predictors of wild turkey (meleagris gallopavo) population

Riva T. Madan Wild Turkey Population Change Spring 2015

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Predictors of Wild Turkey (Meleagris gallopavo) Population Change

in California from 1972 to 2013

Riva T. Madan

ABSTRACT

Wild turkeys never lived in California until humans began introducing them around 1908. The

success of this introduced species was variable at first, but in the 1990s there was a noticeable

increase in their population and range. The reasons behind what drove these population changes

are unknown, especially with only a few studies on wild turkeys in California. With Breeding Bird

Survey, National Land Cover Database, PRISM Climate Group, and California Department of Fish

and Wildlife data, I used a zero-inflated poisson mixed model to determine the influence of

climate, translocations, hunting, and land cover on wild turkey population change. I ran a separate

model for three land cover distances. I calculated population change statewide and at individual

routes using a generalized linear model. Wild turkey populations increased 10% per year statewide

from 1972 to 2013. Hunting was the most influential predictor of population change showing that

populations are increasing even as hunting increases. Land cover was a significant predictor, but

the effects of different land cover types varied with the scale looked at. Urban land cover was

positively related to population change only at 1 and 10 km distances. At the 5 km distance, forest,

grassland, and agriculture were inversely related to population change. This study suggests

increasing urban land cover, which may increase food supply, can increase wild turkey

populations. However, the results also suggest that other factors that are not included in this model

have large influences on wild turkey population change.

KEYWORDS

introduced species, Breeding Bird Survey, zero inflated poisson mixed model, land cover

change, ArcGIS


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INTRODUCTION

Humans often accidentally or intentionally transport species to new locations beyond their

natural range. If the species successfully establishes itself, the species becomes an introduced

species. At this point their populations may grow very large and spread over a large area. If the

introduced species has both expanded in range and is causing ecological impacts, the species is

considered invasive (Duncan et al. 2003). Large populations of introduced species can also be

destructive to human activities, making the species become a pest (Baldwin et al. 2012). Wild

turkeys (Meleagris gallopavo) are an example of an introduced species in California that has

expanded in range and increased in population. Although they are potentially a pest, wild turkeys

are not considered invasive due to lack of research on its ecological impacts.

The wild turkey never originally lived in California. Is native range is east of the

Mississippi River, in the southwest United States (Arizona, New Mexico, and Texas), and Mexico

(Gardner et al. 2004). In the late 1800s, wild turkey populations in their native range declined as a

result of deforestation and hunting (Dickson 1992; Barret and Kucera 2005). As a reaction to the

fear of turkeys going extinct, turkeys were introduced into every state except Alaska (Dickson

1992). The first record of wild turkey introduction in California was in 1877, when private ranchers

introduced the wild turkey on Santa Cruz Island. The next recorded introduction was in 1908 when

the California Department of Fish and Wildlife (CDFW) began releasing turkeys for hunting

purposes. Since then the CDFW has released several thousands of turkeys throughout California,

but many of these introduction attempts failed (Gardner et al. 2004). However, the CDFW noticed

an increase in wild turkey population size and range in the 1990s. As a response, they stopped

releasing turkeys in 1999 (Gardner et al. 2004). The reasons behind the recent spread and growth

of wild turkey populations in California are not well understood, but past studies of wild turkeys

in their native range provide knowledge about what effects their population growth.

Nesting failure and food availability are two main factors that determine wild turkey

population growth. These two factors largely depend on habitat type (Spohr et al. 2004; Lehman

et al. 2008; Fuller et al. 2013). The primary reason for nest failure is from predation and the chance

of nest predation depends on the density of vegetation. In areas with somewhat dense vegetation,

turkeys are able to hide their nests from predators well and therefore reduce nest predation. Open

grasslands would have a high amount of predation due to little nest obscurity. However, if the


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vegetation is too dense, such as a dense shrubland, it reduces the ability of a turkey to flee if a

predator arrives, thereby increasing predation (Keegan and Crawford 1997; Lehman et al. 2008;

Fuller et al. 2013). Urban and agricultural areas have been shown to increase food availability.

Wild turkeys are often reliant on agricultural grains and human feeding to maintain large

populations when other food sources are low (Burger 1954; Barrett and Kucera 2005). Although

urban areas may provide additional food, habitat fragmentation from urbanization has resulted in

increased nest predation (Hogrefe et al. 1998; Sphor et al. 2004). Contrastingly, habitat

fragmentation between forests, open areas and agriculture supports large turkey populations

(Glennon and Porter 1999). A majority of these studies on factors affecting population growth took

place in several states in the wild turkey’s native range, but not in California. There have only been

four studies (Burger 1954; Gardner et al. 2004; Barret and Kucera 2005; Wengert et al. 2009) on

wild turkeys in California and three of the four studies are outdated by at least a decade.

Land use changes increasing habitat fragmentation, urbanization and agriculture bring wild

turkeys closer to humans and make them more likely to become pests. Wild turkeys were shown

to damage twenty-three crops, with corn being the most often damaged crop (Tefft et al. 2005).

Additionally, residents of suburban areas have complained about large numbers of turkeys being

a nuisance on their property (Barrett and Kucera 2005). However, no study has analyzed whether

these land use types have encroached on wild turkey habitat or whether wild turkeys have moved

from their original release sites to these land use types. Besides anecdotal and observational

information, there has not been any studies looking at how the population and range of wild turkeys

has changed in California and the possible factors driving the changes.

To better understand wild turkeys in California and as a step towards determining if they

are pests or invasive species, this study maps and analyzes the changes in population and range to

explain the reasons behind the recent spread and growth of wild turkey populations. Climate and

human involvement, such as hunting and translocation, have often influenced population changes

of species, but I hypothesize that land cover, specifically agriculture and urbanization, had the

greatest influence in increasing wild turkey populations in California. To see whether wild turkey

populations are actually increasing, I quantified how wild turkey population changed and mapped

how range changed in California from 1968 to 2012. To determine what factors could have led to

the changes in population, I ran a model to determine the influence of land cover, climate, and

human involvement on wild turkey population changes.


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METHODS

Study system

The sites I looked at were predetermined by the USGS Patuxent Wildlife Research Center’s

North American Breeding Bird Survey that carries out bird surveys to determine presence and

count of many bird species, one of which being the wild turkey. Wild turkeys are generalists; they

can live in many different types of habitats, such as forests, shrublands, grasslands, woodlands and

around agriculture and urban areas (Dickson 1992). Each study site is surrounded by a combination

of these land types. Percentages of each land type: grassland, shrubland, forest, agriculture, and

urban, are given for each site with wild turkeys present in Appendix A.

Data sources

I obtained georeferenced wild turkey population data from the North American Breeding

Bird Survey (BBS) to determine population change. The BBS conducts point count surveys along

40 km long predetermined, non-random roadsides called routes (Figure 1). Citizens skilled in avian

identification conduct surveys between mid-May and early July. Beginning a half hour before

sunrise, the surveys, conducted by one observer, take 4 to 5 hours to complete. Point counts are

taken every 0.8 km along the route, for a total of 50 stops. The point count is conducted by

recording every bird seen or heard within a 400 m radius over a 3 minute timespan (Link and

Sauer 2002; Sauer and Link 2011). The USGS Patuxent Wildlife Research Center has been

conducting the BBS since 1966, but surveys in California began in 1968. I only included data

beginning in 1972 because I originally planned on using LANDSAT satellite imagery that began

in 1972 for land cover type, but the classification was too inaccurate to use. Only one detection of

a wild turkey in 1969 was left out from my analysis. Although the BBS is conducted annually, not

every route is surveyed each year. Turkeys have been observed in 80 routes in California between

1972 and 2013. I only included 76 routes because there was no geospatial information on the path

of these four routes.


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Figure 1: Routes with and without wild turkeys present

I obtained wild turkey translocation and hunting data in California from the California

Department of Fish and Wildlife (CDFW). Translocation data included translocation within

California and between California and another state. Translocation data is from 1959 to 1999 with

the year 1998 missing. For hunting data, CDFW calculated the amount of turkeys hunted by

extrapolating the results from surveys. Mail survey forms were sent out to randomly selected

hunters until approximately 4% of the hunters returned the survey. CDFW has not documented the

specific methodology of the extrapolation. Data spans from 1949 to 2010, with the year 2009

missing. However, there was still hunting in 2009 and after 2010. Instead of leaving zero values

indicating no hunting, which would be incorrect, I used the same data from 2010 for years 2009 to

2013. This was reasonable given that the counties where wild turkeys were hunted had very little

variation since 2000. Both of these datasets are only specific to county, but routes are at a smaller

scale and sometimes span two counties. Therefore it would be incorrect to directly assign each

route a specific value for this data. To account for this inaccuracy, I only included hunting and

translocation as binary data to show whether it occurred in the county the route in. I assigned a


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county to each route based on which county the majority of the route was located in. For a few

routes that were about evenly split between counties, I combined both of the county’s data.

To obtain land cover data, I used USGS National Land Cover Database (NLCD) for 1992,

2001, 2006 and 2011. I only looked at five land cover classes: forest, urban, grassland, agriculture

and shrubland. Because these datasets contain more than just five land cover classes, I combined

similar land cover classes together. For example, I included both evergreen forest, mixed forest,

and deciduous forest under the forest land cover type. I calculated percentage of land cover within

three buffer distances from the route path: 1, 5 and 10 km. Wild turkey flock home range varies

from 1.5 to 10 km2, therefore I will use multiple distances to account for the various home range

sizes as well as to see if habitat has different effects at different scales (Zeiner et al. 1990; Gardner

et al. 2004; Barrett and Kucera 2005). For years that didn’t have a given NLCD, I used the same

land cover values from the next available dataset with the assumption that the land cover didn’t

change very much between these years. Years prior to and including 1992 had land cover values

from the 1992 dataset, years 1993 to 2001 had values from the 2001 dataset, years 2002 to 2006

had values from the 2006 dataset and years 2007 to 2013 had values from the 2011 dataset.

For climate data, I used temperature and precipitation raster images from Oregon State

University PRISM Climate Group. To obtain average temperature and precipitation for each route,

I used ArcGIS v10.2 (ERSI 2014) to average the temperature and precipitation within 10 km from

each route path. I only used one buffer distance for climate variables because it is unlikely that the

climate will vary significantly under 10km from the route. I squared the average temperature and

precipitation because animals tend to have a quadratic relationship to temperature and precipitation

rather than linear.

Data Analysis

To look at range change, I used kriging, an interpolation technique, in ArcGIS v10.2 to

determine wild turkeys range in California for 6 year intervals. To quantify population change of

wild turkeys at each route and California overall from 1972 to 2013, I used a generalized linear

model (GLM) in R (R Core Team 2014) v3.1.2 using the glm package. I did not run a GLM on

routes that only contained one non-zero data point and I excluded the results of routes where the

generalized linear model didn’t converge due to lack of data. To see the effect of different


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predictors on population change, I used a zero-inflated poisson mixed model in R v3.1.2 using the

glmmADMB package on all 77 routes. I included the percentage of land cover type, temperature,

precipitation, the occurrence of wild turkeys translocation and hunting as covariates. Route was

included as a random effect to account for differences between sites that cannot be explained by

the covariates. I standardized all values, excluding binary and percent values, by subtracting the

mean and dividing by the standard deviation to get more comparable covariate estimates. I ran a

separate zero-inflated model for each of the 3 land cover buffer distances.

RESULTS

Population and Range Change

Wild turkeys throughout California increased in population 10.09% (±0.279%) from 1972

to 2013, according to the generalized linear model (p < 0.0001). Different areas in California

experienced various amounts of population change (Figure 2). Majority of routes experienced less

than a 25% increase in population. More specifically, 31 out of the 61 routes included for this

analysis had an increase between 9% and 20% Route 416 near Meadow Valley in Plumas county

experienced the greatest population change of 108% per year, but was statistically insignificant (p

= 0.0644). Route 210 in Mendocino County and route 12 in Sutter County had increases of 88%

and 67%, respectively, but were also not statistically significant. Route 172, located east of

Berkeley in Contra Costa County, had the greatest statistically significant increase of 39% per year

(p < 0.001). Route 422 located south of Yosemite National Park had the next highest significant

increase of 30% per year (p < 0.01). Route 415 near Crescent Mills in Plumas County and route

202 in Sonoma County had a significant increase of about 28% and 27%, respectively (p < 0.5).

Four routes had a decrease in wild turkey population, but only one route had a significant decrease

in population. This route, route 409 located between Burnt Ranch and Hyampom in Trinity

County, decreased in population of 81% per year (p < 0.05). The percent population change per

year, standard errors and p-values for each route is given in Appendix B.

Range expansion can be seen through a significant (p < 0.0001) increase of 0.266%

(±0.035%) per year in the proportion of wild turkey detected on BBS routes from 1969 to 2013

(Figure 3). This is further supported by kriging maps of six year intervals which show wild turkey


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range, indicated in blue, to be increasing (Figure 4). Based on the maps, the largest expansion was

between 1984 to 1989 interval and the 1990 to 1995 interval. After 1990, the population, indicated

by the darkness of blue, increased more than the range changed. From 1972 to 1989, turkeys

increased 24.8% (±4.56%) per year. However, this increase was actually insignificant (p = 0.489).

In the following period, 1990 to 2013, turkey populations increased by 91.3% (±4.56%) per year

(p < 0.0001). Given that there was no significant increase in population before 1990 and that the

kriging maps show very little change in population and range, I decided to only include years 1990

to 2013 when running the zero-inflated poisson model. Additionally, because the land cover data

only begins in 1992 and all years prior to 1992 would have the same land cover value, the result

of the model would be more accurate after removing the years before 1990.

Figure 2: Geographic distribution of population change. Percent population change, calculated by the generalized

linear model, of wild turkeys from 1972 to 2013. Routes with significant population change are indicated by a black

dot.


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Figure 3: Wild turkey presence. Proportion of Breeding Bird Survey routes in California where wild turkeys were

detected from 1972 to 2013.

Figure 4: Wild turkey range expansion. I mapped the change in wild turkey range and population from 1972 to

2013 by averaging BBS counts for six year intervals for each route and then used kriging (in ArcGIS).

0

0.1

0.2

0.3

0.4

0.5

0.6

1971 1977 1983 1989 1995 2001 2007 2013

Perc

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Year


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Predictors of population change

According to the zero-inflated poisson mixed model for years 1990 to 2013, the most

significant predictor of wild turkey population change for all buffer distances was hunting (p <

0.01) (Table 1). The estimate for hunting was 0.8 for 1 km buffers, 0.74 for 5 km and 0.73 for 10

km. Year was the second most significant predictor than hunting, but still very influential with

estimates of 0.74 for 1 km, 0.78 for 5 km and 0.75 for 10 km. All other factors were much less

influential predictors with estimates of less than 0.1. Precipitation, temperature, translocation and

grassland were not significant for any of the buffer distances. The 95% confidence intervals

included zero, making the effect these covariates neither positive nor negative.

Different factors were significant and had various influences on population change at each

buffer distances. For the 1 km buffer, the percent of urban land cover was significant (p < 0.01)

with an estimate of 0.096. This indicates that areas with increasing urban land cover will

experience increasing population sizes, given that all other factors are held constant. Similarly, for

the 10 km buffer, urban land cover was also significant (p < 0.05) and positively related to

population change (estimate of 0.047). For the 5 km buffer, forest, shrubland and agricultural land

cover were significant covariates, but urban land cover was not (p = 0.078). Forest, shrubland and

agriculture were all inversely related to population change. Forest, with an estimate of -0.077 was

more of an influential predictor than shrubland and agriculture, both with an estimate of -0.023. A

negative estimate indicates that areas that are decreasing with this land cover type have an

increasing wild turkey population.

Table 1: Parameter values for the 3 buffer distances. Summary of each parameter’s influence on wild turkey

populations according to the zero-inflated mixed model. Significant p-values are indicated with *.

Parameter Estimate Standard Error 95% CI p-value

1 km buffer

Intercept -4.0925 2.6214 -9.2303 - 1.0453 0.1185

Year 0.73708 0.06707 0.60563 - 0.86852 <2e-16 *

Precipitation 0.01506 0.04268 -0.06859 - 0.09872 0.7242


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Temperature -0.04387 0.11007 -0.25961 - 0.17186 0.6902

Translocation -0.05994 0.1417 -0.33767 - 0.21778 0.6723

Hunting 0.79648 0.2655 0.27611 - 1.3169 0.0027 *

Urban 0.09639 0.03484 0.02809 - 0.16468 0.0057 *

Forest 0.01566 0.02663 -0.03653 - 0.06785 0.5564

Grassland 0.03881 0.02813 -0.01633 - 0.09395 0.1677

Shrubland 0.03149 0.02788 -0.02315 - 0.08614 0.2587

Agriculture 0.00765 0.02801 -0.04726 - 0.06255 0.7849

Random effect: Variance Standard Deviation

Route 1.323 1.15

AIC: 3562.2

5 km buffer

Intercept 0.95248 0.86066 -0.73438 - 2.6393 0.26843

Year 0.78086 0.0689 0.64582 - 0.91589 <2e-16 *

Precipitation 0.01047 0.04474 -0.07722 - 0.09816 0.81499

Temperature -0.0856 0.10986 -0.30092 - 0.12972 0.43586

Translocation -0.02073 0.14272 -0.30045 - 0.259 0.88454

Hunting 0.84435 0.26364 0.32763 - 1.3611 0.00136 *

Urban -0.0276 0.01568 -0.05833 - 0.00313 0.07837

Forest -0.07653 0.02494 -0.12542 - -0.02764 0.00216 *

Grassland -0.01645 0.01087 -0.03775 - 0.00485 0.13005

Shrubland -0.02385 0.00717 -0.0379 - -0.00981 0.00087 *

Agriculture -0.02371 0.00914 -0.04162 - -0.00581 0.00945 *


Route 1.569 1.253

AIC: 3576

10 km buffer

Intercept -1.45385 0.95128 -3.3183 - 0.41062 0.1264

Year 0.75024 0.06791 0.61713 - 0.88334 <2e-16 *

Precipitation 0.02071 0.04243 -0.06245 - 0.10387 0.6255

Temperature 0.01947 0.11131 -0.1987 - 0.23763 0.8611

Translocation -0.02511 0.14204 -0.30351 - 0.25328 0.8597

Hunting 0.83607 0.27729 0.29259 - 1.3795 0.0026 *


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Urban 0.04671 0.01972 0.00807 - 0.08536 0.0178 *

Forest -0.00489 0.01065 -0.02576 - 0.01598 0.6461

Grassland 0.01008 0.01215 -0.01373 - 0.0339 0.4067

Shrubland 0.00394 0.01232 -0.0202 - 0.02808 0.7489

Agriculture -0.02697 0.01459 -0.05555 - 0.00162 0.0645


Route 1.288 1.135

AIC: 3753

Land Cover

Most wild turkeys were found on routes with majority of forest land cover type. Second

was grassland and then shrubland. For 1 km and 10 km buffers, wild turkeys were common in

land cover types in that same order, but for 5 km the routes with more turkeys tended to be

majority grassland, instead of forest, and followed by shrubland instead of grassland. To get a

perspective of how land cover is changing and predict how wild turkeys should respond, I used

linear regression to calculate the overall change in each land cover type at each buffer distance.

Urban and shrubland both significantly increased for all buffer distances. Forest and grassland

significantly decreased for only the 5 km buffer. Agriculture had no significant changes, but

showed decreases in land cover except for the 10 km, which showed a very small increase

(0.002). All the estimates for agriculture were very small, compared to forest, urban, grassland

and shrub land cover types changes that ranged from 20% to over 40%.


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Figure 5: Wild turkey habitat preference. The number of turkeys found in each land cover type based on the route’s

majority land cover type.

Table 2: Change in land cover. Using linear regression, I calculated the change for each land cover type at each

buffer distance using values from 1992, 2001, 2006 and 2011. Significant p-values are indicated with *.

Land Cover Buffer

Distance

Estimate Std. Error p-value

Urban 1 km 0.19199 0.06022 0.00158 *

5 km 0.23114 0.04801 2.29E-06 *

10 km 0.31237 0.05497 3.02E-08 *

Shrub 1 km 0.4097 0.1218 0.000864 *

5 km 0.4367 0.1457 0.00294 *

10 km 0.3173 0.1274 0.0133 *

Forest 1 km -0.428 0.2323 0.0664

5 km -0.20107 0.09273 0.0309 *

10 km -0.4189 0.2141 0.0513

Grassland 1 km -0.2172 0.1568 0.167

5 km -0.4267 0.1571 0.0069 *

10 km -0.2093 0.1397 0.135

Agriculture 1 km -0.0079 0.149641 0.958

5 km -0.0354 -0.278 0.781

10 km 0.002 0.121 0.988

0

200

400

600

800

1000

1200

1400

Forest Grassland Shrub Urban Agriculture

Nu

mb

er o

f Tu

rkey

s

Land Cover Type

1 km

5 km

10 km


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DISCUSSION

Wild turkeys have significantly increased in population by 10% per year since 1972, which

is consistent with anecdotes and observations of turkey increases. As hypothesized, urban land

cover had a significant factor influencing population change, and climate and translocation were

not significant. Hunting, which I did not hypothesize to be significant, was the most influential

factor. Additionally, I expected there to be similar results at each buffer distance or if there was a

difference, it would be the 10 km distance because it could be too large of an area to accurately

predict turkey population change. Although I expected agriculture to be significant, it was

surprising that agriculture, forest and grassland were all inversely related to population change,

but only at the 5 km distance. Urban land cover was only significant at 1 and 10 km distances.

These differences likely indicate that habitat effects wild turkeys differently at the local and

landscape scale. Given that the results of the model indicate year as the second most influential

factor, the factors I included in the model are probably not enough to explain why wild turkey

populations have increased.

Population and Range Change

Wild turkey population increase is not centralized in one specific region, but the rate of

increase is similar throughout California. Besides Plumas County, all routes with over 20%

increase in population were in different counties. The fairly large increase of 28% per year for

route 202 in Sonoma County is consistent with Barret and Kucera's research in Annadel State park

located less than 10 km from the route (2005). The wild turkey populations grew rapidly in Sonoma

because there were no limiting factors for reproduction, such as lack of food. The food supply was

actually increased due to humans intentionally and unintentionally feeding turkeys high quality

commercial feed. High food quality can then lead to increased reproduction. With a high nutrition

diet, female turkeys tend to lay more eggs and lay eggs earlier in the breeding season, allowing for

attempts at nesting if one fails (Dickson 1992). Adding to that, wild turkeys already have a large

reproductive capacity. They start reproducing at the age of one and each year they can produce a

clutch size, the number of eggs laid in a single nesting, between 10 and 12. And if their first nesting

fails, they often nest a second time (Lehmen et al. 2008). Route 172, with the largest significant


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increase of 39% per year, had the largest amount of urban area for the 5 and 10 km distances and

fourth largest amount at 1 km. Route 172 probably had similar reasons for population increase as

route 202 in Sonoma county. The large amount of urban area increases the likelihood of wild

turkeys getting additional food from humans.

All routes increased in urban land cover. However, urban land cover did not seem to

increase by such a large percent where it seemed to be encroaching on natural turkey habitat. Wild

turkeys were probably either introduced near urban areas or spread there due to the availability of

increased food supply and quality. Therefore, it is more likely that turkeys moved into areas with

humans, rather than humans moving into areas with turkeys. Surprisingly, many turkeys are not

found in routes with a majority of agriculture or urban land cover, especially considering urban

land cover was a significant influence on population change (Figure 5). This is probably because

CDFW didn’t introduce them into areas that are majority urban or agriculture, but some turkeys

spread to these locations.

Route 409, which declined in population, also increased in urban land cover and was not

very different from other routes in terms of land cover composition and change. I suspect that the

change at this route is not representative of the change in that area. However, routes in that area

did have much smaller population changes of less than 5% increase per year. This is likely do to

later introductions by CDFW in this area according to translocation data (Gardner et al. 2004).

Insignificant results and non-convergent GLMs from calculating population change at each

route is probably due to lack of data. Not every route was surveyed every year and some routes

were surveyed as few as 8 to 10 years. Although the results were insignificant or the GLMS did

not converge, the trends they show, usually of an increasing population, are probably close to what

is actually happening. This is likely because translocation and hunting data often indicated turkeys

in counties that the BBS did not detect them. The BBS data is just lacking data to statistically show

the trend and is probably underestimating wild turkey populations. Wild turkeys probably also

have a larger range than what is shown in kriging maps based of BBS data (Figure 4).

Wild turkeys are found more frequently on routes in northern California, with the exception

of route 106 in southern California near San Diego. The majority of routes where turkeys were

found are primarily in the foothills and mountains of California. This distribution resembles the

distribution of CDFW’s wild turkey release sites from 1959 to 1999 (Gardner et al. 2004). This

makes it unclear whether wild turkeys are found in this pattern because CDFW released them in


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these locations, turkeys have a particular habitat preference, or both. CDFW may have also

released them there because studies suggested that these areas are suitable habitat. Consistent with

studies of wild turkey habitat preference, turkeys seem to be found in places that are primarily

forest, followed by grassland, shrubland or a mixture of these three (Figure 5) (Dickson 1992;

Gardner et al. 2004). This consistency shows that wild turkeys have similar preferences and

characteristics regardless of the specific location they are studied. This allows us to reliably make

inferences about wild turkeys in other states they have not been studied in.

Predictors of population change

Hunting is the most influential predictor of wild turkey populations, with coefficient values

ranging from 0.79 to 0.84. This positive value indicates that wild turkey populations increased

with more hunting. However, it is more likely that relationship is reversed. From looking at the

data, the amount of hunting seemed to increase as a response to an increasing wild turkey

population. The years that turkeys show up in counties and the year hunting starts in those counties

often coincides. For example, in San Diego county turkeys did not show up until 1990 to 1995 and

similarly hunting did not start until 1992. This was not a result of hunting data being inputted as

occurrence data in binary rather than number of turkeys actually hunted, which could show an

increase in counties hunted but not an increase in numbers hunted. However, the amount of turkeys

hunted in California usually increased every year according to CDFW Game Take Survey Reports.

Therefore, this result is actually showing that although hunting is increasing, wild turkey

populations are increasing at an even greater rate. The implications of this could be huge,

especially if CDFW expects wild turkey populations to stay under control or stop increasing from

hunting alone.

As predicted, land cover type was an influential predictor of wild turkey population change.

The percent of urban area was the only significant land cover factor at both the 1 and 10 km

distances. In Sonoma County, urbanization is probably increasing food supply. Additional food

sources support larger populations than a habitat would naturally allow (Burger 1954; Barrett and

Kucera 2005; Glennon and Porter 1999). In winter and times when vegetation does not grow well,

food is scarce, but if wild turkeys are not reliant on natural food sources, the food scarcity won’t

have any effect on them. Contradictorily, urbanization may increase predation for wild turkeys due


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to the habitat fragmentation it creates and therefore reduce populations (Hogrefe et al. 1998; Sphor

et al. 2004). It is likely that this is not a huge influence in many places in California because turkey

predator populations are probably low.

Looking at a 5 km scale, forest had a negative influence on population change. Forest is

one of the wild turkey's primary habitat and also is the most preferred habitat in California,

therefore it seems unreasonable that wild turkeys would increase due to a reduction in forest cover

(Dickson 1992) (Figure 5). Instead, given that forests are decreasing in general, it is more likely

that the negative coefficient could be a coincidental result of the two trends occurring at the same

time (Table 2). Similar to the result with hunting, although decreasing forest cover is correlated

with an increasing population, one is not the cause of another. This suggests that another factor

could be influencing wild turkey population increase. The same could be said of agriculture which

decreased overall as well. For shrubland, however, the reason for the inverse relationship with

population change could be because dense shrubland is bad habitat for turkeys because it can

increase predation as well as limits the mobility of them (Keegan and Crawford 1997; Gardner et

al. 2004; Lehman et al. 2008; Fuller et al. 2013). But this is uncertain given that a large amount of

turkeys resides on routes with majority shrubland (Figure 5).

The effects of land cover type at different scales are surprisingly different and

contradictory. The reasons that different factors are significant at different scales is not clear.

Looking at the land cover, it seems these results are attributed to land cover percent values because

the 1 km and 10 km distances often have very similar values that are different from the 5 km

distance. These results show that wild turkeys are effected by habitat at different scales, both local

and landscape. The different results could be also indicating that wild turkeys have a specific home

range size, making two of the three distances inaccurate. If turkeys have a particular home range,

one would expect the results for that distance to best explain the population change. However, the

home range of wild turkeys specifically in California is unknown, therefore it is hard to say which

of the three distances is probably the most accurate. Also, based on other studies, the home range

has a huge range from 1.5 km2 to 10 km2 scales (Zeiner et al. 1990; Gardner et al. 2004; Barrett

and Kucera 2005). Additionally, it is complicated to compare wild turkeys response to habitat at

different scales because turkeys have a huge capacity for range expansion. Hens disperse up to 13

kilometers to find a nest site and individuals can fly up to 64 kilometers in a week (Barrett and

Kucera 2005). Habitat fragmentation and edge effects, which were not taken into account in my


18

model may also be a cause of these odd results. Looking at the land cover at each routes, many of

them did have similar amounts of several land cover types, often about 20%. Wild turkeys are

often associated with edges and fragmentation and can have large populations in these landscapes

(Glennon and Porter 1999).

Temperature and precipitation did not significantly influence population change. It is

reasonable that temperature would not be influential because wild turkeys are habitat generalists.

In addition, they have a large range that spans the entire United States and successfully survive in

various climates (Dickson 1992). Increased precipitation, on the other hand, has been associated

with lower populations due to increased predation. Predation increases in wet weather because the

wild turkey’s scent is more detectable under wet conditions (Palmer et al. 1993, Roberts et al.

1995, Roberts and Porter 1998). However, other studies have shown that the effect of precipitation

is more complex (Lehman et al. 2008; Fuller et al. 2013). In addition to the strength of their scent,

the probability of predation also depends on the type and density of vegetation cover. A large

amount of precipitation over at least a short duration, such as a few days, can increase vegetation

density and therefore reduce predation because more vegetation cover decreases detectability from

predators. The effect of precipitation is varied such that it could either increase or decrease

populations depending on the vegetation type in the area and the amount and duration of

precipitation. The possibility of both positive and negative effects could be a reason why the 95%

confidence interval for precipitation spanned zero, indicating neither a clear positive nor negative

influence on population change. Consequently, precipitation was not a significant predictor of

population change.

Limitations

The biggest limitation of this study is that it relies solely on the accuracy and consistency

of secondary data. Breeding Bird Survey data has a specific methodology and is much more

consistent than other citizen survey data, such as the Christmas Bird Count. However, it still

contains inconsistencies, most notably that not every route is surveyed every year. There is also

surveyor error due to their varying experience in observing and identifying birds. In addition, the

route locations I used for mapping and obtaining land cover and climate data have low accuracy.

According to BBS, if part of the route was not on the map they were digitizing they estimated the


19

route’s location. Start and end points of the routes were also estimated, which often made the

digitized routes longer than the actual routes surveyed. Also, parts of some routes were not mapped

out in the shapefile due to traffic noise or overlap with other routes. The routes BBS uses may also

cause inaccuracies because they follow a road. Road-based surveys are usually biased because

animals are often reluctant to go near a road. Although this hasn’t been shown with wild turkeys,

it has been shown that a turkey flock is less detectable when near roads and populations are

therefore underestimated (Butler et al. 2007). Also, since BBS routes are located on secondary

roads, which are primarily in rural areas, trends in more urban areas are hard to detect (Butcher et

al. 2014).

Additionally, limited number of years for land cover datasets probably lead to less accurate

results. The USGS does not advise the 1992 dataset to be combined with the other datasets due to

different methods in creating it. I still combined the data because the land cover classes had similar

descriptions and did not look too different visually. However, this could still have led to

inaccuracies. For example, urban area seems much smaller for all 1992 years. This could be due

to the different methods used or due to an actual increase in urban area. The only alternative to the

NLCD would have been classifying satellite imagery, but it is not easy when looking at a large

area, such as California, and it may not have been any more accurate.

Future Research

Although vegetation influences the amount of wild turkey predation, the direct effect of

predator population was not taken into account in this study. Adding a predator population as

another covariate to the model in this study will distinguish whether habitat’s effect on population

is due more to its ability to reduce or increase predation or to other factors, such as food

availability.

From this study and the lack of research on wild turkeys in California, the question of

whether an increase in wild turkey populations has a significant ecological impact remains

unanswered. Like many introduced species, large wild turkey populations have the potential to

impact food web dynamics. Invasive and introduced species, including wild turkeys, can influence

native species populations through both apparent and direct competition. Apparent competition

occurs when growing populations cause their predator population to increase leading to a potential


20

decline in population of another species that their predators prey on. Direct competition occurs

when a species has to compete for habitat and food resources with a similar species that shares its

range. Research into other ground-nesting birds, such as the mountain quail and California quail,

would be useful to see if they are experiencing direct or apparent competition from wild turkeys

because they have similar predators and diet.

The maps of wild turkey population growth can also be used to find localized areas for

further research. Areas with the largest population increase can be studied to see if there are any

detrimental effects, either ecological impacts or as pests, because the effects would be most

noticeable in these areas.

Conclusion

The wild turkey has been introduced into every state except Alaska as well as several other

countries, and in general they have been successful in establishing populations. Besides the wild

turkey, there are many species, usually invasive or introduced, that are spreading their range over

huge areas. These species tend to be generalists and often do well in urban or agricultural

environments, such as the wild turkeys and Eurasian collared doves (Fujasaki et al. 2010). As

urbanization and agriculture increases all over the globe, these species have more areas that they

can expand in successfully. These land cover changes are creating homogenous landscapes that

promote non-native species, the loss of native species and therefore less biodiversity (McKinney

2002).

AWKNOWLEDGEMENTS

I would like to greatly thank Steve Beissinger for telling me about the Breeding Bird

Survey and giving me the suggestion to go about answering my thesis questions with a

modelling approach. My thesis would have been very different if I wasn’t able to discuss my

initial research question with him. He also suggested my mentor, Sarah Maclean, who I am very

grateful for helping me with all the little details and problems I ran into, especially discussing

analysis approaches. I would like to acknowledge Reginald Barret, who provided me with

literature about wild turkeys and who to contact in the California Department of Fish and

Wildlife. From the California Department of Fish and Wildlife, I would like to thank Scott


21

Gardner and Levi Sousa for providing data and answering my questions on wild turkey

management. I would also like to Anne Murray for help with the organization and writing of my

thesis. Finally, thank you to Jenny Sholar, Kaela Shiigi and Erik Schmiett for your feedback as

well as the rest of my peers for the support through these semesters.

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California agriculture: Regional variability and subsequent impacts on management. Crop

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California Department of Parks and Recreation.

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Butcher, A. J., B. A. Collier, N. J. Silvy, and J. A. Roberson. 2014. Spatial and temporal patterns

of range expansion of white-winged doves in the USA from 1979 to 2007. Journal of

Biogeography 41:1947-1956.

Butler, M. J., W. B. Ballard, M. C. Wallace, and S. J. Demaso. 2007. Road-Based Surveys for

Estimating Wild Turkey Density in the Texas Rolling Plains. Journal of Wildlife

Management 71:1646–1653.

Dickson, J. G. 1992. The wild turkey: biology and management. Stackpole Books,

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Fuller, A. K., S. M. Spohr, D. J. Harrison, and F. A. Servello. 2013. Nest survival of wild turkeys

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scales. Wildlife Biology 19:138-146.

Gardner, S., T. Blankinship, J. Decker. 2004. Strategic Plan for Wild Turkey Management.

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Glennon, M. J., and W. F. Porter. 1999. Using satellite imagery to assess landscape-scale habitat

for wild turkeys. Wildlife Society Bulletin 27:646–653.


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Hogrefe, T. C., R. H. Yahner, and N. H. Piergallini. 1998. Depredation of artificial ground nests

in a suburban versus a rural landscape. Journal of the Pennsylvania Academy of Science

72:3–6.

Keegan, T. W., and J. A. Crawford. 1997. Brood-rearing habitat use by Rio Grande wild turkeys

in Oregon. Great Basin Naturalist 57:220–230.

Lehman, C. P., M. A. Rumble, L. D. Flake, and D. J. Thompson. 2008. Merriam's turkey nest

survival and factors affecting nest predation by mammals. The Journal of Wildlife

Management 72:1765-1774.

McKinney, M. L. 2006. Urbanization as a major cause of biotic homogenization. Biological

Conservation 127:247–260.

Spohr, S. M., F. A. Servello, D. J. Harrison, and D. W. May. 2004. Survival and reproduction of

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Sauer, J. R., and W. A. Link. 2002. A hierarchical analysis of population change with application

to Cerulean Warblers. Ecology 83: 2832–2840.

Sauer, J. R., and W. A. Link. 2011. Analysis of the North American breeding bird survey using

hierarchical models. The Auk 128:87-98.

Tefft, B. C., M. A. Gregonis, and R. E. Eriksen. 2005. Assessment of crop depredation by wild

turkeys in the United States and Ontario, Canada. Wildlife Society Bulletin 33:590-595.

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Vol. II. California Department of Fish and Game, Sacramento, California.


23

APPENDIX A

Table A1: Percent land cover at each route for a 1km buffer. These results were calculated using tabulate area

function in ArcMap.

1 km

Route Year Urban Forest Grassland Shrub Agriculture

1 1992 0.15 88.52 7.25 1.48 0.23

1 2001 8.45 61.24 1.47 24.38 0.14

1 2006 8.45 61.25 1.42 24.41 0.14

1 2011 8.45 61.22 1.47 24.39 0.14

5 1992 0.10 27.01 66.68 1.40 0.00

5 2001 6.29 40.95 29.13 17.17 0.00

5 2006 6.29 40.95 29.13 17.17 0.00

5 2011 6.29 40.95 29.13 17.17 0.00

6 1992 0.50 75.91 13.32 3.73 0.48

6 2001 6.11 72.07 9.09 6.17 0.02

6 2006 6.11 71.17 8.60 7.28 0.29

6 2011 6.11 70.47 8.70 7.86 0.29

7 1992 2.25 17.31 53.25 8.26 17.36

7 2001 9.56 4.40 50.89 11.91 18.13

7 2006 9.56 4.36 50.96 11.88 18.07

7 2011 9.56 4.81 51.04 11.94 17.47

11 1992 0.64 56.45 15.00 22.22 0.84

11 2001 6.10 51.42 4.08 34.94 0.00

11 2006 6.10 51.37 4.44 34.64 0.00

11 2011 6.10 51.10 4.56 34.78 0.00

12 1992 1.44 1.64 28.93 0.52 59.51

12 2001 6.38 0.00 13.69 0.01 67.68

12 2006 6.38 0.00 13.78 0.01 67.87

12 2011 6.38 0.00 13.78 0.01 67.87

13 1992 0.00 95.86 0.78 2.91 0.00

13 2001 2.46 86.55 0.04 10.79 0.00

13 2006 2.46 85.78 0.16 11.44 0.00

13 2011 2.46 78.46 1.86 17.03 0.00

14 1992 1.57 71.29 13.68 4.14 0.85

14 2001 14.20 63.41 6.52 6.49 0.00

14 2006 14.20 62.91 6.69 6.72 0.00

14 2011 14.20 62.78 6.74 6.81 0.00

15 1992 0.12 45.86 34.84 17.37 0.77

15 2001 4.36 6.54 18.13 68.26 1.49

15 2006 4.36 6.55 18.12 68.26 1.49

15 2011 4.36 6.55 18.12 68.26 1.49

21 1992 4.43 2.92 57.57 0.24 25.60

21 2001 9.66 0.05 28.58 0.11 52.31

21 2006 10.13 0.05 27.98 0.12 52.33

21 2011 10.32 0.03 26.66 0.09 53.50

22 1992 2.94 71.88 5.22 14.58 0.00

22 2001 6.14 59.22 10.19 19.51 0.00

22 2006 6.14 59.18 10.16 19.58 0.00

22 2011 6.14 59.15 10.07 19.70 0.00

25 1992 3.86 2.10 52.78 6.91 32.29

25 2001 17.48 1.92 53.89 4.68 19.87


24

25 2006 17.48 1.92 54.47 4.10 19.87

25 2011 17.48 1.92 54.47 4.10 19.87

27 2001 37.52 0.00 6.74 0.17 51.92

27 2006 38.09 0.00 6.48 0.16 51.95

27 2011 38.81 0.00 7.18 0.17 50.51

27 1992 40.01 2.45 4.39 6.67 42.46

28 1992 0.00 51.14 33.37 14.83 0.25

28 2001 1.86 26.51 45.34 26.23 0.00

28 2006 1.86 26.59 45.36 26.15 0.00

28 2011 1.86 26.59 45.37 26.14 0.00

32 1992 1.02 1.28 76.91 1.86 14.48

32 2001 9.99 1.25 59.13 8.94 15.25

32 2006 10.05 1.16 56.72 8.61 17.97

32 2011 10.40 1.16 55.27 8.95 18.73

34 1992 0.39 1.62 26.39 45.20 17.01

34 2001 4.91 1.58 38.02 46.80 5.58

34 2006 4.91 1.58 38.04 47.11 5.21

34 2011 4.91 1.58 37.99 46.73 5.58

38 1992 2.91 15.28 42.86 18.30 18.18

38 2001 14.18 20.17 25.50 20.20 18.25

38 2006 14.18 20.14 25.15 20.17 18.65

38 2011 14.18 20.14 25.06 20.11 18.62

51 1992 0.15 1.19 14.68 70.18 0.40

51 2001 5.45 0.05 41.20 18.24 0.01

51 2006 5.45 0.05 41.20 18.24 0.01

51 2011 5.45 0.05 41.00 18.45 0.01

52 1992 2.34 0.80 23.43 3.05 69.91

52 2001 16.12 0.56 25.94 0.63 55.81

52 2006 16.12 0.56 25.84 0.73 55.81

52 2011 16.12 0.56 25.84 0.73 55.77

53 1992 0.05 35.42 23.26 36.34 0.12

53 2001 5.04 20.04 12.51 60.21 0.00

53 2006 5.04 19.96 12.56 60.23 0.00

53 2011 5.04 19.95 12.52 60.28 0.00

54 1992 1.70 16.83 37.64 37.86 4.90

54 2001 4.81 7.76 59.30 14.71 12.91

54 2006 4.81 7.76 59.30 14.61 13.01

54 2011 4.96 7.76 59.27 14.61 12.89

57 1992 9.89 35.96 21.69 20.24 5.60

57 2001 22.68 31.30 14.13 21.10 3.85

57 2006 22.77 31.20 14.01 21.24 3.85

57 2011 22.84 31.20 13.93 21.24 3.85

71 1992 2.20 61.41 31.20 2.91 0.36

71 2001 9.34 51.96 23.21 11.79 0.79

71 2006 9.57 52.13 23.21 11.60 0.79

71 2011 9.57 52.13 23.21 11.60 0.79

75 1992 1.81 41.06 46.51 0.71 7.15

75 2001 7.36 44.66 27.87 13.85 2.91

75 2006 7.36 43.70 27.98 14.71 2.91

75 2011 7.36 43.09 28.28 15.00 2.91

77 1992 0.00 76.57 6.12 13.67 0.00

77 2001 2.76 69.11 2.03 21.53 0.00

77 2006 2.76 68.99 2.03 21.67 0.00


25

77 2011 2.76 68.85 2.08 21.76 0.00

80 1992 0.42 0.00 14.83 0.06 84.44

80 2001 6.31 0.00 8.03 0.00 84.46

80 2006 6.31 0.00 7.96 0.00 84.53

80 2011 6.31 0.00 8.76 0.05 83.61

81 1992 0.12 84.33 9.13 3.39 0.79

81 2001 9.00 78.49 3.63 4.02 0.00

81 2006 9.00 78.20 3.78 4.13 0.00

81 2011 9.00 78.10 3.80 4.22 0.00

82 1992 0.08 56.22 10.07 31.83 0.18

82 2001 6.61 40.28 2.45 48.57 0.00

82 2006 6.61 40.13 2.45 48.71 0.00

82 2011 6.61 40.13 2.45 48.71 0.00

83 1992 6.70 56.67 18.53 6.12 0.00

83 2001 14.31 51.53 10.59 12.52 0.00

83 2006 14.31 51.54 10.59 12.52 0.00

83 2011 14.31 51.54 10.59 12.52 0.00

95 1992 0.09 92.98 1.54 3.89 0.00

95 2001 1.82 89.19 0.07 8.92 0.00

95 2006 1.82 88.70 0.07 9.38 0.00

95 2011 1.82 88.66 0.07 9.42 0.00

97 1992 1.69 36.24 41.25 13.69 1.69

97 2001 8.30 11.11 30.70 46.40 1.32

97 2006 8.38 11.11 29.56 46.44 1.79

97 2011 8.38 10.85 29.73 46.48 1.79

98 1992 1.29 52.58 24.16 20.28 0.31

98 2001 8.41 34.94 29.79 25.80 0.00

98 2006 8.87 34.91 29.14 25.61 0.00

98 2011 9.21 34.61 28.87 25.43 0.00

99 1992 0.30 9.05 7.99 70.71 0.15

99 2001 4.47 0.81 10.82 72.22 0.11

99 2006 4.47 0.81 10.82 72.22 0.11

99 2011 4.47 0.81 10.82 72.22 0.11

106 1992 0.04 15.93 24.51 55.23 2.82

106 2001 5.76 7.19 33.73 47.40 0.05

106 2006 5.76 7.16 33.77 47.39 0.05

106 2011 5.76 7.15 33.86 47.31 0.05

108 1992 0.47 40.48 26.58 29.23 1.18

108 2001 5.21 42.57 14.81 36.12 0.10

108 2006 5.21 42.41 14.91 36.19 0.10

108 2011 5.21 42.41 14.92 36.18 0.10

114 1992 0.18 18.32 55.61 15.62 7.33

114 2001 5.39 23.07 30.72 27.28 11.82

114 2006 5.39 23.07 30.30 27.27 12.24

114 2011 5.39 22.94 30.56 27.13 12.24

117 1992 0.04 76.55 10.40 12.59 0.09

117 2001 1.08 43.24 17.11 38.30 0.00

117 2006 1.08 43.56 17.11 37.98 0.00

117 2011 1.08 43.58 17.11 37.96 0.00

123 1992 0.56 18.38 49.70 13.06 14.26

123 2001 6.72 18.57 39.45 24.92 6.42

123 2006 6.72 18.47 39.57 24.90 6.42

123 2011 6.72 18.45 39.46 24.88 6.42


26

136 1992 0.15 38.46 12.01 46.78 1.40

136 2001 3.61 48.43 5.24 38.53 1.71

136 2006 3.61 48.43 5.24 38.53 1.71

136 2011 3.61 48.43 5.24 38.53 1.71

138 1992 0.57 9.94 67.59 11.81 9.05

138 2001 6.24 7.34 49.79 24.35 4.99

138 2006 6.24 7.27 48.28 24.11 6.87

138 2011 6.24 7.27 48.17 24.15 6.98

153 1992 3.64 47.48 28.36 19.86 0.20

153 2001 7.81 38.40 37.46 15.73 0.15

153 2006 7.86 38.37 37.45 15.72 0.15

153 2011 8.01 38.33 37.41 15.65 0.13

157 1992 1.63 42.00 34.99 20.44 0.17

157 2001 3.88 29.56 49.90 15.33 0.09

157 2006 3.96 29.47 50.05 15.18 0.09

157 2011 3.97 29.18 50.44 14.95 0.07

159 2001 15.26 0.06 10.28 0.02 64.26

159 2006 15.52 0.06 10.26 0.02 63.92

159 2011 15.53 0.06 10.23 0.02 63.91

159 1992 15.56 4.07 35.06 3.55 18.67

160 1992 0.00 53.92 33.53 11.25 0.00

160 2001 5.54 39.18 9.50 37.40 6.38

160 2006 5.54 38.72 9.60 37.77 6.38

160 2011 5.54 38.71 9.60 37.78 6.38

161 1992 0.27 59.31 26.28 13.41 0.68

161 2001 4.27 38.04 21.42 35.92 0.27

161 2006 4.27 38.55 21.41 35.27 0.27

161 2011 4.27 37.01 22.20 36.05 0.27

164 1992 0.12 75.76 9.91 10.79 0.00

164 2001 6.85 66.41 1.55 23.32 0.00

164 2006 6.85 65.89 1.59 23.77 0.00

164 2011 6.85 65.28 1.69 24.12 0.00

165 1992 0.57 28.30 14.00 31.97 20.59

165 2001 3.91 28.96 25.83 26.60 12.03

165 2006 3.91 28.95 24.62 27.24 12.34

165 2011 3.91 28.71 24.64 27.46 12.34

172 1992 19.20 32.05 34.24 8.83 1.53

172 2001 29.96 35.41 28.25 1.03 0.15

172 2006 29.98 35.40 28.24 1.03 0.15

172 2011 29.98 35.34 28.23 1.03 0.15

173 1992 0.24 25.29 40.98 10.55 20.49

173 2001 6.33 19.13 34.76 21.35 15.11

173 2006 6.33 19.11 31.32 21.64 15.11

173 2011 6.33 18.84 33.67 22.40 15.11

178 1992 0.72 55.18 34.84 7.45 0.00

178 2001 7.00 52.01 23.06 16.22 0.00

178 2006 7.00 51.94 23.11 16.25 0.00

178 2011 7.00 51.82 23.17 16.31 0.00

179 1992 1.06 21.58 63.28 8.30 5.44

179 2001 4.97 3.50 61.96 24.57 2.28

179 2006 4.97 3.52 61.92 24.56 2.28

179 2011 4.97 3.50 61.98 24.58 2.24

181 1992 0.08 68.81 6.10 13.96 8.42


27

181 2001 2.07 71.17 4.15 15.84 4.76

181 2006 2.07 63.01 4.36 23.33 5.00

181 2011 2.07 62.90 4.36 23.43 5.00

182 1992 1.63 77.97 13.56 4.97 1.28

182 2001 8.99 71.41 8.25 10.25 0.07

182 2006 9.01 71.18 7.97 10.60 0.07

182 2011 9.01 71.30 7.99 10.46 0.07

183 1992 3.19 45.06 22.35 15.24 2.89

183 2001 8.32 23.09 13.57 44.45 0.00

183 2006 8.32 23.01 13.74 44.31 0.00

183 2011 8.32 22.96 13.76 44.34 0.00

184 1992 6.32 83.30 5.44 4.26 0.41

184 2001 12.13 78.59 5.21 3.98 0.00

184 2006 12.28 78.53 5.17 3.93 0.00

184 2011 12.36 78.38 5.25 3.91 0.00

186 1992 10.60 15.63 21.36 2.55 48.86

186 2001 29.81 10.28 31.17 8.60 18.81

186 2006 30.09 10.28 30.89 8.56 18.82

186 2011 30.24 10.25 30.82 8.54 18.72

187 1992 2.20 0.30 11.83 0.32 84.58

187 2001 9.79 0.02 6.61 0.00 81.55

187 2006 10.26 0.02 6.60 0.00 80.98

187 2011 10.26 0.02 6.62 0.00 80.92

189 1992 6.15 33.97 40.12 8.40 8.96

189 2001 19.73 33.10 28.42 8.87 7.22

189 2006 19.83 33.11 28.32 8.86 7.22

189 2011 19.83 33.11 28.32 8.86 7.22

190 1992 3.22 25.87 53.28 12.34 2.16

190 2001 6.62 8.03 77.48 5.01 0.04

190 2006 6.62 8.05 77.48 4.99 0.04

190 2011 6.62 7.97 77.63 4.93 0.04

191 1992 0.20 7.93 87.85 2.77 0.00

191 2001 2.96 3.09 88.61 3.82 0.00

191 2006 2.96 3.09 88.18 3.66 0.00

191 2011 2.96 3.09 88.66 3.64 0.00

193 1992 0.89 44.10 32.53 2.78 1.84

193 2001 6.75 37.93 22.32 14.38 0.00

193 2006 6.75 37.74 22.49 14.29 0.00

193 2011 6.75 37.73 22.49 14.29 0.00

194 1992 2.28 85.72 4.71 6.43 0.00

194 2001 9.17 83.30 2.77 3.83 0.37

194 2006 9.17 83.27 2.75 3.87 0.37

194 2011 9.17 83.20 2.75 3.94 0.37

202 1992 4.13 65.60 17.02 8.52 4.01

202 2001 10.32 42.98 9.56 20.82 15.85

202 2006 10.32 42.47 9.85 20.55 16.21

202 2011 10.32 42.41 9.96 20.51 16.11

203 1992 8.78 4.69 61.68 2.78 21.03

203 2001 31.07 2.26 47.36 1.34 12.42

203 2006 34.15 2.26 45.96 1.34 11.05

203 2011 34.56 2.26 45.35 1.34 10.82

210 1992 1.40 84.82 8.74 5.04 0.00

210 2001 6.27 80.28 3.33 9.94 0.15


28

210 2006 6.30 80.12 3.34 10.05 0.15

210 2011 6.30 79.91 3.39 10.19 0.15

319 1992 0.54 74.68 15.35 5.47 3.88

319 2001 6.38 75.36 11.87 3.25 2.65

319 2006 6.38 75.35 11.76 3.36 2.65

319 2011 6.38 75.35 11.76 3.36 2.65

341 1992 0.01 52.18 22.44 23.84 1.37

341 2001 0.15 28.62 49.06 22.08 0.00

341 2006 0.15 28.78 49.04 21.94 0.00

341 2011 0.15 28.81 49.06 21.89 0.00

357 1992 1.27 48.90 30.80 18.68 0.00

357 2001 8.54 41.69 41.92 7.50 0.00

357 2006 8.54 41.69 41.92 7.50 0.00

357 2011 8.54 41.69 41.92 7.50 0.00

406 1992 0.36 70.52 9.32 15.80 0.01

406 2001 0.58 68.29 11.06 15.48 0.14

406 2006 0.58 68.29 11.06 15.48 0.14

406 2011 0.58 65.95 11.55 17.34 0.14

409 1992 0.15 77.39 8.51 11.81 0.00

409 2001 5.85 75.13 1.68 17.09 0.00

409 2006 5.85 74.61 1.62 17.50 0.00

409 2011 5.85 74.58 1.64 17.51 0.00

411 1992 0.00 37.52 50.46 11.31 0.20

411 2001 3.76 18.48 35.92 41.34 0.00

411 2006 3.76 18.48 35.90 41.34 0.00

411 2011 3.76 18.48 35.95 41.17 0.00

415 1992 0.59 62.09 7.63 8.40 20.92

415 2001 3.69 63.35 7.08 9.91 14.12

415 2006 3.69 63.41 7.03 9.92 14.12

415 2011 3.69 53.32 7.02 19.96 14.17

416 1992 0.19 90.20 3.98 4.79 0.00

416 2001 3.75 85.37 0.29 9.78 0.00

416 2006 3.75 84.55 0.29 10.45 0.00

416 2011 3.75 84.06 0.29 10.94 0.00

419 1992 0.76 89.92 2.12 5.32 0.00

419 2001 4.45 81.60 0.44 11.79 0.00

419 2006 4.45 79.81 0.38 13.60 0.00

419 2011 4.45 78.69 0.36 14.73 0.00

420 1992 0.60 95.65 2.10 1.32 0.00

420 2001 1.10 89.98 1.97 6.92 0.00

420 2006 1.10 89.19 2.14 7.55 0.00

420 2011 1.10 87.10 2.48 9.30 0.00

422 1992 0.91 86.96 4.71 7.36 0.00

422 2001 2.29 88.76 1.38 7.48 0.00

422 2006 2.29 88.79 1.38 7.45 0.00

422 2011 2.29 88.31 1.43 7.88 0.00

431 1992 0.19 81.13 8.99 7.99 0.24

431 2001 4.79 74.81 3.96 16.24 0.16

431 2006 4.79 74.81 3.95 16.24 0.16

431 2011 4.79 74.39 3.97 16.52 0.16

900 1992 0.34 33.21 16.92 35.22 2.73

900 2001 5.07 13.31 23.37 40.04 3.40

900 2006 5.07 13.31 23.36 40.06 3.40


29

900 2011 5.07 13.18 23.40 40.16 3.40

901 1992 0.08 45.84 29.51 16.51 6.32

901 2001 4.05 29.84 18.18 33.20 13.69

901 2006 4.05 29.62 18.11 33.23 13.95

901 2011 4.05 29.58 18.02 33.53 13.78


function in ArcMap.

5 km


1 1992 0.23 38.54 16.02 4.45 0.30

1 2001 8.07 21.02 2.06 38.23 0.24

1 2006 8.06 21.01 1.97 38.35 0.24

1 2011 8.06 21.00 2.01 38.32 0.24

5 1992 0.03 7.00 66.83 3.69 0.00

5 2001 3.78 11.83 27.37 25.01 0.00

5 2006 3.78 11.83 27.39 24.99 0.00

5 2011 3.78 11.83 27.40 24.96 0.00

6 1992 0.59 28.70 24.83 8.84 1.58

6 2001 7.17 25.45 13.85 14.44 0.76

6 2006 7.08 24.79 13.21 16.29 0.94

6 2011 7.02 24.31 13.62 17.13 0.93

7 1992 1.88 3.96 59.72 9.91 16.60

7 2001 7.07 0.89 54.21 13.95 14.25

7 2006 7.06 0.88 54.26 13.92 14.20

7 2011 7.04 0.97 54.25 14.12 13.89

11 1992 0.25 18.37 20.36 37.39 0.31

11 2001 4.60 15.05 3.38 54.07 0.00

11 2006 4.60 15.03 3.41 54.09 0.00

11 2011 4.60 14.94 3.44 54.22 0.00

12 1992 1.65 0.31 33.52 1.36 57.52

12 2001 6.06 0.00 22.61 1.99 61.97

12 2006 6.10 0.00 22.60 1.99 61.88

12 2011 6.11 0.00 22.60 1.99 61.86

13 1992 0.00 43.49 3.45 8.59 0.00

13 2001 1.34 34.12 0.71 27.93 0.00

13 2006 1.30 32.72 0.72 30.83 0.00

13 2011 1.24 28.51 1.92 38.17 0.00

14 1992 0.58 21.09 8.84 3.88 0.87

14 2001 8.10 17.54 3.57 5.81 0.00

14 2006 8.08 17.37 3.63 6.09 0.00

14 2011 8.08 17.33 3.67 6.15 0.00

15 1992 0.07 11.99 45.59 27.57 1.18

15 2001 1.85 1.30 19.29 72.20 1.21

15 2006 1.84 1.30 19.31 72.18 1.20

15 2011 1.84 1.30 19.31 72.18 1.20

21 1992 2.99 0.55 64.49 0.50 20.02

21 2001 7.12 0.01 31.08 0.47 49.25

21 2006 7.32 0.01 30.22 0.47 49.45

21 2011 7.40 0.01 28.86 0.50 50.58

22 1992 3.81 27.87 10.45 23.28 0.00


30

22 2001 5.06 20.40 12.07 34.54 0.01

22 2006 5.03 20.26 11.83 35.13 0.01

22 2011 5.01 20.18 11.89 35.30 0.01

25 1992 3.90 0.38 57.34 11.58 20.89

25 2001 12.21 0.33 56.28 8.94 9.32

25 2006 12.21 0.33 56.27 8.68 9.59

25 2011 12.24 0.33 56.26 8.68 9.56

27 2001 26.08 0.00 10.34 1.09 35.61

27 2006 26.64 0.00 10.23 1.07 34.64

27 2011 27.59 0.00 10.63 1.08 32.32

27 1992 26.56 0.33 8.00 7.52 28.11

28 1992 0.02 14.46 48.99 20.60 0.11

28 2001 0.66 6.29 56.34 29.70 0.00

28 2006 0.66 6.31 56.31 29.69 0.00

28 2011 0.66 6.31 56.32 29.69 0.00

32 1992 2.85 0.23 69.03 2.33 19.44

32 2001 9.92 0.21 51.93 9.38 15.18

32 2006 10.18 0.20 49.63 9.21 17.09

32 2011 10.36 0.19 48.09 9.18 18.34

34 1992 0.11 0.32 27.71 55.23 7.19

34 2001 1.35 0.31 22.85 71.87 1.79

34 2006 1.35 0.31 22.88 71.92 1.70

34 2011 1.35 0.31 22.85 71.88 1.78

38 1992 1.65 3.04 47.40 22.16 10.66

38 2001 9.60 4.03 26.24 28.13 10.23

38 2006 9.60 4.02 26.13 28.12 10.36

38 2011 9.61 4.02 26.10 28.08 10.37

51 1992 0.05 0.23 10.37 77.38 0.10

51 2001 2.03 0.01 24.65 35.30 0.01

51 2006 2.03 0.01 24.65 35.30 0.01

51 2011 2.03 0.01 24.61 35.34 0.01

52 1992 2.58 0.23 37.42 4.29 52.19

52 2001 12.60 0.14 29.11 3.05 42.09

52 2006 12.73 0.14 28.82 2.92 41.88

52 2011 12.81 0.14 28.80 2.92 41.75

53 1992 0.05 8.42 32.12 44.76 0.35

53 2001 2.49 4.01 15.16 70.87 0.00

53 2006 2.49 3.99 15.13 70.88 0.00

53 2011 2.49 3.99 15.23 70.79 0.00

54 1992 1.22 4.11 39.24 43.13 4.35

54 2001 3.17 1.64 57.37 13.69 17.52

54 2006 3.19 1.64 57.39 13.65 17.48

54 2011 3.26 1.64 57.40 13.67 17.32

57 1992 6.38 6.59 25.88 17.15 6.66

57 2001 12.31 5.36 18.40 19.38 4.13

57 2006 12.38 5.34 18.31 19.44 4.08

57 2011 12.46 5.33 18.36 19.34 4.02

71 1992 6.04 16.74 39.60 6.50 0.42

71 2001 11.56 12.92 27.64 15.82 0.81

71 2006 11.71 12.95 27.56 15.59 0.80

71 2011 11.73 12.95 27.56 15.55 0.80

75 1992 1.43 12.89 43.95 1.66 13.15

75 2001 6.99 13.58 23.69 17.21 6.26


31

75 2006 6.95 13.20 23.62 18.13 6.22

75 2011 6.93 12.98 23.76 18.53 6.17

77 1992 0.07 28.40 12.48 27.26 0.00

77 2001 1.92 24.00 2.06 42.30 0.00

77 2006 1.91 23.86 2.18 42.51 0.00

77 2011 1.90 23.70 2.28 42.76 0.00

80 1992 3.27 0.00 21.72 0.23 70.51

80 2001 7.66 0.00 21.96 0.00 60.24

80 2006 7.72 0.00 21.82 0.00 60.22

80 2011 7.72 0.00 21.66 0.10 60.30

81 1992 0.17 31.06 12.46 6.19 0.85

81 2001 10.90 25.95 4.45 6.06 0.00

81 2006 10.83 25.68 4.63 6.55 0.00

81 2011 10.81 25.60 4.68 6.73 0.00

82 1992 0.03 16.42 16.15 46.99 1.83

82 2001 2.43 10.04 4.38 66.28 0.00

82 2006 2.43 10.00 4.49 66.27 0.00

82 2011 2.43 9.99 4.49 66.28 0.00

83 1992 9.38 17.43 13.93 6.75 0.00

83 2001 14.81 14.73 6.84 11.07 0.00

83 2006 14.86 14.72 6.78 11.04 0.00

83 2011 14.86 14.72 6.78 11.04 0.00

95 1992 0.05 42.06 3.36 9.05 0.00

95 2001 2.11 36.28 0.87 21.29 0.00

95 2006 1.99 34.00 0.79 26.16 0.00

95 2011 1.99 33.95 0.79 26.28 0.00

97 1992 4.62 9.21 45.27 20.88 3.48

97 2001 8.65 2.20 26.36 49.08 1.76

97 2006 8.73 2.20 26.22 48.85 1.84

97 2011 8.70 2.12 26.34 48.93 1.81

98 1992 3.73 14.85 29.74 19.24 2.79

98 2001 9.72 8.81 33.98 19.23 0.18

98 2006 12.01 8.56 29.87 18.42 0.14

98 2011 12.78 8.40 28.94 17.93 0.14

99 1992 0.13 1.62 4.05 39.36 0.05

99 2001 1.86 0.14 5.13 40.42 0.02

99 2006 1.86 0.14 5.13 40.42 0.02

99 2011 1.86 0.14 5.13 40.41 0.02

106 1992 0.08 3.35 22.31 67.48 0.80

106 2001 3.38 1.42 32.99 54.78 0.03

106 2006 3.36 1.40 32.86 54.56 0.02

106 2011 3.38 1.41 33.06 54.81 0.02

108 1992 1.32 10.70 34.60 37.27 0.77

108 2001 4.53 10.95 19.58 48.66 0.30

108 2006 4.50 10.83 20.01 48.54 0.30

108 2011 4.50 10.83 19.92 48.63 0.30

114 1992 0.82 4.68 54.87 23.68 6.81

114 2001 5.26 5.80 25.13 41.75 10.53

114 2006 5.26 5.79 25.03 41.73 10.67

114 2011 5.26 5.76 25.12 41.70 10.67

117 1992 0.17 27.19 18.47 25.55 0.28

117 2001 0.90 12.11 22.79 48.96 0.00

117 2006 0.90 12.19 22.62 48.84 0.00


32

117 2011 0.90 12.18 22.60 48.88 0.00

123 1992 0.74 4.11 59.03 14.70 10.51

123 2001 6.23 4.00 45.96 25.19 4.43

123 2006 6.23 3.98 45.59 24.91 4.43

123 2011 6.22 3.97 46.05 25.08 4.52

136 1992 0.27 9.50 14.86 63.59 0.48

136 2001 2.70 13.11 8.12 58.28 0.50

136 2006 2.70 13.11 8.09 58.31 0.50

136 2011 2.70 13.09 8.22 58.21 0.50

138 1992 0.69 2.18 67.62 19.14 5.75

138 2001 4.28 1.64 49.24 33.12 2.18

138 2006 4.28 1.62 47.75 33.00 3.83

138 2011 4.28 1.62 47.54 33.02 3.97

153 1992 5.35 14.64 32.84 23.24 3.37

153 2001 9.38 10.44 41.55 18.32 0.07

153 2006 9.55 10.41 41.42 18.17 0.07

153 2011 9.73 10.36 41.30 18.03 0.07

157 1992 2.52 10.62 44.67 24.64 0.89

157 2001 5.52 6.74 55.50 14.35 1.37

157 2006 5.56 6.70 55.66 14.09 1.34

157 2011 5.55 6.62 55.77 13.96 1.34

159 2001 12.40 0.01 13.21 0.15 55.60

159 2006 12.81 0.01 13.12 0.14 54.82

159 2011 12.83 0.01 13.10 0.14 54.68

159 1992 10.60 0.50 29.33 2.52 35.48

160 1992 0.34 15.99 49.57 16.79 0.00

160 2001 6.06 10.13 14.02 43.57 8.88

160 2006 6.06 10.01 14.11 43.66 8.92

160 2011 6.05 9.99 14.17 43.65 8.87

161 1992 0.14 18.49 41.88 19.71 0.83

161 2001 2.01 9.04 38.56 39.20 0.10

161 2006 2.01 9.15 38.59 38.94 0.11

161 2011 1.98 8.68 38.77 39.76 0.11

164 1992 0.06 28.90 19.44 20.09 0.04

164 2001 4.49 22.64 1.54 43.27 0.00

164 2006 4.40 22.01 1.85 44.37 0.00

164 2011 4.36 21.65 2.08 44.69 0.00

165 1992 0.29 7.64 9.56 28.08 17.94

165 2001 2.08 7.76 14.81 23.03 13.23

165 2006 2.08 7.75 14.22 23.30 13.61

165 2011 2.08 7.69 14.19 23.43 13.67

172 1992 25.71 5.62 28.51 6.56 0.45

172 2001 32.49 5.95 20.45 1.17 0.05

172 2006 32.49 5.94 20.42 1.20 0.05

172 2011 32.56 5.93 20.30 1.20 0.05

173 1992 0.43 5.38 54.82 18.26 10.96

173 2001 5.82 3.79 36.56 37.06 6.10

173 2006 5.82 3.78 33.75 36.90 6.09

173 2011 5.81 3.73 36.06 37.51 6.09

178 1992 0.76 18.88 45.85 12.59 0.00

178 2001 6.55 16.36 27.85 24.30 0.00

178 2006 6.55 16.32 27.97 24.28 0.00

178 2011 6.55 16.29 28.01 24.30 0.00


33

179 1992 3.15 4.61 68.58 10.28 4.90

179 2001 5.73 0.63 58.18 26.65 1.72

179 2006 5.80 0.64 58.07 26.63 1.72

179 2011 5.83 0.63 58.06 26.63 1.68

181 1992 0.22 24.95 9.50 23.89 12.53

181 2001 1.98 27.45 6.24 25.84 7.51

181 2006 1.75 21.51 5.49 39.86 6.72

181 2011 1.69 20.72 5.40 41.92 6.52

182 1992 0.65 26.89 22.19 7.50 1.74

182 2001 7.91 22.14 11.09 16.43 0.40

182 2006 7.89 21.99 11.01 16.80 0.40

182 2011 7.89 22.02 11.01 16.77 0.40

183 1992 2.47 12.81 24.03 19.15 2.17

183 2001 7.32 5.04 9.29 48.01 0.07

183 2006 7.32 5.02 9.45 47.86 0.07

183 2011 7.32 5.01 9.49 47.87 0.07

184 1992 13.37 26.92 10.09 8.20 0.36

184 2001 19.21 22.35 8.90 7.77 0.00

184 2006 19.29 22.32 8.83 7.75 0.00

184 2011 19.42 22.21 8.85 7.70 0.00

186 1992 11.86 2.89 24.35 2.64 43.13

186 2001 23.26 1.63 30.13 8.34 11.51

186 2006 23.50 1.63 29.65 8.26 11.59

186 2011 23.56 1.62 29.61 8.24 11.55

187 1992 2.09 0.05 10.00 0.21 85.23

187 2001 7.33 0.00 7.05 0.00 77.73

187 2006 7.63 0.00 7.02 0.00 77.09

187 2011 7.72 0.00 7.03 0.00 76.89

189 1992 14.37 7.75 38.40 9.57 6.75

189 2001 23.66 7.01 24.43 10.15 3.38

189 2006 23.69 7.01 24.39 10.14 3.38

189 2011 23.69 7.01 24.38 10.14 3.38

190 1992 2.09 5.64 62.02 17.81 0.64

190 2001 3.25 1.56 76.92 9.10 0.31

190 2006 3.24 1.56 76.58 8.84 0.31

190 2011 3.24 1.54 76.63 8.82 0.31

191 1992 0.16 1.67 91.44 4.19 0.00

191 2001 1.78 0.61 89.29 4.93 0.13

191 2006 1.78 0.61 88.94 4.82 0.18

191 2011 1.78 0.61 89.29 4.81 0.18

193 1992 0.52 10.38 30.55 2.50 0.80

193 2001 4.18 8.57 18.75 15.05 0.00

193 2006 4.18 8.53 18.84 15.01 0.00

193 2011 4.18 8.53 18.85 15.01 0.00

194 1992 4.65 30.63 13.15 12.99 0.52

194 2001 14.00 27.57 7.96 5.68 1.63

194 2006 14.00 27.55 3.71 9.97 1.63

194 2011 14.00 27.53 3.72 9.99 1.63

202 1992 3.61 17.38 33.25 13.41 8.84

202 2001 8.81 8.49 15.37 31.03 17.80

202 2006 8.80 8.38 14.95 30.68 18.70

202 2011 8.80 8.36 14.99 30.62 18.70

203 1992 7.12 0.83 62.38 3.72 17.21


34

203 2001 18.29 0.34 47.75 1.73 12.05

203 2006 19.67 0.34 46.34 1.74 10.81

203 2011 19.89 0.34 46.05 1.74 10.64

210 1992 4.45 29.82 21.30 9.88 0.21

210 2001 11.94 22.96 7.42 19.37 3.33

210 2006 12.01 22.79 7.21 19.68 3.43

210 2011 12.03 22.68 7.25 19.82 3.42

319 1992 0.67 22.75 28.97 10.50 4.73

319 2001 6.89 24.31 18.74 6.06 4.17

319 2006 6.89 24.30 18.60 6.21 4.17

319 2011 6.89 24.30 18.60 6.21 4.17

341 1992 0.00 15.34 36.48 31.84 0.88

341 2001 0.21 7.59 62.50 21.89 0.00

341 2006 0.21 7.63 62.40 21.91 0.00

341 2011 0.21 7.64 62.40 21.90 0.00

357 1992 0.33 39.47 10.12 10.24 0.00

357 2001 2.27 33.53 16.98 11.39 0.00

357 2006 2.28 33.46 17.54 10.95 0.00

357 2011 2.28 33.45 17.86 10.66 0.00

406 1992 0.28 21.99 13.18 32.68 0.06

406 2001 0.68 20.06 20.84 28.08 0.23

406 2006 0.68 20.06 20.83 28.07 0.23

406 2011 0.66 18.93 21.65 29.77 0.22

409 1992 0.09 32.87 13.01 18.16 0.00

409 2001 5.66 28.81 1.86 27.41 0.00

409 2006 5.67 28.65 1.85 27.66 0.00

409 2011 5.65 28.51 1.90 27.95 0.00

411 1992 0.00 9.48 60.92 17.97 0.67

411 2001 1.99 3.91 35.29 52.58 0.13

411 2006 1.99 3.91 35.26 52.61 0.13

411 2011 1.99 3.90 35.29 52.45 0.13

415 1992 0.34 32.36 8.41 14.51 10.38

415 2001 2.74 32.17 3.94 19.73 6.00

415 2006 2.74 32.24 3.90 19.65 6.01

415 2011 2.52 24.88 3.67 35.64 5.51

416 1992 0.12 39.45 5.16 11.12 0.31

416 2001 2.02 35.89 0.39 20.57 0.00

416 2006 2.03 35.77 0.35 20.49 0.00

416 2011 2.02 35.37 0.38 21.31 0.01

419 1992 1.96 38.52 5.10 12.26 0.00

419 2001 5.51 30.44 0.73 26.51 0.00

419 2006 5.37 29.05 0.68 29.64 0.00

419 2011 5.36 28.57 0.68 30.64 0.00

420 1992 0.86 41.78 5.33 4.75 1.40

420 2001 2.75 35.86 3.82 18.57 0.00

420 2006 2.73 35.30 3.96 19.58 0.00

420 2011 2.72 34.33 4.16 21.35 0.00

422 1992 0.91 35.22 11.12 15.63 0.00

422 2001 2.00 37.05 4.26 16.52 0.00

422 2006 2.00 37.03 4.28 16.53 0.00

422 2011 1.98 36.57 4.34 17.43 0.00

431 1992 0.55 32.37 15.35 16.82 0.10

431 2001 6.58 25.75 3.73 31.26 0.14


35

431 2006 6.57 25.71 3.76 31.26 0.14

431 2011 6.52 25.38 3.78 31.96 0.14

900 1992 0.29 8.81 33.06 42.91 2.24

900 2001 2.24 2.91 29.35 53.05 1.31

900 2006 2.24 2.90 29.26 53.15 1.33

900 2011 2.23 2.87 29.24 53.27 1.32

901 1992 0.26 14.79 40.24 20.54 8.35

901 2001 2.78 7.80 20.13 42.58 13.21

901 2006 2.77 7.72 20.51 43.03 13.01

901 2011 2.77 7.70 20.52 43.17 12.96


function in ArcMap.

10 km


1 1992 0.15 88.79 7.56 2.77 0.11

1 2001 4.30 72.74 1.31 20.60 0.08

1 2006 4.30 72.65 1.27 20.69 0.08

1 2011 4.30 72.51 1.38 20.72 0.08

5 1992 0.01 46.84 31.09 3.27 0.00

5 2001 1.36 54.40 11.75 13.59 0.00

5 2006 1.36 54.30 11.79 13.64 0.00

5 2011 1.36 54.23 11.82 13.66 0.00

6 1992 0.21 78.65 13.94 4.72 0.37

6 2001 3.03 76.94 8.31 9.15 0.17

6 2006 3.03 75.86 8.27 10.12 0.22

6 2011 3.03 75.23 8.50 10.55 0.22

7 1992 2.15 26.26 49.16 9.57 11.30

7 2001 7.18 6.14 53.53 18.35 10.58

7 2006 7.26 6.10 53.52 18.34 10.54

7 2011 7.27 6.01 53.69 18.46 10.34

11 1992 0.08 63.55 13.28 21.64 0.13

11 2001 2.94 56.83 3.33 35.71 0.11

11 2006 2.94 56.15 3.62 36.04 0.11

11 2011 2.94 56.09 3.60 36.16 0.11

12 1992 1.15 3.70 18.36 0.86 59.14

12 2001 5.44 0.81 13.47 1.73 62.44

12 2006 5.55 0.81 13.48 1.73 62.34

12 2011 5.55 0.81 13.48 1.73 62.31

13 1992 0.27 91.71 1.99 4.90 0.00

13 2001 0.99 83.23 0.46 14.66 0.00

13 2006 0.99 81.02 0.48 16.80 0.00

13 2011 0.99 78.11 0.98 19.19 0.00

14 1992 0.17 57.16 6.84 2.54 0.35

14 2001 3.48 54.73 2.99 5.44 0.00

14 2006 3.48 54.56 3.02 5.55 0.00

14 2011 3.48 54.53 3.04 5.57 0.00

15 1992 0.07 42.93 35.99 18.97 1.50

15 2001 1.36 7.40 23.30 65.73 1.55

15 2006 1.36 7.37 23.33 65.73 1.55

15 2011 1.36 7.37 23.33 65.73 1.55


36

21 1992 7.27 1.62 57.58 0.79 26.95

21 2001 10.65 0.05 34.75 0.84 48.61

21 2006 11.16 0.05 33.71 0.88 49.01

21 2011 11.50 0.05 32.51 0.89 49.81

22 1992 1.36 80.72 5.73 10.74 0.00

22 2001 2.18 72.23 6.40 17.61 0.00

22 2006 2.18 71.87 6.44 17.92 0.00

22 2011 2.18 71.67 6.50 18.10 0.00

25 1992 2.09 11.66 50.73 14.68 19.45

25 2001 9.28 10.88 52.42 14.94 10.98

25 2006 9.29 10.88 52.32 14.78 11.09

25 2011 9.31 10.88 52.32 14.78 11.08

27 2001 28.43 0.25 20.24 2.21 47.37

27 2006 29.41 0.25 20.14 2.20 46.50

27 2011 30.77 0.25 20.51 2.22 44.73

27 1992 29.15 2.45 19.01 11.73 33.67

28 1992 0.18 46.27 35.39 14.62 1.62

28 2001 1.11 27.36 47.95 20.41 2.68

28 2006 1.11 27.45 47.89 20.37 2.68

28 2011 1.12 27.43 47.91 20.39 2.64

32 1992 1.79 4.07 67.73 5.31 18.07

32 2001 8.34 3.18 59.04 14.83 10.94

32 2006 8.48 3.16 57.18 14.72 12.75

32 2011 8.58 3.15 56.30 14.71 13.45

34 1992 0.10 22.12 22.71 44.62 3.62

34 2001 0.94 19.75 14.66 62.59 0.72

34 2006 0.94 19.72 14.58 62.59 0.68

34 2011 0.94 18.71 14.56 63.55 0.72

38 1992 0.97 17.63 37.45 21.54 6.47

38 2001 6.29 21.48 20.78 29.59 6.96

38 2006 6.31 21.42 20.83 29.47 7.05

38 2011 6.32 21.40 20.83 29.45 7.04

51 1992 0.03 6.98 8.26 73.17 0.09

51 2001 1.80 2.54 16.45 43.30 0.02

51 2006 1.80 2.54 16.45 43.30 0.02

51 2011 1.80 2.54 16.46 43.29 0.02

52 1992 3.83 23.67 47.38 4.75 20.17

52 2001 12.84 20.38 35.37 7.10 23.79

52 2006 13.05 20.39 35.36 7.08 23.61

52 2011 13.16 20.39 35.33 7.08 23.52

53 1992 0.08 29.91 29.63 31.86 2.47

53 2001 2.38 14.12 20.62 60.41 0.89

53 2006 2.38 14.11 20.61 60.42 0.89

53 2011 2.38 14.08 20.65 60.41 0.89

54 1992 0.90 32.32 27.00 31.92 5.26

54 2001 2.99 17.34 43.96 11.93 21.63

54 2006 3.01 17.34 43.97 11.92 21.62

54 2011 3.04 17.30 44.08 11.95 21.48

57 1992 4.74 14.23 25.52 11.27 4.56

57 2001 10.20 12.73 19.90 13.89 3.42

57 2006 10.23 12.70 19.79 14.02 3.39

57 2011 10.33 12.67 19.81 13.96 3.37

71 1992 7.75 38.90 29.31 4.24 1.66


37

71 2001 12.86 33.40 22.45 11.20 1.20

71 2006 13.12 33.43 22.29 11.08 1.20

71 2011 13.15 33.44 22.24 11.06 1.20

75 1992 1.00 47.47 19.14 1.14 11.78

75 2001 5.11 48.00 10.59 8.43 7.85

75 2006 5.12 47.23 10.64 9.06 7.87

75 2011 5.12 46.70 10.80 9.39 7.85

77 1992 0.26 71.81 8.51 16.70 0.00

77 2001 1.32 65.87 2.16 27.16 0.20

77 2006 1.32 65.72 2.19 27.29 0.20

77 2011 1.32 65.22 2.31 27.67 0.20

80 1992 1.76 1.23 39.64 0.42 52.60

80 2001 5.92 0.04 37.84 0.04 50.26

80 2006 5.97 0.05 37.77 0.05 50.23

80 2011 5.97 0.05 37.70 0.18 50.20

81 1992 0.11 72.36 6.78 3.15 0.22

81 2001 5.88 68.77 3.13 3.28 0.10

81 2006 5.88 68.32 3.21 3.62 0.10

81 2011 5.88 68.24 3.25 3.67 0.10

82 1992 0.01 54.63 10.51 31.21 1.64

82 2001 1.14 39.85 3.24 51.05 0.31

82 2006 1.14 39.75 3.28 51.12 0.32

82 2011 1.14 39.72 3.28 51.16 0.32

83 1992 11.22 15.21 9.41 3.85 0.02

83 2001 16.75 12.29 3.83 7.73 0.01

83 2006 16.78 12.26 3.82 7.73 0.00

83 2011 16.81 12.26 3.82 7.73 0.00

95 1992 0.06 90.32 2.94 5.37 0.00

95 2001 1.33 85.00 0.38 12.69 0.00

95 2006 1.33 81.06 0.36 16.51 0.00

95 2011 1.33 80.75 0.35 16.82 0.00

97 1992 4.30 47.01 30.12 15.12 1.91

97 2001 9.32 26.17 22.76 39.60 1.07

97 2006 9.47 26.01 22.70 39.56 1.11

97 2011 9.52 25.03 23.08 40.01 1.10

98 1992 7.20 40.17 28.08 13.72 4.92

98 2001 16.48 31.06 32.94 13.08 0.81

98 2006 18.73 30.91 30.29 12.89 0.76

98 2011 19.68 30.63 29.75 12.80 0.68

99 1992 0.11 3.23 1.75 17.68 0.03

99 2001 0.96 0.27 2.30 19.19 0.01

99 2006 0.96 0.27 2.30 19.19 0.01

99 2011 0.96 0.27 2.30 19.19 0.01

106 1992 0.42 22.99 13.24 61.00 0.72

106 2001 3.40 14.26 22.40 53.03 0.03

106 2006 3.47 14.24 22.31 53.05 0.03

106 2011 3.47 14.23 22.39 53.01 0.03

108 1992 0.90 46.61 25.33 24.44 0.41

108 2001 2.81 44.85 13.92 36.99 0.24

108 2006 2.81 44.62 14.06 37.08 0.24

108 2011 2.81 44.45 14.10 37.22 0.24

114 1992 0.85 21.13 44.91 24.51 5.75

114 2001 4.83 20.10 17.17 46.55 9.69


38

114 2006 4.83 20.06 17.12 46.53 9.83

114 2011 4.83 20.04 17.16 46.50 9.83

117 1992 0.05 71.25 11.42 14.23 0.15

117 2001 0.69 48.80 15.71 28.53 0.01

117 2006 0.69 48.84 15.46 28.48 0.01

117 2011 0.69 48.71 15.46 28.60 0.01

123 1992 1.47 22.75 51.75 12.33 6.65

123 2001 7.41 22.05 41.10 20.86 3.01

123 2006 7.41 21.96 40.91 20.73 3.11

123 2011 7.42 21.82 41.22 20.81 3.14

136 1992 0.74 34.65 13.44 48.64 0.81

136 2001 2.77 45.42 6.04 42.56 1.26

136 2006 2.77 45.42 6.05 42.55 1.26

136 2011 2.77 45.38 6.10 42.53 1.26

138 1992 1.61 19.10 54.27 18.97 4.22

138 2001 5.65 20.22 35.33 30.53 4.17

138 2006 5.66 20.15 34.72 30.43 4.96

138 2011 5.68 20.14 34.55 30.39 5.11

153 1992 3.36 49.63 30.84 13.11 2.74

153 2001 6.53 40.70 38.95 13.42 0.04

153 2006 6.58 40.67 38.75 13.36 0.04

153 2011 6.70 40.41 39.00 13.25 0.05

157 1992 1.85 44.68 33.59 15.22 1.27

157 2001 4.93 34.25 43.77 11.18 1.20

157 2006 4.97 34.05 43.87 11.02 1.19

157 2011 4.98 33.87 44.01 10.92 1.19

159 2001 14.86 3.44 21.23 0.19 53.16

159 2006 15.30 3.42 21.25 0.18 52.73

159 2011 15.34 3.41 21.23 0.19 52.63

159 1992 10.34 7.20 27.39 1.85 46.68

160 1992 0.15 58.10 30.29 10.86 0.05

160 2001 4.17 45.83 12.38 32.80 3.99

160 2006 4.17 45.68 12.48 32.83 4.01

160 2011 4.17 45.63 12.50 32.85 3.99

161 1992 0.10 57.61 28.25 12.97 0.93

161 2001 1.49 31.13 37.88 29.30 0.14

161 2006 1.49 30.70 37.87 29.72 0.14

161 2011 1.49 29.14 38.23 30.92 0.14

164 1992 0.21 77.41 8.88 11.72 0.01

164 2001 2.72 71.33 1.48 23.76 0.00

164 2006 2.72 70.50 1.61 24.44 0.00

164 2011 2.72 69.71 1.69 25.04 0.00

165 1992 0.15 31.75 6.27 20.10 9.47

165 2001 1.26 34.00 8.45 16.72 8.35

165 2006 1.26 33.77 8.22 16.83 8.44

165 2011 1.26 33.72 8.20 16.87 8.45

172 1992 41.70 19.69 22.74 4.93 0.24

172 2001 51.10 22.20 15.51 1.21 0.03

172 2006 51.14 22.19 15.46 1.22 0.03

172 2011 51.22 22.17 15.39 1.22 0.03

173 1992 0.39 29.59 39.61 17.96 4.41

173 2001 4.32 24.05 25.18 36.78 2.44

173 2006 4.32 23.97 23.86 36.66 2.43


39

173 2011 4.32 23.87 25.01 36.89 2.43

178 1992 0.23 69.30 22.31 7.27 0.00

178 2001 3.06 67.19 14.00 14.79 0.00

178 2006 3.06 67.10 14.03 14.85 0.00

178 2011 3.06 67.08 14.05 14.85 0.00

179 1992 1.68 28.91 54.10 8.57 5.89

179 2001 5.17 9.88 51.40 25.61 6.25

179 2006 5.23 9.85 51.34 25.60 6.24

179 2011 5.24 9.83 51.56 25.59 6.04

181 1992 0.12 76.54 4.70 13.04 4.36

181 2001 0.71 79.47 2.60 13.63 2.48

181 2006 0.71 74.16 2.50 19.01 2.50

181 2011 0.71 70.64 2.58 22.44 2.51

182 1992 0.20 70.42 8.43 3.29 0.78

182 2001 4.08 67.02 4.60 7.03 0.13

182 2006 4.08 66.61 4.59 7.41 0.13

182 2011 4.08 66.57 4.60 7.44 0.13

183 1992 1.90 46.35 17.31 15.79 1.62

183 2001 7.35 20.33 7.63 46.15 1.87

183 2006 7.35 20.25 7.71 46.04 1.96

183 2011 7.35 20.23 7.71 46.07 1.96

184 1992 4.34 82.00 5.98 5.30 0.21

184 2001 8.23 77.21 6.32 6.62 0.00

184 2006 8.26 77.05 6.40 6.67 0.00

184 2011 8.30 76.30 6.80 6.96 0.01

186 1992 9.51 21.64 30.57 3.02 34.24

186 2001 22.08 15.42 38.42 11.59 10.86

186 2006 22.46 15.33 37.95 11.41 11.17

186 2011 22.53 15.31 37.93 11.40 11.15

187 1992 4.71 0.58 11.60 0.23 82.23

187 2001 10.81 0.01 9.26 0.00 78.32

187 2006 11.07 0.01 9.16 0.00 78.05

187 2011 11.21 0.01 9.16 0.01 77.88

189 1992 17.52 39.88 28.13 9.03 4.46

189 2001 27.61 38.81 17.73 12.39 2.06

189 2006 27.76 38.79 17.67 12.41 1.97

189 2011 27.80 38.78 17.66 12.41 1.94

190 1992 1.10 27.80 50.64 14.09 0.57

190 2001 2.45 12.53 66.08 11.37 0.16

190 2006 2.45 12.46 65.57 10.92 0.17

190 2011 2.45 12.42 65.62 10.91 0.17

191 1992 0.95 21.03 72.24 4.85 0.19

191 2001 2.40 14.71 74.03 6.08 1.57

191 2006 2.40 14.59 73.85 6.01 1.65

191 2011 2.40 14.50 74.06 6.01 1.66

193 1992 0.29 34.47 17.36 1.71 0.25

193 2001 2.79 31.01 10.95 8.92 0.13

193 2006 2.79 30.85 11.03 8.98 0.13

193 2011 2.79 30.84 11.05 8.97 0.13

194 1992 4.85 67.23 7.90 7.34 0.25

194 2001 10.45 68.70 4.49 3.40 0.89

194 2006 10.47 68.68 2.13 5.77 0.88

194 2011 10.47 68.66 2.10 5.82 0.88


40

202 1992 2.13 61.53 20.36 10.18 4.87

202 2001 6.51 38.08 13.14 31.37 9.88

202 2006 6.53 37.88 13.03 31.20 10.26

202 2011 6.53 37.84 13.06 31.17 10.27

203 1992 8.83 10.69 57.37 3.79 17.07

203 2001 19.45 7.74 52.70 2.75 14.20

203 2006 20.93 7.75 52.37 2.82 13.03

203 2011 21.22 7.75 52.18 2.82 12.90

210 1992 1.77 70.97 19.04 6.80 0.21

210 2001 7.05 56.89 8.80 22.36 3.76

210 2006 7.10 56.72 8.75 22.46 3.82

210 2011 7.12 56.60 8.77 22.54 3.82

319 1992 0.42 27.53 24.66 6.64 3.27

319 2001 4.15 36.20 16.10 3.58 2.60

319 2006 4.15 36.19 15.97 3.72 2.60

319 2011 4.15 36.19 15.97 3.72 2.60

341 1992 0.01 52.61 26.86 18.18 2.00

341 2001 0.36 40.11 41.44 17.47 0.36

341 2006 0.36 40.09 41.41 17.50 0.34

341 2011 0.36 40.08 41.41 17.50 0.34

357 1992 2.28 76.17 9.24 12.28 0.01

357 2001 3.76 63.73 18.04 14.44 0.04

357 2006 3.76 55.30 18.63 22.28 0.04

357 2011 3.76 56.44 18.63 21.14 0.04

406 1992 0.10 60.14 5.30 26.63 0.83

406 2001 0.64 54.67 14.86 21.75 0.56

406 2006 0.64 54.64 14.86 21.78 0.56

406 2011 0.64 53.20 15.31 22.76 0.56

409 1992 0.06 85.14 5.35 7.56 0.08

409 2001 3.72 81.02 0.96 13.25 0.00

409 2006 3.72 79.31 1.80 14.01 0.01

409 2011 3.72 78.94 1.79 14.38 0.01

411 1992 0.00 46.25 36.18 15.93 0.36

411 2001 1.65 25.01 21.96 50.79 0.02

411 2006 1.65 25.00 21.94 50.81 0.02

411 2011 1.65 24.87 22.01 50.73 0.02

415 1992 0.12 85.30 5.94 6.94 0.57

415 2001 0.98 86.05 1.11 10.04 0.17

415 2006 0.98 85.87 1.10 9.99 0.17

415 2011 0.98 77.34 1.29 18.32 0.17

416 1992 0.11 90.31 2.40 5.35 0.48

416 2001 1.02 86.86 0.62 10.31 0.14

416 2006 1.02 86.81 0.58 10.32 0.14

416 2011 1.02 86.31 0.61 10.80 0.14

419 1992 0.93 86.54 3.30 7.43 0.00

419 2001 2.77 71.96 1.40 22.97 0.00

419 2006 2.77 69.09 1.43 25.69 0.00

419 2011 2.77 68.73 1.42 26.07 0.00

420 1992 1.03 91.11 2.51 2.66 1.10

420 2001 2.61 83.92 2.33 10.73 0.00

420 2006 2.61 82.91 2.42 11.17 0.00

420 2011 2.61 81.44 2.52 12.54 0.00

422 1992 0.42 78.44 8.25 12.43 0.00


41

422 2001 0.93 82.13 3.78 12.27 0.00

422 2006 0.93 82.08 3.82 12.28 0.00

422 2011 0.93 81.54 3.86 12.79 0.00

431 1992 0.16 81.30 8.02 9.61 0.12

431 2001 3.50 73.22 2.13 20.89 0.08

431 2006 3.50 73.04 2.16 20.97 0.08

431 2011 3.50 72.61 2.24 21.22 0.08

900 1992 0.13 53.87 21.45 21.77 1.45

900 2001 1.34 33.87 20.07 40.03 2.08

900 2006 1.34 33.19 20.33 40.41 2.12

900 2011 1.34 32.55 20.39 41.00 2.11

901 1992 0.16 59.01 22.88 10.20 5.62

901 2001 1.93 44.03 13.79 28.64 8.89

901 2006 1.93 43.46 14.03 29.30 8.87

901 2011 1.93 43.34 14.11 29.38 8.84

APPENDIX B

Table B1: Population change at each individual route. These results were calculated using a GLM for each route.

Route Latitude Longitude Estimate Standard Error P-value Significance

416 39.94937 -121.038 1.0886 0.5887 0.0644

12 39.27543 -121.734 0.881 0.6503 0.175

210 39.16346 -123.227 0.6697 0.4459 0.133

172 37.81835 -122.146 0.39204 0.0583 1.87E-11 ***

422 37.46188 -119.794 0.3064 0.105 0.00351 **

415 40.04023 -120.996 0.2834 0.1393 0.042 *

202 38.37951 -122.514 0.27467 0.08571 0.00135 **

34 35.73357 -118.078 0.24659 0.08571 0.00402 **

181 40.20176 -120.624 0.2237 0.04547 8.63E-07 ***

13 39.12397 -120.760 0.212 0.1787 0.235

357 38.35393 -120.680 0.20584 0.08126 0.0113 *

194 37.21153 -122.157 0.19729 0.03155 3.99E-10 ***

138 35.31838 -120.508 0.18429 0.03321 2.88E-08 ***

15 38.97873 -122.240 0.18139 0.03442 1.37E-07 ***

80 39.76334 -122.239 0.17938 0.08392 0.0326 *

114 34.73757 -120.235 0.16884 0.02712 4.76E-10 ***

173 35.92226 -121.064 0.16613 0.05157 0.00128 **

14 38.80359 -123.585 0.16292 0.03754 1.42E-05 ***

190 37.98349 -120.645 0.15338 0.02342 5.78E-01 ***

83 37.98512 -122.591 0.14388 0.01753 2.26E-16 ***

51 32.90395 -116.233 0.14257 0.07376 0.0533

75 40.52807 -124.164 0.1412 0.1082 0.192

106 33.19909 -116.786 0.1323 0.01307 <2e-16 ***


42

186 38.54987 -122.719 0.12701 0.02487 3.27E-07 ***

183 39.18442 -122.709 0.12407 0.06137 0.0432 *

81 39.16123 -123.582 0.12135 0.01785 1.06E-11 ***

160 39.56649 -123.398 0.12128 0.02138 1.40E-08 ***

184 39.06869 -121.010 0.11862 0.02577 4.16E-06 ***

189 37.25217 -121.677 0.11788 0.01927 9.58E-10 ***

71 38.04535 -122.867 0.1157 0.0292 7.42E-05 ***

153 38.76876 -120.925 0.1089 0.007789 <2e-16 ***

178 40.17327 -123.604 0.10574 0.02104 5.01E-07 ***

179 40.02796 -122.639 0.10572 0.01459 4.23E-13 ***

182 39.68784 -123.485 0.09925 0.01116 <2e-16 ***

159 39.49790 -121.609 0.09821 0.06706 0.143

193 38.34077 -123.048 0.09727 0.0229 2.15E-05 ***

117 36.46643 -118.655 0.09492 0.06132 0.122

5 40.24319 -124.123 0.09425 0.01644 9.96E-09 ***

54 36.63160 -119.305 0.09215 0.05083 0.0698

25 36.76236 -121.175 0.09107 0.04062 0.0249 *

53 36.22517 -120.519 0.0872 0.02509 0.00051 ***

191 37.12012 -119.701 0.08258 0.04579 0.0713

98 38.72022 -121.036 0.07863 0.009435 <2e-16 ***

108 36.45308 -121.696 0.06964 0.01332 1.70E-07 ***

161 40.33571 -121.916 0.0684 0.02314 0.00312 **

6 40.41509 -123.962 0.06403 0.03081 0.0377 *

95 38.55626 -120.291 0.06008 0.06699 0.37

341 35.89089 -118.850 0.05952 0.07675 0.438

123 35.63783 -120.855 0.03829 0.01361 0.0049 **

900 41.90242 -122.563 0.02538 0.04168 0.543

420 38.79958 -120.724 0.01042 0.10213 0.919

97 40.51649 -122.382 0.009377 0.037764 0.804

901 41.52092 -122.520 0.001347 0.027552 0.961

99 33.36574 -118.374 -0.1031 0.1129 0.361

411 40.18032 -122.860 -0.1951 0.1696 0.25

419 38.73968 -120.573 -0.2315 0.1984 0.243

409 40.80313 -123.483 -0.8109 0.3727 0.0296 *

predictors of wild turkey (meleagris gallopavo) population

Documents