Thesis Defense

SNAKE COMMUNITY ECOLOGY SNAKE COMMUNITY ECOLOGY WITHIN THE FLINT HILLS WITHIN THE FLINT HILLS NATIONALNATIONALWILDLIFE REFUGEWILDLIFE REFUGE

By: Bryan MaherBy: Bryan Maher

Emporia State University, Department of Biological Sciences,Emporia State University, Department of Biological Sciences,1200 Commercial Street, Emporia, KS 668011200 Commercial Street, Emporia, KS 66801

Chapter 1: Snake Assemblage Related to the Vegetation Structure on Native Prairies

IntroductionIntroduction

Midwest – grasslands: millions of acres Midwest – grasslands: millions of acres scattered scattered remnants remnants

Anthropogenic disturbance: conversion to agricultural Anthropogenic disturbance: conversion to agricultural crops, rangeland, & residential developmentcrops, rangeland, & residential development

Enhances natural differences among patches within Enhances natural differences among patches within heterogeneous landscapeheterogeneous landscape

Profound effect on plant & animal communitiesProfound effect on plant & animal communities


Habitat selection/presence & absence studied widely Habitat selection/presence & absence studied widely

Assemblages = species populations (immigrated or Assemblages = species populations (immigrated or already present) within a patchalready present) within a patch

Influenced by environment, interaction with other Influenced by environment, interaction with other species, & properties associated with assemblage species, & properties associated with assemblage groupingsgroupings

Natural history important to determine appropriate Natural history important to determine appropriate scales to investigatescales to investigate


Heterogeneity effects community assemblage/structure Heterogeneity effects community assemblage/structure by increasing species richness by increasing species richness

Vegetation structure diversity, climate, diversity of Vegetation structure diversity, climate, diversity of microhabitat, morphological, & behavioral traitsmicrohabitat, morphological, & behavioral traits

Habitat selection by any species depends on structural Habitat selection by any species depends on structural features of the patch or surrounding areas features of the patch or surrounding areas must be must be suitable for reproductive purposessuitable for reproductive purposes

Available forage, den/nest sites, predators, Available forage, den/nest sites, predators, competitors, etc.competitors, etc.


Few conservation efforts focus on prairie amphibians & Few conservation efforts focus on prairie amphibians & reptiles…much to learn about snake habitat selection reptiles…much to learn about snake habitat selection in Midwest in Midwest

Why study snakes?Why study snakes?- specific habitat requirements for daily - specific habitat requirements for daily activities activities- great for ecological research- great for ecological research

Numbers are declining…must focus on identifying & Numbers are declining…must focus on identifying & protecting critical habitatprotecting critical habitat

• Identify distinct patterns that may Identify distinct patterns that may exist between snake assemblages & exist between snake assemblages & vegetation structure within native vegetation structure within native prairie patchesprairie patches

• Account for differences that may exist Account for differences that may exist between study sites using local between study sites using local vegetation compositionvegetation composition

Drift Fence Arrays Within Each Site

Figure 1. Drift fence array consisting of three arms spaced 120º apart, three single-ended funnel traps, and six double-ended funnel traps.

• • Biomark PIT tags Biomark PIT tags inserted on left lateral, inserted on left lateral, about 5 cm from cloaca about 5 cm from cloaca

• Scale-clipped ventral • Scale-clipped ventral scales on juvenile snakesscales on juvenile snakes

• Pocket Reader to • Pocket Reader to identify unique number identify unique number codescodes

Vegetation SamplingVegetation Sampling

5-cm by 100-cm pole to measure height of visibility 5-cm by 100-cm pole to measure height of visibility (Robel et al. 1970) (Robel et al. 1970)

1-m1-m² ² quadrat to determine % bare ground, woody quadrat to determine % bare ground, woody vegetation, forbs, and grassesvegetation, forbs, and grasses

ArcView 3.3 to select random coordinates from the ArcView 3.3 to select random coordinates from the native patches…used GPS device to find random native patches…used GPS device to find random pointspoints

Took three fixed samples between arms of each array Took three fixed samples between arms of each array at all sites x 2at all sites x 2

Data AnalysisData Analysis

Bray-Curtis measure of dissimilarity with LNMDS Bray-Curtis measure of dissimilarity with LNMDS ordination to find clustersordination to find clusters

Simpson’s Index of Diversity for abundance patterns Simpson’s Index of Diversity for abundance patterns and species richness for each siteand species richness for each site

Randomized block to determine significant differences Randomized block to determine significant differences for species diversity between site and yearfor species diversity between site and year

Randomized block to determine significant differences Randomized block to determine significant differences for vegetation composition between site and yearfor vegetation composition between site and year

ResultsResults

Prairie consisted of warm-season grasses: Big Prairie consisted of warm-season grasses: Big bluestem (bluestem (Andropogon gerardiiAndropogon gerardii)), , little bluestem little bluestem ((Andropogon scopariumAndropogon scoparium), Indian grass (), Indian grass (Sorghastrum Sorghastrum nutansnutans) and switchgrass () and switchgrass (Panicum virgatumPanicum virgatum); and ); and various forbs various forbs

13 snake species (brownsnake, copperhead, 13 snake species (brownsnake, copperhead, gartersnake, kingsnake, massasauga, racer, ratsnake, gartersnake, kingsnake, massasauga, racer, ratsnake, ribbonsnake, ringneck snake, and watersnake) in ribbonsnake, ringneck snake, and watersnake) in 29,268 trap nights 29,268 trap nights

Few recaptures (Few recaptures (Thamnophis sirtalis parietalisThamnophis sirtalis parietalis))

Total number of unique individuals by species captured Total number of unique individuals by species captured at the Blackbird, Fitch Hill, Garner Lane, and Slead sites at the Blackbird, Fitch Hill, Garner Lane, and Slead sites from June 2004 to November 2004 at FHNWR, Kansas. from June 2004 to November 2004 at FHNWR, Kansas.

SpeciesSpeciesBlackbirBlackbir

dd FitchFitch GarnerGarner SleadSlead TotalTotal

Agkistrodon contortrixAgkistrodon contortrix 00 00 00 22 22

Coluber constrictor flaviventrisColuber constrictor flaviventris 1818 1616 1010 1515 5959

Diadophis punctatus arnyiDiadophis punctatus arnyi 00 00 00 11 11

Elaphe o. obsoletaElaphe o. obsoleta 11 22 22 33 88

Lampropeltis c. calligasterLampropeltis c. calligaster 11 00 00 11 22

Lampropeltis g.holbrookiLampropeltis g.holbrooki 00 00 00 00 00

Nerodia e. transversaNerodia e. transversa 33 00 11 00 44

Nerodia r. rhombiferNerodia r. rhombifer 00 00 00 00 00

Sistrurus catenatus tergeminusSistrurus catenatus tergeminus 00 22 11 00 33

Storeria dekayi texanaStoreria dekayi texana 00 00 00 00 00

Thamnophis radixThamnophis radix 11 00 00 11 22

Thamnophis proximusThamnophis proximus 2323 22 22 1212 3939

Thamnophis sirtalis parietalisThamnophis sirtalis parietalis 4848 1717 1717 1919 101101

TotalTotal 9595 3939 3333 5454 221221

Total number of unique individuals by species captured Total number of unique individuals by species captured at the Blackbird, Fitch Hill, Garner Lane, and Slead sites at the Blackbird, Fitch Hill, Garner Lane, and Slead sites from April 2005 to November 2005 at FHNWR, Kansas.from April 2005 to November 2005 at FHNWR, Kansas.

SpeciesSpecies BlackbirdBlackbird FitchFitch GarnerGarner SleadSlead TotalTotal

Agkistrodon contortrixAgkistrodon contortrix 00 00 00 55 55

Coluber constrictor flaviventrisColuber constrictor flaviventris 2424 1111 1515 2222 7272

Diadophis punctatus arnyiDiadophis punctatus arnyi 00 00 00 00 00

Elaphe o. obsoletaElaphe o. obsoleta 11 00 11 11 33

Lampropeltis c. calligasterLampropeltis c. calligaster 00 00 00 11 11

Lampropeltis g.holbrookiLampropeltis g.holbrooki 00 00 00 11 11

Nerodia e. transversaNerodia e. transversa 00 00 00 11 11

Nerodia r. rhombiferNerodia r. rhombifer 11 00 00 00 11

Sistrurus catenatus tergeminusSistrurus catenatus tergeminus 00 11 22 00 33

Storeria dekayi texanaStoreria dekayi texana 11 00 00 00 11

Thamnophis radixThamnophis radix 00 00 00 11 11

Thamnophis proximusThamnophis proximus 88 44 22 33 1717

Thamnophis sirtalis parietalisThamnophis sirtalis parietalis 2121 88 99 3535 7373

TotalTotal 5656 2424 2929 7070 179179

Total number of unique individuals captured each month Total number of unique individuals captured each month from June 2004 to November 2005 for Yellow-bellied from June 2004 to November 2005 for Yellow-bellied Racers, Red-sided Gartersnakes, and Western Racers, Red-sided Gartersnakes, and Western Ribbonsnakes within the Blackbird, Slead, Fitch Hill, and Ribbonsnakes within the Blackbird, Slead, Fitch Hill, and Garner Lane sites at FHNWR, Kansas. Garner Lane sites at FHNWR, Kansas.

Total number of unique individuals captured from June Total number of unique individuals captured from June 2004 to November 2005 for Massasauga Rattlesnakes 2004 to November 2005 for Massasauga Rattlesnakes and Osage Copperheads within the Blackbird, Slead, and Osage Copperheads within the Blackbird, Slead, Fitch Hill, and Garner Lane sites at FHNWR, Kansas. Fitch Hill, and Garner Lane sites at FHNWR, Kansas.

-1.0 -0.8 -0.6 -0.4 -0.2 0.0 0.2 0.4 0.6 0.8

-0.6

-0.4

-0.2

0.0

0.2

0.4

0.6

0.8

1.0

BlackbirdSleadGarnerFitch

LNMDS depicting the clustering affect of similar communities on each array for all sites from June 2004 to November 2004 at FHNWR, Kansas.

-1.0 -0.5 0.0 0.5 1.0 1.5

-0.4

-0.2

0.0

0.2

0.4

0.6 BlackbirdSleadGarnerFitch

LNMDS depicting the clustering affect of similar communities on each array for all sites from April 2005 to November 2005 at FHNWR, Kansas.

-1.5 -1.0 -0.5 0.0 0.5 1.0 1.5

-1.5

-1.0

-0.5

0.0

0.5

1.0

1.5

BlackbirdSleadGarnerFitch

LNMDS depicting the clustering affect of similar communities on each array for all sites from June 2004 to November 2005 at FHNWR, Kansas.

ResultsResults

No significant difference for snake No significant difference for snake diversity between all sites (Fdiversity between all sites (F = 2.207, = 2.207, P P = = 0.266) or between years (F = 1.42,0.266) or between years (F = 1.42, P P = = 0.319)0.319)

Significant difference between year (F = Significant difference between year (F = 46.84, 46.84, PP = 0.006) for the average height = 0.006) for the average height of visibility through the floraof visibility through the flora

Paired T-test indicating significant differences Paired T-test indicating significant differences in grass cover from June 2004 to November in grass cover from June 2004 to November 2005 on several sites at FHNWR, Kansas. 2005 on several sites at FHNWR, Kansas.

Site Variance df t Stat P-value

Fitch Hill 0.0156 8 2.931 0.0189

Garner Lane 0.0219 8 3.962 0.0042

Slead 0.0232 8 -2.564 0.0334

Blackbird 0.0230 8 0.285 0.7826

Paired T-test indicating significant differences Paired T-test indicating significant differences in forb cover from June 2004 to November 2005 in forb cover from June 2004 to November 2005 on several sites at FHNWR, Kansas. on several sites at FHNWR, Kansas.

Site Variance df t Stat P-value

Fitch Hill 0.0148 8 2.525 0.035

Slead 0.3833 8 3.231 0.012

Blackbird 0.0043 8 -3.039 0.016

Garner Lane 0.0198 8 0.377 0.716

-0.5 0.0 0.5 1.0 1.5 2.0

-1.5

-1.0

-0.5

0.0

0.5

1.0

1.5

2.0

2.5BlackbirdSleadGarnerFitch

LNMDS of Bray-Curtis dissimilarities depicting changes in the vegetation composition from June 2004 to November 2004 at the FHNWR, Kansas.

DiscussionDiscussion

Controlled spring burns in 2004 vs. uncontrolled burns in Controlled spring burns in 2004 vs. uncontrolled burns in 2005 on several sites2005 on several sites

Spring flood in 2005 restricted access to several sitesSpring flood in 2005 restricted access to several sites

Species utilizing multiple habitat patchesSpecies utilizing multiple habitat patches

Individuals learned to avoid trapsIndividuals learned to avoid traps

Longer sampling period = higher censusLonger sampling period = higher census


Trend of increased richness with increased Trend of increased richness with increased heterogeneityheterogeneity

Species richness increased as plant richness Species richness increased as plant richness increased, then decreased when warm-season grasses increased, then decreased when warm-season grasses dominateddominated

Decrease activity as thatch layer increasesDecrease activity as thatch layer increases

As vegetative cover increased, plant diversity As vegetative cover increased, plant diversity decreased creating lower quality habitatdecreased creating lower quality habitat

Snake populations depend upon lushness of ground Snake populations depend upon lushness of ground vegetation & environmental conditionsvegetation & environmental conditions


The Garner Lane & Fitch Hill sites indicated the lack of The Garner Lane & Fitch Hill sites indicated the lack of plant cover may repel speciesplant cover may repel species

The amount of plant cover instead of vegetation type The amount of plant cover instead of vegetation type has been shown to influence habitat preferencehas been shown to influence habitat preference


Vegetation structure within a patch is important, but so Vegetation structure within a patch is important, but so are habitat features around a patchare habitat features around a patch

Over-wintering den sites and rookeries (pregnancy Over-wintering den sites and rookeries (pregnancy rocks) very important featuresrocks) very important features

Some patches may not be utilized if not in close Some patches may not be utilized if not in close proximity to hibernaculum, even though it has suitable proximity to hibernaculum, even though it has suitable featuresfeatures

Management ImplicationsManagement Implications

Improved conservation efforts that focus on expanding Improved conservation efforts that focus on expanding upon national wildlife refuge systemupon national wildlife refuge system

Certain biogeographical principles apply: distance from Certain biogeographical principles apply: distance from main species pool, and areamain species pool, and area

Link landscape on multiple scales: within refuge and Link landscape on multiple scales: within refuge and outside refugeoutside refuge

Most importantly: educate the public!!!Most importantly: educate the public!!!

Chapter 2: Abundance Patterns for Chapter 2: Abundance Patterns for Coluber constrictor flaviventrisColuber constrictor flaviventris, , Thamnophis proximusThamnophis proximus, and , and Thamnophis sirtalis parietalis Thamnophis sirtalis parietalis at at the Flint Hills National Wildlife the Flint Hills National Wildlife RefugeRefuge


Anthropogenic disturbance has not only had profound Anthropogenic disturbance has not only had profound effect on plant animal communities, but with ecological effect on plant animal communities, but with ecological interactions as well. interactions as well.

Population and community level caused by which Population and community level caused by which species are present, which habitat they occupy, and the species are present, which habitat they occupy, and the type of forage available. type of forage available.

Species populations within a patch are limited by Species populations within a patch are limited by suitable conditions or resources, size, proximity of suitable conditions or resources, size, proximity of other patches, length of time the patch is habitable, other patches, length of time the patch is habitable, and the rate of population growth.and the rate of population growth.

Anthropogenic disturbances enhances natural Anthropogenic disturbances enhances natural differences within a population.differences within a population.


Reptiles and amphibians constantly interact with their Reptiles and amphibians constantly interact with their physical environment.physical environment.

- pursuing prey, vocalizing, defending territory, - pursuing prey, vocalizing, defending territory, thermoregulation and breeding thermoregulation and breeding

Indicators of changing environmental conditions.Indicators of changing environmental conditions.


Investigating the abundance patterns of several Investigating the abundance patterns of several species within a stable community can help identify species within a stable community can help identify conditions within a habitat patch which species prefer, conditions within a habitat patch which species prefer, and to determine the distribution and abundance of and to determine the distribution and abundance of similar patches where a species would be expected to similar patches where a species would be expected to occur. occur.

The presence of diverse prey items can influence The presence of diverse prey items can influence whether enough individuals to form a population will whether enough individuals to form a population will inhabit a patch. inhabit a patch.

Kansas snake species such as yellow-bellied racers Kansas snake species such as yellow-bellied racers ((Coluber constrictor flaviventrisColuber constrictor flaviventris) and red-sided ) and red-sided gartersnakes (gartersnakes (Thamnophis sirtalis parietalisThamnophis sirtalis parietalis) prey ) prey mostly on small rodents or amphibians.mostly on small rodents or amphibians.


Social interactions may be just as important as prey Social interactions may be just as important as prey availabilityavailability

In New Hampshire, social interactions between racers In New Hampshire, social interactions between racers and gartersnakes influenced activity patterns and body and gartersnakes influenced activity patterns and body sizesize

Individuals that could not compete (small body size or Individuals that could not compete (small body size or less active) were forced to occupy lower quality habitat less active) were forced to occupy lower quality habitat (lower veg. height, more bare ground, & lower plant (lower veg. height, more bare ground, & lower plant diversity)diversity)

Predation can cause smaller species/individuals to Predation can cause smaller species/individuals to occupy marginal quality habitat (juveniles moving from occupy marginal quality habitat (juveniles moving from natal areasnatal areas))


What interactions or factors influence these species’ What interactions or factors influence these species’ abundance patterns? abundance patterns?

Recapture rates would be low and the abundance of Recapture rates would be low and the abundance of the most frequently captured species to not vary from the most frequently captured species to not vary from site to site site to site

The larger of the two species would suppress the The larger of the two species would suppress the smaller species’ abundance. smaller species’ abundance.

Adult body size for both species would be comparable Adult body size for both species would be comparable

Data AnalysisData Analysis

Randomized block to determine significant differences Randomized block to determine significant differences for species between site and yearfor species between site and year

Randomized block to determine significant difference Randomized block to determine significant difference for prey species between site and yearfor prey species between site and year

Collapsed ANOVA to determine significant site effects Collapsed ANOVA to determine significant site effects for snake speciesfor snake species

Collapsed ANOVA to determine significant site effects Collapsed ANOVA to determine significant site effects for prey species; Least Squared Means to detect for prey species; Least Squared Means to detect abundance patterns within each site for speciesabundance patterns within each site for species

Fisher’s Least-Significant Difference Test to determine Fisher’s Least-Significant Difference Test to determine significant differences within populations between sitessignificant differences within populations between sites

ResultsResults

400 unique individuals from 13 snake species; 29,268 400 unique individuals from 13 snake species; 29,268 trapnightstrapnights

The three most abundant species accounted for 90 The three most abundant species accounted for 90 percent of unique individuals captured: 44 % percent of unique individuals captured: 44 % Thamnophis s. parietalis,Thamnophis s. parietalis, 33 % 33 % Coluber c. flaviventrisColuber c. flaviventris, , and 14 % and 14 % Thamnophis proximusThamnophis proximus

Prey items regurgitated include: Prey items regurgitated include: Sigmodon hispidus, Sigmodon hispidus, adult Microtus ochrogaster, adult Microtus ochrogaster, grasshoppersgrasshoppers, , juvenile and juvenile and adultadult Rana blairi, Rana blairi, andand Acris crepitans blanchardi Acris crepitans blanchardi

87% of87% of Rana blairi Rana blairi captured were on Blackbird and captured were on Blackbird and Slead sites; 59% of Slead sites; 59% of Sigmodon hispidusSigmodon hispidus were captured were captured on the Garner Lane siteon the Garner Lane site

ResultsResults

The number of unique individuals for The number of unique individuals for Thamnophis s. Thamnophis s. parietalisparietalis (F = 4.12, (F = 4.12, P P = 0.048) and = 0.048) and Thamnophis Thamnophis proximusproximus (F = 18.45, (F = 18.45, P P = 0.00059) and = 0.00059) and Sigmodon Sigmodon hispidushispidus (F = 8.63, (F = 8.63, P P = 0.0068) were significantly = 0.0068) were significantly different by site different by site

Thamnophis s. parietalisThamnophis s. parietalis population on Blackbird was population on Blackbird was significantly different from the Fitch Hill (significantly different from the Fitch Hill (P P = 0.018) and = 0.018) and the Garner Lane (the Garner Lane (P P = 0.022) sites; = 0.022) sites; Coluber c. Coluber c. flaviventrisflaviventris population on Blackbird also was population on Blackbird also was significantly different from the Fitch Hill (significantly different from the Fitch Hill (P P = 0.027) and = 0.027) and the Garner Lane (the Garner Lane (P P = 0.027) sites= 0.027) sites

Fisher’s Least-Significant-Difference Test with Fisher’s Least-Significant-Difference Test with pairwise comparison probabilitiespairwise comparison probabilities

Thamnophis sirtalis parietalisThamnophis sirtalis parietalis

BlackbirdBlackbird Fitch HillFitch Hill Garner LaneGarner Lane SleadSlead

BlackbirdBlackbird 1.0000001.000000

Fitch HillFitch Hill 0.0180360.018036 1.0000001.000000

Garner LaneGarner Lane 0.0220730.022073 0.8984480.898448 1.0000001.000000

SleadSlead 0.3227310.322731 0.0925130.092513 0.1132270.113227 1.0000001.000000


Coluber constrictor flaviventrisColuber constrictor flaviventris





SleadSlead 0.3705550.370555 0.1201810.120181 0.1201810.120181 1.0000001.000000


Thamnophis proximusThamnophis proximus





SleadSlead 0.0061060.006106 0.0392070.039207 0.0183080.018308 1.0000001.000000

Least Squares Means from Collapsed ANOVA analysis Least Squares Means from Collapsed ANOVA analysis indicating a similar abundance pattern for three snake indicating a similar abundance pattern for three snake species and one amphibian species at the FHNWR, species and one amphibian species at the FHNWR, Kansas. Kansas.

T. proximus

C. c. flaviventris

T. s. parietalis


Yellow-bellied racers suppress smaller-bodied species Yellow-bellied racers suppress smaller-bodied species populationpopulation. .

September 2004 smaller September 2004 smaller Thamnophis sirtalis parietalis Thamnophis sirtalis parietalis individuals were present when individuals were present when Coluber constrictor Coluber constrictor flaviventrisflaviventris individuals were absent. individuals were absent.

First-years moving from lower quality habitat to fill void First-years moving from lower quality habitat to fill void in higher quality habitat.in higher quality habitat.

Could be last individuals to reach hibernaculum Could be last individuals to reach hibernaculum avoiding predation and competition.avoiding predation and competition.


Grassland snakes utilize wide variety of prey items.Grassland snakes utilize wide variety of prey items.

In the past, these snakes followed trends of rodent In the past, these snakes followed trends of rodent prey.prey.

Population would decline as species diversity declined Population would decline as species diversity declined (grass replaced forbs & woody vegetation replaced (grass replaced forbs & woody vegetation replaced grass).grass).

Overgrazed pastures and cultivated fields support Overgrazed pastures and cultivated fields support fewer species fewer species Fitch Hill and Garner Lane. Fitch Hill and Garner Lane.


Blackbird and Slead exhibited larger populations due to Blackbird and Slead exhibited larger populations due to proximity to wetlands proximity to wetlands 2005 flood could have 2005 flood could have reduced those populationsreduced those populations

Juvenile Juvenile Thamnophis s. parietalis Thamnophis s. parietalis prefer mesic, lowland prefer mesic, lowland habitats habitats

Flood forced smaller individuals to upland habitats Flood forced smaller individuals to upland habitats which were inhabited by larger individuals which were inhabited by larger individuals increased increased predation and competition.predation and competition.

Interactions in 2004 much different than 2005Interactions in 2004 much different than 2005


Breeding activities influences abundance patternsBreeding activities influences abundance patterns

Physiological characteristics also influence abundance Physiological characteristics also influence abundance patterns by determining habitat use (thermoregulation patterns by determining habitat use (thermoregulation & activity)& activity)

Individuals shift home ranges depending on Individuals shift home ranges depending on thermodynamics and active foragingthermodynamics and active foraging

Feasible that individuals compete for habitat with Feasible that individuals compete for habitat with abundant prey, why not compete for habitat with abundant prey, why not compete for habitat with

suitable thermodynamics?suitable thermodynamics?

Ecological research is complex; Ecological research is complex; therefore, difficult to pin down one therefore, difficult to pin down one factor within a natural environment factor within a natural environment that solely influences snake that solely influences snake assemblage patterns…too many assemblage patterns…too many factors that could all contribute!factors that could all contribute!

ACKNOWLEDGMENTSACKNOWLEDGMENTS

I would like to thank my committee members (L. Sievert, B. I would like to thank my committee members (L. Sievert, B. Thomas, G. Sievert, and T. Menard) for their comments on each Thomas, G. Sievert, and T. Menard) for their comments on each draft of my thesis. My deepest gratitude for the financial support draft of my thesis. My deepest gratitude for the financial support from the Harold Durst Graduate Research Award Committee and from the Harold Durst Graduate Research Award Committee and Emporia State University Department of Biological Sciences. The Emporia State University Department of Biological Sciences. The generous support of the United States Fish and Wildlife Service generous support of the United States Fish and Wildlife Service (USFWS) and the Flint Hills National Wildlife Refuge (FHNWR) for (USFWS) and the Flint Hills National Wildlife Refuge (FHNWR) for allowing me to set up trapping arrays. I was also like to thank G. allowing me to set up trapping arrays. I was also like to thank G. Smith, and D. Zelmer for their comments on earlier drafts of my Smith, and D. Zelmer for their comments on earlier drafts of my thesis. J. Campbell, B. Smart, C. Dobbs, L. Simpson, K. Jackson, thesis. J. Campbell, B. Smart, C. Dobbs, L. Simpson, K. Jackson, J. Phillis, A. Parsons, W. Dunn, and K. Shepherd for their J. Phillis, A. Parsons, W. Dunn, and K. Shepherd for their assistance in the field. The support and love of my family and assistance in the field. The support and love of my family and friends. friends.

Date post:	21-Jan-2015
Category:	Education
Upload:	bmaher7
View:	1,466 times
Download:	0 times

Thesis Defense

Education