Status and Reproductive Outcome of the Breeding Population of Striated Caracaras (Phalcoboenus australis) at Franklin Bay, Staten Island, Argentina

Abstract

The Striated Caracara (Phalcoboenus australis) is a poorly known raptor associated with seabird colonies, restricted to the islands of southern South America, and with an estimated extant population of <2,500 mature individuals. We evaluated the number of breeding pairs, described the characteristics and spatial pattern of nest sites, and estimated breeding output of a population of Striated Caracaras in Franklin Bay, Staten Island, Argentina. We found one of the lower breeding density values reported for this species, although this population is associated with one of the biggest colonies of Rockhopper Penguins. The main material used for the construction of Striated Caracaras' nests was tussac grass, though only half of nests were placed in grassland. The spatial pattern for nest sites corresponds with global and local clustering. All successful Striated Caracaras' nests were at least 250 m from the nearest neighboring nest, were generally closer to the colony of Rockhopper Penguins than failed nests, and had more Rockhopper patches around them. Breeding success was 0.73 successful nests/active nests, productivity was 1.27 ± 1.01 young/active nest, and brood size was 1.75 ± 0.71 young/successful nest. The presence of invasive wild goats and red deer is proposed as a factor that could be restricting nest site availability in the study area.

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Status and Reproductive Outcome of the Breeding Population
of Striated Caracaras (Phalcoboenus australis) at Franklin Bay,
Staten Island, Argentina
Author(s): Ulises Balza, Nicolás A. Lois, and Andrea Raya Rey
Source: The Wilson Journal of Ornithology, 129(4):890-898.
Published By: The Wilson Ornithological Society
https://doi.org/10.1676/16-189.1
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The Wilson Journal of Ornithology 129(4):890–898, 2017
Status and Reproductive Outcome of the Breeding Population of Striated Caracaras
(Phalcoboenus australis) at Franklin Bay, Staten Island, Argentina
Ulises Balza,
1,4
Nicola´s A. Lois,
1,2
and Andrea Raya Rey
3
ABSTRACT.—The Striated Caracara (Phalcoboenus
australis) is a poorly known raptor associated with seabird
colonies, restricted to the islands of southern South America,
and with an estimated extant population of ,2,500 mature
individuals. We evaluated the number of breeding pairs,
described the characteristics and spatial pattern of nest sites,
and estimated breeding output of a population of Striated
Caracaras in Franklin Bay, Staten Island, Argentina. We
found one of the lower breeding density values reported for
this species, although this population is associated with one
of the biggest colonies of Rockhopper Penguins. The main
material used for the construction of Striated Caracaras’
nests was tussac grass, though only half of nests were placed
in grassland. The spatial pattern for nest sites corresponds
with global and local clustering. All successful Striated
Caracaras’ nests were at least 250 m from the nearest
neighboring nest, were generally closer to the colony of
Rockhopper Penguins than failed nests, and had more
Rockhopper patches around them. Breeding success was
0.73 successful nests/active nests, productivity was 1.27 6
1.01 young/active nest, and brood size was 1.75 60.71
young/successful nest. The presence of invasive wild goats
and red deer is proposed as a factor that could be restricting
1
Laboratorio de Ecolog´
ıa y Conservaci ´
on la Vida
Silvestre. Centro Austral de Investigaciones Cient´
ıficas
(CADIC-CONICET). Bernardo Houssay 200, Ushuaia,
Argentina.
2
Laboratorio de Ecolog´
ıa y Comportamiento Animal.
Instituto de Ecolog´
ıa, Gen ´
etica y Evoluci´
on (IEGEBA-
CONICET). Facultad de Ciencias Exactas y Naturales,
Universidad de Buenos Aires. Intendente G¨
uiraldes 2160,
Buenos Aires, Argentina.
3
Instituto de Ciencias Polares, Ambiente y Recursos
Naturales (ICPA), Universidad Nacional de Tierra del
Fuego (UNTdF). Leandro N. Alem 1036. Ushuaia,
Argentina.
4
Corresponding author; e-mail:
ulisesbalza@cadic-conicet.gob.ar
890 THE WILSON JOURNAL OF ORNITHOLOGY Vol. 129, No. 4, December 2017

nest site availability in the study area. Received 24 October
2016. Accepted 29 March 2017.
Key words: breeding output, invasive species, nest site
availability, spatial analysis.
The Striated Caracara (Phalcoboenus australis)
is a large subantarctic bird of prey restricted to
isolated shores and islands south of latitude 508S
in South America (Ferguson-Lees and Christie
2001). The species is classified as Near Threatened
(IUCN 2015) with an extant population of ,2,500
mature individuals. They reside mostly on Staten
Island (Isla de los Estados) in the Fuegian region
and in the Malvinas/Falkland Islands (BirdLife
International 2013), with fragmented breeding
records in southern Chile (Mar´
ın et al. 2006).
The species is largely associated with seabird
breeding colonies (i.e., Rockhopper Penguins
[Eudyptes chrysocome], Black-browed Albatross
[Thalassarche melanophris]). Its breeding season
extends from October to February during which it
lays 1–4 eggs in a single clutch. It primarily nests
on the ground in tussac grasslands of Poa
flabellata ,10 m from seabird nests and feeds its
offspring on eggs, chicks, and adults from the
nearby colony (Strange 1996, Catry et al. 2008,
Liljesthr¨om et al. 2008).
Striated Caracaras on the Malvinas/Falklands
declined as a consequence of direct persecution
during the 1900s, and their numbers have not
recovered despite the legal protection being
formalized in 1999 (Woods 2007). In this
archipelago, breeding territories have been record-
ed as close as possible to the seabird colony, and at
the same time as far as possible from the nearest
conspecific territory, following an apparently
regular pattern (Strange 1996). The published
mean distance between breeding pairs during the
breeding season ranges 47–700 m, depending on
the island studied (Strange 1996, Catry et al.
2008).
It is unclear whether the population on Staten
Island is threatened. Unlike populations on the
Malvinas/Falkland Islands, persecution by humans
has not been recorded on Staten Island, which has
long been a Provincial Reserve, and has recently
been declared as a National Reserve. However,
two factors could potentially affect the viability of
the population of Striated Caracaras on Staten
Island in the long term: a) the negative population
trend of their apparent main food resource during
the breeding season, the globally threatened
Rockhopper Penguins (IUCN 2015), which has
been declining in numbers in the area for the last
decades at a yearly rate of 2%(Raya Rey et al.
2014); and b) the probable, but unknown, impact
on the tussac grassland produced by two exotic
herbivores, the wild goat (Capra hircus) and the
red deer (Cervus elaphus), introduced in 1856 and
1973, respectively (Valenzuela et al. 2014).
In this context, the main objective of this work
is to increase our knowledge on the current status
and breeding biology of the Striated Caracaras’
population in Franklin Bay, Staten Island, the only
breeding population known to coexist with large
exotic herbivores, as far as we know. In this study
we, a) estimate the number of breeding pairs and
describe the characteristics of accessible nests; b)
analyze the spatial pattern of nests and productiv-
ity; c) estimate the breeding success, productivity,
and brood size and; d) evaluate the variation of
breeding success in relation to the presence of
nearest neighbors, and the distribution and density
of the main seabird reproductive patches.
METHODS
We carried out this study between 14 November
and 19 December 2014, which corresponds with
late incubation period and early chick rearing of
Striated Caracaras (Strange 1996).
Study Site.—Our study took place in Franklin
Bay, Staten Island, Argentina (Fig. 1; 548530S,
648390W), where the mean annual temperature
varies from 2.7–9.0 8C, and the mean annual
precipitation is 1,500 mm. Strong winds, mainly
from the SW are predominant throughout the year
(Dudley and Crow 1983). The island is character-
ized by tussac grassland on the shores and
subantarctic forests inland, dominated by ever-
green beech (Nothofagus betuloides) and winter’s
bark (Drimys winteri). The understory is rich in
shrubs, ferns, lichens, and mosses, including
diddle dee (Empetrum rubrum) which is one of
the most abundant shrubs in the area (Niekisch and
Schiavini 1998). Several species of seabirds and
marine mammals breed on the shores of Staten
Island (Chebez and Bertonatti 1994, Schiavini
2000, Schiavini and Raya Rey 2001, Schiavini et
al. 2004). Franklin Bay holds one of the biggest
891SHORT COMMUNICATIONS

colonies of Rockhopper Penguins (127,000 breed-
ing pairs; Eudyptes chrysocome), a colony of
Imperial Shags (4,600 breeding pairs; Leucocarbo
atriceps), and a colony of Magellanic Penguins
(1,600 breeding pairs; Spheniscus magellanicus;
Raya Rey et al. 2014). Nests of Rockhopper
Penguins are arranged in 133 discrete patches
around the bay which vary largely in area (range
30–5,848 m
2
) and density of breeding pairs within
each patch (range 0.22–1.74 nest/m
2
), while
Magellanic Penguins’ nests are scattered in a
colony in the southern area of Franklin Bay.
Imperial Shags breed in 24 discrete patches which
overlap with the colony of Rockhopper Penguins,
ranging 11–1,442 breeding pairs per patch (ARR,
unpubl. data) (see Fig 1).
Breeding Population Survey.—To obtain a
census of breeding pairs of Striated Caracaras,
we surveyed 24 transects in a SW-NE direction,
perpendicular to the coastline, with variable
lengths (range: 250–1,250 m), and a 200-m
separation between each one (Fig 1). The study
area (i.e., the area bounded by the band transects)
included all the main seabird colonies on the site
and its total area was 3.69 km
2
. We followed
Woods (2007) to identify active territories, and
when territorial behavior was observed, we
searched exhaustively for the active nest and
aged the members of the breeding pairs following
Strange (1996). Nests were georeferenced, and
when accessible, standard variables were mea-
sured to record basic characteristics of emplace-
ment, dimensions, and construction materials
following Tapia et al. (2007). In some cases,
nests were inaccessible because of the steepness
of cliffs where they were placed. We visited each
accessible nest every 7–10 days to record its
breeding output, and depending on the time of
encounter, each nest was visited 2–4 times during
the study.
Spatial Analysis of Nest Sites.—We analyzed
the spatial pattern of nest sites in the study area
using three Ripley’s functions, which have the
advantage that all point-to-point distances are
FIG. 1. Study site showing the location of Striated Caracaras’ nests sites, the main seabird colonies, and the transects
surveyed during the breeding season of 2014–2015.
892 THE WILSON JOURNAL OF ORNITHOLOGY Vol. 129, No. 4, December 2017

evaluated to show spatial clustering at different
scales (Ripley 1977, Getis and Ord 1992, de la
Cruz Rot 2008). We used Global-K and Local-K
functions to identify global or local clustering of
events (i.e., nest sites). By contrast, Weighted-K
function measures the degree of clustering of
points by some mark, in this case by productivity
(i.e., number of young produced/nest) at the time
of the last visit of each nest. For both global
analyses, we used radial distances ranging between
50–800 m with distance categories every 50 m.
For the local analysis, we used radial distances
ranging between 20–500 m with distance catego-
ries every 20 m. In each case, we ran 99 Monte
Carlo simulations to build a confidence envelope
over the null hypothesis of complete spatial
randomness of the points. In all cases, if the
statistical L(d) (for global and local analysis) and
L
w
(d) (for global analysis of weighted points) lie
within the confidence envelope at a certain
distance, then the points (or the weighted points)
are randomly distributed. If the statistic lies in the
upper confidence interval, the events tend to be
clustered and conversely, if it lies below the
confidence envelope, the events follow a regular
pattern. We compared the distance between nesting
sites and the closest breeding patches or areas of
colonies of Rockhopper Penguins, Magellanic
Penguins, and Imperial Shags in the study area
using paired t-tests and also measured the distance
between nest sites of Striated Caracaras.
Breeding Output.—Finally, we estimated the
apparent breeding success as the proportion of
breeding pairs that produced at least one young by
the end of the study. We also measured apparent
productivity as the mean number of young
produced per active nest and the apparent brood
size, as the mean number of young produced per
successful nest at the same time. Breeding success,
productivity, and brood size are interpreted as
apparent because we were not able to monitor each
nest until fledging, which is the traditional way to
estimate these parameters (Newton 1979, Steenhof
and Newton 2007). The values obtained are an
overestimation of the real values, because we
assume no more breeding attempts occurred
between the end of the study and the end of the
breeding season. We also analyzed breeding
success in relation to the distance to the nearest
conspecific neighbor and variables relative to the
distribution and breeding density of seabird
colonies.
RESULTS
We found 16 nests of Striated Caracaras in the
study area, yielding a density of 4.34 breeding
pairs/km
2
. Half of these nests were placed on cliffs
of variable height (mean 39.8 m, range 6–100 m)
at different heights from the cliff bases (mean 30.6
m, range 2–90 m). The other nests were built on
the ground, in most cases associated with one
individual tussac grass (Poa flabellata). All
breeding birds presented full-adult plumage (.5
years old, Strange 1996). In accessible nests (n¼
11), pairs were found either incubating eggs or
raising nestlings. All accessible nests were con-
structed mainly with tussac grass fibers. However,
two nests also contained branches of diddle dee
and one nest included anthropogenic elements
(pieces of rope and fishing lines). Nest area was
0.34 60.14 m
2
(mean, SD), ranging 0.20–0.69
m
2
. Nests were nearer patches of Rockhopper
Penguins (68 658 m, mean, SD) than patches of
Imperial Shags (376 6222 m, t¼5.74, P,
0.001) and the colony of Magellanic Penguins
(1,822 61,126 m, t¼6.18, P,0.001). Indeed,
three nests were placed at the edge or inside
Rockhopper Penguins’ breeding patches. The
distance between nest sites of Striated Caracaras
in this study was 298 6184 m (mean, SD).
The global spatial pattern analysis indicated
scale dependent clustering (Fig. 2A). However,
productivity did not correlate with spatial location,
as its pattern showed no difference with the
complete spatial randomness (Fig. 2B). In the
latter case, the statistical L
w
(d) takes non-null
values starting from a radial distance of 250 m.
Given that L
w
(d) ¼0 for null values of the mark
(i.e., a failed nest), all successful nests (i.e., 1
young produced) were separated at least 250 m
from the nearest neighbor. For the local spatial
pattern analysis, three nests located in the center of
the study area (see Fig. 1) presented significant
local clustering at radial distances of 140–240 m,
280 m, and 360–440 m respectively. All other
nests showed values of L(d) which corresponded
with randomly distributed events.
The apparent breeding success for the accessible
nests was 0.73 successful nest/active nest, apparent
893SHORT COMMUNICATIONS

productivity was 1.27 61.01 young/active nest,
and apparent brood size was 1.75 60.71 young/
successful nest (range: 1–3 young/successful nest).
When analyzing the distance to the nearest
neighbor, failed nests were consistently closer to
other nests of Striated Caracaras compared to
successful nests (Fig. 3A). Also, successful nests
were generally closer to reproductive patches of
Rockhopper Penguins and had more patches in a
200-m radius around the nest, with respect to the
failed ones (Fig. 3B, C). We did not carry out any
statistical inference in any of these data sets
because of small sample sizes.
DISCUSSION
Breeding density in birds of prey is mainly
limited by food and nest site availability (Newton
1979). In this case, food limitation seems not
likely, since this population of Striated Caracaras
is associated with one of the largest Rockhopper
Penguin colonies in the world. The breeding
density estimate in our study area is similar to that
reported by Catry et al. (2008) for a growing
population on New Island but lower than the
value reported for a stable population on Beau-
chene Island (Lewis Smith and Prince 1985,
Strange 1996), both in the Malvinas/Falkland
Islands (Table 1). The values found for apparent
breeding success, productivity, and brood size do
not seem particularly large, and we found the
lowest mean brood size reported until now for the
species.
Nest placement in this study differs from that
previously reported. In our case, half of the nests
were built in cliffs, in contrast with the report of
FIG. 2. Spatial patterns analysis using K-Ripley’s functions. A) Global pattern analysis using Global-K function for
events (presence of a Striated Caracaras’ nest) in the study area. B) Weighted-K function for a subsample of accessible
Striated Caracaras’ nests using productivity as a mark (see text).
894 THE WILSON JOURNAL OF ORNITHOLOGY Vol. 129, No. 4, December 2017

Strange (1996), which found 95%(n¼53) of the
nests on the ground. Simultaneously, even though
the nests of Striated Caracaras were relatively
close to patches of Rockhopper Penguins, separa-
tion between the two seems to be greater than the
distance previously described. Strange (1996)
reports that 74%of his surveyed nests (n¼19)
were 10 m or less from seabird colonies, while in
TABLE 1. Reproductive parameters of Striated Caracaras published in the literature and presented in this work.
Season Location
Density
(territorial
pairs/km
2
)
Surveyed
active nests
Breeding success
(successful nests/
total nests)
Productivity
(young/
active nests)
Brood size
(young/
successful nests) Source
1979/80 Beauchene Island
(Malvinas/Falkland Is.)
35.83
a
64 0.42
a
0.88 61.16
a
2.07 60.83
a
Lewis Smith and
Prince 1985
2005/06 New Island
(Malvinas/Falkland Is.)
- 18 - - 2.50 60.51 Catry et al. 2008
2006/07 New Island
(Malvinas/Falkland Is.)
4.31 36 - - 2.36 60.59 Catry et al. 2008
2014/15 Franklin Bay,
Staten Island (Argentina)
4.34 11 0.73 1.27 61.01 1.75 60.71 This study
a
Indicate values calculated from original data.
FIG. 3. Relationship between reproductive success and ecological variables in the study area for a subsample of
accessible nests.
895SHORT COMMUNICATIONS

our study only 19%(n¼16) were consistent with
that metric. It could be possible that the difference
in nest placement along with the low breeding
density found could be a response to the absence
of suitable nesting sites near penguin patches. The
spatial pattern analysis showed that nesting sites
tended to be clustered in our study area, while in
other sites the distribution of breeding pairs tended
to be regular (Strange 1996), which in birds of
prey is expected when availability of nesting sites
is not a limitation and the population has reached
carrying capacity (Newton 1979). Nests showing
local clustering could be influencing the global
pattern, as they represented 19%of the nest sites
we studied.
Populations that reach their carrying capacity
show non-breeding or non-territorial adults during
the breeding season, which reproduce only when a
given site becomes available as a consequence of
the death of one or both members of a breeding
couple (Newton 1979). Catry et al. (2008)
reported several flocks of non-breeding individu-
als (up to 65 birds) foraging near seabird colonies,
but none of these individuals presented full-adult
plumage. Moreover, the authors reported three
territorial pairs with at least one of the members of
the couple presenting immature plumage, which is
a typical sign of a growing population with
available breeding sites (Newton 1979). In our
case, not only did all breeding birds present full-
adult plumage, but we anecdotally recorded
several non-breeding adults foraging or eating
carrion in flocks with juveniles and immature
birds. Even though we could not estimate the
abundance of Striated Caracaras, it seems likely
that most adult birds in the study area are non-
reproductive, and thus we hypothesize this
population has reached its carrying capacity
limited by nest site availability.
We propose that population dynamics of the
Striated Caracaras in Franklin Bay could be
complex and include the effect of the presence of
introduced herbivore species known for their
detrimental effect on native flora communities,
the negative effect over the reproductive outcome
of seabirds and other vertebrates in oceanic
islands, and the promotion of local extinctions in
areas where similar species are absent (Cronk
1989, Veblen et al. 1989, Takatsuki 2009, Pafilis et
al. 2013). In our case, two main effects of the
exotic goats and deer are proposed.
On the one hand, these herbivores could be
affecting the abundance and/or cover of tussac
grassland. In ungrazed islands of the Malvinas/
Falkland Islands, this grass covers .60%of the
land, forming a monospecific community with
some individuals 3.5 m high and .300 years old
which provide favorable conditions for nest
placement (e.g., temperature on the ground
surrounded by tussac is between 4–20 8C higher
than in uncovered soil; Lewis Smith and Prince
1985). The reduction or the lack of this resource
could be affecting the caracaras in three ways.
First, direct reduction of potential nesting sites
could be limiting the maximum number of
breeding pairs in the study area. Second, the
relative distance to their main food resource, the
nearest reproductive patch of Rockhopper Pen-
guins, may be increased thus augmenting their
foraging energy expenditure. Third, nesting in
other emplacements, like cliffs, could be in this
context, suboptimal for egg incubation, chick
development, or both.
On the other hand, during the winter, while
seabirds are not present, both goats and deer could
potentially be an important component of the
carrion portion of Striated Caracaras’ diet, a well-
documented situation for raptors in other invaded
scenarios (Speziale and Lambertucci 2013). Stri-
ated Caracaras in the Malvinas/Falklands Islands
apparently move in winter to farmland settlements,
where they feed partly on livestock as carrion
(Rexer-Huber and Bildstein 2013). If something
similar happened on Staten Island, it would be
expected that winter survival will be higher than in
a non-invaded system. These two situations would
be resulting in a lower population recruitment rate
because of fewer breeding pairs and more
intraspecific competition.
The total eradication of the invasive species is
proposed for the restoration of island environ-
ments (Glen et al. 2013), but in this case it seems
important to have more information to assess the
consequences of this intervention, as Striated
Caracaras’ population could be suddenly losing
an important winter-time subsidy, thus abruptly
changing its dynamics with unknown conse-
quences. It would be important to know the
current status of other populations of Striated
Caracaras in other potential breeding locations of
Staten Island, A ˜
no Nuevo Island, and in Tierra del
Fuego main Island, including the seasonal
896 THE WILSON JOURNAL OF ORNITHOLOGY Vol. 129, No. 4, December 2017

movements of individuals, where seabird colonies
are not so important in numbers, but goats and
deers are absent (Schiavini 2000, Schiavini et al.
2005).
ACKNOWLEDGMENTS
This study was funded through a research project of the
Agencia Nacional de Promoci´
on Cient´
ıfica y Tecnol´
ogica
(PICT 2012 n81832 – ANPCyT), and with funding provided
by the Inter-American Development Bank (IDB) and
Wildlife Conservation Society. This work would have not
been possible without the field assistance provided by R.
Sa´ enz Samaniego and S. Harris, the background information
on seabird populations on Staten Island collected in previous
campaigns by V. Bruno, N. Rosciano, and P. Petracci, and
the technical support of A. E. Capdevielle and G. M.
Wiemeyer from Cabure-´
ı Foundation. We are also very
grateful to L. I. Rodr´
ıguez Planes, who provided us valuable
help in the design of the analysis and drafting of this
manuscript, and to M. G. Pizzarello, M. Dom´
ınguez, and S.
Dodino, who provided important comments improving our
study. And last but not least, we thank J. C. Reboreda, R. F.
B´
o, and J. H. Sarasola, members of the panel in the graduate
thesis presentation that led to this work, for their feedback
and precious advice provided.
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The Wilson Journal of Ornithology 129(4):898–902, 2017
An Indigo Bunting (Passerina cyanea) Transporting Snails During Spring
Migration
Theodore J. Zenzal Jr.,
1,2,3,5
Emily J. Lain,
1,4
and J. Michael Sellers Jr.
1
ABSTRACT.—Organisms with limited motility may use
animal transport as a mechanism for dispersal. Migratory
birds can provide a vehicle to move small organisms great
distances, which may allow them access to areas that are
otherwise inaccessible. During normal mist netting
operations at a spring migration banding station along the
northern Gulf of Mexico coast in Louisiana, USA, we
encountered an Indigo Bunting (Passerina cyanea) with
numerous snails, possibly Galba cubensis, underneath its
breast feathers. While encounters of songbirds carrying
snails appear rare, long-distance migrating songbirds
represent a possible mechanism to transport small snails
great distances to expand ranges, colonize new areas, or
maintain genetic continuity. Received 12 October 2016.
Accepted 28 February 2017.
Key words: animal transport, chenier, Gulf of Mexico,
Louisiana, migration, snails, stopover.
Migratory animals are capable of traveling
considerable distances, often traversing diverse
habitats along their journey (Rappole 1995, Dingle
1996, Newton 2008). Through these large-scale
movements, migratory animals may act as dis-
persal agents for both plants and animals (e.g.,
Raven 1963, Rees 1965, Mukherjee et al. 2014).
Migratory birds can provide a mechanism for
dispersal over great distances to habitats that some
organisms may not otherwise reach, such as
habitats located on isolated mountain summits,
islands, and across ecological barriers (e.g.,
Cockerell 1921, Rees 1965, Mukherjee et al.
2014). Additionally, many migratory birds stop to
rest and refuel en route (Rappole 1995, Newton
2008) as they encounter a number of habitats on a
single journey.
Animals like snails might seem limited in their
dispersal abilities, but birds have been shown as a
mechanism to transport them across great distanc-
es (Rees 1965, Dundee et al. 1967, Vagvolgyi
1975, van Leeuwen et al. 2012). Migratory
landbirds, shorebirds, and waterfowl have been
found to transport snails either internally or
externally (e.g., Rees 1965, Dundee et al. 1967,
Wesselingh et al. 1999, van Leeuwen et al. 2012,
1
Department of Biological Sciences, University of
Southern Mississippi, Hattiesburg, MS 39406, USA.
2
Department of Entomology and Wildlife Ecology,
University of Delaware, Newark, DE 19716, USA.
3
Current address: Department of Natural Resources
and Environmental Sciences, University of Illinois,
Urbana, IL 61801.
4
Current address: Illinois Natural History Survey,
University of Illinois, Champaign, IL 61820.
5
Corresponding author; e-mail: tjzenzal@gmail.com
898 THE WILSON JOURNAL OF ORNITHOLOGY Vol. 129, No. 4, December 2017