: 03/01/2012
Publication: Name: The Wilson Journal of Ornithology Publisher: Wilson Ornithological Society Audience: Academic Format: Magazine/Journal Subject: Biological sciences Copyright: COPYRIGHT 2012 Wilson Ornithological Society ISSN: 1559-4491
Issue: Date: March, 2012 Source Volume: 124 Source Issue: 1
Topic: Event Code: 310 Science & research
Geographic: Geographic Scope: Brazil Geographic Code: 3BRAZ Brazil
Accession Number: 285207250
Full Text: Several studies have addressed avian life history adaptations of northern temperate versus tropical and southern temperate habitats. Broad latitudinal patterns of reproductive traits have been proposed, i.e., Southern Hemisphere species have smaller clutch sizes (Moreau 1944, Lack 1947, Skutch 1949, Murray 1985), lay more clutches per year (Lack and Moreau 1965, Ricklefs 1969), and have longer incubation and nestling periods (Skutch 1949). The occurrence of latitudinal differences in clutch size has been tested (Yom-Tov et al. 1994, Young 1994, Geffen and Yom-Toy 2000, Martin et al. 2000, Ghalambor and Martin 2001), but the claim that incubation and nestling periods, as well as number of renesting attempts, differ between Northern and Southern hemispheres is still not accepted (Geffen and Yom-Tov 2000). Only a few studies have compared species paired by phylogeny and ecology, and attempted to isolate the latitudinal effect from the phylogenetic influence (Yom-Toy et al. 1994, Martin et al. 2000, Ghalambor and Martin 2001, Martin 2002). Passerines of the genus Anthus (Motacillidae) are globally distributed, and occur on every continent except Antarctica (Ridgely and Tudor 1994, Tyler 2004). This makes them well suited for studying breeding trait diversification across latitudes. However, many motacillids have not been studied in detail, especially those in South America (Tyler 2004).

We present the first comprehensive description of the reproductive life history traits of the South American Yellowish Pipit (A. lutescens). Our objectives were to: (1) provide information on phenology and duration of breeding season, clutch size, length of incubation and nestling periods, nesting success, renesting attempts, and parental care for a Sao Paulo State population, southeast Brazil; and (2) compare these life history traits with data from the literature for a set of northern temperate congeners.

METHODS

Study Area.--Observations of Yellowish Pipits were conducted along the Sorocaba River and in an adjacent urban park (16 ha), including an artificial lake of 2.8 ha. The study area is in the suburbs of the city of Sorocaba, State of Sao Paulo in southeast Brazil (23[degrees] 28' 99" S, 47[degrees] 26' 17" W). The area is characterized by the presence of large patches of exotic grasses (mainly Zoysia japonica and Cynodon dactylon). Trees and bushes are widely spaced and the grass is kept relatively short by employees of Sorocaba city prefecture, which provided suitable breeding habitat for Yellowish Pipits. The climate is tropical with two well-marked seasons: a humid, hot season from October through March (average rainfall = 919 mm, temperatures varied from 15.7 to 32.4 [degrees]C) and a dry, cold season from April through September (average rainfall = 294 mm, temperatures varied from 11.4 to 30.6[degrees]C).

Study Species.--The Yellowish Pipit is a monomorphic passerine widely distributed in South America from western Panama south to Uruguay and Argentina (south to La Pampaand Buenos Aires). It inhabits grasslands, open Cerrado, pastures, and cultivated lands often near water or marshes (Ridgely and Tudor 1994, Sick 1997, Tyler 2004). The Yellowish Pipit is a common species, but data on its breeding biology are scattered and many aspects are poorly documented. Data on incubation and nestling periods, breeding phenology, and parental care are presented here for the first time.

Field Procedures.--We conducted systematic nest searches from July to January during three breeding seasons (2008, 2009, and 2010). We searched for nests at least three times per week by covering 1 km of each margin of the Sorocaba River and the entire adjacent park. Nests were located by following adults in their territories when they were carrying nest material or feeding young, and were checked every 1-3 days. We used metal calipers accurate to [+ or -] 0.01 mm to measure nests and eggs, and a spring scale accurate to [+ or -] 0.1 g to weigh eggs.

The incubation period was from the first day of incubation to the day before hatching, and nestling period was from hatching day to the day before fledging. Observations were performed daily during the laying stage, and we could detect females that began incubation before and after the set of eggs was complete. We also checked if eggs were warm to detect the beginning of incubation. We did not touch or handle young to avoid shortening the nestling period (Skutch 1945). Clutch initiation dates were obtained from nests found in the construction stage (i.e., we observed the first egg in the nests) (n = 15), and by backdating for nests for which hatching or fledging dates were known, based on mean incubation and nestling periods (n = 12).

We estimated the frequency at which adults brought materials to build nests, the proportion of time females spent incubating the eggs, as well as the frequency of provisioning visits during the nestling period from 1-hr focal observation sessions every 1-3 days using 8 x 40 binoculars. These observations were made early in the morning (0600-0900 hrs).

We assumed predation to have occurred when eggs or nestlings younger than fledging age (with poorly developed feathers) disappeared from a nest or when eggs or young were found partially eaten near nests. Abandonment was considered when adults were not seen near the nests during the observation sessions for at least 3 consecutive days, and eggs were cold (Pletschet and Kelly 1990).

Statistical Analysis.--We used the maximum likelihood method implemented in Program MARK, Version 6.1 (White and Burnham 1999, Dinsmore et al. 2002) to estimate nest success based on 20 nests for which information met the criteria for Program MARK (pooling all years). We calculated daily survival rate (DSR) using the null model of constant DSR, S(.), which is similar to that of Mayfield (1961). The cumulative probability of overall nest success was estimated by raising DSR to the power corresponding to the mean duration of the nesting cycle obtained in our study (incubation + nestling periods). We evaluated a set of three Program MARK candidate DSR models that used continuously varying covariates: (1) S(nest age) (number of days since incubation onset date), (2) S(date) (date within the breeding season), and (3) their combinations, S(date + nest age). These models were compared to the null model of constant DSR using Akaike's Information Criterion for small samples (AI[C.sub.c]). The model with the lowest AI[C.sub.c] value was considered the best fit of the data, but models with [DELTA]AI[C.sub.c] [less than or equal to] 2 were also considered as presenting substantial support for explaining the data (Burnham and Anderson 1998). Akaike weight ([w.sub.i]) was used to measure the relative support for each candidate model. We used logit-link function to convert all DSR values to an interval between 0 and one. Model-averaged estimates of DSR were calculated for early (arbitrarily defined as the 10th day after the encounter of the first nest), middle (50th day), and late (90th day) breeding season. Egg-laying stage was not considered in our DSR analyses due to lack of parental activities during this stage.

The clutch size of Yellowish Pipit was compared to populations of: Buff-bellied (American) Pipit (A. rubescens) from Wyoming, USA (Verbeek 1970); Sprague's Pipit (A. spragueii) from southern Saskatchewan, Canada (Davis 2003, 2009), and England and Northern Europe populations of Meadow Pipits (A. pratensis) (Coulson 1956, Davies 1958). Incubation (laying to hatching of the last egg) and nestling periods (hatching to fledging of the last young) were compared to England populations of Meadow Pipit. These analyses were performed by bilateral t-tests using BioEstat 2.0 (Ayres et al. 2000). There are divergences in habitat types among these species, but we believe they are well suited for comparison, as they are all ground insectivores, occur in open habitats, and build open to domed nests on the ground.

[FIGURE 1 OMITTED]

RESULTS

We analyzed nests during three breeding seasons, 2008 (n = 9), 2009 (n = 8), and 2010 (n = 15). Nests were domed with a small side entrance, often invisible from above. Nests were exclusively on the ground, where the grass was sufficiently tall to conceal them, usually in slight depressions in the soil under dense vegetation. Fourteen nests were on flat terrain, while 18 were in crevices in the banks of the Sorocaba River, 2-5 m from water. Nest material consisted of dry grasses, and the nest chamber was lined with finer grass leaves and grass stems (Fig. 1). Nest measurements varied (Table 1) and one nest had a tubular entrance ~13 cm in length. Males and females shared nest building activities with both carrying and placing nest materials, often one after the other at similar rates (MSF, pers. obs.). Adults brought nest materials on average 12.6 [+ or -] 8.4 times/hr (range = 4-24) during 7 hrs of focal observation (n = 6 nests). Time gaps between visits were 3.6 [+ or -] 4.7 min in length (range = 0.1-25, n = 88 observations). Two nests found at the beginning of construction took 3 days to complete.

The earliest clutch initiation dates (laying of the first egg) varied among years: 20 August 2008, 6 September 2009, and 28 July 2010 (Fig. 2). The latest clutch initiation was on 19 October 2008, and the latest nesting activity (the last young observed in a nest) was on 4 November 2008. The average breeding season (pooling the 3 yrs together) spanned almost 5 months, but nests found in July and November were exceptions (only 1 nest each). Laying initiations were highly concentrated in August and September, and the average number of active nests was concentrated in August through October (Fig. 3). However, clutch initiations were clearly concentrated in only 1 month (Fig. 2) when each year was analyzed separately.

The basic egg color was pale white and markings varied remarkably, even within a single nest. Some eggs were predominantly white with pale to intense brown spots and blotches more concentrated at the larger end, while others were heavily spotted over the entire surface (Fig. 1). Eggs measured 18.2 [+ or -] 0.82 mm in length (range = 17.1-19.7) and 13.7 [+ or -] 0.3 mm in width (range = 13.3-14.2), and weighed 1.7 [+ or -] 0.12 g (range = 1.5-2.0) (n = 16). Clutch sizes were two (n = 1), three (n = 18), or four (n = 2) eggs or young (3.05 [+ or -] 0.4), and eggs were invariably laid on consecutive days (n = 15 nests). Incubation of 10 observed nests, started on the third day after onset of laying, even when clutch sizes were two (n = 1 nest) or four eggs (n = 1 nest). One egg took 14 days to hatch, but the incubation period was 13 days (29 eggs from 10 nests) (13.03 [+ or -] 0.2). We believe that only females incubated for two reasons: (1) we did not observe adults taking turns to incubate; and (2) the non-incubating individual often remained near the nest singing vigorously, suggesting it was the male. Males did not feed females in the nests in 31 hrs of focal observations at nine different nests, but usually escorted them during incubation recesses. Females spent from 23.2 to 55.8 min incubating the eggs per hr (38 [+ or -] 7.1 min). They left the nests 1-3 times per hr (2.2 [+ or -] 0.6), and incubation recesses were 3.8 to 26.3 min in length (9.4 [+ or -] 4, n = 53). When incubating females were flushed from nests, they performed a 'broken-wing' display, vocalizing and dragging their wings along the ground in an attempt to distract the observers.

[FIGURE 2 OMITTED]

[FIGURE 3 OMITTED]

Hatching was synchronous (nests were checked from 0900 to l000 hrs) in 12 of 16 nests. Nestlings had yellowish skin and were covered with gray down at hatching. The bill and swollen flanges were yellow and the mouth lining was bright orange. Adults gathered a large number of small insects at the tip of their beaks, which were offered intact to the young. Both parents provisioned nestlings, often at similar rates, arriving near the nest together and feeding the young one after the other. Young were provisioned on average 13.3 [+ or -] 7.9 times/hr (range = 3-35) in 29 hrs of observation at 11 nests and feeding recesses lasted 3.8 [+ or -] 3.54 min (range = 0.05-24.4, n = 346). Both males and females were observed removing fecal sacs directly from the young, which were either swallowed or dropped a few meters from nests. Fledging was synchronous in 13 of 14 nests, and young departed from nests with well developed tail and wings and flying well, differently from many altricial young passerines that leave the nests before they can fly. Nestling periods of 26 young from 11 different nests were 13 (n = 3), 14 (n = 12), 15 (n = 9) or 17 days (n = 2) (14.5 [+ or -] 1.0). Nests were not reused.

Three of 28 nests for which fate was known were abandoned in late construction stage (11%), two were abandoned in the incubation stage (7%), two were predated during the incubation stage (7%), two were predated during the nestling stage (7%), and 19 successfully fledged young (68%). Three of 55 eggs that were not predated or abandoned were infertile (5.4%), each in a different nest. Nests for which we had evidence of human predation (i.e., during grass cutting) were not considered. The average nest exposure period was 27.5 days (13 days of incubation and 14.5 days of nestling stage). Estimated nest daily survival rate using the null model of constant DSR was 0.995 with a 95% confidence interval (Cl) of 0.979-0.999. Estimated overall nesting success was 87% (95% Cl, 56-97%) (419 nest days and 4 nest failures).

Two models of DSR received substantial support ([DELTA]AICc [less than or equal to] 2). The best-fit model was S(nest age), being 1.4 AICc units better than the second best model, S(nest age + date). These models together accounted for 57% of the data variation ([w.sub.i] values = 0.38 and 0.19), and the null model of constant DSR received less support ([DELTA]AICc = 2.3, and [w.sub.i] = 0.12). The S(nest age) model indicated a negative correlation between DSR and nest age ([beta] = -0.245, Cl = -0.553-0.062), and the S(nest age + date) model indicated DSR was negatively correlated with both nest age ([beta] = -0.227, CI = -0.544-0.089) and date ([beta] = -0.043, CI = -0.146-0.059). Model averaging revealed DSRs of 0.999 (CI = 0.994-1.004) in early, 0.999 (CI = 0.993-1.003) in middle, and 0.433 (CI = 0.006-0.989) in late breeding season.

Comparisons of breeding life history traits of Yellowish Pipit and northern temperate congeners (Sprague' s, Meadow, and American pipits) varied (Table 2). The clutch size of Yellowish Pipit was significantly smaller when compared to Sprague's Pipit (t = -12.5, P = 0.000), American Pipit (t = - 14.4, P = 0.000), and England (t = -13.4, P = 0.000), and northern Europe populations (t = -15.9, P = 0.000) of Meadow Pipits. Incubation periods of our study population and England populations of Meadow Pipits did not differ significantly (t = 0.33, P = 0.74), but the nestling period was significantly shorter for Meadow Pipits (t = 6.45, P = 0.000).

DISCUSSION

The breeding seasons of central-south Brazilian birds follow a general pattern, beginning in August/September (at the end of the dry season), reaching the highest peak from October to December (early and middle rainy season), and diminishing or ending in January-February (late rainy season) (Davis 1945, Sick 1997, Piratelli et al. 2000, Marini and Duraes 2001). Yellowish Pipits had an earlier pattern (i.e., 89% of the clutch initiations occurred during the cold/dry season), concentrating its nesting activities in dry months (Aug-Sep), and ceasing in early rainy season (Oct). Nests of this species are poorly adapted to heavy rains. All nests that were abandoned (n = 6) occurred after the loose nest walls became moist and collapsed (but we did not detect flooding in nests in the fiver banks). Five abandonments occurred in 2009, an exceptionally rainy year with frequent storms even in July and September. Clutch initiation was also later in 2009 with the number of clutch initiations smaller than in 2008 and 2010.

Clutch initiation of Yellowish Pipit lasted 2 to 3 months when each of the three study years was analyzed separately, but were highly concentrated in a single month. Our data reveal a short breeding season length when compared to congeners from the Northern Hemisphere, and suggests that if double-brooding occurs in this population, it must be rare. Laying periods lasted from April to July for central Europe Meadow Pipits (but middle Jun to Jul in northern Europe), from early May to late July in Sprague's Pipits (Coulson 1956, Davies 1958, Jones et al. 2010), but mid June to mid July for American Pipits in Wyoming alpine tundra (Verbeek 1970). The double-brooding rate seems low for Sprague's Pipit (Sutter et al. 1996, Davis 2009), but common in central Europe populations of Meadow Pipits (Davies 1958).

Breeding season length has been often positively correlated with the number of broods per season and, consequently, to annual fecundity (Cooper et al. 2005). It has been theorized that tropical and southern temperate birds produce fewer offspring than northern temperate species by having smaller clutch sizes (Lack 1947, Skutch 1949, Geffen and Yom-Tov 2000, Ricklefs 2000, Auer et al. 2007). Recent reviews have shown that while clutch size tends to increase away from the tropics, nesting attempts tend to decrease, resulting in similar annual fecundity across latitudes (Martin 1995, Boehning-Gaese et al. 2000, Cooper et al. 2005). Thus, reproductive investment would be better estimated by the combination of clutch size and number of broods per season (Cooper et al. 2005). Our study population must be typically single-brooded, and clutch sizes are smaller than northern temperate pipits; thus, our study corroborates the findings that not only clutch sizes are smaller for the tropical representative (Yellowish Pipit) population, but also total annual egg productivity. Thus, the hypothesis that annual fecundity can be similar across latitudes due to a negative correlation between clutch size and number of renesting attempts was not supported (Martin 1995, Boehning-Gaese et al. 2000, Cooper et al. 2005). Our data also contradicts the commonly claimed, but poorly tested hypothesis, that smaller clutch sizes in the tropics can be explained by a longer breeding season that permits more opportunities to renest within the same breeding season (Martin 1996). Skutch (1954) also found that ground-nesting parulids were single-brooded in both Central and North America. Auer et al. (2007) found similar breeding season lengths of guilds of turdids, emberizids, and flycatchers from subtropical Argentina compared with their North American counterparts. However, numbers of within-season renesting attempts were not recorded and the annual fecundity comparisons between Central-South and North American birds remain poorly evaluated.

Jones et al. (2010) found much lower values of model-averaging nest success (27%) in a long-term study on Sprague's Pipit than obtained for our study population of Yellowish Pipit. This refutes Skutch's (1949) premise that higher nest predation in the tropics is a key factor explaining smaller clutch sizes in the Southern Hemisphere through constraining the rate at which parents can deliver food to the young. It supports the findings of Martin et al. (2000) that nest predation may not account for clutch size differences between eight pairs of phylogenetically close and ecologically similar passerines from northern and southern temperate latitudes. However, DSR in our study area could be overestimated due to the presence of people visiting the park. Yellowish Pipits were very tolerant of human presence, but park visitors may have repulsed potential nest predators.

Developmental rates are usually thought to be slower in the tropics and Southern Hemisphere habitats, causing longer incubation and nestling periods compared to north temperate populations (Skutch 1949, 1985; Ricklefs 1968, 1976; Mason 1985; Martin 2002). Geffen and Yom-Tov (2000) analyzed a large sample of tropical and northern temperate passerine species from the Old and New World; they concluded incubation and nestling periods actually tend to be similar, and the differences reported in previous studies could be due to phylogenetic influence. These authors, to control for phylogeny, analyzed the representatives of different infra-orders and continents separately (New World Deutro-Oscines, New World Oscines, and Old World Oscines), and ecological aspects, such as habitat and nest types, were not considered. The study performed by Martin (2002) compared nine pairs of passerines (3 of them intrageneric) from Arizona and subtropical Argentinean Yunga forests, and corroborated the premises of longer incubation and nestling periods in Southern Hemisphere species. We did not find support for a longer incubation period for Yellowish Pipit in relation to Northern Hemisphere congeners, being similar to Meadow and Sprague's pipits. The incubation period for populations of American Pipit from Wyoming alpine tundra, seem to be longer than that of Yellowish Pipit. Our findings corroborated the tendency for longer nestling periods in Yellowish Pipits (although quite close to that of American Pipits). Our comparisons must be viewed with caution because incubation and nestling periods presented in many of the studies are based on small sample sizes, and statistical tests were performed only between Yellowish and England Meadow pipits.

The proposition of the major patterns of bird breeding life history adaptations to varying latitudes can be traced to the 1940s (Moreau 1944, Lack 1947, Skutch 1949), but the premise of slower developmental rates and lower annual fecundity in Southern Hemisphere passerines still relies on a limited number of comparisons that controlled for phylogeny and ecology (Martin et al. 2000, Martin 2002). Many other passerine genera are distributed in both Northern and Southern hemispheres, and they should be the focus in future studies. Providing basic life history data for the poorly studied tropical representatives, and inclusion of raw data in papers (clutch sizes, and incubation and nestling periods) will permit more accurate comparisons.

ACKNOWLEDGMENTS

We are grateful to Secretaria do Meio Ambiente de Sorocaba for authorizing field work in the study area. We especially thank S. L. Jones, S. K. Davis, M. N. Schlindwein, A. J. Piratelli, and an anonymous referee for important comments on the previous versions of this manuscript. M. S. Freitas was supported by a fellowship from PIBIC/CNPq.

Received 14 February 2011. Accepted 15 August 2011.

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MAIKON S. FREITAS (1) AND MERCIVAL R. FRANCISCO (1,2)

(1) Universidade Federal de Sao Carlos, Campus de Sorocaba, Rod. Joao Leme dos Santos, km 110, Sorocaba, SP 18052-780, Brazil.

(2) Corresponding author; e-mail: mercival@ufscar.br
TABLE 1. Measurements (mm) of Yellowish Pipit nests (n = 11):
outside high (OH), outside width (OW), outside length (OL), inside
high (IH), inside diameter (ID), entrance width (EW), and
entrance high (EH).

           OH         OW         OL            IH          ID

Mean       88         114.4      130.25        62          82.4
[+ or -]   [+ or -]   [+ or -]   [+ or -]      [+ or -]    [+ or -]
SD         8.3        11.7       15.6          8           16.7
Range      71-104     93-127.5   104.5-157.3   50.1-74.7   55.8-108.5

           EW         EH

Mean       56         46.5
[+ or -]   [+ or -]   [+ or -]
SD         9.9        98.8
Range      37.7-68    30-58.6

TABLE 2. Average values, range, and sample sizes (n) of breeding
life history traits of Yellowish Pipit and northern temperate congeners
from Europe and North America, and their references.

Species           Clutch size (range)       Incubation period (range)

Yellowish Pipit   3.05 [+ or -] 0.4 (2-4)   13.03 (13-14)
                           n = 21              n = 10

Sprague's Pipit   4.8 [+ or -] 0.84 (2-6)   13.4 (12-15)
                           n = 57              n = 9

American Pipit    4.7 [+ or -] 0.7 (3-6)    14.4 (not provided)
                           n = 87              n = 9

Meadow Pipit      5.4 [+ or -] 0.58 (4-6)   13 (13)
                           n = 23              n = 2

                  4.3 [+ or -] 0.64 (3-6)   13 (11-15)
                           n = 246             n = 19

Species           Nestling period (range)   Reference

Yellowish Pipit   14.5 (14-17)              Present work
                      n = 11

Sprague's Pipit   12.1 (11-14)              Davis (2003, 2009)
                      n = 39

American Pipit    14.4 (not provided)       Verbeek (1970)
                      n = 41

Meadow Pipit      12.3 (12-13)              Davies (1958)
                      n = 6                 (northern Europe)

                  12.3 (10-14)              Coulson (1956)
                      n = 36                (England)
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