12/11/2015
Amanda
Spencer
RH:
Spencer × Productivity
Index from Creamer's Field Migratory Waterfowl Refuge
Temporal Patterns in
Passerine Productivity Index in Fairbanks, Alaska.
AMANDA
SPENCER University of Alaska Fairbanks
ABSTRACT Passerines,
Dark-eyed Juncos (Junco hyemalis), Lincoln
Sparrows (Melospiza lincolnii), and Orange-crowned
Warblers (Vermivora celata), breed in
interior Alaska’s boreal forest. Climate change is dramatically affecting the
boreal forest (Hinzman 2005). Bird abundance can indicate changes in vegetation
and changes in weather (Tauzer 2013). This study used the Creamer’s Field
Migration Station (CFMS) in Fairbanks, Alaska banding bird data from 1992. These
data were used to determine temporal patterns for productivity index and relate
the productivity index to species first arrival and weather patterns. The
annual productivity indices of these passerines, number of juveniles per 100
mist net hours, were related by linear regression to first arrival and weather of
the Fairbanks area. This study found the indices’ related over time and have a negative
relationship to mean annual temperature. Other explanatory variables’ linear
regression relationship had various and magnitudes and directions. The
passerine’s productivity various productivity responses are likely due to the
diversity of foraging techniques. Understanding passerine’s productivity
responses to weather can predict future passerine abundance. This is important
for managers in areas where range shift may occur in the future.
KEYWORDS Alaska,
boreal forest, climate change, passerine, productivity index.
Climate
change is dramatically and disproportionately affecting polar regions globally
(Hinzman 2005). The state of Alaska, which is comprised of arctic and
sub-arctic regions, is among areas experiencing rapid change, especially in the
vegetation composition of habitats, and the bird species that rely on them (Butler
2003, Markus 2004, and Tauzer 2013,). The interior Alaska is primarily composed
of boreal forest with massive expanses of spruce (Picea spp.) forest (Boreal 2012). Bird abundance can indicate changes
in vegetation and changes in weather (Tauzer 2013). Productivity is an
effective indicator of changes in abundance.
As
a relatively high proportion of autumn caught birds are juveniles, using a metric
consisting of the number of juveniles caught in relation to mist net catch effort
can represent a productivity index (Benson 2012). Many variables have been
found to affect this productivity index including first arrival of the species
(Butler 2003, Huppop 2003, Markus 2004), annual snowfall (DeSante 1987, Dybala
2013), annual mean temperature (Dickey 2008), and annual precipitation (Dickey
2008). This study considers the annual number of juveniles caught to catch
effort (mist net hours) as it relates to each of these factors to explain
temporal patterns. These relationships can help to understand the productivity
index and find likely models to explain temporal patterns. Boreal nesting
passerines have a variety of migration patterns and diets. To account for the
various life histories, three representative migratory passerine species were chosen
from the watch list of the Neotropical Migratory Bird Conservation Act. These
species were also chosen because of their various migration patterns, diets and
high abundances in Interior Alaska. The species of choice are Dark-eyed Juncos (Junco hyemalis), Lincoln Sparrows (Melospiza lincolnii), and Orange-crowned
Warblers (Vermivora celata).
This
study aims to improve understanding of the major factors impacting productivity
of passerines in the boreal forest. The results of this study will benefit
Alaska Department of Fish and Game, Creamer’s Field Waterfowl Migration Refuge,
land managers of boreal forest (Department of Natural Resources, Borough of
Land Management, US Fish and Wildlife Service), ornithologists, and fellow
volunteer bird banders.
Using
the proportion of juveniles captured in autumn at CFMS, the objectives of this
study are to 1) determine temporal patterns for productivity index and 2)
relate the productivity index to species first arrival and local weather.
STUDY AREA
This
study took place in interior Alaska’s Creamer’s Field Migratory Waterfowl
Refuge, located in Fairbanks, Alaska (Fig. 1, 2). The 1,800 acre land functioned as a dairy
during the Alaska gold rush in the early 1900s. It was purchased by the
community in 1966 to preserve the migratory birds to create Creamer’s Migratory
Waterfowl Refuge managed by Alaska Department of Fish and Game for human use
and recreation (Creamer’s 2015). The long-term
migration bird banding station started in the autumn of 1992 at Creamer’s Field
Migration Station (CFMS), the farthest north migration station (64 50’N, 147
50’W). Bird banding takes place in the spring and autumn to catch migrating
passerines.
Fairbanks
has an annual temperature of -2.45C with variation from 3.3C to -8.2C. The
average precipitation is 275mm with 112 rain days. The average annual snowfall
is 165mm (U.S. Climate Data 2015).
CFMS
consistently maintained mist nets in the same locations beginning in 1992 with little
variation. These mist nets are in old growth spruce forest, deciduous birch forests,
shrub lands dominated by willow and alder, edge habitats and seasonal wetlands (Fig.
3).
The
refuge is used and managed for human experiences with wild spaces. There are
numerous dog walking trails, dog mushing trails, hiking and snow machining
trails. Hunting and trapping is permitted in the area.
METHODS
To
determine temporal patterns for productivity index, juveniles were captured at CFMS
for 24 years. This study used years 1992-2004, 2006-2011. Methods for banding
the songbirds were established for CFMS and are written in the Creamers Field
Migration Station Operations Manual (Guers 2002). CFMS adheres to this manual
to reduce variability among observers over many years. All nets are open for
six hours each day from approximately sunrise or 5:30 am (whichever is later).
Many nets in the seasonal wetlands are not opened if the flooding is above 1.5
meters which occurs in spring. Nets are closed early in heavy or constant rain,
snow, intense wind or excessively hot or cold weather. Daily records of open
net duration are used to calculate total net hours for each season. Nets are checked
at 45 minute intervals to collect birds from the nets and the following
information is recorded: species, age, date, and morphometric data are
collected.
Age
is determined by ‘skulling’ (determining ossification of skull) and by plumages,
and presence of breeding characteristics (i.e. brooding patches and cloacal
protuberances). In this study young of the year and fledglings are considered
juveniles. The productivity index is calculated using the number of juveniles
caught divided by 100 mist net hours of effort.
To
relate the productivity index to species first arrival and weather patterns, the
average annual productivity is compared to weather data collected from Fairbanks
International Airport. The productivity
can be related using linear regression to first arrival of the species, annual
snowfall, annual mean temperature, annual brooding season temperature and annual
precipitation to find relationships between arrival, local weather and
productivity.
Weather
data was collected the Fairbanks International Airport. The maximum daily
temperature, minimum daily temperature, amount of snowfall and amount of
precipitation was recorded each day. The mean annual temperature was an average
of daily maximum and minimums averaged over the course of the year. The annual brooding
season temperature was an average of daily maximums and minimums for June
through August. The total annual snowfall was the total sum of all fallen snow
for the calendar year. The total annual precipitation was the total sum of all
rainfall and snowfall over the course of a calendar year (Fairbanks 2015).
Microsoft
Excel was used to make linear regressions of the annual productivity index to
weather data and arrival data for each year and for each species to best
explain productivity temporal patterns. Each explanatory variable was graphed
against each species productivity index. A linear regression, best supported by
the data, gave the change in the explanatory variable for each unit of
productivity, one juvenile per 100 mist net hours. This linear regression also
supplied the multiple correlation (R2) to determine how close the
data correlated. This was done for each explanatory variable for each passerine
species.
RESULTS
Our
sample determined temporal patterns for productivity index using 23,245 juveniles
of the three species caught over a series 189,099.2 mist net hours from
1992-2004 and 2006-2011 (Fig. 4). The data for year 2005 was not available for
analysis at this time of the study. The Dark-eyed Juncos’, Orange-crowned Warbler’s
and Lincoln Sparrow’s productivity indices were related by linear regression to
first arrival and weather patterns. Linear regression compares one dependent
variable, productivity, to one explanatory variable. Due to the single
explanatory variable, no result was statistically significant, however some trends
were observed.
Our
survey included 11,671 juvenile Dark-eyed Junco. The first arrival of Dark-eyed
Juncos at CFMS had a slight positive relationship of 0.062 Julian days per productivity
with a multiple correlation (R2) value of 0.0025. The annual mean
temperature had a negative effect on productivity with -0.673 degrees Celsius
per productivity with a R2 of 0.3328 (Fig. 5). The annual brooding
season temperature had a slightly negative impact in productivity with -0.0136
degrees Celsius per productivity with an R2 of 0.026. Total annual
precipitation had a slight negative relationship with productivity with a
relationship of -0.0468 centimeters of precipitation per productivity with a R2
of 0.0132. Total annual snowfall had a negative effect on productivity with -1.1688
centimeters of snowfall per productivity with a R2 of 0.0332.
Our
survey included 7,309 juvenile Orange-crowned Warblers. The first arrival of Orange-crowned
Warblers at CFMS had a negative relationship of -0.4545 Julian days per productivity
with a R2 value of 0.0779 (Fig. 6). The annual mean temperature had
a negative effect on productivity with -0.5226 degrees Celsius per productivity
with a R2 of 0.1406. The annual brooding season temperature had a
slightly positive impact in productivity with .0449 degrees Celsius per
productivity with an R2 of 0.016. Total annual precipitation had a
slight negative relationship with productivity with a relationship of -0.0017
inches of precipitation per productivity with a R2 of 0.000001.
Total annual snowfall had a negative effect on productivity with -1.9743
centimeters of snowfall per productivity with a R2 of 0.2122.
Our
survey included 4,265 juvenile Lincoln Sparrows. The first arrival of Lincoln Sparrows
at CFMS had a positive relationship of 2.1291 Julian days per productivity with
a R2 value of 0.0084 (Fig. 7). The annual mean temperature had a
negative effect on productivity with -1.6123 degrees Celsius per productivity
with a R2 of 0.1246. The annual brooding season temperature had a
slightly negative impact in productivity with -0.0042 degrees Celsius per
productivity with an R2 of 0.00001. Total annual precipitation had a
slight negative relationship with productivity with a relationship of -0.1057
centimeters of precipitation per productivity with a R2 of 0.0036.
Total annual snowfall had a slightly negative effect on productivity with -3.7836
centimeters of snowfall per productivity with a R2 of 0.0185.
DISCUSSION
In
determining the temporal patterns of the productivity index, there is a visual
relationship between the three species productivity each year. While the
productivity of each species is different than the others and reacts
differently to environmental factors, all the species have joint peaks of
productivity at 1994-1995, 1997-1998 and 2009. They all have productivity
crashes at 1996, 2002-2003 and 2008 (Fig. 4).
While
the peaks and crashes in productivity appear to be related, each species has a
different relationship to first arrival and weather patterns. These different relationships
vary in magnitude and direction. All species in this study have lower numbers
of juveniles during fall migration when the annual mean temperatures are high.
As temperatures are expected to increase in interior Alaska due to climate
change (Dybala 2013), these species may experience reduced productivity. This
negative relationship between passerine productivity with temperature could be
attributed to a variety of factors including food abundance, heat affects or the
increased abundance of parasites. Interestingly, the annual brooding season
temperature had little effect on the productivity index compared to annual mean
temperature. This possibly could be because the forage available each summer is
dependent on the over-wintering conditions or other in-direct affects.
Interestingly
the Lincoln Sparrow productivity remained more consistent than the other
species’ productivity indices. This shows that Lincoln Sparrows are less
affected by positive or negative changes. This may be because of the foraging behavior
or nesting behavior of Lincoln Sparrows being different than the other species.
This may also be because the Lincoln Sparrows are able to use more alternatives
to use available forage. More research exploring Lincoln Sparrows resilience
should be explored.
The
Dark-eyed Junco and Orange-crowned Warbler have higher productivity indexes in
years with less snowfall. This may be a direct cause of decreasing available
forage or nest sites, but it maybe indirect with snowfall decreasing early spring
forage by other methods or it could be a correlation. More research exploring
snowfall depth’s impacts on passerines should be explored. Doing a productivity
study for a wider range of passerines with stratification by foraging behaviors
would allow more insight on particular weather’s effects of various foraging
behaviors.
Orange-crowned
Warblers’ and Lincoln Sparrows’ productivity indices have opposite relations to
arrival dates. Orange-crowned Warblers have lower productivity when they arrive
late (Fig. 6). Conversely, Lincoln Sparrows benefit from earlier arrivals (Fig.
7). Arrival timing is expected to continue to alter with climate changes
(Butler 2013). Arrival dates have not been correlated to breeding dates, therefore
arrival does not necessarily mean the birds are beginning the breeding season.
A more definitive way to find the beginning of the breeding season is checking
cloacal protrusions on males and brooding patches on females to note active
breeding behaviors.
Annual
precipitation had little relation to productivity for any of the species
possibly because the annual precipitation is not as important as precipitation
during the brooding season itself (DeSante 1987). A further study looking at
consecutive rain days during the brood season may have a larger effect on nest
survival.
Juveniles
of most passerine species depart their breeding grounds in Alaska significantly
earlier than adults (Benson 2012). The juvenile to mist net hours is not
accounting for departure of juveniles, this biases the data because an early
autumn migration will reduce the total juveniles caught. This will lead to
underestimating the productivity index in those years. To correct for this, the
last date of juvenile capture should mark the end of the season.
This
study likely consistently underestimates productivity. However, because this
index is used with explanatory variables to find influential factors of
productivity, the question is if the productivity index a good representation
of actual productivity and not biased by detection probability variation. In
this study, detection bias is not considered an issue because the same methods
for detection with the same nets were used each year. The variation of
detection probability between years would be minimal. A future passerine study
of productivity could be coupled with juvenile mist net effort to determine if
indeed there is a detection bias.
MANAGEMENT IMPLICATIONS
Modeling
passerine productivity index’s response to various weather, allows predictions
of passerine recruitment due to weather. The correlation between annual mean temperature
and productivity index shows a decrease in productivity during warmer years.
This can be used to predict passerine abundance changes due to potential annual
temperatures. As recruitment to a population declines, the population abundance
will decline. To stabilize recruitment into the population, passerines may nest
in more northern latitudes or high altitudes to avoid this ecological trap.
Acknowledgments
I
thank T. J. Brinkman, J. A. Curl, A. Harding Scurr for review comments and
contributions to this manuscript.
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Figure
3. Mist net locations of the Creamer’s Field Migration Station.
Figure
4. Relationships between annual productivity indices of Dark-eyed Juncos, Orange-crowned
Warblers, and Lincoln Sparrows and time in years.
Figure
5. Relationship between annual mean temperature in Celsius and productivity
index of Dark-eyed Juncos.
Figure
6. Relationship between date of first arrival and productivity index of Orange-crowned
Warblers.
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