EV-1
Occupancy of yellow-billed and Pacific loons: evidence for
interspecific competition and habitat mediated co-occurrence
Trevor B. Haynes, Joel A. Schmutz, Mark S. Lindberg, Kenneth G. Wright, Brian D. Uher-Koch
and Amanda E. Rosenberger
T. B. Haynes (t.haynes@alaska.edu), School of Fisheries and Ocean Sciences, Univ. of Alaska Fairbanks, Fairbanks, AK 99775, USA. – J. A.
Schmutz, K. G. Wright and B. D. Uher-Koch, US Geological Survey, Alaska Science Center, Anchorage, AK 99508, USA. – M. S. Lindberg,
Inst. of Arctic Biology, Univ. of Alaska Fairbanks, Fairbanks, AK 99775, USA. – A. E. Rosenberger, US Geological Survey, Missouri Cooperative
Fish and Wildlife Research Unit, Univ. of Missouri, Columbia, MO 65211, USA.
Interspeciic competition is an important process structuring ecological communities, however, it is diicult to observe
in nature. We used an occupancy modelling approach to evaluate evidence of competition between yellow-billed (Gavia
adamsii) and Paciic (G. pacifica) loons for nesting lakes on the Arctic Coastal Plain of Alaska. With multiple years
of data and survey platforms, we estimated dynamic occupancy states (e.g. rates of colonization or extinction from
individual lakes) and controlled for detection diferences among aircraft platforms and ground survey crews. Results
indicated that yellow-billed loons were strong competitors and negatively inluenced the occupancy of Paciic loons
by excluding them from potential breeding lakes. Paciic loon occupancy was conditional on the presence of yellow-
billed loons, with Paciic loons having almost a tenfold decrease in occupancy probability when yellow-billed loons
were present and a threefold decrease in colonization probability when yellow-billed loons were present in the current
or previous year. Yellow-billed and Paciic loons co-occurred less than expected by chance except on very large lakes or
lakes with convoluted shorelines; variables which may decrease the cost of maintaining a territory in the presence of
the other species. hese results imply the existence of interspeciic competition between yellow-billed and Paciic loons
for nesting lakes; however, habitat characteristics which facilitate visual and spatial separation of territories can reduce
competitive interactions and promote species co-occurrence.
A central theme in community ecology is that competition,
particularly among sympatric but taxonomically similar
species, plays a key role in species’ ecology (Korpimäki
1987, Samraoui et al. 2012) and distributional patterns
(Hutchinson 1957, Hardin 1960). Competition is, in fact,
considered not only a primary driver of species’ ecology,
but also an evolutionary force, leading sympatric species to
use new resources and promoting phenotypic divergence
and less competitive sympatry (Schluter 2000). For inter-
speciic competition to occur, one species must limit the
resource use of another. Investigating competition, there-
fore, often takes the form of niche overlap studies (e.g.
diet; Korpimäki 1987). However, under many conditions,
even with similar resource preferences, competing species
can co-exist with minimal in situ overlap – largely due to
the potential for species to shift spatial and/or dietary
preferences in the presence of competitors (Pontin 1982,
Baxter et al. 2004).
Interspeciic competition can be reduced by factors
such as temporal and spatial separation among species,
habitat heterogeneity (allowing more potential for niche-
shifts), or when the shared resource is not limited due to
other limiting factors for the species (Ritchie 2002).
Further, competition may afect distributions at diferent
scales with the stronger competitor excluding the other at
the patch scale but whose distributions overlap at broader
spatial scales (Yackulic et al. in press). Manifestations of
competition, therefore, can be subtle and more inter-
mittent than other ecological interactions (Wiens 1977),
and thus competition has been diicult to demonstrate
outside of experimental settings (Connor and Simberlof
1979, Connell 1980, 1983). When experiments are not
possible or practical, as is the case with many taxa, data on
species distributions can be assessed for evidence of
competition by examining whether species co-occur less
than would be expected by chance. However, interpreta-
tion of the results of this approach can be ambiguous
because distributional patterns can be inluenced not
only by competitive interactions, but also by difering
ecological requirements among species or imperfect detec-
tion during sampling (i.e. false absences).
Recent advances in occupancy modeling allow for
the investigation of species co-occurrence patterns
while accounting for difering ecological requirements
Journal of Avian Biology 45: 001–009, 2014
doi: 10.1111/jav.00394
Published 2014. his article is a US government work and is in the public domain in the USA
Journal of Avian Biology © 2014 Nordic Society Oikos
Subject Editor: Wesley M. Hochachka. Accepted 8 January 2014