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2007 SOUTHEASTERN NATURALIST 6(2):283–292
Nest Success of Grassland Birds in Florida Dry Prairie
Dustin W. Perkins1,2,* and Peter D. Vickery3
Abstract - This paper represents the first study of reproductive success for Sturnella
magna (Eastern Meadowlark) and Columbina passerina (Common Ground-dove) in
Florida dry prairie and, to our knowledge, the first published study of reproductive
rates of Chordeiles minor (Common Nighthawk) in North America. We located 34
Eastern Meadowlark, 13 Common Ground-dove, and 14 Common Nighthawk nests
during the 1997 and 1998 breeding seasons. We estimated daily nest success (standard
error) to be 0.93 (0.01), 0.94 (0.02), and 0.93 (0.02) for Eastern Meadowlarks,
Common Ground-doves, and Common Nighthawks, respectively. Subsequently,
total nest success was 0.16, 0.22, and 0.28 for Eastern Meadowlarks, Common
Ground-doves and Common Nighthawks. Predation was the most common cause for
nest failure. Our estimates of nest success for Eastern Meadowlarks and Common
Ground-doves are generally lower than reported for other regions, which could be
due to the small and fragmented nature of Florida dry prairie.
Grassland birds have received increased attention in recent years as
their populations have declined. Grassland bird declines in North America
are well documented (Brennan and Kuvlesky 2005, Peterjohn and Sauer
1999), with the best-supported explanation being the loss of breeding habitat
(Goriup 1988, Herkert et al. 1996, Vickery et al. 1999). Florida dry
prairie has similarly suffered substantial declines. Historically, dry prairie
was the dominant grassland in central Florida, occurring from the southwest
shoreline of Lake Okeechobee, north and east to Lake Kissimmee and
Osceola County (Howell 1932). Florida dry prairie is endemic to the state
and is ranked as a G2 (globally imperiled) community type (FNAI and
FDNR 1990, Grossmann et al. 1994) due to its rarity. Shriver and Vickery
(1999) reported that at least 81% of the dry prairie habitat in central
Florida has been converted to other uses, primarily improved pasture and
Perkins et al. (1998, 2003) have previously reported nest success rates in
Florida dry prairie for 2 rare species Aimophila aestivalis Lichtenstein
(Bachman’s Sparrow) and the federally endangered Ammodramus
savannarum floridanus Mearns (Florida Grasshopper Sparrow). While conducting
fieldwork on these 2 rare species we monitored nests of 3 other
1Department of Forestry and Wildlife Conservation, University of Massachusetts,
Amherst, MA 01003. 2Current address - 1175 Texas Avenue, Biology Department,
Mesa State College, National Part Service, Grand Junction, CO 81501 3Current
address - Center for Ecological Research, PO Box 127, Richmond, ME 04357.
*Corresponding author - firstname.lastname@example.org.
284 Southeastern Naturalist Vol. 6, No. 2
ground-nesting grassland species, Sturnella magna L. (Eastern Meadowlark),
Chordeiles minor Forster (Common Nighthawk), and Columbina
passerina L. (Common Ground-dove). While none of these species are
restricted to dry prairie in Florida, this ecosystem was historically a major
breeding habitat for these species, but one for which there are no reported
reproductive rates. In addition, studies of reproductive rates for Common
Nighthawks and Common Ground-doves are rare. The objective of this
study was to document nest success rates for these 3 ground-nesting grassland
birds in Florida dry prairie.
Native dry prairie is characterized as flat, unforested, fire-dependent grasslands
with scattered shrubs (Abrahamson and Hartnett 1990). Dominant
graminoids include Aristida beyrichiana Trin. & Rupr. (wiregrass), Ctenium
aromaticum (Walt.) Wood (toothache grass), Andropogon spp. (bluestem), and
Rhynchospora spp. (beak rush). Dominant shrubs include Serenoa repens
(Bartr.) Small (saw palmetto), Quercus minima (Sarg.) Small (dwarf oak),
Lyonia lucida (Lam.) K. Koch (fetterbush), and Ilex glabra (L.) Gray
(gallberry). Dominant forbs include Polygala spp. (bachelor’s button), Xyris
spp. (yellow-eyed grass), Eriocaulon decangulare L. (hat pin), Rhexia spp.
(meadow beauty), and a variety of milkweeds, orchids, and asters.
We used 3 dry prairie sites for this study: Delta Trail/OQ Range at Avon
Park Air Force Range in Highlands and Polk counties, (27°37'N, 81°19'W),
Three Lakes Wildlife Management Area in Osceola County
(27°47'N, 81°06'W), and Kissimmee Prairie Preserve State Park (27°34'N,
81°02'W) in Okeechobee County, hereafter referred to as Delta/OQ, Three
Lakes, and Kissimmee Preserve. The 3 study sites are less than 30 km from
each other. To simulate the natural fire frequency, all thee sites conduct
prescribed fires on a 2–3 year rotation throughout the prairie, resulting in a
mosaic of burn classes available in each year. Delta/OQ, owned by the
Department of Defense, is approximately 700 ha in size, and conducted only
winter prescribed fires during this study. Three Lakes, owned by the Florida
Fish and Wildlife Conservation Commission, contains approximately 4000 ha
of dry prairie, and was burned with summer and winter prescribed fires.
Kissimmee Preserve, owned by the Florida Park Service, is approximately
19,000 ha, and was burned with summer and winter fires.
We located nests opportunistically from March through August, during the
1997 and 1998 breeding seasons, as we conducted spot-mapping surveys and
banding exercises for Florida Grasshopper Sparrows and Bachman’s Sparrows
(Perkins and Vickery 2001, 2005; Perkins et al. 2003). We usually found
nests after flushing birds from the nest or after watching them land with food.
After we located nests, we marked them with flags 3 m from the nest and
checked nests every 2 to 3 days. We classified a nest as successful if at least 1
nestling left the nest. We also calculated mean clutch size for each species.
2007 D.W. Perkins and P.D. Vickery 285
We used Mayfield’s (1961, 1975) method to determine nest survival
rates and Johnson’s (1979) method to calculate standard errors and 95%
confidence intervals around nest success rates. A nest was considered a
failure only if the whole brood was lost. Losses were assumed to have
occurred at the midpoint between 2 visits. We considered nests successful if
young were expected to leave the nest between nest visits, and on the later
visit, we found the nest empty with no evidence of disturbance. In order to
increase sample size, we combined both sites and years to give an estimate
of nest success for each species for Florida dry prairie. This method assumes
that rate of nest loss is constant.
Eastern Meadowlark incubation lasts 13–14 days (Saunders 1932), with
nestlings remaining in the nest for an additional 10–12 days before fledging
(Lanyon 1995). Common Ground-doves have a 12–14 day incubation period
(Nicholson 1937) and 12-day nestling period (Bowman and Woolfenden
1997). Common Nighthawks have a 16–20 day incubation period (Campbell
et al. 1990, Dexter 1977), with hatchlings being semi-precocial and capable
of self-locomotion in response to female’s call during the first day after
hatching (Weller 1958). They generally move an average of 0.5 m per day
just 2 days after hatching (Fowle 1946). We used nesting periods of 24, 26,
and 18 days for Eastern Meadowlarks, Common Ground-doves, and Common
We determined annual productivity rates and estimated the survival rates
necessary in order for these populations to be sustainable. We fully acknowledge
that these estimates have wide confidence intervals due to the small
number of nests. However, we still think this provides additional useful
information beyond reporting only nest success rates. Juvenile survival rates
for birds are one of the hardest parameters to estimate. Based on demographic
data and extrapolation, Perkins and Vickery (2001) estimated
juvenile Florida Grasshopper Sparrow survival to be 35% (66–73% of adult
survival) at the same study sites as this study. Kershner et al. (2004a)
estimated juvenile Eastern Meadowlark survival at 56–69% during the first
90 days after fledging. Others have estimated juvenile survival to be between
25% and 50% of adult survival (Donovan et al. 1995, McCoy et al.
1999, Ricklefs 1973). We assumed that juvenile survival would be 50% of
adult survival and then estimated the adult survival rates that would be
needed to maintain stable populations assuming both 2 and 3 nest attempts.
Double broods (17%, 4 of 23 nests) have been reported for Eastern Meadowlarks
in Wisconsin (Lanyon 1957), and they can have as many as 4 nest
attempts (Lanyon 1995). Kershner et al. (2004b) followed radio-marked
Eastern Meadowlarks in Illinois and found only 9% (3 of 34) to have double
broods, and only 6% (2) attempted 3 nests. Common Ground-doves are
thought to breed year round in Florida (Baynard 1909 in Howell 1932,
Johnston 1962, Landers and Buckner 1979), with Bailey (1925) and Bowman
and Woolfenden (1997) reporting that 3–4 successful broods are
possible. Pairs of Ground-doves have been observed re-nesting in the same
286 Southeastern Naturalist Vol. 6, No. 2
nest within 20 days of successfully fledging young (Nicholson 1937). There
is little information on Common Nighthawks, but Weller (1958) and
Walbeck (1989) noted that 2 broods are possible.
We found 35 Eastern Meadowlark, 13 Common Ground-dove, and 14
Common Nighthawk nests. We estimated daily survival rate to be 0.927
(SE = 0.013), 0.944 (SE = 0.020), and 0.932 (SE = 0.023), for Eastern
Meadowlarks, Common Ground-doves, and Common Nighthawks, respectively.
With a nesting cycle length of 24, 26, and 18 days for these 3
species, we calculated nest success rates over the entire nesting cycle to be
0.16 (95% confidence interval = 0.08–0.32), 0.22 (0.07–0.68), and 0.28
(0.11–0.68) for Eastern Meadowlarks, Common Ground-doves, and Common
Nighthawks, respectively (Table 1). Nest predation was the major
cause of nest failure (40 of 42 failed nests, 95%); one Eastern Meadowlark
nest was abandoned and 1 Common Ground-dove nest was flooded. We
observed no Molothrus ater Boddaert (Brown-Headed Cowbird) or
Molothrus bonariensis Smelin (Shiny Cowbird) parasitism, which was
consistent with our findings for Florida Grasshopper and Bachman’s Sparrows
at the same study sites during the same time period (Perkins et al.
1998, 2003). Mean clutch size was 3.65 ± 0.73 ( ± SD, n = 20), 2 ±
0 (n = 6), and 1.93 ± 0.28 (n = 14) for Eastern Meadowlarks, Common
Ground-doves, and Common Nighthawks, respectively.
If each species had 2 nesting attempts 1.14 (95% Confidence interval
0.57–2.28), 0.81 (0.26–2.44), and 0.85 (0.34–2.03) young would be produced
Table 1. Eastern Meadowlark, Common Ground-dove, and Common Nighthawk nesting estimations
derived from nests at Avon Park Air Force Range, Kissimmee Prairie Preserve State
Park, and Three Lakes Wildlife Management Area during the 1997 and 1998 breeding seasons.
Eastern Common Common
Estimation Meadowlark Ground-dove Nighthawk
Nests 35 13 14
Nest failures 27 7 8
Nest exposure days 371.5 125.5 118
Daily nest survival (SE)A 0.927 (0.013) 0.944 (0.020) 0.932 (0.023)
Nest successB 0.16 (0.08–0.32) 0.22 (0.07–0.68) 0.28 (0.11–0.68)
Fledglings/successful nest (SE) 3.5 (0.19) 1.8 (0.17) 1.5 (0.22)
Productivity/nest attemptC 0.57 (0.28–1.14) 0.40 (0.13–1.22) 0.42 (0.17–1.01)
Productivity—2 nest attemptsC 1.14 (0.57–2.28) 0.81 (0.26 –2.44) 0.85 (0.34–2.03)
Productivity—3 nest attemptsC 1.72 (0.85–3.41) 1.21 (0.38–3.66) 1.27 (0.51–3.04)
Adult survival—2 nest attemptsC 0.64 (0.47–0.78) 0.71 (0.45–0.89) 0.70 (0.50–0.86)
Adult survival—3 nest attemptsC 0.54 (0.37–0.70) 0.62 (0.35–0.84) 0.61 (0.40–0.80)
ACalculated following Johnson (1979).
BNumbers in parentheses are 95% confidence intervals calculated based on the daily nest
survival rate ± one standard error.
CNumbers in parentheses are 95% confidence intervals calculated from the lower and upper
confidence limits on nesting success.
2007 D.W. Perkins and P.D. Vickery 287
for Eastern Meadowlarks, Common Ground-doves, and Common Nighthawks,
respectively. In order to be sustainable at 2 nest attempts, adult
survival would need to be 64% (47–78%), 71% (45–89%), and 70% (50–86%)
for Meadowlarks, Ground-doves, and Nighthawks, respectively (Table 1). For
3 nesting attempts, 1.72 (0.85–3.41), 1.21 (0.38–3.66), and 1.27 (0.51–3.04)
young would be produced for Eastern Meadowlarks, Common Ground-doves,
and Common Nighthawks, respectively. In order to be sustainable at 3 nest
attempts, adult survival would need to be 54% (37–70%), 62% (35–84%), and
61% (40–80%) for the Meadowlarks, Ground-doves, and Nighthawks, respectively
Our estimate of nest success for Eastern Meadowlarks (16.4%) was
generally lower than estimates for this species in other parts of the country,
but was similar to those found for Florida Grasshopper Sparrows (10–33%)
and Bachman’s Sparrows (7–38%) from the same study sites (Perkins et al.
1998, 2003). For Eastern Meadowlarks, Lanyon (1957) reported 33% in
Wisconsin and Kershner et al. (2004b) reported 46% in Illinois, while
Knapton (1988) reported 30% for monogamous females and 52% for polygynous
females in Ontario. Granfors et al. (1996) reported Eastern
Meadowlark nest success rates for 1990 and 1991 of 20 and 25%, respectively,
in Kansas rangelands, and 17 and 10%, respectively, in Kansas
Conservation Reserve program fields. McCoy et al. (1999) reported Eastern
Meadowlark nest success rates of 25–42% in Missouri CRP fields. Two
studies found low Meadowlark nest success rates similar to our study;
however, they both attributed the low numbers to anthropogenic reasons.
Roseberry and Klimstra (1970) attributed low nest success of 18% in Illinois
to high predator densities, and Kershner and Bollinger (1996) suggested that
low nest success rates (14%) at Illinois airports were due to mowing.
Our nest success estimate for Common Ground-doves (22.5%) was also
lower than has generally been reported. Mitchell et al. (1996) reported nest
success rates of 25% and 38% for Common Ground-doves nesting in trees in
2 citrus groves along the Florida coast. Bowman (2002) reported nest success
varying from 25 to 48% in the Florida Keys. Passmore (1984) reported
42% nest success in Texas. One published estimate was lower than ours; in
dry coastal Puerto Rican forests, only 17% of nests fledged at least 1 young
We did not find any published estimates to compare with our Common
Nighthawk nest rate of 28.3%. Poulin et al. (1996) reports that no information is
known about annual reproductive success or annual survival for this species,
and suggest that studying reproductive effort is a priority for future research.
Our nest success rates may be low due to the small and fragmented
nature of the study sites. Florida dry prairie once occupied 5000 km2 in
central Florida (Orzell and Bridges 2006), but 81% of the dry prairie has
been converted to other land uses such as citrus plantations and improved
288 Southeastern Naturalist Vol. 6, No. 2
pastures seeded with exotic grasses (Shriver and Vickery 1999). At 700
and 4000 ha, Delta/OQ and Three Lakes are relatively small compared to
the historic extent of dry prairie. Perkins et al. (2003) reported higher nestsuccess
rates for ground-nesting Florida Grasshopper Sparrows in core
areas (> 400 m from edge) than in edge areas in the same dry prairies as
this study. Herkert et al. (2003) studied 39 prairie fragments (24 to
> 40,000 ha) in Illinois, Kansas, Missouri, North Dakota, and Oklahoma
and reported that Eastern Meadowlarks, as well as Grasshopper Sparrows,
Ammodramus henslowii Audubon (Henslow’s Sparrows), and Spiza
americana Smelin (Dickcissels), all experienced increasing nest predation
as the size of the prairie fragment declined.
Predation is considered to be the most common cause of nest failure for
many songbirds (Martin 1992). Predation was the major cause of nest failure
in common Ground-doves in Florida citrus groves (Mitchell et al. 1996),
Eastern Meadowlarks in Illinois (Kershner et al. 2004b), and Florida Grasshopper
Sparrows and Bachman’s Sparrows on Florida dry prairie (Perkins et
al. 2003). There are at least 12 bird species, 10 mammalian species, and 12
snake species present in the dry prairie of Florida that could be predators of
ground nests. We only observed 1 nest that failed due to flooding, but this
could be an additional major, but infrequent, reason for nest failure in
Florida dry prairie (Perkins and Vickery 2005, Pranty 2000).
We do not think that rates of nest predation were artificially inflated by
our activities at the nests because our study sites had extensive human
activity throughout the breeding season, therefore scent trails would be of
little use to mammalian predators. Nest flags would not likely have provided
clues to visual predators such as Corvus brachyrhynchos Brehm (American
Crows) because the same flags were used to mark 50-m grids across almost
the entire study area.
Our Eastern Meadowlark clutch sizes (3.65) were lower than those reported
in other areas of the country. Clutch sizes for Eastern Meadowlarks
have been reported at 4.8 in Wisconsin (Lanyon 1957), 4.2 in Illinois
(Roseberrry and Klimstra 1970), and 5.2 in Kansas (Johnston 1964). Common
Ground-doves (Bowman 2002) and Common Nighthawks (Dexter
1952, 1956, 1961; Fowle 1946; Howell 1959; Kantrud and Higgins 1992)
almost always lay 2 eggs. All 6 Common Ground-dove nests had two eggs.
We found 13 Common Nighthawk nests with 2 eggs and 1 nest with 1 egg. It
is possible that depredation had occurred at this nest prior to our detection.
Assuming that fledgling survival to the first breeding season is 50% of
adult survivorship, Eastern Meadowlarks, Common Ground-doves, and
Common Nighthawks would need adult survival rates of 0.64, 0.71, and 0.70
with an average of 2 attempts, and 0.54, 0.62, and 0.61 with an average of 3
nest attempts, respectively, in order to be sustainable in dry prairie. Kershner
et al. (2004b) similarly estimated that Eastern Meadowlark annual survival
would need to be between 59–61% for a stable population in Illinois. Little is
known about the annual survival rates for Eastern Meadowlarks (Lanyon
2007 D.W. Perkins and P.D. Vickery 289
1995), Common Ground-doves (Bowman 2002),and Common Nighthawks
(Poulin et al. 1996). Perkins and Vickery (2001) reported annual survival
rates for Florida Grasshopper Sparrows as 0.48 and 0.53 at Delta/OQ and
Three Lake's respecively. While an average of 2 or 3 nest attempts per
breeding season is high, it may be possible in Florida dry prairie due to the
long breeding season. We acknowledge that these numbers reflect assumptions
of juvenile survival and uncertainties in the nest success data due to
limited sample sizes. However, these types of data are rare, and we present
them in hope that it will encourage other researchers to build on our data.
The authors thank the Department of Defense and the Environmental Flight at
Avon Park Air Force Range for financial and logistical support. The Massachusetts
Cooperative Fish and Wildlife Research Unit assisted in administrative and logistical
support. We greatly appreciate permission for access and logistic support at Three
Lakes Wildlife Management Area from the Florida Fish and Wildlife Conservation
Commission. C. Griffin, J. Pederson, and M. Delany were especially helpful. B.
Pranty, C. Collins, M. Bakermans, A. Vitz, M. Scheuerell, D. Updegrove, N. Hamel,
D. Perkins, C. O’Brien, K. McKay, and T. Dean all aided in field work. This
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