Polygyny and male parental care by Sprague's Pipit.
Abstract: Sprague's Pipit (Anthus spragueii) is a ground-nesting passerine of the northern mixed-grass prairie. Few studies have examined the natural history and demography of the species, and little information exists on its mating system or parental care. We installed video monitoring systems at 11 randomly selected pipit nests to study parental care and nestling growth. We recorded one male providing parental care to nestlings in two nests active during the same time period. The male began delivering prey to nestlings and removing fecal sacs at both nests on day 5 of the nestling period. The male re-nested with one of his original mates 11 days after the nests failed, and began providing parental care to nestlings from day 2 until fledging. This is the first case of polygyny and male parental care documented for Sprague's Pipit. Future studies should examine the extent of polygyny in Sprague's Pipit using marked adults and genetic analyses.
Article Type: Report
Subject: Polygamy (Observations)
Passeriformes (Behavior)
Courtship of animals (Research)
Familial behavior in animals (Research)
Authors: Dohms, Kimberly M.
Davis, Stephen K.
Pub Date: 12/01/2009
Publication: Name: The Wilson Journal of Ornithology Publisher: Wilson Ornithological Society Audience: Academic Format: Magazine/Journal Subject: Biological sciences Copyright: COPYRIGHT 2009 Wilson Ornithological Society ISSN: 1559-4491
Issue: Date: Dec, 2009 Source Volume: 121 Source Issue: 4
Topic: Event Code: 310 Science & research Canadian Subject Form: Mating behaviour; Animal familial behaviour
Geographic: Geographic Scope: Canada Geographic Code: 1CANA Canada
Accession Number: 216267554
Full Text: Social monogamy is the predominant mating system for the majority of bird species. This mating system is characterized by one male and one female forming a pair bond, and raising a brood together. Polygyny is an alternative mating system in which one male pairs with multiple females (Lack 1968). Polygyny has been documented for 71 North American passerines (Ford 1996), many of which nest in marsh or grassland habitats with high primary production and abundant food resources (Verner and Willson 1966, Ford 1983). Polygyny is the typical mating system of some grassland passerines (Pleszczynska and Hansell 1980; Wittenberger 1980; Post and Greenlaw 1982; Lanyon 1994, 1995; Temple 2002), and has been occasionally documented for others (Welsh 1975, With 1994, Jones and Comely 2002).

Sprague's Pipit (Anthus spragueii) is a ground-nesting passerine of the northern mixed-grass prairie and has been described as one of the least-known species in North America (Robbins and Dale 1999). The species is recognized globally as Vulnerable (BirdLife International 2008), and is listed as threatened in Canada (COSEWIC 2000). Sprague's Pipit is believed to be monogamous and males are thought to provide little, or no parental care (Robbins and Dale 1999). However, like many grassland passerines, details of the mating system and male parental care for the species have not been well studied.

We used video monitoring to study parental care and nestling ecology of Sprague's Pipit. Video monitoring allows researchers to record nests continually with minimal disturbance to the birds (McQuillen and Brewer 2000). Continual investigator disturbance at the nest can cause reduced parental attendance (Westmoreland and Best 1985), increase the likelihood of nest abandonment (Gotmark 1992), and increase predation risk (Westmoreland and Best 1985, Major 1990, Verboven et al. 2001). Video monitoring has been used successfully to examine parental care in passerines including Gray Catbird (Dumetella carolinensis) (Hanley et al. 2007) and Blue Tit (Cyanistis caeruleus) (Dickens and Hartley 2007). The objective of our paper is to report the first documented case of polygyny and male parental care for Sprague's Pipit.

METHODS

We located and monitored Sprague's Pipit nests from 2004 to 2008. Study sites were at the Last Mountain Lake National Wildlife Area, and Nokomis and Wreford Agriculture and Agri-Food Canada community pastures at the north end of Last Mountain Lake, Saskatchewan (51[degrees] 20' N, 105[degrees] 15' W). Our study area consisted of tracts of native mixed-grass pasture, planted grasslands used for hay and pasture, wetlands, and cropland (Davis and Fisher 2009). We located nests from 14 May to 26 August between 0600 and 1400 hrs CST. Each pair of researchers dragged a weighted 25 m polypropylene rope with metal cans attached every 0.5 m across the vegetation to flush adults from nests (Davis 2003). We also located nests fortuitously while conducting other activities.

We installed small (37 x 86 mm) color/ infrared video cameras (National Electronics Bullet C/IR Low Light Color Bullet Camera, Brookvale, NSW, AU) in 2006 at 11 randomly selected nests. Cameras were mounted on small metal stands ~30 cm from the nest entrance. Sprague's Pipits returned to the nest within 20 min of camera placement (unpubl. data) and we assume the cameras had little influence on their behavior. Each camera was connected via coaxial cable (RG6) to a time-lapse 24-hr videocassette recorder (VCR, Sanyo SRT 2400DC or 4040DC, Concord, ON), and a 12-volt deep cycle marine battery concealed beneath vented boxes at least 50 m from the nest. We changed the videocassette each morning and continued recording video footage starting late in incubation (12-15 days) or on hatch day until nestlings fledged (11-14 days) (Davis 2009) or the nest failed.

We captured incubating females by flushing them into 6-m mist nets placed near the nest entrance. Females were banded with a single Federal aluminum band and marked with non-toxic permanent marker (Newell Rubbermaid Sharpie fine point, Sandy Springs, GA, USA) on their breast or flanks to facilitate identification. We placed a pair of 6-m mist nets in a V-formation with a wooden pipit decoy inserted into the ground 1-2 m from the intersection point of the mist nets to capture males. We placed a digital caller (Western Rivers Predation MP3 game caller, Lexington, TN, USA) next to the decoy and broadcast a territorial male song. Males were banded with a Federal aluminum band and a unique combination of color bands. Unique band combinations and markings allowed us to identify individuals, but nest-site vegetation at times precluded us from seeing the markers clearly.

The three nests we discuss in this paper (#'s 9, 38, and 93) were in the same territory, cared for by the same male during the 2006 breeding season, and were recorded on video from hatch day until the nests failed (#'s 9 and 38) or nestlings fledged (# 93). We recorded the gender of the attending adult, the adult's behavior (e.g., prey delivery, brooding, and vigilance), and nest arrival and departure times while reviewing the video footage. We calculated the number of prey deliveries from days 1 to 7 of the nestling period and for each bird when the gender of the adult could be identified. We limited our calculations to this period as nests #'s 9 and 38 failed on day 7 of the nestling period. Data are presented as mean [+ or -] SE.

RESULTS

Nest # 9 was initiated on 20 May (Table 1). We captured and banded male # 380 75 m from the nest on 25 May. We documented male # 380 at the nest for the first time on 4 June from video footage, delivering prey to 5-day old nestlings. Male # 380 continued to provision nestlings and remove fecal sacs up to and including day 7 (12 Jun) when the nest failed due to cold temperatures and heavy precipitation. Neither adult returned to the nest after nest failure.

Nest # 38 (70 m from nest # 9) was initiated by female # 383 and male # 380 on 30 May (Table 1). We observed male # 380 delivering prey to 5-day old nestlings for the first time on 17 June. We found three dead nestlings and one live nestling in the nest on 21 June. Mortality was due to cold temperatures and heavy precipitation. Male # 380 continued to provide parental care up to and including day 7 until the remaining nestling was depredated on 22 June. Neither adult was seen attending the nest after that time.

Female # 383 (from nest # 38) and male # 380 re-nested (nest # 93) on 2 July in male # 380's territory, 54 m from their previous nest (# 9) and 58 m from the site of nest # 38. Male # 380 provided parental care for the first time on 15 July, delivering prey to 2-day old nestlings. We documented male # 380 delivering prey to nestlings up to and including 25 July when one nestling fledged (day 11). We observed both adults attending the young until the fledgling left the camera view.

We documented male # 380 providing prey items and removing nestling fecal sacs at all three nests, but did not observe him brooding nestlings at any time. Parents provisioned nestlings at similar rates at nest # 9 (1.2 provisionings/ nestling/hr) and nest # 38 (1.1 provisionings/ nestling/hr). Provisioning rates in the third recorded nesting attempt were similar to nests #'s 9 and 38 (1.1 provisionings/nestling/hr) (Table 2).

DISCUSSION

Our study is the first to document polygyny and male parental care in Sprague's Pipit. We cannot presently ascertain whether polygyny is the typical mating system of the species, but it is consistent with other studies that found polygyny to be relatively common among grassland species (Verner and Willson 1966, Ford 1983). Furthermore, between 2004 and 2008, we found 11 territories with two or more nests initiated within 11 days of each other, 15-60 m apart (41 [+ or -] 5 m). These circumstances are consistent with those associated with the polygynous nest, but we lack direct evidence linking these territories to polygynous males. Polygynous mating systems may be more common in grassland habitats because of the availability of key resources such as abundant or accessible food (Welsh 1975, Wittenberger 1980), or availability of nest sites (Pleszczynska and Hansell 1980, Temple 2002).

Predation is a driving factor in the rapid development of altricial young in many species (Remes and Martin 2002), and predators are often cited as the primary reason for low nest success rates in passerines (Heske et al. 2001). Providing intensive parental care is essential for raising altricial young (Silver et al. 1985, Starck and Ricklefs 1998), and bi-parental care is expected for species where nest success is low (Clutton-Brock 1991). Sprague's Pipit is prone to high rates of nest failure in some areas as a result of nest predation and inclement weather (Davis 2003, Davis and Fisher 2009). Bi-parental care is common in Anthus species (Halupka 1994, Verbeek and Hendricks 1994, Frey-Roos et al. 1995, Rauter et al. 2000), and may allow females to fledge more young than without male parental care (Webster 1991, Johnson et. al 1993, Lynn and Wingfield 2003). Male parental effort is important for nesting success; serial polygyny (male with an active nest pairing with another mate) (Miskelly 1990, Yosef 1992) may allow male pipits to invest parental care without dividing effort between same age clutches.

Male Sprague's Pipits have been assumed to be monogamous and to not provide parental care (Robbins and Dale 1999). Video monitoring, and marking individual males and females, allowed us to document a single case of polygyny and confirm that males assist females in brood rearing. We encourage future studies to consider using marked birds and genetic analysis to examine if Sprague's Pipits are typically monogamous or polygynous.

ACKNOWLEDGMENTS

We thank R. J. Fisher, K. A. Martin, L. D. Parker, D. J. Sawatzy, J. M. Szarkowicz, and a number of volunteers for assistance during the 2006 field season. We are grateful to D. G. McMaster (Saskatchewan Watershed Authority) and T. I. Wellicome (Canadian Wildlife Service) for generous loan of video monitoring systems. A. E. Dohms and two anonymous reviewers provided valuable comments on previous versions of this manuscript. This study was made possible by financial support from the National Science and Engineering Research Council (KMD), Saskatchewan Environment Fish and Wildlife Fund (KMD), Canadian Wildlife Service-Environment Canada, and the Government of Canada Interdepartmental Recovery Fund. All procedures were approved by the Canadian Wildlife Service Animal Care Committee (#EC-PN-06-001).

Received 4 December 2008. Accepted 15 June 2009.

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Kimberly M. Dohms (1,3) and Stephen K. Davis (1,2)

(1) Department of Biology, University of Regina, 3737 Wascana Parkway, Regina, SK S4S 0A2, Canada.

(2) Canadian Wildlife Service, 300-2365 Albert Street, Regina, SK S4P 2K1, Canada.

(3) Corresponding author; e-mail: dohmskk@uregina.ca
TABLE 1. Events in the nesting cycle for three Sprague's Pipit nests
(#'s 9, 38, and 93) initiated and cared for by the same male in 2006.

                     Initiation   Hatch    Chicks
Nest #   Female ID      date       date    hatched         Fate

9        unhanded      20 May      3 Jun      5      Failed 12 Jun
38         # 383       30 May     13 Jun      4      Failed 22 Jun
93         # 383        2 Jul     14 Jul      4      1 chick
                                                       fledged 25 Jul

TABLE 2. Number of nestling prey deliveries by male # 380 and mates
for the first 7 days of the nestling period.

       Prey deliveries
                          Total prey
                         deliveries by   Total prey
                          identified     deliveries   Observation
Nest   Male     Female      adults        observed    effort (hrs)

9       59       166          225           429            72
38      17        9           26            305            71
93      20       341          361           425            93

Totals  96       516          612          1,159          236
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