Gray, 2002 PDF

EFFECT OF ANTHROPOGENIC DISTURBANCE AND
LANDSCAPE STRUCTURE ON BODY SIZE, DEMOGRAPHICS, AND
CHAOTIC DYNAMICS OF SOUTHERN HIGH PLAINS AMPHIBIANS
by
MATTHEW JAMES GRAY, B.S., M.S.
A DISSERTATION
IN
WILDLIFE SCIENCE
Submitted to the Graduate Faculty

of Texas Tech University in
Partial Fulfillment of
the Requirements for
the Degree of
DOCTOR OF PHILOSOPHY
Approved
May, 2002
©2002, Matthew J. Gray
ACKNOWLEDGEMENTS
Foremost I thank my major advisor, Dr. Loren M. Smith, for providing the
opportunity to continue my education and professional development at Texas Tech
University (TTU). Dr. Smith not only provided me with this great academic opportunity:
his guidance honed me as a scientist. I now am capable of perceiving the edge of science
and conducting research independently. Dedicating my dissertation to him is not enough
(see page v). Without question, my future research endeavors will be a product of his
mentorship. I hope my science will make him proud. Thank you so much, Loren!
Next, I thank my minor professor. Dr. Linda J. S. Allen of the Department of
Mathematics and Statistics at TTU. Dr. Allen sculptured my quantitative abilities and
burgeoned my interest in biomathematics. I now understand and appreciate the
usefulness of mathematics in explaining ecological phenomena. Indeed, her tutelage
changed my perception of science. Thank you. Dr. Allen, for making me quantitatively
literate. I also want to thank my other instructors in the math department: Drs. Kamal
Chanda, Wijesuriya Dayawansa, William Gustafson, Ira Wayne Lewis, and James Surles.
Each of these individuals helped me develop important quantitative skills.
In addition to Drs. Smith and Allen, I thank my committee members: Drs. Lou
Densmore (Department of Biological Sciences, TTU), David Haukos (U.S. Fish and
Wildlife Service), and David Wester (Department of Range, Wildlife and Fisheries,
TTU). These individuals provided incredible personal and professional guidance during
my four years at TTU. My dissertation is a testament of their devotion and support.
Thank you for making my doctoral pursuit a stimulating and intellectual experience!
n
I also want to thank Dr. James Surles of the Department of Mathematics and
Statistics at TTU for quantitative guidance on se\'eral anal> ses in my dissertation and
numerous stimulating discussions on experimental design and statistics. I am grateful to
Drs. Kevin Pope, Mark Wallace, and Gene Wilde of the Department of Range. Wildlife,
and Fisheries at TTU and Dr. Ray Semlitsch of the University of Missouri (Di\ision of
Biological Sciences) for editorial comments on my dissertation. I also thank the
following TTU graduate students for reviewing chapters: J. Avey, J. Brunjes. K. Brunjes,
and S. Sebring. I thank Stephanie Dupree of the Geospatial Technology Lab at TTU for
helping construct Figure 2.1. I am grateful to Dr. Ernest Fish, Cherise Ginsburg. and
Raquel Leyva for instructing me in ERDAS®, ESRI®, and FRAGSTATS® software.
I am deeply indebted to the landowners of my study playas: A. Arajo, L. Crum, K.
Gabel, B. Harrell, R. Hunter, B. Kimbrough, M. Marley, S. McPhearson, J. Merritt, G.
Olson, S. Olson, D. Schulte, E. Schulte, C. Sellers, E. Verett, and M. Wilmeth. Their
gratefulness allowed me to take my research thoughts to the field. I also thank M. Cerda.
T. Grand, L. Lloyd, and A. Wallace of the U.S. Department of Agriculture (USDA) Farm
Service Agency (FSA) for assistance in locating aerial slides and FSA Farm Folders for
my study playas.
This study would not have been possible without field and lab assistance from
many individuals. I gratefully thank: A. Anderson, J. Anderson, J. Avey, R. Brenes, W.
Dayawansa, E. Edwards, J. Emrich, D. Garza, D. Ghioca, W. Huskisson, A. Kauffman,
R. Kostecke, P. Lemons, A. Marshall, R. McCaffrey, D. Miller, K. Offill, J. Pautz, C.
Perchellet, R. Phillips, A. Quarels, D. Ray, W. Redell, S. Sebring, C. Shavlik, and C.
ill
Smith. I especially am indebted to Roberto Brenes (now at the University of Texas-
Tyler)—my right-hand man and savior. Roberto assisted me in the field for two years.
Honestly, I do not understand how he tolerated me—and, I am still quite amazed that we
survived. Roberto was more than a technician to me—he was a colleague and great
friend. I am confident my research would have suffered without him. I thank him for all
his dedication, patience, and herpetological insight. I doubt if I will ever be able to repay
him equally. Muchisimo gracias y pura vida, Sementall
This study was financially supported primarily by the Caesar Kleberg Foundation
for Wildlife Conservation and the Department of Range, Wildlife, and Fisheries
Management at TTU. Student research grants also were provided by the Society of
Wetland Scientists and Ducks Unlimited, Inc., and Home Depot donated buckets for
pitfall traps. I thank Dr. Gene Wilde of TTU for use of various sampling equipment and
Program CANOCO for multivariate ordination analyses. I also am very grateful to Dr.
Rob Mitchell of TTU for allowing me use his ATV on days when the west Texas roads
were impassable. Lastly, I thank Dr. Nick Parker of TTU and Dick Lubke of the Texas
Parks and Wildlife Department for use of aquariums and a PIT tag scanner, respectively.
Finally, I am grateful for the incredible support of my family: James S. Gray
(father), Scott H. Gray (brother), Sharon L. Malloy (mother), and Carly L. Pratt (sister).
No one gets anywhere by himself or herself My family has been an amazing nucleus of
love and support. Without their encouragement, my 12-year academic journey would
have been impossible. I also thank my very dear friend, Jessica L. Emrich. Her love and
patience gave me the strength to succeed.
IV
I dedicate my dissertation to my four academic mentors. In order of academic
interaction: Drs. Harold H. Prince (Michigan State Universit>). Richard M. Kaminski
(Mississippi State Universit>'). Francisco J. Vilella (Mississippi Cooperati\ e Fish and
Wildlife Research Unit), and Loren M. Smith (Texas Tech University). Dr. Prince
trained me in the scientific methods. Dr. Kaminski fueled m\ passion for wetlands and
research and de\ eloped my wxiting skills. Dr. \ ilella sparked my interest in conservation
biology, and Dr. Smith directed me in ecological theor>' and landscape ecology. These
individuals made me the person and scientist that I am toda}'. Indeed, I am indebted.
Thank \ ou so much gentlemen!
\'
TABLE OF CONTENTS
ACKNOWLEDGEMENTS ii
ABSTRACT ix
LIST OF TABLES xi
LIST OF FIGURES xiii
CHAPTER
I. INTRODUCTION 1
II. EFFECT OF LANDUSE AND RAINFALL ON POSTMETAMORPHIC

BODY SIZE OF PLAYA AMPHIBIANS 5
Abstract 5
Introduction 6
Study Area and Amphibians 9
Methods 13
Experimental Design and Units 13
Sampling Techniques 14
Terrestrial capture 14
Biological processing 15
Statistical Analyses 15
Results 16
Landuse Type and Year Interaction Effects 16
Adults 16
Subadults 17
Metamorphs 17
Landuse Type Main Effect 19
Adults 19
Subadults 19
Metamorphs 19
Year Main Effect 21
Adults 21
Subadults 21
Metamorphs 23
Discussion 23
Landuse Effect 23
Year Effect 27
Competing Body Size Hypotheses 29
VI
Conclusions ^^
III. EFFECT OF LANDUSE AND YEAR ON POPULATION

DEMOGRAPHICS OF SOUTHERN HIGH PLAINS
AMPHIBIANS 32
Abstract 32
Introduction 33
Methods 36
Results 41
Abundance and Diversity 41
Source-Sink Dynamics 43
Discussion 45
Landuse Effect 47
Year Effect 52
Source-Sink Dynamics 53
Conservation Implications 54
IV. INFLUENCE OF LANDSCAPE STRUCTURE ON COMMUNITY

COMPOSITION AND RELATIVE ABUNDANCE OF
AMPHIBIANS 56
Abstract 56
Introduction 57
Methods 59
Results 63
Discussion 76
Conservation Implications 78
V. TEMPORAL NICHE PARTITIONLNG IN SOUTHERN HIGH PLAINS

AMPHIBIAN POPULATIONS 80
Abstract 80
Introduction 81
Methods 83
Results 85
Discussion 91
VI. CHAOS AND REGULATING MECHANISMS IN A SOUTHERN HIGH

PLAINS AMPHIBIAN ASSEMBLAGE 106
Abstract 106
Introduction 107
Methods 110
Vll
Results 117
Discussion 128
Ecological and Mathematical Implications 133
VII. CONCLUSION 135
LITERATURE CITED 140
APPENDIX
A. PERCENT COVERAGE OF COVER TYPES IN LANDSCAPES

SURROUNDLNG PLAYA WETLANDS 167
B. ANOVA TABLES FOR CHAPTER III 169
C. AGE CLASS ANOVA TABLES FOR CHAPTER III 172
D. CROPLAND AND GRASSLAND LANDSCAPES 175
Vlll
ABSTRACT
Amphibian populations are declining globally. Anthropogenic disturbance of
landscapes surrounding wetlands may affect fitness, demographics, and dynamics of
amphibian populations. Spatial positioning and relati\'e connectedness of wetlands also
may influence population demographics. Thus, I examined the effect of anthropogenic
landscape use (cultivation vs. grassland) and structure on postmetamorphic body size (a
fitness correlate), demographics, and dynamics of amphibians at 16 playa wetlands on the
Southern High Plains (SHP) of Texas during 1999 and 2000. Amphibian populations
were monitored using drift fence and pitfall traps, landscape structure was quantified
using spatial analysis software, and dynamics were assessed using difference equations.
Postmetamorphic body size of all amphibian species and age classes generally
was greater at grassland than cropland playas, and in 1999 (i.e., a wetter year) than 2000.
Abimdance of New Mexico and plains spadefoots (Spea multiplicata and S. bombifrons)
generally was greater at cropland than grassland playas, and greater for barred tiger
salamanders {Ambystoma tigrinum mavortium) in 1999 than 2000. Mean daily
abundance of amphibians also was positively related to landscape structure indices
representing geometric complexity and spatial positioning of wetlands. In general, as
landscapes became more complex (e.g., numbers of edges increased) and inter-playa
distance decreased, mean daily abundance of amphibians increased. Additional
demographic anah ses indicated that temporal niche partitioning existed in SHP
amphibian populations; however, no differences existed between landuses. Lastly,
biological chaos in the amphibian assemblage existed at 1 of 8 cropland and 7 of 8
ix
grassland playas. A stochastic density-dependent Ricker function predicted chaotic
dynamics most accurately.
Anthropogenic disturbance surrounding wetlands affects body size,
demographics, and dynamics of SHP amphibians. Spatial positioning of wetlands and
landscape complexity may be as or more important than general landuse in affecting
amphibian demographics. Armual differences in body size and abimdance suggest
rainfall may be important in influencing amphibian populations. Although spadefoot
abundance was positively influenced by anthropogenic disturbance, I recommend
retention and restoration of grasslands surrounding playa wetlands because landscape
cultivation decreased body size and altered amphibian demographics and dynamics from
an undisturbed state. These results have important implications in conservation biology,
landscape ecology, and basic ecological and mathematical theory.

LIST OF TABLES
2.1 Adult body size of New Mexico spadefoot {Spea multiplicata, NSF), plains
spadefoot {S. bombifrons, PSF), Great Plains toad {Bufo cognatus, GPT),
and barred tiger salamander {Ambystoma tigrinum mavortium, BTS) between
landuse types and years (fixed factorial effects) at 16 playa wetlands (random
nested effect) on the Southern High Plains of Texas, April-September 1999
and 2000 18
2.2 Subadult body size of New Mexico spadefoot {Spea multiplicata, NSF), plains
spadefoot {S. bombifrons, PSF), and Great Plains toad {Bufo cognatus, GPT)
between landuse types and years (fixed factorial effects) at 16 playa wetlands
(random nested effect) on the Southern High Plains of Texas, April-
September 1999 and 2000 20
2.3 Metamorph body size of New Mexico spadefoot {Spea multiplicata, NSF), plains
spadefoot {S. bombifrons, PSF), Great Plains toad {Bufo cognatus, GPT), and
barred tiger salamander {Ambystoma tigrinum mavortium, BTS) between
landuse types and years (fixed factorial effects) at 16 playa wetlands (random
nested effect) on the Southern High Plains of Texas, April-September 1999
and 2000 22
3.1 Relative daily abundance and diversity of amphibians between landuse types
and years and among species at 16 playa wetlands on the Southern High
Plains, Texas, May-October 1999 and April-August 2000 42
3.2 Relative daily abundance of age classes between landuse types and years for
amphibian species that differed significantly in daily abundance (see Table
C.2) at 16 playa wetlands on the Southern High Plains, Texas, May-October
1999 and April-August 2000 44
4.1. Pearson coefficients of correlations and p-values (located beneath coefficients)

from parametric testing {t distribution) of linear associations between
landscape metrics^ at 16 playa wetlands on the Southern High Plains,
Texas, 1999 and 2000 64
4.2 Intra-set correlations of landscape* metrics associated with the first two axes
generated by a canonical correspondence analysis of mean daily abundance of
New Mexico spadefoot {Spea multiplicata), plains spadefoot {S. bombifrons),
barred tiger salamander {Ambystoma tigrinum mavortium), and Great Plains toad
{Bufo cognatus) at 16 playa wetlands on the Southern High Plains, Texas, 1999
and 2000 66
XI
4.3 Univariate Pearson coefficients of correlations between mean daily abundance
of amphibians and landscape* metrics at 16 playa wetlands on the Southern
High Plains, Texas, 1999 and 2000 70
4.4 Multiple linear regression between mean daily abundance of amphibians and
landscape* metrics at 16 playa wetlands on the Southern High Plains, Texas,
1999 and 2000 74
6.1 Stability of amphibian assemblage trajectories at 16 playa wetlands during 16

May-17 October 1999 and 19 April-18 August 2000 on the Southern High
Plains, Texas 122
6.2 Occurrence of chaos between landuses and models in amphibian assemblages

at 16 playa wetlands during 16 May-17 October 1999 and 19 April-18
August 2000 on the Southern High Plains, Texas 126
6.3 Least-squares estimates of parameters for the discrete stochastic Ricker

equation* from amphibian assemblages at 16 playa wetlands during 16
Plains, Texas 127
6.4 Iterative and least-squares estimates of parameters for deterministic difference

equations used to model amphibian assemblages at 16 playa wetlands during 16
Plains, Texas 129
6.5 Estimates of parameters between landuses for difference equations used to

model amphibian assemblages at 16 playa wetlands during 16 May-17
October 1999 and 19 April-18 August 2000 on the Southern High Plains,
Texas 130
A. 1 Percent coverage of cover types in landscapes* surrounding 16 playa wetlands

between landuses on the Southern High Plains, Texas, 1999 and 2000 168
B. 1 Statistics for ANOVAs testing differences in relative daily abundance and

diversity of amphibians between landuse types (cropland vs. grassland) and
years and among species at 16 playa wetlands on the Southern High Plains,
Texas, May-October 1999 and April-August 2000 170
C.l Statistics for ANOVAs for species (by age class) that differed significantly in
relative daily abundance between landuse types (cropland vs. grassland) or years
(see Table B.l) at 16 playa wetlands on the Southern High Plains, Texas,
May-October 1999 and April-August 2000 173
xii
LIST OF FIGURES
2.1 Location of study playa wetlands on the Southern High Plains of Texas,
U.S.A 12
3.1 Effect of landuse (top graph) and year (bottom graph) on frequency of directional
movement of amphibian species at 16 playa wetlands on the Southern High
Plains, Texas, May-October 1999 and April-August 2000 46
4.1 Canonical correspondence analysis of mean daily capture (natural-log transformed)

of amphibians and uncorrelated metrics in landscapes (i.e., 2,830-ha circular plot)
associated with 16 study playas on the Southern High Plains, Texas, 1999 and
2000 67
4.2 Simple linear regression and 95% confidence bands of mean daily abundance
(natural-log transformed) of New Mexico spadefoot {Spea multiplicata) and
percent aerial coverage of playa wetlands (PP), interspersion/juxtaposition index
(IJI, McGarigal and Marks 1995:103), edge density (ED, McGarigal and Marks
1995:106), playa edge density (PED [i.e., mean number of edges to cross from
study playa to surrounding playas), landscape shape index (LSI, McGarigal and
Marks 1995:109), and landuse richness (LR, McGarigal and Marks 1995:119) in
landscapes (i.e., 2,830-ha circular plot) associated with 16 study playas on the
Southern High Plains, Texas, 1999 and 2000 71
4.3 Simple linear regression and 95% confidence bands of mean daily abundance
(natural-log transformed) of plains spadefoot {Spea bombifrons) and percent
aerial coverage of playa wetlands (PP), interspersion/juxtaposition index (IJI,
McGarigal and Marks 1995:103), edge density (ED, McGarigal and Marks
1995:106), and landscape shape index (LSI, McGarigal and Marks 1995:109) in
landscapes (i.e., 2,830-ha circular plot) associated with 16 study playas on the
Southern High Plains, Texas, 1999 and 2000 73
5.1 Time series plots for pairs of amphibian species residing at 16 playa wetlands
in 2 landuses and years on the Southern High Plains of Texas 87
5.2 Percent occurrence pairs of standardized species-specific linear trajectories (see

Figure G.l) exceeded each other during 3 sequential and equal time intervals
{n=22 and 17 days for 1999 and 2000) within 2 landuses and years on the
Southern High Plains, Texas 92
6.1 Time series plots of amphibian assemblage dynamics at 8 playa wetlands during
16 May-17 October 1999 on the Southern High Plains, Texas 118
Xlll
6.2 Time series plots of amphibian assemblage dynamics at 8 playa wetlands during
19 April-18 August 2000 on the Southern High Plains, Texas 120
D.l Anthropogenic and natural cover types in landscapes (i.e., 2,830-ha [3-km radius]
circular plots) surrounding 16 playa wetlands on the Southern High Plains, 1999
and 2000 176
xiv
CHAPTER I
INTRODUCTION
Amphibians are declining globally and anthropogenic disturbance of landscapes
surrounding wetlands has been implicated as a primary cause (Blaustein et al. 1994.
Houlahan et al. 2000). Deforestation and agricultural cultivation in watersheds of
wetlands have been considered ultimate reasons for decline (Alford and Richards 1999).
Interestingly, few data exist in the United States that suggest amphibian populations are
negatively affected by cultivation. Two of the three studies (Knutson et al. 1999,
Kolozsvary and Swihart 1999) indicate that amphibian populations can be elevated in
wetlands surrounded by cropland, and one (Anderson et al. \999a) documented no
differences between disturbed and undisturbed wetlands. In Canada and Europe, most
studies suggest that landscape cultivation significantly decreases amphibian abundance
and species richness, and negatively affects fitness correlates (Laan and Verboom 1990,
Oldham and Swan 1991, Hecnar and M'Closkey 1996, Bonin et al. I997a,b). However,
most of these studies were conducted using call surveys during the breeding season,
which have less demographic resolution than other methods (e.g., pitfall sampling, Heyer
et al. 1994). Drift fence and pitfall traps facilitate the most comprehensive examination
of dynamics in non-arboreal amphibian populations (Heyer et al. 1994). Thus, previous
studies may not have sufficiently documented population responses to anthropogenic
disturbance, particularly considering that few sampling efforts were performed outside
the breeding season.

It has been hypothesized that anthropogenic disturbance affects amphibian
populations by altering abiotic and biotic conditions in and around wetlands (Semlitsch
2000). Anthropogenic disturbance can negatively affect water qualit>, which might
affect trophic structuring in the aquatic environment, and ultimately growth and survival
of larvae (Boone and Semlitsch 2001). Agricultural chemicals also can reduce
immunocompetence, mobility, and survival of larvae (Carey et al. 1999, Bridges and
Semlitsch 2000). Sedimentation in wetlands can decrease hydroperiods (Luo et al. 1997).
thus presumably breeding opportunities for adults and de\ elopmental time of larvae.
Finally, landscape disturbance may cause direct mortality, reduce terrestrial habitat and
subterranean retreats, and affect movement of postmetamorphic amphibians among
spatially structured populations (Semlitsch 2000).
Amphibians are important ecological entities of aquatic and terrestrial ecosystems
(Wilbur 1984). Larval amphibians can significantly affect trophic structuring and
nutrient cycling in wetlands via consumption and excrement of fine and course
particulate organic matter and micro- and macro-invertebrates (Scale 1980, Hoff et al.
1999, Petranka and Kennedy 1999). Also, amphibian larvae are prey for various
invertebrate and vertebrate species (Skelly 1997). Postmetamorphic amphibians consume
numerous invertebrates, some of which are agricultural pests, and they are food items for
various terrestrial predators (Duellman and Trueb 1994, Anderson et al. 19996). Larval
and postmetamorphic amphibians also readily absorb chemicals via osmoregulation
(Boutilier et al. 1992, Shoemaker et al. 1992), which can affect their physiology and
bioaccumulate (Semlitsch 2000). Consequently, amphibians are considered good

indicators of ecosystem health (Beebee 1996). Thus, it is important to discern the effect
of human disturbance on dynamics, demographics, and fitness correlates of amphibians
(Beebee 1996).
Amphibians exist primarily in spatially structured playa wetlands on the Southern
High Plains (Bolen et al. 1989). Cultivation and grassland surround most playas; thus,
they are ideal systems to comparatively and simultaneously investigate the effect of
human disturbance and landscape structure on amphibian populations. Therefore, I
proposed the following research. My goal was to determine the effect of anthropogenic
disturbance and landscape structure on body size and population demographics of
Southern High Plains amphibians. I accomplished this goal through 5 simultaneous
studies at 16 playa wetlands in 2 anthropogenic landuse types (cultivation and grassland)
over 2 years (1999 and 2000, 4 playas per landuse per year). My specific research
objectives were to determine the effect of (1) landuse and yearly rainfall on
postmetamorphic body size (a fitness correlate) of amphibians (Chapter II), (2) landuse
and year on demographics of amphibians (Chapter II), (3) landscape structure on
community composition and relative abundance of amphibians (Chapter IV), (4) landuse
on temporal niche partitioning and chaotic dynamics in amphibian communities
(Chapters V and VI), and to (5) identify potential regulating mechanisms in an amphibian
assemblage (Chapter VI). I used pitfall sampling, the geographic information system and
associated software, and difference equations to investigate these objectives. Results
presented herein provide insight into the effect of anthropogenic disturbance and
landscape structure on amphibian populations and mechanisms structuring their

communities. These results have important implications in amphibian and conservation
biology, landscape ecology, applied and theoretical mathematics, and basic ecological
theo^>^
CHAPTER II
EFFECT OF LANDUSE AND RAINFALL ON POSTMETAMORPHIC
BODY SIZE OF PLAYA AMPHIBIANS
Abstract
Postmetamorphic body size of amphibians is positively related to evolutionary
fitness. Landuse surrounding wetlands and rainfall may affect body size by influencing
proximate factors (e.g., hydroperiod, food resources) that affect larval and
postmetamorphic growth rates. I hypothesized that cultivation of terrestrial habitats
surrounding wetlands would affect body size of amphibian, but the magnitude of its
effect would depend on yearly differences in rainfall. I compared postmetamorphic body
size between two landuse types (cultivation versus grassland) surrounding playa wetlands
and two years (1999 versus 2000) with different rainfall (1999 > 2000) for 4 species
{Spea multiplicata, S. bombifrons, Bufo cognatus, and Ambystoma tigrinum mavortium)
and 3 age classes (metamorph, subadult, and adult) of amphibians on the Southern High
Plains of Texas. Sixteen playas (4 per landuse type per year) were partially enclosed with
drift fence and pitfall traps, and mass and snout-vent length (SVL) measured from a
subsample of captured individuals {n = 5,188). Mass and SVL generally were greater at
grassland than cropland playas for all species and age classes. Mass and SVL also were
greater in 1999 (i.e., the wetter year) than in 2000 for most species and age classes.
Moreover, the effect of landuse type and year on postmetamorphic body size generally
was non-additive; cultivation had a greater negative effect on body size in 2000 (i.e., the
drier year) than in 1999 for some of the age classes and species. Differences in body size
between landuse types and > ears may have been related to densit>' independent and
dependent factors, such as reduced wetland hydroperiod, food availabilit\\ and predators
or increased chemicals and density of conspecifics in cropland pla>as. Anthropogenic
modifications of landscapes surrounding wetlands may negati\eh influence
postmetamorphic body size of amphibians, especially during drier \ears.
Introduction
Postmetamorphic body size of amphibians can influence e\olutionary fitness b>
affecting survi\'al and reproduction (Wilbur 1984, Werner 1986, Semlitsch et al. 1988).
Probability of survival and reproduction in amphibians is positiveh related to
postmetamorphic body size (Smith 1987, Berven 1990. Goater 1994, Morey and Reznick
2001). Large individuals may have greater survival than small indi\ iduals, because they
can use a greater moisture range of microhabitats (Newman and Dunham 1994). can
consume a greater range of prey sizes (Flowers and Graves 1995. Newman 1999). are
more efficient foragers (Newman 1999), and have lower specific metabolic rates
(Hutchison et al. 1968, Ultsch 1973, 1974). Large individuals also take more time to
reach critical levels of dehydration than small individuals (Newman and Dunham 1994).
In addition, large individuals have greater sprint speeds, jumping ability, and endurance
than small individuals (Taigen and Pough 1981, John-Alder and Morin 1990, Goater et
al. 1993, Beck and Congdon 2000). Mating success of male and female amphibians also
may be positively related to body size (Berven 1981, Verrell 1982, 1985. Howard and
Young 1998, Marco et al. 1998). For territorial amphibians, territory size and quality
(i.e., density of food resources) can be positively correlated with body size (Howard
1978, Mathis 1990, Gabor 1995). Large females also may exhibit greater fecundity than
small females and spawn multiple times (Clarke 1974, Howard 1978, Berven 1982,
Semlitsch 1985a, Krupa 1986, 1994, Fontenot 1999). Finally, large individuals may
reproduce at an earlier age than small individuals (Berven and Gill 1983, Smith 1987.
Bervenl990, Scott 1994).
Postmetamorphic body size of amphibians can be affected by duration of larval
development and pre- and post-metamorphic conditions that differentially affect growth
and mortality rates of individuals in aquatic and terrestrial habitats (Wilbur and Collins
1973, Werner 1986). Generally, postmetamorphic body size is correlated with body size
at metamorphosis (Smith-Gill and Berven 1979. Semlitsch et al. 1988, Scott 1994).
Wetland hydroperiod, water quality, and density of food resources, conspecifics, and
predators can individually or synergistically affect body size at metamorphosis (Travis
1984, Wilbur 1984, Semlitsch 1987fl, Alford and Harris 1988, Pfennig et al. 1991,
Bradford et al. 1992, Werner and Anholt 1993). Postmetamorphic body size also is
affected by density of food resources and conspecifics in the terrestrial landscape (Goater
1994, Morey and Reznick 2001). Indeed, small body size at metamorphosis may be
compensated for by high density of terrestrial food resources (as theoretically suggested
by Werner 1986), allowing small individuals to attain a body size at first reproduction
similar to individuals that were larger at metamorphosis (e.g.. Goater 1994, Morey and
Reznick 2001).
Landuse type surrounding wetlands can affect larval and postmetamorphic
amphibians (Hecnar and M'Closkey 1996, Bonin et al. 1997a,b, Dodd 1997, Alford and
Richards 1999, Semlitsch 2000), presumably by influencing some of the aforementioned
mechanisms that affect growth and mortality rates of individuals in aquatic and terrestrial
environments (Werner 1986). Landscape cultivation (i.e., arable cropland) may localize
amphibians in wetlands, resulting in unexpected positive species associations or high
densities (Knutson et al. 1999, Kolozsvary and Swihart 1999, Chapter III).
Consequently, landscape cultivation may affect postmetamorphic body size of
amphibians by affecting density of conspecifics (Oldham 1985). Cultivation of the
watershed also can increase sedimentation in wetlands, which decreases hydroperiods
(Martin and Hartman 1987, Corn and Bury 1989, Luo et al. 1997), and may reduce
duration of larval development (Brady and Griffiths 2000). Insecticides and herbicides
associated with agricultural practices may affect postmetamorphic body size by reducing
available food resources. Similarly, agricultural chemicals (e.g., nitrates, ammonia,
organophosphates) can bioaccumulate and reduce foraging activity and growth of larval
amphibians (Hall and Kolbe 1980, Baker and Waights 1993, 1994, Hecnar 1995,
Semlitsch et al. 1995), and perhaps postmetamorphic body size.
Rainfall can affect growth rates of larval and postmetamorphic amphibians
(Tinsley and Tocque 1995, Camp et al. 2000) and potentially postmetamorphic body size
and condition (Bruce and Hairston 1990, Reading and Clarke 1995). Rainfall can
increase pond hydroperiod hence growth rates and duration of larval development
(Semlitsch et al. 1996, Reading and Clark 1999, Camp et al. 2000). Rainfall also can
increase prey availability in the terrestrial landscape (Dimmitt and Ruibal 1980a, Jaeger
8
1980). Moreover, rainfall reduces probability of desiccation and increases foraging
activity for amphibians (Jaeger 1980, Grover 1998, Newman 1999).
No studies have been conducted in North America examining the effect of
landuse type on postmetamorphic body size of amphibians. I hypothesized that
cultivation of terrestrial landscapes surrounding wetlands would negatively affect body
size of amphibians, but the magnitude of its effect would depend on rainfall. Thus, my
objective was to measure the effect of two landuse types (i.e., cultivation and grassland
[control]) over two years (i.e., 1999 and 2000) on postmetamorphic body size of
amphibians. I quantified the effect of landuse and year for 3 age classes (i.e., metamorph,
subadult, and adult) and 4 species (i.e., Spea multiplicata, S. bombifrons, Bufo cognatus,
and Ambystoma tigrinum mavortium) of amphibians using playa wetlands on the
Southern High Plains. This information is particularly important for species currently in
decline, such as amphibians (Houlahan et al. 2000), because body size can potentially
influence local population persistence.
Study Area and Amphibians
My study was conducted on the Southern High Plains (SHP) of Texas. The SHP
(i.e., the Llano Estacado) is the largest continuous plateau in the United States (Sabin and
Holliday 1995), encompassing ca. 8.9 million ha, of which 5.9 million ha are in
northwestern Texas (Haukos and Smith 1994). Primary land uses on the SHP are
agricultural cultivation (2.7 million ha, 46%)) and grassland (2.9 million ha, 49%) [native
and replanted lands combined], Haukos and Smith 1994). The most significant
topographic formations on the SHP are playa wetlands (Bolen et al. 1989).
Playa wetlands (or simply playas) occur at the highest density (ca. 19,340) in
Texas, comprising ca. 2% of the Texas SHP landscape (Haukos and Smith 1994). Playas
are circular wetlands (xsize=6.3 ha) that form dynamically and continually via an
interaction of eolian and alluvial processes (Osterkamp and Wood 1987, Gustavson et al.
1994). Playas are the lowest points on the relatively level SHP landscape, and their
watersheds are not interconnected (Guthery and Bryant 1982). Playas also are not
directly connected to the underlying Ogallala Aquifer (Bolen et al. 1989). Consequently,
playa hydrology is dependent on precipitation and watershed runoff (Haukos and Smith
1994). Because the climate on the SHP is semi-arid (i.e., Xrainfaii=33-45 cm/year,
><evaporation^200-250 cm/ycar) and playa basins are shallow, playa hydroperiods are
variable (Bolen et al. 1989). Playas usually have intermittent to seasonal or annual
hydroperiods, and they are oases of biodiversity in a predominately agricultural landscape
(Bolen et al. 1989, Haukos and Smith 1994).
Amphibians can be the most abundant vertebrate in playas during summer (Bolen
et al. 1989). Thirteen species exist on the SHP; I selected the 4 most abundant for this
study: New Mexico spadefoot {Spea multiplicata Cope), plains spadefoot {S. bombifrons
Cope), Great Plains toad {Bufo cognatus Say), and barred tiger salamander {Ambystoma
tigrinum mavortium Green) (Rose and Armentrout 1974, Anderson 1997:74, Chapter III).
The above anurans are explosive breeders (i.e., complete breeding in 1-3 d, Sullivan and
Fernandez 1999), although Great Plains toads may breed for several weeks and spawn
multiple times if ambient conditions are favorable (Krupa 1986, 1994). Barred tiger
salamanders can be more prolonged breeders, taking ca. 2-4 weeks to breed (Petranka
10
1998:111). Breeding for these species is concentrated during and after seasonal rains
March-August; however, barred tiger salamander may breed during any month on the
SHP if sufficient water is present (Webb and Roueche 1971. Rose and Armentrout 1976).
Duration of larval development for these amphibians is rapid (i.e.. 2-6 weeks) and likely
depends on phenotypic responses to competition, predation, hydroperiod, and water
quality (Rose and Armentrout 1976, Newman 1992, 1994a. Werner and McPeek 1994,
Skelly 1997, Denver et al. 1998, Morey and Reznick 2000).
Sixteen playas (4 per landuse type per year) were selected via aerial and ground
reconnaissance in March 1999 and 2000 according to surrounding landuse (cultivated or
grassland), presence of water (i.e., all playas were inundated initially), landowner
cooperation, and study playa juxtaposition (i.e., clusters of playas within the same
landuse type were sought to increase sampling efficiency). Playas were considered to be
in grassland if >75%) of the surrounding landscape (i.e., <3 km from the playa center) was
undisturbed and vegetated, and cultivation if >75% of the surrounding landscape was
arable cropland (Anderson et al. 1999(^:760, Chapter III). Grassland could have included
native, replanted (via the Conservation Reserve Program, CRP), or grazed lands with
intact grass. I chose the 3-km threshold for landuse designation, because it probably was
the near maximum dispersal distance for my study species (cf Gehlbach 1967, Gehlbach
et al. 1969, Sinsch 1990, 1997, Miaud et al. 2000).
Grassland playas were located in Castro County northeast of Dimmitt, Texas USA
(Figure 2.1). Playas surrounded by cropland were located in Hale and Floyd counties
11
34M5'0CrN
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amoiitt*©
A ^
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Ca stre <\ ••••' lllv»rWfiV_,
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1
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BHn»M ^J-.
-A
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o %
Hale F10 y d ^'••^.
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^
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33*22-3 (TN 1 •;:•
102*37'30W
•*• Major Qties Rateau Escarpment

o Cropland Playas Study Area Counties
© Grassland Playas
20 10 0 20 40 60 80 100
Kilometers
Figure 2.1. Location of study playa wetlands on the Southern High Plains of Texas,
U.S.A.
12
near Plainview, Texas in 1999 and north of Ralls, Texas in Crosby County in 2000.
Although grassland playas were located northwest (x=97 km, SD=11) of cropland playas,
soils and historical vegetation were similar between locations (Allen et al. 1972, D. A.
Haukos and L. M. Smith, U. S. Fish and Wildlife Service and Texas Tech University,
unpublished data). Moreover, rainfall (and I assume other ambient variables because
elevation and latitudinal change is minimal) was similar between sites (x=5.5 cm, SE=1.8
[cultivation]; x=6.9 cm, SE=2.7 [grassland]) during field sampling. Thus, I assumed a
possible landuse effect would not be confounded by geographic differences in sampling
location. Average rainfall at the study playas during sampling was greater in 1999
(x=15.1 cm, SE=1.6) than in 2000 (x=7.3 cm, SE=3.5).
Methods
Experimental Design and Units
Independent factors were arranged as a 3-way factorial-nested design
(Montgomery 2001:569-573). Landuse types and years were crossed factors and
considered fixed effects. Playas were nested within landuses and years and treated as a
random effect. I used individuals captured at nested playas as experimental units of
landuse and year. Individuals might be considered pseudoreplicates of the crossed effects
(Hurlbert 1984); thus, I restricted my inferences to the population of amphibians residing
in these 2 landuse types and years on Southern High Plains. I also used an unrestricted
mixed model, which allows covariance (i.e., dependency) between observations within
levels of the nested random effect (Hocking 1973). Consequently, results of ratio
significance tests using this model remained unbiased and accurate, despite potential
13
interdependencies between observations (Hocking 1973, Montgomer> 2001:526-527).
The aforementioned further supports my use of individuals as experimental units, because
potential interdependencies between individuals within nested playas were modeled in
the response.
Recent research (e.g., Morey and Reznick 2001) also suggests that terrestrial food
resources might differentially affect postmetamorphic body size of genders. Thus. 1 also
tested the effect of gender and landuse type on body size of adults in preliminary
multivariate analyses. These analyses were not conducted on the other age classes (i.e.,
metamorph and subadult), because captured individuals were not dissected for gender
determination. Body size of females was greater than males for adult New Mexico
spadefoot, plains spadefoot and Great Plains toads. However, because the gender effect
did not interact with landuse type and mean sex ratio across species was similar between
landuses, I combined genders for final statistical analyses (discussed later).
Sampling Techniques
Terrestrial capture. Playas were partially enclosed (i.e., 25% of circumference)
with 60-cm high drift fence (xiength= 330 m per playa) and 19-L pitfall traps (Dodd and
Scott 1994). Pitfall traps were placed on alternate sides of the fence at 10-m intervals
with openings flush to the ground (Dodd and Scott 1994). Pitfall traps in each landuse
type were checked alternate days for captures from 16 May-17 October 1999 and 19
April-18 August 2000. Pitfall traps within one landuse type were opened near dusk (e.g.,
1800-2100 hrs) before the night of intended capture and processed during the next
14
afternoon (e.g., 1400-1800 hrs). Traps were subsequently closed and reopened in 24 hrs.
Simultaneous with closing of pitfall traps in one landuse type, pitfall traps in the other
landuse type were opened.
Biological processing. Captured individuals were enumerated by species and age
(metamorph [juveniles <1 yr], subadult [>1 yr but not displaying secondary sexual
characters], adult [>1 yr and possessing reproductive morpholog) such as vocal sacs,
nuptial excrescences, or eggs for anurans, and enlarged cloacae for the salamander];
Duellman and Trueb 1994:33-38, 52-60). I did not designate a subadult category for
barred tiger salamander because distinction is based on color (Webb and Roueche 1971).
which can be hydroperiod-dependent in playas (Rose and Armentrout 1976). Snout-vent
length (i.e., tip of snout to posterior of vent; SVL) and mass were measured using calipers
and a spring scale for the first 5 individuals captured per playa per species per age class
per day (« = 2816 and 2372 in cultivation and grassland [years combined]). Individuals
were toe-clipped uniquely using sterilized scissors (soaked in 0.01% chlorhexidine
gluconate); recaptured individuals were not measured. Sampling techniques in this and
subsequent chapters followed approved Texas Tech University Animal Use and Care
Committee protocol (permit #99843).
Statistical Analyses
Multivariate analysis of variance (MANOVA) was used to test for differences (a
= 0.05) in body size within species and age classes between landuse types and years and
among levels of the random nested effect (i.e., playas). I assumed the matrix composed
of SVL and body mass response vectors represented a multivariate description of body
15
size. I used an unrestricted mixed MANOVA model with an unstructured covariance
matrix for factorial-nested designs to account for between- and within-subject
heterogeneity (Littell et al. 1991:120-126, Littell et al. 1996:269, 293, Montgomery
2001:526-527). Consequently, it was not necessary to assume sphericity or other
covariance structures (Littell et al. 1996:99). Because sample size generally was large
(i.e., «>30 individuals per level per effect), I assumed average body size per effect
followed approximately a bivariate normal density (Milton and Arnold 1995:241,
Johnson and Wichem 1998:187). Despite these precautions, I also used Pillai-Bartlett
trace test statistic, because it is more powerful and robust than other statistics (e.g.,
Wilk's likelihood ratio) when multivariate assumptions are violated (Olson 1974, 1976,
1979). I tested additivity of fixed main effects by including an interaction term in the
mixed model (Montgomery 2001:526); analyses were sorted by simple main factors (i.e.,
landuse type by year and year by landuse) with playas nested when this assumption was
violated. Westfall-Young (1993:113-121) multivariate multiple comparison method
(hereafter STEPBOOT) was used to test for simple effect differences in body size
between levels of fixed factors when MANOVA was significant (Westfall et al.
1999:227-239). I did not present results of the random effect (i.e., differences in body
size among playas nested in landuse and year) herein.
Results
Landuse Type and Year Interaction Effects
Adults. Landuse and year effects were not additive for adult Great Plains toad
(MANOVA F=12.6; 2, 593 df; P<0.001), New Mexico spadefoot (MANOVA F=49.8; 2.
16
1541 df; P<0.001), and barred tiger salamander (MANOVA F=7.5; 2, 361 df; i'<0.001);
therefore, I separated analyses by main factors (see next 2 sections). Landuse and year
effects were additive for aduft plains spadefoots (MANOVA F=2.5; 2, 240 df; ^^=0.09);
thus, analyses were not separated by main factors for this species. Body size of adult
plains spadefoots was greater at grassland than at cropland playas (MANOVA F=15.0; 2,
240 df; /'<0.001) and greater in 1999 than in 2000 (MANOVA F=4.1; 2, 240 df;
P=0.02): both SVL and mass were significant (STEPBOOT P<0.001, Table 2.1).
Subadults. Landuse and year effects were not additive for subadult Great Plains
toad (MANOVA F=\9A; 2, 325 df; P<0.001) and New Mexico spadefoot (MANOVA
F=12.3; 2, 383 df; P<0.001); therefore, I separated analyses by main factors. Additivity
of main effects was not tested for subadult plains spadefoot because no individuals were
captured in cropland in 1999; thus, these analyses were separated immediately by main
factors. Analyses were not performed for barred tiger salamander, because there was no
subadult category for this species.
Metamorphs. Landuse and year effects were not additive for metamorph New
Mexico spadefoot (MANOVA F=12.2; 2, 577 df; P<0.00\) and barred tiger salamander
(MANOVA F=7.8; 2, 462 df; P<0.001); therefore, I separated analyses by main factors.
Additivity of main effects was not tested for metamorph Great Plains toad and plains
spadefoots, because no individuals were captured in grassland in 2000. Consequently, I
separated analyses immediately by main factors for these species.
17
Table 2.1. Aduk body size of New Mexico spadefoot {Spea multiplicata. NSF). plains
spadefoot {S. bombifrons. PSF), Great Plains toad {Bufo cognatus. GPT). and barred tiger
salamander {Ambystoma tigrinum mavortium, BTS) between landuse types and \ears
(fixed factorial effects) at 16 playa wetlands (random nested effect) on the Southern High
Plains of Texas, April-September 1999 and 2000.
Size Landuse
Variable Species Year' Cultivation Grassland
n X SE P<0.05' n X SE P<0.05
Snout- NSF 1999 536 42.1 0.1 Aa 409 44.4 0.2 Ba
vent 2000 401 38.7 0.2 Ab 206 43.7 0.2 Bb
length PSF 1999 45 46.1 0.6 Aa 57 49.2 0.7 Ba
(mm) 2000 77 43.9 0.4 Ab 72 46.9 0.4 Bb
GPT 1999 256 72.4 0.5 Aa 164 80.2 0.6 Ba
2000 76 64.9 0.9 Ab 108 75.3 0.8 Bb
BTS 1999 63 92.6 1.8 Aa 107 101.4 0.9 Ba
2000 66 81.3 0.9 Ab 100 86.5 1.2 Bb
Mass NSF 1999 536 8.5 0.1 Aa 409 9.8 0.1 Ba
(g) 2000 401 7.2 0.1 Ab 206 9.4 0.2 Bb
1999 45 11.4 0.5 Aa 57 12.2 0.4 Ba
PSF 2000 77 9.7 0.3 Ab 72 11.5 0.3 Bb
1999 256 51.5 1.1 Aa 164 70.1 1.7 Ba
GPT 2000 76 40.5 1.7 Ab 108 53.8 1.8 Bb
1999 63 39.7 2.8 Aa 107 48.1 1.6 Ba
BTS 2000 66 22.3 0.9 Ab 100 27.1 1.4 Bb
Ânalyzed by year because landuse and year main effects were not additi\e (P<0.001) for
all species except PSF (P=0.09); PSF statistics were presented by year for tabular
parsimony.
Within-species means in the same row with unlike uppercase letters are different (i.e..
simple effect test between landuses within years); means in the same column within size
variables and species with unlike lowercase letters are different (i.e., simple effect test
between years within landuses); all analyses performed using Westfall-Young
(1993:113-121) multivariate multiple comparison test after the simple main effect
MANOVA for factorial-nested designs (Montgomery 2001:569-573) was significant
(P<0.05).
18
Landuse Type Main Effect
Adults. Body size was greater at grassland than at cropland playas for adult Great
Plains toad (MANOVA F=47.2; 2, 411 df; P<0.00\), New Mexico spadefoot (MANOVA
F=9A; 2, 936 df; P<0.001), and barred tiger salamander (MANOVA F=12.5; 2, 161 df;
P<0.001) in 1999; SVL and mass were significant for all species (STEPBOOT P<0.005,
Table 2.1). Body size also was greater at grassland than at cropland playas for adult
Great Plains toad (MANOVA F=18.3; 2, 175 df; P<0.001), New Mexico spadefoot
(MANOVA F=90.6; 2, 598 df; P<Om\), and barred tiger salamander (MANOVA
F=5.4; 2, 157 df; /*<0.005) in 2000; SVL and mass were significant for all species
(STEPBOOT P<0.03, Table 2.1).
Subadults. Body size was greater at grassland than at cropland playas for
subaduh Great Plains toad (MANOVA F=\3.7; 2, 256 df; P<0.001), New Mexico
spadefoot (MANOVA F=\ 12.2; 2, 298 df; P<0.001), and plains spadefoot (MANOVA
F=24.8; 2, 64 df; P<0.00\) in 2000; SVL and mass were significant for all species
(STEPBOOT P<0.001, Table 2.2). Body size did not differ between landuses for
subadult Great Plains toad (MANOVA F=0.7; 2, 62 df; P=0.49) and New Mexico
spadefoot (MANOVA F=2.9; 2, 80 df; P=0.07) in 1999.
Metamorphs. Body size was greater at grassland than at cropland playas for
metamorph Great Plains toad (MANOVA F=l 14.1; 2, 335 df; P<0.001), New Mexico
spadefoot (MANOVA F=19.2; 2, 494 df; P<0.00\), and plains spadefoot (MANOVA
F=21.9; 2, 156 df; P<0.001) in 1999; SVL and mass were significant for all species
19
Table 2.2. Subadult body size of New Mexico spadefoot {Spea multiplicata, NSF), plains
spadefoot {S. bombifrons, PSF), and Great Plains toad {Bufo cognatus, GPT) between
landuse types and years (fixed factorial effects) at 16 playa wetlands (random nested
effect) on the Southern High Plains of Texas, April-September 1999 and 2000.
Size Landuse
Variable Species Year' CLiltivation Grassland
n X SE P<0.05'' n X SE P<0.05
Snout- NSF 1999 26 34.1 0.9 Aa 61 36.2 0.5 Aa
vent 2000 221 32.5 0.2 Ab 86 39.1 0.4 Bb
length PSF 1999 0 0 0 NT 15 38.1 0.8 a
(mm) 2000 55 34.2 0.5 A 18 43.2 0.7 Bb
GPT 1999 41 58.4 0.9 Aa 30 56.5 1.3 Aa
2000 64 47.1 0.8 Ab 201 59.0 0.6 Ba
Mass NSF 1999 26 4.8 0.3 Aa 61 5.3 0.3 Aa
(g) 2000 221 4.6 0.2 Aa 86 6.6 0.2 Bb
PSF 1999 0 0 0 NT 15 6.5 0.4 a
2000 55 4.7 0.2 A 18 7.8 0.4 Bb
GPT 1999 41 23.3 0.9 Aa 30 24.4 1.8 Aa
2000 64 14.7 0.7 Ab 201 26.9 0.7 Ba
Analyzed by year because landuse and year main effects were not additive (P<0.001) for
NSF and GPT; PSF analyzed for 2000 only.
Within-species means in the same row with unlike uppercase letters are different (i.e.,
(/*<0.05); NT indicates no test performed because SE=0 for >1 level of the effect.
20
(STEPBOOT P<0.001, Table 2.3). Body size did not differ between landuses for
metamorph barred tiger salamander (MANOVA F-0.5; 2, 267 df; P=0.62) in 1999.
Body size was greater at grassland than at cropland playas for metamorph barred tiger
salamander (MANOVA F=18.4; 2, 188 df; P<0.001) in 2000; SVL and mass were
significant (STEPBOOT P<0.005, Table 2.3). Body size was greater at cropland than at
grassland playas for metamorph New Mexico spadefoot (MANOVA F=7.9; 2, 78 df;
P<0.001) in 2000; SVL and mass were significant (STEPBOOT P<0.006, Table 2.3).
Year Main Effect
Adults. Body size was greater in 1999 than in 2000 for aduk Great Plains toad
(MANOVA F-15.6; 2, 323 df; P<0.001), New Mexico spadefoot (MANOVA F=l 10.6;
2, 928 df; P<0.001), and barred tiger salamander (MANOVA F=22.5; 2, 120 df;
P<0.001) at cropland playas; SVL and mass were significant for all species (STEPBOOT
P<Om\, Table 2.1). Body size also was greater in 1999 than in 2000 for aduU Great
Plains toad (MANOVA F=63; 2, 263 df; 7^=0.002), New Mexico spadefoot (MANOVA
F=3.1; 2, 606 df; P=0.05), and barred tiger salamander (MANOVA F-65.1; 2, 198 df;
P<Q.OO\) at grassland playas; SVL and mass were significant for all species
(STEPBOOT P<0.05, Table 2.1).
Subadults. Body size was greater in 1999 than in 2000 for subaduh Great Plains
toad (MANOVA F=12.4; 2, 96 df; /'<0.001) and New Mexico spadefoot (MANOVA
F=3.7; 2, 239 df; P=0.03) at cropland playas; SVL and mass were significant for Great
Plains toad (STEPBOOT P<0.001), and SVL was significant for New Mexico spadefoot
(STEPBOOT P=0.04, Table 2.2). Body size was greater in 2000 than in 1999 for
21
Table 2.3. Metamorph body size of New Mexico spadefoot {Spea multiplicata. NSF).
plains spadefoot {S. bombifrons, PSF), Great Plains toad {Bufo cognatus, GPT). and
barred tiger salamander {Ambystoma tigrinum mavortium, BTS) between landuse types
and years (fixed factorial effects) at 16 playa wetlands (random nested effect) on the
Southern High Plains of Texas, April-September 1999 and 2000.
Size Landuse
Variable Species Year" Cultivation Grassland
n X SE P<0.05'' n X SE P<0.05
Snout- NSF 1999 384 28.1 0.2 Aa 119 31.9 0.4 Ba
vent 2000 51 24.5 0.4 Ab 34 22.9 0.4 Bb
length PSF 1999 76 31.8 0.4 Aa 89 36.5 0.4 B
(mm) 2000 27 25.9 0.4 b 0 0 0 NT
GPT 1999 142 25.1 0.7 Aa 202 38.2 0.7 B
2000 39 25.5 1.1 a 0 0 0 NT
BTS 1999 103 77.4 1.0 Aa 173 78.5 0.6 Aa
2000 76 72.4 0.7 Ab 121 79.4 0.7 Ba
Mass NSF 1999 384 2.8 0.1 Aa 119 3.7 0.1 Ba
(g) 2000 51 2.3 0.1 Ab 34 1.2 0.1 Bb
1999 76 3.6 0.2 Aa 89 5.6 0.2 B
PSF 2000 27 2.7 0.1 b 0 0 0 NT
1999 142 2.5 0.2 Aa 202 8.6 0.5 B
GPT 2000 39 2.8 0.3 a 0 0 0 NT
1999 103 21.4 0.9 Aa 173 22.0 0.6 Aa
BTS 2000 76 15.4 0.9 Ab 121 18.8 0.8 Bb
Ânalyzed by year because landuse and year main effects were not additive (PÔ.OOl) for
NSF and BTS; PSF and GPT analyzed for 1999 only.
Within-species means in the same row with unlike uppercase letters are different (i.e.,
(/*<0.001); NT indicates no test performed because SE=0 for >1 level of the effect.
22
subaduh New Mexico spadefoot (MANOVA F=3.2; 2. 139 df: P-0.04) and plains
spadefoot (MANOVA F=3.5; 2, 24 df; P=0.05) at grassland playas; SVL and mass were
significant (STEPBOOT P<0.01, Table 2.2). Body size did not differ between years for
subadult Great Plains toad (MANOVA F=0.2; 2, 222 df; P=0.78) at grassland playas.
Metamorphs. Body size was greater in 1999 than in 2000 for metamorph New
Mexico spadefoot (MANOVA F=17.9; 2, 427 df; P<0.00\), plains spadefoot (MANO\'A
F=37.5; 2. 96 df; P<0.001), and barred tiger salamander (MANOVA F=10.8: 2. 170 df
P<0.00\) at cropland playas: SVL and mass were significant for all species (STEPBOOT
P<0.006, Table 2.3). Body size did not differ between years for metamorph Great Plains
toad at cropland playas (MANOVA F=0.3; 2, 174 df; ^=0.73). Body size was greater in
1999 than in 2000 for metamorph New Mexico spadefoot (MANOVA F=16.4; 2. 145 df:
P<0.001) and barred tiger salamander (MANOVA F=20.7; 2, 285 df; P<0.001) at
grassland playas; SVL and mass were significant for New Mexico spadefoot
(STEPBOOT P<0.00\), and mass was significant for barred tiger salamander
(STEPBOOT P=0.002, Table 2.3).
Discussion
Landuse Effect
Body size of amphibians generally was greater at grassland playas than at playas
surrounded by cultivation for all age classes (metamorph, subadult, and adult) on the
Southern High Plains. To my knowledge, this is the first documentation of the potential
effect of anthropogenic landuse on body size of amphibians in North America. Beebee
(1983) and Oldham (1985) documented in the United Kingdom that body size of Bufo
23
bufo and two species of newts {Triturus vulgaris, T. helveticus) at wetlands in cultivation
was greater than at wetlands with an uncultivated watershed (i.e.. opposite of my results).
Oldham (1985) related his resuhs to postmetamorphic density, where spring influx of
adults was 14X greater at the uncultivated wetland than at the wetland surrounded by
cropland. Similarly, with respect to relative abundance, mean daily capture of
individuals in pitfall traps (all species combined) at cropland playas (x=648. SE=226
[1999]: x=157. SE=29 [2000]) was greater than (P<0.03, Wilcoxon 2-sample
nonparametric test, Conover 1980:216) at grassland playas (x=135, SE=20 [1999]; x=90.
SE=23 [2000]) both years (also see Chapter III). Density of conspecifics, and possibly
congeners, can affect postmetamorphic growth rate and body size of amphibians (Goater
1994, Pechmann 1994. Morey and Reznick 2001) by increasing competition for food
resources.
Beebee (1983) and Oldham (1985) also speculated that body size might be a
consequence of differential food abundance. Although I did not quantify invertebrate
abundance, capture of beetles, springtails, and spiders appeared greater in pitfall traps at
grassland playas than at cropland playas (M. J. Gray and R. Brenes. Texas Tech
University, personal observation). Moreover, mean total mass (g) of invertebrates in
stomachs {n=70) of metamorph Great Plains toads was greater (P=0.005, Wilcoxon test)
for individuals I collected at grassland playas (x=0.047, SE=0.008) than at cropland
playas (x=0.009, SE=0.002). Runoff or aerial drift of insecticides and scarification of
vegetation and soil may negatively affect terrestrial food resources available for
24
amphibians at wetlands surrounded by cultivation (Freemark and Boutin 1995); however,
this h\ pothesis needs to be tested for playa wetlands.
Postmetamorphic body size also may have been affected by conditions in the
larval em ironment (Wilbur and Collins 1973, Werner 1986, Brady and Griffiths 2000,
More> and Reznick 2001). Hydroperiod can affect postmetamorphic body size b}
reducing duration of larval development (e.g., Newman 1988, 1989. Denver et al. 1998).
Mean h\ droperiod (i.e., measured from the date amphibian sampling started until playas
dried) of grassland playas (x=139 d, SE=14) was greater than (P=0.05, Wilcoxon test)
cropland playas (x=89 d. SE=22) in 1999. Although 1 did not detect differences (F=0.18.
Wilcoxon test) between landuse types, mean hydroperiod in grassland playas (x=107 d.
SE=13) was greater than in cropland playas (x=69 d, SE=18) during 2000. Because
rainfall at playas was similar {P>0.32, Wilcoxon test) between landuses (x=6.4 cm,
SE=1.3 [cultivation, 1999]; x=9.2 cm, SE=1.5 [grassland. 1999]; x=4.4 cm, SE=2.2
[cultivation, 2000]; x=4.5 cm, SE=3.8 [grassland, 2000]) both years, hydroperiod
probably was shorter in cropland playas because of differential sedimentation (Luo et al.
1997).
Although I did not quantify potential herbaceous (e.g., nektonic algae. Savage
1952) and animal (e.g., anostracan shrimp. Pfennig 1992) foods between landuse types,
amphibian densities may have been lower in cropland playas because of increased
mortality of these organisms associated with agricultural runoff or drift (Freemark and
Boutin 1995). Reduced aquatic food resources can negatively influence larval growth
(e.g.. Wilbur 1977, Steinwascher 1979, Newman 19946, Walls 1998), and consequently
25
postmetamorphic body size. Sediments and agricultural chemicals also ma> have altered
the percent composition of aquatic invertebrate taxa in cropland playas (Hall 1997:68-69.
85-87), which could have influenced potential predators (Skelly 1997). If predator)
insect density was reduced in cropland playas, competition for food resources may have
been greater than in grassland playas (e.g., Werner and McPeek 1994). perhaps
contributing to reduced postmetamorphic body size. Lastly, agricultural chemicals can
directh reduce foraging acti\'ity and growth of larval amphibians, which can decrease
body size at metamorphosis (Semlitsch et al. 1995, Fioramonti et al. 1997. Boone and
Semlitsch 2001).
Amphibian predator density and size also may have been related positively to
hydroperiods (Skelly 1996). Mean daily capture of barred tiger salamander larvae and
neonates (a potential predator and cannibal. Rose and Armentrout 1976, Collins and
Holomuzki 1984) in seine net samples (0.002 ha sample area) was greater (P<0.01.
Wilcoxon test) in grassland playas (x=108, SE=16 [1999]; x=l 1. SE=4 [2000]) than in
cropland playas (x=26, SE=6 [1999]; x=0.2, SE=0.1 [2000]). Predators can positiveh
influence postmetamorphic body size of anurans by reducing density of conspecific
larvae, thus competition for food resources (Wilbur 1984). Alternatively, predators may
negati\'ely affect postmetamorphic body size of individuals by decreasing foraging
activity and size at metamorphosis (Skelly and Werner 1990), yet this effect may be
species-dependent (Werner 1991, Werner and McPeek 1994) and secondary. I
hypothesize, if predators are influencing postmetamorphic body size in playas, their
presence is primarily reducing competition (e.g., Morin 1983), thus resulting in large
26
postmetamorphic body size associated with predator-rich wetlands (i.e., grassland
playas).
One different resuU existed in my landuse results; body size of metamorph New
Mexico spadefoots was greater at playas surrounded by cultivation than at grassland
playas in 2000. I believe this was a consequence of timing at metamorphosis. New
Mexico spadefoot tadpoles metamorphosed ca. one month earlier from playas in
cultivation (06/28/00) than from grassland playas (07/25/00), which was ca. 25 and 52 d
following breeding, respectively. Thus, metamorph New Mexico spadefoots had more
time to grow postmetamorphically in cropland than in grassland, which likely positiveh
affected body size. Developmental plasticity in response to wetland hydroperiod is well
documented for spadefoot tadpoles (see references in Newman 1992 and Denver 1997).
Presumably, New Mexico spadefoot tadpoles assayed the ratio between their size-specific
mortality and growth rates (Wilbur 1984, Werner 1986) and delayed metamorphosis in
grassland playas, perhaps because water levels were more stable than in cropland playas
(as suggested by my hydroperiod data). Indeed, body size (e.g., SVL) of newly
metamorphosed individuals (i.e., captured the first day) at grassland playas (x=21.9 mm,
SE=0.4) was slightly greater than at cropland playas (x=20.7 mm, SE=0.4), but it
apparently did not compensate for the longer duration of postmetamorphic growth of
individuals in cultivation.
Year Effect
Postmetamorphic body size of amphibians was greater in 1999 (i.e., the wetter
year) than in 2000 generally for all species and age classes. Moreover, the effect of
27
landuse type and year on body size generally was non-additive. Differences in body size
between landuse treatments increased 33-545% between 1999 and 2000 for adult and
subadult New Mexico spadefoot and Great Plains toads, and metamorph barred tiger
salamanders. Rainfall can affect postmetamorphic growth rates (Tinsley and Tocque
1995) and presumably body size (Bruce and Hairston 1990, Reading 1990) by reducing
probability of desiccation (Newman and Dunham 1994) and increasing foraging (Jaeger
1980) and movement activity (e.g., Hurlbert 1969, Semlitsch \9S5b, Sinsch 1988). Mean
number of individuals immigrating and emigrating daily (all species and movement
directions combined) at my playas was greater {P<0.00\, Wilcoxon test) in 1999 (x=391,
SE=115) than in 2000 (x=129, SE=20) for both landuses. Increased opportunity for
movement may increase foraging success of amphibians (Jaeger 1980). Prey abundance
also may increase with rainfall (Dimmitt and Ruibal \9S0a). Diet of metamorphs and
adults of my study species was primarily composed of beetles on the Southern High
Plains (Anderson et al. 19996, L. M. Smith and M. J. Gray, unpublished manuscript).
Coleoptera reproduction in semi-arid environments often is correlated with rain (Crowson
1981:380-381,391-396).
Rainfall also can affect postmetamorphic body size by influencing duration of
larval development (Reading and Clarke 1999, Camp et al. 2000). Mean larval
emergence at my playas was 43.5 and 38.5 d after breeding commenced (determined as
per ancillary call surveys) in 1999 and 2000. Interestingly, this difference in larval
duration between years is attributed to the reduction in developmental time in cropland
(x=36 d, SE=4 [1999], 25 d, SE=2 [2000]) not in grassland playas (x=51 d, SE=2 [1999],
28
52 d, SE=2 [2000]). Thus, postmetamorphic body size may have been different between
years, and landuse and year main effects non-additive, because of greater available food
resources and duration of larval development associated with 1999 and grassland playas.
Body size of subadult New Mexico and plains spadefoots was greater in 2000
than in 1999 at grassland playas, which was contradictory to the above results. Because
body size of subadult anurans reflects growth during the previous year (see Turner 1960),
body size of subadults may have been elevated in 2000 because of additional growth
experienced during more favorable ambient conditions in 1999. However, if this was
true, then subadults captured at cropland playas should have been larger in 2000 than in
1999 (which was not observed). Possibly, the negative effect of cultivation on
postmetamorphic body size of subadults during both years counteracted the positive
benefits of higher rainfall in 1999.
Competing Body Size Hypotheses
Wilbur and Collins (1973) suggested that body size at metamorphosis would be
maintained through postmetamorphic development thus confer adult fitness. Several
empirical studies (e.g., Semlitsch et al. 1988, Berven 1990, Scott 1994) have supported
their prediction. Werner (1986) suggested that body size at metamorphosis (and
ultimately adult body size and fitness) would be a consequence of size-specific mortality
(u) and growth rates {g) in aquatic and terrestrial habitats. Thus, an individual that
metamorphosed at a small body size due to a high larval \i/g ratio might exhibit ''catch-
up" growth because of a low postmetamorphic \i/g ratio. Morey and Reznick (2001) and
Goater (1994) supported this prediction (at least in part). Although I did not measure the
29
same indi\'iduals through time, because individuals were random samples from the
population residing within main effect treatments and body size of amphibians can var>'
seasonally in response to environmental conditions (Reading and Clarke 1995 and
references therein). I can assume that postmetamorphic bod\ size for each age class
represented mean population response of age-specific growth rate to main effects. Thus,
my results seemingly provide support for U'ilbur and Collins' (1973) h\ pothesis. because
differences in body size between levels of main effects generally were maintained
throughout all age classes (i.e.. metamorphs, subadults, and adults were consistenth
larger at grassland playas [thus no catch-up growth exhibited in cultivation]).
Conclusions
Landuse type surrounding wetlands and rainfall can affect postmetamorphic body
size of amphibians on the Southern High Plains of Texas. Individuals captured at
grassland playas generally were larger than those caught at playas surrounded by
agricultural cultivation, and individuals in 1999 (i.e., a wetter year) usually were larger
than those captured in 2000. Moreover, I found the effect of landuse type and year
usually was non-additi\'e; cultivation negati\ eh affected body size greater during the
drier year. Thus, my results suggest that anthropogenic modification of terrestrial
landscapes surrounding wetlands may negatively influence postmetamorphic bod> size of
amphibians, especially during drier years. Density independent and dependent factors
both may be related to landuse and year effects, and proximally responsible for
influencing body size of amphibians. This information is important because
postmetamorphic bod> size can be correlated with individual survival and fitness, thus
30
population persistence of amphibians. Indeed, retention of grasslands surrounding
wetlands may positively benefit amphibians.
31
CHAPTER III
EFFECT OF LANDUSE AND YEAR ON POPULATION
DEMOGRAPHICS OF SOUTHERN HIGH PLAINS AMPHIBIANS
Abstract
Antliropogenic disturbance of landscapes surrounding wetlands is considered a
factor in local and global amphibian declines. Few data exist on the effects of landscape
cultivation on amphibians, and results from previous studies are contradictory. My
objective was to test the effect of landuse (cultivation vs. grassland [control]) on
population demographics of 7 species and 3 age classes of amphibians during 2 years
(1999 vs. 2000) on the Southern High Plains of Texas. I partially enclosed 16 playa
wetlands (4 per landuse per year) with drift fence and pitfall traps, and monitored relative
abundance and diversity daily from 16 May-17 October 1999 and 19 April-18 August
2000. I also monitored immigration and emigration rates to investigate the effect of
landuse and year on possible source-sink dynamics. In general, abundance (i.e., daily
capture rate) of New Mexico and plains spadefoots {Spea multiplicata, S. bombifrons)
was greater at cropland than grassland playas; abundance of other species and diversity of
the amphibian assemblage was not affected by landuse. Also, abundance generally was
greater in 1999 than 2000 for metamorph spadefoots and barred tiger salamanders
{Ambystoma tigrinum mavortium). Frequency of days emigration exceeded immigration
(i.e., source dynamics) was greater at grassland than cropland playas and in 1999 than
2000. Differences in spadefoot abundance between landuse types may have been related
to low species-specific vagility, resulting in increased nestedness within disturbed
32
landscapes and absence of a keystone intraguild predator (i.e.. neotenic or large larval
tiger salamanders) in cropland playas. Yearly variation in metamorph abundance and
frequency of direcfional movements likely was related to armual rainfall (i.e., 1999 >
2000). Landscape disturbance surrounding wetlands influences demographics and
source-sink dynamics of amphibian populations.
Introduction
Amphibian populations are declining globally (Houlahan et al. 2000), and
identifying factors associated with declines is a necessary prelude to conservation efforts.
Anthropogenic modifications of landscapes surrounding wetlands can affect resident
organisms (Findlay and Houlahan 1997). Most research suggests species richness and
abundance of amphibians are negatively associated with anthropogenic disturbance (e.g.,
timber harvesting, agricultural cultivation) of uplands adjacent to wetlands (e.g., Petranka
et al. 1993, Hecnar and M'Closkey 1998, Scribner et al. 2001). However, some recent
studies have documented unexpected positive or neutral associations of amphibians in
disturbed landscapes (Anderson et al. 1999a, Knutson et al. 1999, Kolozsvary and
Swihart 1999). Presumably, landscape disturbance affects amphibians by influencing
probability of survival, reproduction, or interdemic movement.
Landscape disturbance probably affects amphibian population parameters by
physically altering the aquatic (i.e., wetland) and terrestrial (i.e., upland) environments
(Wilbur 1980, Semlitsch 2000). For example, agricultural cultivation can affect water
quality and increase sedimentation in wetlands (Hanson et al. 1994, Luo et al. 1997).
Pesticides in drift and runoff can directly increase mortality of amphibian larvae or
33
indirectly affect survivorship by reducing swimming mobility, thus probability of
predator avoidance and food item acquisition (Bridges 1997, Bridges and Semlitsch
2000, Relyea and Mills 2001). Pesticides also may reduce invertebrate densities in
wetlands (Boone and Semlitsch 2001), which can function as predators, competitors, and
food for amphibian larvae (Morin et al. 1988, Skelly 1997). Herbicides in runoff can
reduce densities of phytoplankton and periphyton (Freemark and Boutin 1995), which
could reduce foraging carrying capacity of amphibian larvae (Wilbur 1997). In contrast,
nitrogen fertilizers can increase net primary productivity in wetlands (Hanson et al.
1994), which might increase foraging opportunities for larval amphibians (Leibold and
Wilbur 1992). Finally, sedimentation decreases wetland volume and hydroperiod (Luo et
al. 1997), which may decrease time available for larvae to develop (Chapter II).
Landscape disturbance also may affect metamorphosed amphibians by
mechanically or chemically altering the terrestrial environment. For example, soil
cultivation can cause direct mortality of fossorial amphibians or it can unbury and expose
them to inhospitable ambient conditions (M. J. Gray, personal observation). Mechanical
disturbance also reduces live and detrital vegetation, which can be important in retaining
soil moisture and ftjnction as foraging, retreat, and borrowing sites for amphibians (Dodd
1996, deMaynadier and Hunter 1998, Herbeck and Larsen 1999, Naughton et al. 2000).
Additionally, mechanical manipulations of the landscape may increase soil compaction
(Welsh 1990), thus decrease burrowing efficiency of amphibians (Jansen et al. 2001),
which can be particularly important for fossorial species inhabiting xeric environments
(McClanahan et al. 1994).
34
Landscape disturbance also can affect connectivity of spatially structured
populafions by influencing permeability to movement (Turner et al. 1989, Wiens 1997,
Gibbs 1998a). For example, agricultural landscapes may be more complex (e.g., contain
greater number of edges to cross. Chapter IV), thus be less permeable to interdemic
movement (Stamps et al. 1987). Agricultural fields also may contain vegetation with less
vertical structure than grasslands or forests thus be less viscous and more permeable to
movement (Wiens et al. 1993, Vos and Chardon 1997). Additionally, pesticides applied
to crops or silvicultural plots may reduce invertebrate abundance for postmetamorphic
amphibians. Finally, the aforementioned factors in the aquatic and terrestrial
environments may individually or synergistically reduce postmetamorphic body size of
amphibians in disturbed landscapes (Chapter II). Populations composed of smaller
individuals can have a greater probability of extinction, because smaller individuals are
less evolutionarily fit than larger individuals (Semlitsch et al. 1988, Berven 1990).
Several studies have been conducted in North America examining the effects of
forest disturbance (e.g., clearcutting, timber thinning) on amphibian populations (e.g.,
deMaynadier and Hunter 1995, Gibbs \99Sb, Herbeck and Larsen 1999, Naughton et al.
2000). Interestingly, few data exist (cf Knutson et al. 1999, Kolozsvary and Swihart
1999) on the effects of agricultural cultivation on amphibians (Dodd 1997:178). It
generally is assumed that agriculture negatively influences populations (Semlitsch 2000).
However, recent evidence suggests that some amphibian populations may become nested
in wetlands within disturbed landscapes (Hecnar and M'Closkey 1997, Wright et al.
1998), thus be positively associated with agricultural cultivation on a local scale
35
(Knutson et al. 1999. Kolozsvary and Swihart 1999). Addifionally. some species (e.g.,
American toad, Bufo americanus Holbrook) may actually benefit from disturbance
(Kolozsvary and Swihart 1999).
Two primary landuse types exist on the Southern High Plains (SHP) of Texas:
cultivation and intact grassland (Haukos and Smith 1994). Amphibians on the SHP
inhabit spatially structured playa wetlands (Bolen et al. 1989). Because playas are
numerous, similar structurally, and exist in their own watershed (Smith and Haukos
2002), they are ideal systems to test the effects of anthropogenic disturbance on
amphibians. My objective was to determine the effect of landscape cultivation around
playa wetlands on amphibian demographics during 1999 and 2000. In agreement with
the status quo (e.g., Berger 1989, Laan and Verboom 1990, Oldham and Swan 1991:147.
Jung 1993, Bonin et al. 1997a, Hecnar and M'Closkey 1998), I hypothesized that playa
amphibian populations would be negatively affected by agricultural cultivation. I also
hypothesized that populations in playas surrounded by cultivation would exhibit sink
dynamics (i.e., immigration > emigration) more frequently than source dynamics (i.e..
emigration > immigration, PuUiam 1988). Thus, I examined whether relative daily
abundance, diversity, immigration, and emigration of 7 species and 3 age classes of
amphibians residing in playa wetlands on the SHP varied by landuse (cultivation vs.
grassland) and between years (1999 vs. 2000) that had different rainfall (Chapter II).
Methods
This study was conducted on the Southern High Plains of Texas (Sabin and
Holliday 1995). I used playa wetlands as experimental units of landuse and year effects.
36
On average, playas located in grassland and cropland were surrounded by >75% natural
or replanted grass and agricultural vegetation, respecfively (Appendix A). I located 16
playas (4 per landuse per year) via aerial and ground reconnaissance in March 1999 and
2000. Because wildlife population dynamics can be highly variable among playas (Smith
and Haukos 2002), I restricted experimental inferences to these landuses and years and
my study playas. A more detailed description of my study area, playas, and years was
presented in Chapter II.
Playas were partially enclosed (i.e., 25% of circumference) with 60-cm high drift
fence and 19-L pitfall traps (Dodd and Scott 1994); a cardinal quadrant was randomly
selected for placement. Fence and pitfalls were placed ca. 10 m upslope from and
parallel to the playa edge (i.e., clay-silt loam line, Luo et al. 1997). One 10-m
perpendicular lead centered at the main fence existed at each end. Vegetation underneath
the main fence and leads was removed, and the bases covered with soil to reduce
probability of trespass (Dodd and Scott 1994). Pitfalls were placed on alternate sides of
the fence at 10-m intervals with openings flush to the ground (Dodd and Scott 1994).
Water and sponges were placed in pitfalls to reduce desiccation of captured individuals
(Daoust 1991). Pitfall traps were checked alternate days in each landuse for captures
from 16 May-17 October 1999 and 19 April-18 August 2000. Pitfalls were open for
23,520 and 12,460 trap nights in 1999 and 2000, respectively.
Captured individuals were enumerated by species and age (i.e., metamorph,
subadult, and adult). Morphological characters used to distinguish among age classes
have been described (Chapter II). Also, I did not designate a subadult category for
37
captured barred tiger salamanders (Chapter II). Individuals were toe clipped uniquely
with respect to study playa and direction of movement (i.e., immigration, emigration)
using sterilized scissors (i.e., soaked in 0.01% chlorhexidine gluconate). Individuals
were assumed to be immigrating and emigrating if captured on the upslope and wetland
side of the fence, respectively (Dodd and Scott 1994). Individuals were released on the
opposite side of the fence after marking (Dodd and Scott 1994).
I used mean daily capture per playa as an estimate of relative daily abundance,
and captures on the upslope and wetland side of the fence as an estimate of relative daily
immigration and emigration of amphibians (Dodd and Scott 1994 and references therein.
Heyer et al. 1994:77). I assumed if any bias existed by using mean daily capture to
estimate relative daily abundance, immigration, and emigration, it w as constant between
landuses and years. It follows that results of ratio tests of significance between levels of
effects would not be affected as per the rules for expectation (Milton and Arnold
1995:53). Mean daily diversity was calculated using the Shannon-Weaver algorithm
(Hair 1980:273). Because individuals were marked uniquely with respect to direction of
movement at capture, trespass rate per direction could be estimated from subsequent
captures. For example, if an individual emigrated during its first and second captures, it
trespassed in the immigrating direction between captures. This enabled me to estimate
percent trespass per direction (M. J. Gray, unpublished data) and correct immigration and
emigration rates prior to analysis. Frequency of days that emigration exceeded
immigration (for all species combined), immigration exceeded emigration, and
emigration equaled immigration was recorded per playa to examine source-sink dynamics
38
of the amphibian assemblage (Pulliam 1988, 1996). I assumed any bias associated with
combining species to examine source-sink dynamics was constant between landuses and
years, thus did not affect resuhs of ratio tests of significance as discussed previously.
The experiment was arranged as a 3-factor factorial design (Montgomery
2001:194-196), where landuse, year, and species captured were crossed-fixed effects. 1
analyzed population demographics for 7 species {Spea multiplicata Cope [New Mexico
spadefoot, NSF], S. bombifrons Cope [plains spadefoot, PSF], Bufo cognatus Say [Great
Plains toad, GPT], B. woodhousii Girard [Woodhouse's toad, WHT], Pseudacris clarkii
[spotted chorus frog, SCF], Rana blairi Mecham et al. 1973 [plains leopard frog, PLF],
Ambystoma tigrinum mavortium Green [barred tiger salamander, BTS]). Two additional
species were captured {B. debilis Girard [green toad], Gastrophryne olivacea Hallowell
[Great Plains narrowmouth toad]); however, there was insufficient capture data for
analysis so they were not used.
I used a 3-factor analysis of variance (ANOVA) model with days as subsamples
to test for differences (a = 0.05) in relative abundance of individuals (i.e., daily capture)
between landuse types and years and among species (Montgomery 2001:194-196). I
used a 2-factor ANOVA model (landuse and year main effects) with days as subsamples
to analyze diversity, because species were combined to calculate the Shannon-Weaver
index thus it was not available as an effect. Nonadditivity was tested by including all 2-
way and the 3-way interaction in the models (Montgomery 2001:194). When
nonadditivity was violated (i.e., >1 interaction significant), analyses were separated by
species to test for differences between landuses and years (i.e., 2-way ANOVAs) and by
39
landuse and year (i.e., 1-way ANOVAs) to test the species effect. For species that
differed in response between levels of landuse or year, I performed a post-hoc set of 2-
way ANOVAs using landuse and year as crossed factors, but by age class, to discern
which demographic category was responsible for the observed abundance differences for
that species. Age was not included as a factor for the initial analyses, because there was
no subadult category for BTS (i.e., design was not balanced) and I assume levels of age
were not independent. Also, this analytical approach conserved experimentwise error
rate at the lowest probability for the main effect ANOVAs, because only significant
species were analyzed in the post-hoc tests for each age class. Tukey's HSD multiple
comparison test was used to test for differences among species when the main effect
ANOVA was significant (Milliken and Johnson 1992:36-38, Westfall et al. 1999:179). 1
natural-log transformed all data to meet linear model assumptions of normality and
homoscedasticity {P > 0.08 as per Shapiro-Wilk and Levene's tests, Milliken and Johnson
1992); means and standard errors were back-transformed for presentation.
I used logistic regression with a generalized logit model to test for differences (a
= 0.05) in frequency distributions of « days that emigration (EM) exceeded immigration
(IM), IM exceeded EM, and EM equaled IM between landuses and years; nonaddivity
was tested by including an interaction term in the model (Agresti 1990:307, Stokes et al.
2000:257- 259). I assumed the categorical response of « days followed a multinomial
distribution per effect, and frequency distributions were independent (Agresti 1990:306).
Normalized chi-square tests of homogeneity (i.e., Z-tests), Bonferroni corrected at a =
0.015, were used to test pairwise differences in frequency distributions when the main
40
effect logit model was significant (Zar 1984:395-396, Milton and Arnold
1995:667-670).
Results
Abundance and Diversity
Mean daily abundance of amphibians was different between landuses and years
and among species (Appendix B). Cropland playas (x=71.5, SE=1.4) had greater
abundance of amphibians than grassland playas (x=22.4, SE=1.4), and abundance was
greater in 1999 (x=67.8, SE=1.4) than 2000 (x-23.5, SE=1.4). However, the species
effect interacted with landuse and marginally interacted with year (Appendix B);
therefore, analyses were separated by species.
Mean daily abundance of NSF and PSF was greater at cropland than grassland
playas, and abundance was greater in 1999 than 2000 for BTS (Appendix B, Table 3.1).
Mean daily abundance also differed among species across and within levels of landuse
and year main effects. New Mexico spadefoot were more abundant than all species at
cropland playas. Mean daily abundance of BTS, GPT, and PSF was greater than SCF,
PLF, and WHT at cropland playas. New Mexico spadefoot, BTS, GPT, and PSF were
more abundant than SCF, PLF, and WHT at grassland playas. Mean daily abundance of
NSF was greater than all other species except BTS in 1999 and 2000. Barred tiger
salamanders were more abtmdant than all other species except NSF and GPT in 1999,
and they were more abundant than other species except NSF, GPT, and PSF in 2000.
Mean daily abundance of GPT was greater than SCF, PLF, and WHT in 1999 and 2000.
41
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Plains spadefoot were more abundant than PLF and WHT in 2000. No differences were
detected in amphibian diversity between landuses and years (Appendix B, Table 3.1).
Mean daily abundance of adult NSF was greater at cropland than grassland
playas; landuse and year effects interacted for metamorph and subadult NSF thus
analyses were separated by main effects (Appendix C, Table 3.2). Metamorph NSF were
more abundant at cropland than grassland playas in 1999, and they were more abundant
in 1999 than 2000 at cropland and grassland playas. Mean daily abundance of subadult
NSF was greater at cropland than grassland playas in 2000, and abundance was greater in
2000 than 1999 at cropland playas. Abundance of metamorph PSF was greater in 1999
than 2000, and adult PSF were more abundant in 2000 than 1999; landuse and year
effects interacted for subadult PSF thus analyses were separated by main effects. Mean
daily abundance of subadult PSF was greater at cropland than grassland playas in 2000,
and abundance was greater in 2000 than 1999 at cropland playas. Metamorph BTS were
more abundant in 1999 than 2000. No additional differences were detected between
landuses and years in mean daily abundance for age classes of significant species
(Appendix C, Table 3.2).
Source-Sink Dynamics
Frequency distributions of n days for the amphibian assemblage that emigration
(EM) exceeded immigration (IM), IM exceeded EM, and EM equaled IM were different
between landuses (xV26.1, P<0.001) and years (xV27.1, P<0.001); main effects did
not interact (% 2^2.8, P=0.25). Percent days EM exceeded IM (i.e., source dynamics)
43
Table 3.2. Relafive daily abundance of age classes between landuse types and years for
amphibian species that differed significantly in daily abundance (see Appendix B) at 16
playa wetlands on the Southern High Plains, Texas, May-October 1999 and
April-August 2000.
Landuse
Species^ Age Year^ Cultivation Grassland
X SE' P<0.05^ X SE P<0.05
NSF Metamorph 1999 100.5 1.7 Aa 3.1 1.8 Ba
2000 1.6 1.5 Ab 2.2 2.1 Aa
Subadult 1999 1.2 1.0 Aa 1.3 1.2 Aa
2000 11.8 1.4 Ab 2.1 1.1 Bb
Aduh 1999 8.9 1.2 Aa 3.5 1.5 Ba
2000 12.2 1.2 Aa 3.2 1.5 Ba
PSF Metamorph 1999 2.6 1.2 Aa 1.9 1.3 Aa
2000 1.3 1.2 Ab 1.0 1.0 Ab
Subadult 1999 1.0 1.0 Aa 1.1 1.0 Aa
2000 3.3 1.4 Ab 1.2 1.0 Ba
Adult 1999 1.3 1.2 Aa 1.3 1.1 Aa
2000 2.9 1.3 Ab 1.6 1.1 Ab
BTS Metamorph 1999 8.9 1.7 Aa 14.5 1.1 Aa
2000 2.6 1.2 Ab 4.7 1.5 Ab
Adult 1999 2.2 1.2 Aa 2.4 1.3 Aa
2000 2.7 1.3 Aa 3.2 1.4 Aa
^ S F = New Mexico spadefoot {Spea multiplicata), PSF = plains spadefoot {S.
bombifrons), and BTS = barred tiger salamander {Ambystoma tigrinum mavortium).
''Analyzed by year for metamorph NSF and subadult NSF and PSF because landuse and
year main effects interacted (Appendix C); statistics for remaining ages and species
presented by year for tabular parsimony.
'^n=A playa wetlands per landuse per year; xs and SEs were back-transformed from
natural-logs for presentation; transformation was necessary to meet linear model
assumptions of ANOVA.
''within-species means in the same row (i.e., within ages and years) with unlike
uppercase letters are different (i.e., landuse effect test); means in the same column within
species and ages with unlike lowercase letters are different (i.e., year effect test).
44
was greater at grassland than cropland playas (Zi=2.7, /'=0.007, Figure 3.1), and greater
in 1999 than 2000 (Zi=2.7, P<0.001). Percent days IM exceeded EM (i.e., sink
dynamics) was greater at cropland than grassland playas (Zi=4.9, P<0.00\), and greater
in 2000 than 1999 (Zi=2.4, P=0.015). Percent days EM equaled IM (i.e., neutral
dynamics) was greater at grassland than cropland playas (Zi=3.1, P=0.002), and greater
in 2000 than 1999 (Zi=2.9, P=0.004). At cropland playas, percent days IM exceeded EM
was greater than percent days EM exceeded IM and EM equaled IM (Zi=8.4-13.5,
P<0.001). Also, percent days EM exceeded IM was greater than percent days EM
equaled IM at cropland playas (Zi=5.6, /*<0.001). At grassland playas and in 1999,
percent days EM exceeded IM and IM exceeded EM were greater than percent days EM
equaled IM (Zi=5.4-10.6, P<0.001), but the former did not differ (Zi=0.9-1.5,
P=0.147-0.322) between each other. In 2000, percent days IM exceeded EM was greater
than percent days EM exceeded IM and EM equaled IM (Zi=8.3-9.3, P<0.00\), but latter
did not differ (Zi=l.l, P=0.292) between each other (Figure 3.1).
Discussion
Mean daily abundance of amphibians was greater at playa wetlands surrounded
by cultivation than at grassland playas, and abundance was greater in 1999 than 2000.
However, when I combined data for all species, frequency of days emigration exceeded
immigration for the amphibian assemblage was greater at grassland than cropland playas;
no yearly differences existed. In contrast, frequency of days immigration exceeded
emigration was greater at cropland than at grassland playas and greater in 2000 than
1999. In general, New Mexico spadefoots were more abundant than all other species,
45
0.6
0.5 :
!
ays 0.4 Ba
Q
Aa Cropland
0.3
Percei
Grassland
#»f#
0.2
0.1
4 •, * ]
0 -—^ * *
EM IM EQ
Directional Movement
11999
12000
EM IM EQ
Directional Movement
Figure 3.1. Effect of landuse (top graph) and year (bottom graph) on frequency of
directional movement of amphibians at 16 playa wetlands on the Southern High Plains,
Texas, May-October 1999 and April-August 2000; EM = percent days emigration
exceeded immigration (i.e., source dynamics), IM = percent days immigration exceeded
emigration (i.e., sink dynamics), EQ = percent days emigration equaled immigration (i.e.,
neutral dynamics); unlike uppercase letters within directional movements and unlike
lowercase letters within landuses are different (P<0.007) by a main effect logistic
regression (generalized logit model) and pairwise chi-square tests of homogeneity
Bonferroni corrected at a = 0.015 (Milton and Arnold 1995:667-670, Stokes et al.
2000:257- 259).
46
followed numerically by barred tiger salamander and Great Plains toad. No difference in
mean daily diversity of amphibians was detected between landuses and years.
Landuse Effect
Abundance of spadefoots may have been elevated at cropland playas, because
landscape disturbance may have confined individuals to the remnant available habitat
(i.e., the playa, Kolozsvary and Swihart 1999). Spadefoots may have become more
nested in cropland wetlands than other species, because of their relative vagility and
perception to patch viscosity and edge permeability in adjacent agricultural fields (Wiens
1997). Indeed, geometric complexity (i.e., mean fractal dimension, FD) and edge density
(i.e., m edge/ha, ED) were greater {P<0.0\, Wilcoxon 2-sample nonparametric test,
Conover 1980:216) in cropland (XFD=1.32, S E = 0 . 0 1 ; XED=64.3, S E = 2 . 0 ) than grassland
(XFD=1-28, S E = 0 . 0 1 ; XED=40.6, S E = 3 . 5 ) landscapes (also see Chapter IV). Road density
also was greater in cropland than grassland landscapes (Appendix A), which can decrease
permeability (Yanes et al. 1995). Thus, dispersing spadefoots may have been unable to
penetrate cropland landscapes, resulting in increased nestedness and abundance at natal
wetlands (Kolozsvary and Swihart 1999).
Spadefoot vagility may have been less than other species on the Southern High
Plains, because of their relatively small postmetamorphic body size (With and Crist
1995). Species-specific perception to landscape context and migration distance is
positively correlated with body size for various organisms (e.g., Peters 1983:89-91, Crist
et al. 1992, With 1994). Excepting the spotted chorus frog, spadefoots were smaller
physically than other species that I monitored (Degenhardt et al. 1996:61, 81, Chapter II).
47
I would expect the spotted chorus frog to be affected similarly by landscape disturbance
if population demographics in playa wetlands were exclusively dictated by the relative
connectivity between them. Differences may not have been detected in demographics
between landuses for spotted chorus frog, because they have relatively low abundance on
the Southern High Plains (Anderson et al. 1999fl, this study). Alternatively, other
proximate factors may be individually or synergistically responsible for demographic
responses to landuse.
Elevated spadefoot abundance in cropland playas also may have been a
consequence of changes in the trophic structure of the aquatic environment. Barred tiger
salamander may fiinction as a keystone intraguild predator {sensu Paine 1969, Polls and
Holt 1992) in playas similar to different species in other wetland systems (e.g.,
Ambystoma opacum. Walls and Williams 2001), because they can establish larval and
neotenic populations prior to anurans if water is present during winter or early spring
(Rose and Armentrout 1974, 1976). Indeed, density of larval and neotenic barred tiger
salamanders was greater in playas surrounded by grassland than in cropland playas,
presumably due to longer hydroperiods associated with grassland playas (Chapter II).
Grassland playas generally have greater volume and longer hydroperiods than cropland
playas because of differential sedimentation (Luo et al. 1997). Consequently, barred tiger
salamander populations became established in grassland playas before anurans bred both
years (M. J. Gray, unpublished data), which may have resulted in top-down control on
spadefoot populations via predation on their eggs and larvae (Henrickson 1990, Walls
and Williams 2001). Insect predators (e.g., Anax, Dytiscus) also may have been abundant
48
in grassland playas due to their longer hydroperiods (Schneider and Frost 1996). In
contrast, cropland playas had shorter hydroperiods and were dry until intense rains early
summer both years, resulting in simultaneous breeding and larval development of
salamanders, anurans, and insects. Thus, salamander and insect larvae likely interacted
with spadefoot larvae as competitors not predators in cropland playas (Morin 1983.
Morin et al. 1988). Other predators of amphibian larvae (e.g., wading birds, fish) occur
infrequently in playa wetlands during summer (Bolen et al. 1989); thus, cropland and
grassland playas may have represented relatively predator-free and -rich environments,
respectively (Heyer et al. 1975). The aforementioned may have facilitated greater
recruitment of metamorphs into spadefoot populations located in cropland than in
grassland.
Spadefoot larvae may have been more susceptible to predation in grassland pla\ as
than other species on the Southern High Plains, because of their relative palatabilit\ and
escape probability (Alford 1999:260-264). Spadefoots typically inhabit xeric
environments worldwide and exploit ephemeral water sources with few vertebrate or
invertebrate predators (Bragg 1965, Dimmitt 1975). Consequently, spadefoots have
evolved physiological mechanisms for rapid growth (Richmand 1947, Newman 1992) but
lack the ability (e.g., toxicity, cryptic coloration, altered microhabitat use) to efficienth
escape predators (e.g., Spea bombifrons, Kruse and Francis 1977). For example,
spadefoot tadpoles are more active than tadpoles of certain Bufo, Rana, and Pseudacris
species; thus, they are more conspicuous and vulnerable to predators (Morin 1983.
Woodward 1983, Dayton and Fitzgerald 2001). Spadefoot tadpoles also lack dermal
49
toxins and cryptic coloration unlike many species of Bufo, Rana, and Pseudacris (e.g.,
Walters 1975, Formanowicz and Brodie 1982, Peterson and Blaustein 1991, Smith and
VanBuskirk 1995).
Spadefoots also may be superior competitors to other species on the Southern
High Plains (e.g., Morin 1983, Wilbur 1987, Dayton and Fitzgerald 2001). Thus, in the
absence of predation, spadefoots might outcompete other species for food and other
resources resulting in increased recruitment (Woodward 1982, Wilbur 1987, Dayton and
Fitzgerald 2001). Indeed, several elegant lab experiments have documented that
spadefoot tadpoles are competitively dominant in anuran communities when aquatic
predators are absent (e.g., Morin 1981, 1983, Wilbur 1987). Unfortunately, no studies
have been performed on the relative palatability, escape probability, or competitive
ability among all my species simultaneously; thus, I only can speculate that
aforementioned hypotheses are responsible for differential demographics in spadefoot
populations between landuses on the Southern High Plains.
Additional modifications of the aquatic and terrestrial envirormients also may
have positively influenced spadefoot populations at cropland playas. Smith and Haukos
(2002) noted that plant diversity and structure could be greater in cropland playas, which
could increase food resources and escape cover for larvae. Nitrogen influx from
fertilizers may have increased food resources for larval amphibians in cropland playas
(Leibold and Wilbur 1992). Additionally, pesticide drift and runoff into cropland playas
may have reduced aquatic insect densities (Boone and Semlitsch 2001), which can
compete with amphibian larvae for food resources (Morin et al. 1988). Similarly,
50
cultivafion and chemical application may have reduced invertebrate densities and cover
in the surrounding landscape near cropland playas (Freemark and Boutin 1995), inducing
greater nestedness in remnant available habitat within disturbed landscapes as mentioned
previously (Kolozsvary and Swihart 1999). However, if any of the aforementioned
suggestions were strictly true, I should have observed similar demographic responses by
species other than spadefoots. Thus, these hypotheses probably are not ultimate factors
responsible for observed demographics in our disturbed and undisturbed landscapes.
Lastly, subadults were the only age class of plains spadefoots that was greater in
cropland than grassland playas. I am uncertain why the other age classes did not respond
similarly considering the aforementioned logic concerning spadefoot response; however,
it may have been a consequence of delayed development or competition. Subadults were
individuals >1 year that did not exhibit sexual characteristics thus probably did not breed
(Chapter II). Amphibians may forego breeding if resource conditions are not adequate
(Semlitsch et al. 1996). Perhaps, cropland landscapes contained less food and other
resources than grassland landscapes (Freemark and Boutin 1995), resulting in greater
number of individuals at cropland playas that did not breed. However, if this was true, I
should have observed similar elevated subadult levels with other species. Alternatively,
increased New Mexico spadefoot abundance at cropland playas may have induced
delayed breeding of plains spadefoots, because these species are known to compete for
resources (Pfermig and Murphy 2000). Thus, abundance of subadult plains spadefoots
may have been a consequence of interspecific competition with New Mexico spadefoot
(see Chapter V).
51
Year Effect
Yearly differences in abundance occurred because of a greater number of
metamorph New Mexico and plains spadefoots, and barred tiger salamanders in 1999
than 2000. Demographic differences between years likely were related to rainfall
(Berven 1995, Semlitsch et al. 1996). Average monthly rainfall at my playas was greater
in 1999 than 2000 (Chapter II). Rainfall stimulates amphibian emergence (Dimmitt and
Ruibal \9S0b) and positively affects intra- and inter-demic movement (Hurlbert 1969,
Sinsch 1988). Adult survival also can be positively related to rainfall (e.g., Berven
1990), presumably because density of food resources increases (Dimmitt and Ruibal
1980a) and probability of desiccation decreases (Jaeger 1980, Newman and Dunham
1994). Probability of breeding, larval survival, and juvenile recruitment also increases
with rainfall, because wetland hydroperiods are positively correlated with rainfall
(Pechmann et al. 1989, Sinsch and Seidel 1995, Semlitsch et al. 1996). Indeed, mean
duration of hydroperiods at my study playas was greater in 1999 than 2000 (Chapter II).
Lastly, the difference in yearly abundance of metamorph spadefoots occurred only at
cropland playas, suggesting that disturbance may affect recruitment greater during drier
years (Lannoo 1998).
Three inconsistencies existed in my year results; abundance of subadult New
Mexico and plains spadefoots was greater in 2000 than 1999 at cropland playas, and adult
plains spadefoots were more abundant in 2000 than 1999. Subadult spadefoot abundance
probably was elevated in 2000 because of increased recruitment experienced in 1999 at
cropland playas (Semlitsch et al. 1996, Meyer et al. 1998). Also, food densities may have
52
been lower during the drier year (i.e., 2000) at cropland playas (Dimmitt and Ruibal
1980a), resulting in more individuals not breeding and consequently classified as
subadults. I am uncertain why adult plains spadefoots were more abundant during 2000
in both landuses, but perhaps drier conditions during this year facilitated competitive
dominance over some species.
Source-Sink Dynamics
Frequency of days emigration exceeded immigration (i.e., source dynamics) for
the assemblage was greater at grassland than cropland playas and in 1999 (i.e., the wetter
year) than 2000. Grassland playas may have served as sources more often than cropland
playas, because of their longer hydroperiods (Chapter II). Consequently, larvae may
have metamorphosed more continuously from grassland than cropland playas, facilitating
the former to function as a more constant source of emigrating propagules. Also, abiotic
conditions (e.g., relative humidity) may have been more ideal in landscapes surrounding
grassland than cropland playas for interdemic movement, particularly the exodus of
individuals (Bardsley 1998).
Source dynamics likely were greater in 1999 than 2000, because favorable
ambient conditions resulted in greater production of individuals (Semlitsch et al. 1996).
Lannoo (1998) suggested that source dynamics would occur more frequently during
relatively wet years in seasonal and semipermanent wetlands. Indeed, catastrophic
mortality or reduced recruitment of individuals may occur in ephemeral wetlands if they
dry before larvae can attain a critical body-size threshold for metamorphosis (Semlitsch
19876, Newman 1992, Denver 1997).
53
Conservation Implications
Anthropogenic disturbance around wetlands can affect demographics and
community structure of amphibian populations. Less vagile species, such as spadefoots
and chorus frogs, may be affected more by disturbance, resulting in greater nestedness of
their populations near natal wetlands. Thus, I caution exclusive interpretation of my
abundance results, because elevated values could be a consequence of area-restricted
movement or reduced home ranges. Moreover, more individuals does not necessarily
imply better conditions for a population (Yeargers et al. 1996). Populations with elevated
abundance may be less stable, particularly if they exceed their carrying capacity or have a
high intrinsic rate of increase (Edelstein-Keshet 1988). Pathogenic incidence also can be
positively correlated with population density (Hastings 1996). Additionally, landscape
disturbance may negatively affect genetic structure of populations (Reh and Seitz 1990)
and fitness correlates, such as postmetamorphic body size of amphibians (Chapter II).
My source-sink results for combined species also suggest that playas in cultivated
landscapes may serve as sinks more often than sources, perhaps because of wetland
hydroperiod and connectivity is less than in undisturbed landscapes. Considering the
aforementioned, amphibian assemblages in disturbed landscapes may have a greater
probability of local or metapopulation extinction than those positioned in natural
landscapes (see Chapter VI also).
Additional research is needed to discern proximate and ultimate mechanisms
driving population responses to anthropogenic disturbance. For example, the effect of
chemical and mechanical disturbance in and around wetlands on the trophic structure of
54
invertebrate and amphibian communities needs to be investigated. Pathogenic incidence
in amphibians (i.e., viruses, fungi, bacteria, and parasites) also should be quanfified in
disturbed and undisturbed wetland systems. Estimates of landuse type and boundary
permeability to amphibian movement also are needed. This latter suggestion would
enable landscape ecologists to assign permeability and hardness indices to cover types
and boundaries in spatial analysis programs (e.g., FRAGSTATS, McGarigal and Marks
1995), and subsequently estimate relative connectivity of habitat patches and extinction
probabilities for local populations. Finally, my year results suggest that annual rainfall
may be an important variable to include in extincfion probability models for amphibians.
55
CHAPTER IV
INFLUENCE OF LANDSCAPE STRUCTURE ON COMMUNITY
COMPOSITION AND RELATIVE ABUNDANCE OF AMPHIBIANS
Abstract
Landscape structure can influence local dynamics of spatially structured
populations, particularly with less \agile organisms such as amphibians. I examined the
effect of landscape structure on communit}' composition and relative abundance of
amphibians inhabiting the Southern High Plains of Texas. Amphibian populations were
monitored using pitfall traps and drift fence located at 16 playa wetlands (8 playas/year)
in 1999 and 2000. I quantified landscape structure surrounding each playa via estimating
13 spatial metrics that represented playa isolation and landscape complexity using remote
sensing from aerial photographs and geocorrection and spatial analysis software.
Multivariate ordination and univariate correlations and regressions indicated that
landscape structure influenced {P<0.05) community composition and mean daily
abundance of amphibians. Spadefoots {Spea multiplicata, S. bombifrons) generally were
positively associated with metrics representing playa juxtaposition and landscape
complexity. Great Plains toads {Bufo cognatus) and barred tiger salamanders
{Ambystoma tigrinum mavortium) usually were negatively associated with spadefoots but
not affected by landscape structure. Differences in community composition and relati\ e
abundance probably were related to species-specific vagility and perception to patch
viscosity and edge permeability. Spatial separation of these species in the ordination also
may have been a consequence of differential competitive ability for food or other
56
resources in aquatic and terrestrial environments. Results in Chapter III suggested
general landuse (i.e., cultivation or grassland) could influence demographics of
amphibians. These additional findings indicate that landscape structure may be as or
more important than general anthropogenic landuse in influencing demographics of
amphibian populations. I also present multiple regression models for predicting relative
species-specific daily abundance using field estimates of spatial metrics. This
information can be used by landscape ecologists to locate potential sites for amphibian
conservation initiatives.
Introduction
Landscape structure can influence demographics of spatially structured
populations via influencing probability of interdemic movement (Fahrig and Paloheimo
1988). Components of landscape structure include spatial positioning of habitat patches
and the geometric matrix in which they are embedded (Fahrig and Merriam 1994).
Relative abundance of wildlife generally declines as habitat patches become isolated or
are not optimally juxtaposed (Lefkovitch and Fahrig 1985, Burel 1989). Moreover, as
landscapes become more geometrically complex, probability of successful dispersal may
decrease because boundary density increases (Stamps et al. 1987, Wiens 1997).
Consequently, isolated populations embedded in complex landscapes may have a greater
probability of extinction than those more optimally positioned within simpler landscapes
(Fahrig and Merriam 1985, Taylor et al. 1993).
Amphibian populations are declining worldwide (Houlahan et al. 2000).
Anthropogenic modifications of habitat patches and the landscape matrix have been
57
implicated in many declines (Blaustein et al. 1994). Various studies have demonstrated
amphibian abundance is negatively related to wetland or terrestrial habitat isolation (e.g.,
Marsh and Pearman 1997. Pope et al. 2000, Scribner et al. 2001). Anthropogenic
modifications of the landscape (e.g., cuhivation) may induce isolation by disrupting
connectivity among patches (Marsh and Trenham 2001). Connectivity in disturbed
landscapes might be negatively affected because of differential viscosity of
anthropogenic cover types or permeability of edges (Durelli et al. 1990. With 1994,
Wiens 1997). Consequently, amphibian abundance and probability of persistence in
wetlands surrounded by disturbance may be lower than in undisturbed wetlands (Sjogren
1991, Sjogren-Gulve 1994). Alternatively, cultivated landscapes may possess greater
edge density and less terrestrial habitat, resulting in increased nestedness and abundance
of amphibians in remaining remnant wetlands (Knutson et al. 1999, Kolozsvary and
Swihart 1999, Chapter III).
On the Southern High Plains (SHP), amphibians exist primarily in playa wetlands
(Bolen et al. 1989). General landscape use (i.e., cultivation vs. grassland) can affect
postmetamorphic body size and demographics of amphibian populations in playas
(Chapters II and III); however, the effect of landscape structure on community
composition and relative abundance of amphibians in playas is unknown. Because
amphibians can interact among playas via dispersal (M. J. Gray, unpublished data),
landscape structure may be an important factor influencing local population dynamics.
Thus, I examined the influence of landscape structure on amphibian populations via
estimating various spatial metrics and relating them to relative daily abtmdance of 4 SHP
species {Spea multiplicata Cope [New Mexico spadefoot], S. bombifrons Cope [plains
58
spadefoot], Bufo cognatus Say [Great Plains toad], Ambystoma tigrinum mavortium
Green [barred figer salamander) at 16 playas (8/year) during 1999 and 2000. I
hypothesized that relative abundance of amphibians would be negatively influenced b>'
playa isolation (Loman 1988, Laan and Verboom 1990, Vos and Stumpel 1995, Lehfinen
et al. 1999). I also hypothesized that landscape complexity would be positively related to
abundance (Knutson et al. 1999, Kolozsvary and Swihart 1999). Understanding the
effects of landscape structure on population dynamics is a ftjndamental construct of
landscape ecology, and a critical prelude to conservation initiatives for amphibians
(Marsh and Trenham 2001).
Methods
This study was conducted at 16 playa wetlands on the Southern High Plains of
Texas. Playas were partially enclosed (i.e., 25% of circumference) with 60-cm high drift
fence and 19-L pitfall traps (Dodd and Scott 1994). Pitfall traps were checked alternate
days for captures from 16 May-17 October 1999 and 19 April-18 August 2000.
Captured amphibians were enumerated by species, marked uniquely, and released. A
more detailed account of my study area, amphibians, and sampling protocol was
presented in Chapters II and III.
I quantified landscape structure surrounding each study playa using remote
sensing from aerial photos and geographical information system and spatial analysis
software. Landscapes («=16) were 2,830-ha circular plots (i.e., 3-km radius) with their
origins positioned at the center of each study playa (Appendix D). I chose a 3-km radius
to delineate the landscape plot, because this distance probably was the near maximum
59
dispersal distance for my species (cf Gehlbach 1967, Gehlbach et al. 1969, Sinsch 1990,
1997, Miaud et al. 2000). Aerial images of my landscapes in summers 1999 and 2000
were obtained from the Farm Service Agency (FSA) of the U.S. Department of
Agriculture in Crosby, Floyd, Hale, and Castro counties, Texas. Ground control points
were identified on images and U.S. Geological Survey 7.5-minute quadrangle maps then
geo-corrected using ERDAS® software. Geo-referenced images were mosaicked in
ERDAS® and cover types (i.e., crop type, replanted Conservation Reserve Program
grass, and natural grass) were identified in each landscape plot using 1999 and 2000 FSA
farm folders. Boundaries of cover types were digitized in ERDAS® then exported as an
ESRI® ARC/INFO coverage. Coverages were cleaned, built, and polygons classified as
per cover types in ESRI® ARC/INFO. Landscape structure was quantified per plot using
FRAGSTATS* ARC® {www.innovativegis.com, McGarigal and Marks 1995); I used the
following 13 spatial metrics. Shape index of the study playa (PSI), study playa size (PS),
mean nearest-neighbor distance from the study playa to surrounding playas (PNN), mean
nearest-neighbor distance from all playas to each other (MNN), number of playas (NP),
percent aerial coverage of playas (PP), and an interspersion/juxtaposition index of playas
(IJI) were used to quantify relative spatial positioning and isolation of playa wetlands
(McGarigal and Marks 1995). Mean number of edges to cross from the study playa to
surrounding playas (PED), edge density (m edge/ha, ED), landscape shape index (i.e.,
measure of geometric complexity, LSI), landuse (i.e., cover type) richness (LR), Shannon
evenness index of landuses (SEI), and Shannon diversity index of landuses (SDI) were
used as measures relative landscape complexity (McGarigal and Marks 1995). I direct
readers to McGarigal and Marks (1995) for spatial metric formulae and algorithms.
60
Unity was assigned to cover type and edge permeability in FRAGSTATS* ARC®
(McGarigal and Marks 1995), because relafive viscosity for my species is unknown.
I used mean daily capture per species per playa as an index of relative daily
abundance (Dodd and Scott 1994). Canonical correspondence analysis (CCA) was used
to examine the effect of landscape structure metrics on relative abundance of species in
the amphibian assemblage (ter Braak 1986, 1994). Correlated metrics (P<0.05) were
removed prior to CCA to reduce probability of an arch effect (ter Braak 1995:139).
Because CCA is sensitive to outliers and bimodally distributed data, mean daily
abundance per species was natural-log transformed prior to analysis (ter Braak 1995); all
resulting distributions were unimodal with <2 outliers. A global Monte Carlo
permutation test was performed to test for existence of a relationship between landscape
V
metrics and species composition (ter Braak and Smilauer 1998:47-49, 124-125). A
dimensionless species-landscape metric biplot was constructed using CANOCO® to
graphically examine the pattern of variation in relative species abundance with landscape
metrics (ter Braak 1995, ter Braak and Smilauer 1998). Biplot axes were interpreted by
examining the sign and magnitude of their respective intra-set correlations and using
subject-matter knowledge (ter Braak 1995:140). Length of metric-specific eigenvectors
(i.e., the arrows) in the biplot was interpreted as the strength of correlation between the
metric and species abundance (ter Braak 1995:141-142). Therefore, long eigenvectors
were most important in affecting the species assemblage (ter Braak 1995:141). Relative
correlative ranking of species with respect to the metrics was graphically represented by
extending each eigenvector through the origin of the biplot and intersecting it with
orthogonal lines drawn from the species (ter Braak 1995:143). Species positioned near
61
the arrow- and blunt-end of the eigenvector were most positively and negafively
correlated with the metric, respecfively (ter Braak 1995:141-143).
Canonical correspondence analysis in CANOCO® does not provide univariate
measures of association between individual metrics and species-specific abundance (ter
Braak and Smilauer 1998). Thus, I also calculated Pearson coefficients of correlafion and
tested the relationship of each metric with species-specific abundance (Milton and Arnold
1995:425-430). Metrics that were correlated (P<0.05) with species-specific abundance
also were regressed linearly using least-squares estimation, and univariate prediction
models developed (Milton and Arnold 1995:386-391). Finally, I used multiple linear
regression with stepwise selection (entry and stay significance level at a=0.15) to develop
models for predicting relative daily abundance per species using all landscape metrics
and 2 indicator variables (i.e., general landuse [O=cropland, l=grassland] and year [0=wet
year, l=dry year]) as potential explanatory variables (Milton and Arnold 1995:497-501,
504-506). These indicator variables were included in prediction model development,
because results in Chapter III suggested that general landuse and annual rainfall could
affect population demographics of amphibians. Linear dependency (i.e., coUinearity
[intercept adjusted]) among predictor variables in the final models was examined via
estimating variance inflation factors (VIF) and condition numbers (CN); VIF > 10 and
CN > 30 were suggestive of coUinearity between >1 variable (Freund and Littell
2000:98-101). Standardized and unstandardized parameter estimates were presented for
linear interpretation and predicting relative daily abundance of amphibians using field
estimates of metrics or indicator variables, respecfively (Myers 1990:384-385).
Coefficients of determination adjusted for number of variables in the model (i.e., R adj)
62
were presented as a measure of model performance (Freund and Littell 2000:23). Mean
daily abundance per species were natural-log transformed for all correlation and
regression analyses to satisfy (P>0.05 as per Shapiro-Wilk tests) linear model
assumpfions (Milton and Arnold 1995:391-392).
Results
Six of the 13 landscape metrics (PNN, NP, PED, ED, LSI, and SDI) were not
used in the canonical correspondence analysis, because they were correlated with other
metrics (Table 4.1). The global Monte Carlo permutation test based on 199 permutations
revealed that species composition was associated (F=2.201, P=0.05) with landscape
metrics. The first two canonical axes were most important in explaining variation in the
species assemblage (i.e., as per respective eigenvalues, ?ii=0.061, ?t2=0.022, >.3=0.001,
X4=0.013); they collectively explained 67.9% of variation in mean daily abundance of
these SHP amphibian species. Intra-set correlations suggested that axes 1 and 2
explained variation in species abundance with respect to wetland positioning and
landscape complexity, respectively, because of the large absolute magnitude of PP and IJI
for axis 1 and SEI for axis 2 (Table 4.2). The dimensionless species-environmental biplot
of axes 1 (ordinate) and 2 (abscissa) suggested that SEI was most positively correlated (as
per eigenvector length) with the pattern of species abundance (Figure 4.1a). Playa size,
PP, IJI, and LR were moderately correlated and PSI and MNN least correlated with
species composition patterns. Orthogonal inferred ranking of species against
eigenvectors suggested that both spadefoot species were most positively associated with
PS, PP, IJI, and LR; barred tiger salamander and Great Plains toad generally were
63
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65
Table 4.2. Intra-set correlations of landscape^ metrics associated with the first two axes
generated by a canonical correspondence analysis of mean daily abundance of New
Mexico spadefoot {Spea multiplicata), plains spadefoot {S. bombifrons). barred tiger
salamander {Ambystoma tigrinum mavortium). and Great Plains toad {Bufo cognatus) at
16 playa wetlands on the Southern High Plains, Texas, 1999 and 2000.
Metric^ Correlations^
Axis 1 Axis 2
PSI 0.0202 -0.1854
PP -0.6970 0.1859
IJI -0.7625 -0.0788
MNN 0.1489 -0.0134
LR -0.5559 -0.2485
SEI 0.0092 -0.7729
PS -0.3993 0.4379
Landscapes {n=\6) were 2830-ha circular plots (i.e., 3-km radius) with their origins
positioned at the center of each study pla\a.
PSI = shape index of study playa. PP = percent aerial coverage of playas. IJI =
interspersion/juxtaposition index of playas, MNN = mean nearest-neighbor distance from
all playas to each other. LR = landuse richness, SEI = Shannon evenness index of
landuses, and PS = study playa size (McGarigal and Marks 1995).
'Întra-set correlations are standardized and dimensionless thus the\ can be interpreted as
the strength and direction of metric-specific correlation and species abundance in the
presence of all other metrics (ter Braak 1995:140): I infer that axes 1 and 2 explained
variation in species abundance with respect to wetland positioning and landscape
complexity because of the large absolute magnitude of PP and IJI for axis 1 and SEI for
axis 2.
66
67
Figure 4.1. Canonical correspondence analysis of mean daily capture (natural-log
transformed) of amphibians and uncorrelated metrics in landscapes (i.e., 2,830-ha circular
plot) associated with 16 study playas on the Southern High Plains, Texas, 1999 and 2000.
a) Species-environmental (i.e., -landscape metric) biplot (ter Braak 1995:142); axes 1
(ordinate) and 2 (abscissa) are dimensionless and represent wetland positioning and
landscape complexity, respectively (see corresponding intra-set correlations in Table 2);
length of eigenvectors (i.e., their respective eigenvalue) indicates the strength of
correlation between the variable and pattem of variation in species composition (ter
Braak 1995:141); species more close to ends of eigenvectors are more positively
correlated with it; PS = study playa size, SEI = Shannon evermess index of landuses, PP
= percent aerial coverage of playas, PSI = shape index of study playa, MNN = mean
nearest-neighbor distance from all playas to each other, IJI = interspersion and
juxtaposition index of playas, and LR = landuse richness (McGarigal and Marks 1995).
b) Inferred ranking of species along variables based on biplot interpretation of Part a of
figure (ter Braak 1995:143); ranking constructed by extending eigenvectors through the
origin and intersecting with orthogonal lines from the species to the vector; the vertical
segment represents the origin (i.e., centroid) of the biplot and is the grand mean of each
variable; species more close to the arrow or blunt end are positively or negatively
correlated with the variable, respectively. NSF = New Mexico spadefoot {Spea
multiplicata), PSF = plains spadefoot {S. bombifrons), GPT = Great Plains toad {Bufo
cognatus), and BTS = barred tiger salamander {Ambystoma tigrinum mavortium).
a)
PS
PP
' ..^^^pspy i GPT
•
•
IJI ^,..-'^^^?.Y
I
PSI
LR
-
' jSEI
I r — - - - " - T — - t f T • T ---—,-- f 1 f T T
-1.0 + 1.0
p N G B P G B N
s S P T S P T S
PS F T S F T S F SEI
b)
P N G B p G B N
S S P T s P T S
PP F F T S F T S F PSI
^ - •
N P G B P N G B
S S P T S S P T
IJI F F T S F F T S MNN
^4- —•
N P B G
S S T P
LR F F S T
68
negatively associated with these spatial metrics (Figure 4.\b). In contrast, barred tiger
salamander and Great Plains toad were most positively and the spadefoots most
negatively related with MNN. Plains spadefoot also appeared to be negatively associated
with SEI and PSI (Figure 4.1^?).
Mean daily abundance was correlated linearly with PP, IJI, ED, LSI, PED. and
LR for New Mexico spadefoot and the first 4 aforementioned metrics for plains
spadefoot; significant correlations were not detected for Great Plains toad or barred tiger
salamander (Table 4.3). Approximately 26%, 27%, 45%, 47%, 48%, and 53% of the
variation in mean daily abundance of New Mexico spadefoots was explained by PP
(Fi,14=4.82, P=0.046), PED (Fi,,4=5.14, P=0.04), LR (Fi.,4=11.41, 7^=0.005), LSI
(Fi.14=12.49, P=0.003), ED (Fi,i4=13.04, P=0.003), and IJI (Fi,i4=15.6, F=0.001),
respectively (Figure 4.2). Approximately 30%, 33%, 35%, and 35% of the variation in
mean daily abundance of plains spadefoots was explained by ED (Fi,14=6.03, F=0.028),
LSI (Fi,14=6.78, F=0.021), IJI (Fi,i4=7.46, F=0.016), and PP (Fi,i4=7.55, F=0.016),
respectively (Figure 4.3).
Multiple linear regression models explained 38-78% of the variation in mean
daily abundance for these amphibian species (Table 4.4). Approximately 78% of the
variation in mean daily abundance of plains spadefoot was explained by PP, SEI, PSI, IJI,
and PNN (F5,io=l 1 -62, F<0.001). Approximately 73% of the variation in mean daily
abundance of New Mexico spadefoot was explained by LU, SDI, MNN, and PSI
(F4,i 1=11.02, F<0.001). Forty-one percent of the variation in mean daily abundance of
Great Plains toad was explained by MNN, PSI, NP, and PNN (F4,ii=3.59, F=0.042).
69
Table 4.3. Univariate Pearson coefficients of correlation between mean daily abundance
of amphibians and landscape^ metrics at 16 playa wetlands on the Southern High Plains.
Texas, 1999 and 2000.
Species'^
Variable'' NSF PSF BTS GPT
r P r P r P r P
PSI -0.138 0.612 -0.175 0.515 -0.102 0.708 -0.284 0.286
PS -0.011 0.970 0.274 0.304 -0.477 0.062 -0.138 0.611
PNN -0.199 0.459 -0.284 0.287 0.102 0.708 -0.166 0.538
MNN -0.405 0.119 -0.299 0.259 -0.322 0.224 -0.414 0.111
NP 0.432 0.095 0.388 0.138 -0.023 0.933 0.082 0.763
PP 0.506 0.046 0.592 0.016 -0.352 0.182 -0.073 0.787
IJI 0.726 0.002 0.589 0.016 -0.036 0.896 0.063 0.816
PED 0.518 0.039 0.457 0.075 -0.353 0.180 -0.186 0.492
ED 0.694 0.003 0.548 0.028 -0.234 0.383 0.048 0.858
LSI 0.687 0.003 0.571 0.021 -0.256 0.338 0.046 0.863
LR 0.670 0.005 0.249 0.351 -0.085 0.754 0.247 0.356
SEI 0.219 0.414 -0.473 0.065 0.104 0.701 -0.077 0.777
SDI 0.421 0.105 -0.297 0.263 0.055 0.838 0.032 0.905
landscapes (« = 16) were 2830-ha circular plots (i.e., 3-km radius) with their origins
positioned at the center of each study playa.
PSI = shape index of study playa, PS = study playa size, PNN = mean nearest-neighbor
distance from study playa to surrounding playas, MNN = mean nearest-neighbor distance
from all playas to each other, NP = number of playas, PP = percent aerial coverage of
playas, IJI = interspersion/juxtaposition index of playas, PED = mean number of edges to
cross from study playa to surrounding playas, ED = edge density (m edge/ha), LSI =
landscape shape index (i.e., measure of geometric complexity), LR = landuse richness,
SEI = Shannon evenness index of landuses, and SDI = Shannon diversity index of
landuses (McGarigal and Marks 1995).
"^NSF = New Mexico spadefoot {Spea multiplicata), PSF = plains spadefoot {S.
bombifrons), BTS = barred tiger salamander {Ambystoma tigrinum mavortium), and GPT
= Great Plains toad {Bufo cognatus).
70
71
Figure 4.2. Simple linear regression and 95% confidence bands of mean daily abundance
(natural-log transformed) of New Mexico spadefoot {Spea multiplicata) and percent
aerial coverage of playa wetlands (PP), interspersion/juxtaposition index (IJI, McGarigal
and Marks 1995:103), edge density (ED, McGarigal and Marks 1995:106), playa edge
density (PED [i.e., mean number of edges to cross from study playa to surrounding
playas), landscape shape index (LSI, McGarigal and Marks 1995:109), and landuse
richness (LR, McGarigal and Marks 1995:119) in landscapes (i.e., 2,830-ha circular plot)
associated with 16 study playas on the Southern High Plains, Texas, 1999 and 2000.
6 —r
0}
o a>
c o
ca c
^—^ OJ
• • . ^ — - ' • ' ' ' ^
•D
c ^^^-^"''''^ ' . C
13
jQ
ailyA
< 3 —
ro
2 —
c Q
to
c
nj
Y = 0 6905 + 0.7074X
R-square = 0.256 Y = 0 4447 + 0.0534X
R-square = 0.527
0 —
~~\ I
10 20 30 40 50 60
PP
IJI
6 —r
(V 5 —
o <u
c o
ro c
c T3
Q
< c
3 —
TO
<
2 —
TO 2 —
C
CD Q
i? 1
Y = -0.92 + 0 0726X c 1 — Y = 0.2842 + 0 3078X
TO R-square = 0.269
R-square = 0 482
"T" —r 0 —
T 1 r I
30 40 so 60 70 6 7 8 13
9 10 11 12 14
ED PED
6 —r
5 —
0) 5 —
O
c
CO
4 — •D
c c
< 3 —
<
CO
:o Q
c
(0 1 —
1 —
Y =-1.3868+ 0 3254X
Y = -0.84 + 0.4482X 0 — R-square = 0.449
R-square = 0.472
I I I r
9 10 11 13 1S
LSI LR
72
25 — /! S —
O •
C 2.0 —
O 20 — CO
C TD
(0 C
•D 3
C 15 —
13 1.5 —
n <
^^,-^^^
. . * • • • '
<
'co 1.0 —
= 10 Q
(0 •• .
c
Q CO
c CU 05 — • ,.-"
CO 05
0) •
Y = -0.033 + 0 3498X Y = 0.2139+ 0.0184X
00 — R-Square = 0 348
R-Square = 0 35
0.0 — 1 1 1 1
10 20 30 40 50 60 70
PP IJI
0) 2 — (U 2 —
O
o c
c
m CO
T3
"D C
c 3
3 XI
.Q <
< CO
Q Q
c c
CO CO
0)
Y =-0 22 + 0.0243X o
Y = -0.26 + 0 1577X
0 — R-Square = 0.301
R-Square = 0.326
"T" "T
30 40 50 60 70 10 11
ED LSI
Figure 4.3. Simple linear regression and 95% confidence bands of mean daily abundance
(natural-log transformed) of plains spadefoot {Spea bombifrons) and percent aerial
coverage of playa wetlands (PP), interspersion/juxtaposition index (IJI, McGarigal and
Marks 1995:103), edge density (ED, McGarigal and Marks 1995:106), and landscape
shape index (LSI, McGarigal and Marks 1995:109) in landscapes (i.e., 2,830-ha circular
plot) associated with 16 study playas on the Southern High Plains, Texas, 1999 and 2000.
73
Table 4.4. Multiple linear regression between mean daily abundance of amphibians and
landscape^ metrics at 16 playa wetlands on the Southern High Plains, Texas, 1999 and
2000.
Estimates*^ Collinlearity'
Species^ Variable"'^ Parameter Standardized / P VIF CN
NSF Intercept 9.869 0 2.35 0.039 0 0
LU -2.077 -0.703 -4.91 <0.001 1.13 1.0
SDI 1.203 0.319 2.24 0.046 1.12 1.14
MNN -0.002 -0.278 -1.98 0.073 1.08 1.39
PSI -5.634 -0.244 -1.67 0.124 1.19 1.58
PSF Intercept 9.199 0 4.53 0.001 0 0
PP 0.190 0.322 2.23 0.049 1.418 1.0
SEI -2.299 -0.458 -3.56 0.005 1.125 1.453
PSI -6.469 -0.665 -4.23 0.002 1.642 1.546
IJI 0.012 0.379 2.48 0.033 1.598 2.175
PNN -0.0005 -0.372 2.26 0.047 1.843 2.587
BTS Intercept 2.619 0 11.51 <0.001 0 0
YR -0.939 -0.615 -2.92 0.011 1.0 1.0
GPT Intercept 17.36 0 3.72 0.003 0 0
MNN -0.003 -0.759 -2.74 0.019 1.957 1.0
PSI 9.220 -0.756 -2.87 0.015 1.754 1.231
NP -0.157 -0.599 -2.14 0.056 1.990 2.285
PNN -0.001 -0.516 -1.92 0.082 1.838 2.731
^Landscapes {n= 16) were 2830-ha circular plots (i.e., 3-km radius) with their origins
positioned at the center of each study playa.
hSlSF = New Mexico spadefoot {Spea multiplicata), PSF = plains spadefoot {S.
bombifrons), BTS = barred tiger salamander {Ambystoma tigrinum mavortium), and GPT
= Great Plains toad {Bufo cognatus).
"^Landscape metrics retained as per stepwise selection using default entry and stay
significance level at a=0.15; all overall F-tests on final models were significant
{P<Q.042) and coefficients of determination adjusted for number of variables in the
model (i.e., /?%) = 0.728, 0.779, 0.378,and 0.409 for NSF, PSF, BTS, and GPT,
respectively.
PSI = shape index of study playa, PNN = mean nearest-neighbor distance from study
playa to surrounding playas, MNN = mean nearest-neighbor distance from all playas to
each other, NP = number of playas, PP = percent aerial coverage of playas, IJI =
interspersion and juxtaposition index of playas, SEI = Shannon evenness index of
landuses, and SDI = Sharmon diversity index of landuses (McGarigal and Marks 1995);
LU (landuse) and YR (year) were indicator variables using 0 = cropland watershed and 1
= grassland watershed, and 0 = 1999 (wet year) and 1 = 2000 (dry year), respectively.
74
Table 4.4. Continued.
^Relative natural-log (LN) abundance per species can be estimated via multiplying field
estimates of landscape metrics or indicator variables by parameter estimates, and
summing them and the intercept within species; predictions can be back-transformed to
original units by exponentiation; magnitude and sign of species-specific standardized
estimates can be interpreted as the strength and relation of the variable with LN
abundance in the presence of the other retained explanatory variables (Myers
1990:384-385).
CoUinearity diagnostics (intercept adjusted) were variance inflation factors (VIF) and
condition numbers (CN); VIF > 10 and CN > 30 are suggestive of a linear dependency
between >1 variable (Freund and Littell 2000:98-101).
75
Finally, 38% of the variation in mean daily abundance of barred tiger salamander was
explained by YR (Fi,14=8.52, P=0.011). No linear dependencies existed among variables
(Table 4.4).
Discussion
Canonical correspondence analysis (CCA) revealed that landscape structure
influenced the composition of the amphibian species assemblage at playa wetlands.
Subsequent univariate analyses indicated New Mexico and plains spadefoots generally
were positively associated with spatial metrics representing optimal spatial positioning of
playas (i.e., interspersion/juxtaposition index, percent aerial coverage of playas) and
landscape complexity (i.e., landscape shape index, edge density). Great Plains toad and
barred tiger salamander usually were negatively associated with spadefoots (as per CCA),
but corresponding univariate analyses indicated they were not associated with spatial
metrics. Thus, these results suggest Great Plains toad and barred tiger salamander may
be less affected by spatial positioning of playas and complexity of the landscape matrix
but perhaps are poorer competitors than spadefoots because of their negative association
with these species.
Spadefoots probably are associated with landscape structure more than the other
species because of their relatively small body size (Chapter II). Body size has been
shown to connote vagility (Peters 1983:89-91, With and Crist 1995), seemingly due to
differential perception to patch viscosity and edge permeability (Crist et al. 1992, With
1994, Wiens et al. 1997, Mclntyre 2000). Although relative vagility has not been
compared among these amphibians, dispersing spadefoots may have been unable to
76
penetrate geometrically complex landscapes, resulting in increased abundance in natal
wetlands. Indeed, several recent studies have documented positive associations of
amphibians at wetlands embedded in complex agricultural landscapes (Knutson et al.
1999, Kolozsvary and Swihart 1999, Chapter III). Similarly, abundance of spadefoots
may have been positively related with less isolated and more optimally juxtaposed playas
because of their lower relative vagility. Probability of successful interdemic dispersal
can increase with decreasing inter-patch distance and increasing patch juxtaposition
(Bascompte and Sole 1996, Hess 1996). Consequently, abundance and persistence of
less vagile species can be positively related to habitat patch positioning thus the exchange
of individuals among spatially structured local populations (Ritchie 1997, Stacey et al.
1997). Therefore, these results support my hypotheses and previous applied and
theoretical research on the relationship of abundance, spatial positioning of populations,
and landscape complexity.
Great Plains toad and barred tiger salamander may have been negatively
associated with spadefoots because of differential competitive ability of their larvae
(Wilbur 1984). Spadefoot larvae have been shown to be competitively dominant over
various genera (e.g., Bufo, Gastrophryne, Hyla, Rana, Morin 1983, Wilbur 1987, Dayton
and Fitzgerald 2001). Increased recruitment of spadefoots also could increase
postmetamorphic interactions of individuals among our species (Wilbur 1980), which
may compete for food resources as suggested by diet composition studies (Anderson et
al. 1999^, L. M. Smith and M. J. Gray, unpublished data). Thus, Great Plains toad and
barred tiger salamander may have been displaced or experienced reduced survival in
77
terrestrial landscapes with greater spadefoot abundance, because of potential diet overlap
and relative competitive ability. Also, if microhabitat characteristics (e.g., vegetation
composition and structure) were correlated with landscape structure, differential
microhabitat use might explain species separation. Spadefoots seem to prefer playas with
less vegetation for breeding unlike Great Plains toad and barred tiger salamander
(Anderson et al. 1999).
Conservation Implications
Although regulating mechanisms are unknown, these results suggest that
landscape structure influences species composition and abundance of amphibians.
Landscape structure may favor certain species as per species-specific vagility and their
relative perception to patch viscosity and edge permeability (Wiens 1997). Competitive
ability also may interact individually or synergistically with landscape structure and
species-specific vagility to affect community composition and abundance of amphibians
in wetlands. Therefore, landscape ecologists should consider species-specific mobility
differences when developing conservation initiatives for spatially structured amphibian
populations (DriscoU 1997, Szacki 1999). Existing software packages (e.g.,
FRAGSTATS* ARC®, RAMAS® GIS) have options for inclusion of relative patch
viscosity, boundary permeability, and species vagility data; however, generally unity or
educated estimates are used because the aforementioned data do not exist. Future
research directives need to focus on estimation of species-specific vagility in
anthropogenic and natural cover types and the relative permeability of their boundaries
for realistic estimation of extrinsic connectivity among habitat patches (e.g., With and
78
Crist 1995, Wiens et al. 1997). Indeed, various theoretical studies (e.g., O'Neill et al.
1988, With and Crist 1995) that used percolation models have shown this information is
critical in estimating viability and persistence spatially structured local populations.
Finally, my prediction models can be used to estimate relative species-specific abundance
of amphibians on the SHP using field estimates of landscape metrics (e.g., Knutson et al.
1999, Scribner et al. 2001). Landscape ecologists may use this and ground-truthing
information to locate potential locations for conservation endeavors.
79
CHAPTER V
TEMPORAL NICHE PARTITIONING IN SOUTHERN
HIGH PLAINS AMPHIBIAN POPULATIONS
Abstract
Temporal niche partitioning reduces interactions among species by decreasing
probability of interference and exploitative competition. Although temporal segregation
of resource use has been recorded among amphibian species, it has not been examined for
amphibians in semi-arid and prairie environments. Moreover, anthropogenic landuse
may affect temporal resource use of amphibians. My objective was to test for temporal
niche partitioning in Southern High Plains amphibians, and examine if landuse
(cultivation vs. grassland) affected temporal resource use among 4 common species {Spea
multiplicata, S. bombifrons, Bufo cognatus, and Ambystoma tigrinum mavortium).
Standardized species-specific daily abundance from pitfall traps at 16 playa wetlands
during 2 years (1999 and 2000) was compared among 3 within-year time periods and
between 2 landuses. Inspection of time series plots and hypothesis testing indicated that
temporal niche partitioning existed in Southern High Plains amphibian populations.
Differences in temporal activity may be a consequence of interspecific competition
associated with dietary or microhabitat niche overlap. No differences were detected in
temporal niche partitioning between landuses, suggesting that phenotypic plasticity in
temporal resource use is unlikely for these amphibian species. Thus, amphibians that use
cropland wetlands later in a year may have greater probability of extinction in grassland
wetlands, because hydroperiods are shorter in wetlands surrounded by disturbed
80
landscapes. It may be beneficial to preserve grassland playas and restore hydroperiods in
cropland landscapes to ensure habitat for later acti\'e amphibian species.
Introduction
Resource partitioning in ecological communities facilitates species coexistence
and allows biological diversity (Schoener 1974). The way in which species partition
resources defines their ecological niche (Pianka 1976). Hutchinson (1957) suggested that
an ecological niche could be conceptualized as an ^-dimensional hypervolume, where
each axis represented abiotic and biotic resources necessary for survival and
reproduction. Niche dimensions also may include axes for time and space (Cody 1968.
Schoener 1974. Rosenzweig 1995), resulting in a pulsating hypervolume (Pianka 1994).
The competitive exclusion principle {sensu Gause 1934, Hardin 1960) states that species
coexistence depends on the dissimilarity of niche hypervolumes (Pianka 1994:249, 272-
273).
Niche hypervolumes must be dissimilar along at least one axis to facilitate species
coexistence (Schoener 1974). The most basic hyper\olume includes axes for a resource,
space, and time (Pianka 1974). Temporal segregation is necessary for coexistence if
species use a common limiting resource, such as food, in the same location (Schoener
1974). Use of shared resources at different times is called temporal niche partitioning
(Huey and Pianka 1983), and a fundamental construct to niche complementarit}' and
biodiversity (Loreau 2000. Loreau and Hector 2001). Presumably, temporal niche
partitioning facilitates species coexistence by reducing interference and exploitative
competition (MacArthur and Levins 1967, Case and Gilpin 1974). Indeed, it has been
81
suggested that temporal niche partitiomng in^lies competition along space or resource
(e.g., food) axes (Huey and Pianka 1983, Schoener 1983).
Temporal niche partitioning is most common in poikilotherms among \'ertebrates
(Schoener 1974). Several studies have documented temporal differences in activit\'
patterns among amphibian species (Blair 1%1, Murphy 1963, Creusure and UTiitford
1976, Eraser 1976flA Bowker and Bov^êr 1979, Toft and Duelhnan 1979. W iest 1982).
Following Pianka (1976) and others (e.g., Schoener 1974), it has been hypothesized that
temporal niche partitioning exists in amphibian communities to reduce competition for
breeding sites or food resources (Wilbur 1980, Toft 1985). For amphibians that
e>q)losively breed in wetlands, temporal niche partitiomng probably reduces competition
most for food resources, because few species establish breeding territories (Duellman and
Trueb 1994). Indeed, dietary overly has been documented in adults and juveniles of
several coexisting explosive breeding species (Whitaker et al. 1977, Dimmitt and Ruibal
1980a, Anderson et al. 1999ft, Newman 1999).
Southern High Plains amphibians exist primarily in spatially structured playa
wetlands and tbey are explosive breeders (Bolen et al. 1989, CHiapters II and HI).
Because diets of Southern High Plains amphibians overlap (Anderson et al. 1999ft, L. M.
Smith and M. J. Oray, itnpublished data), species may exhibit temporal niche partitioning
to reduce exploitative competition. Several Southem High Plains species also use similar
habitat (e.g., vegetation structure, percent water coverage) for breeding (Anderson et al.
1999a); thus, species may temporally segregate to reduce interference competition (albeit
explosive breeders). Anthropogenic landuse also affects demographics and d>Tiamics of
82
Southem High Plains amphibian populations (Chapters III and VI). Therefore, my
objective was to test for the existence of temporal niche partitioning in Southem High
Plains amphibian populations, and examine the effect of landuse (cultivation vs.
grassland) on species-specific temporal activity pattems. I hypothesized that Southern
High Plains amphibians would temporally partition resources, and landuse would affect
temporal activity pattems.
Methods
This study was conducted at 16 playa wetlands (8/year) on the Southern High
Plains of Texas during 1999 and 2000 (Bolen et al. 1989). Playas were considered to be
in grassland if on average >75% of the surrounding landscape (2830-ha circular plot) was
undisturbed and vegetated, and cropland if >75% of the landscape was cultivated
(Chapter III). Playas were partially enclosed (i.e., 25% of the circumference) with 60-cm
high drift fence and 19-L pitfall traps (Dodd and Scott 1994), and pitfalls checked
altemate days from 16 May-17 October 1999 and 19 April-18 August 2000. Captured
amphibians were enumerated by species, marked uniquely, and released. After releasing
individuals, pitfalls were closed for 24 hr to reduce probability of immediate recapture
and possible biasing of species-specific temporal abundance (Turchin 1998). Temporal
niche partitioning was examined for the 4 most common species of amphibians on the
Southem High Plains (New Mexico spadefoot [NSF, Spea multiplicata], plains spadefoot
[PSF, S. bombifrons]. Great Plains toad [GPT, Bufo cognatus], and barred tiger
salamander [BTS, Ambystoma tigrinum mavortium]. Chapter III). A more detailed
83
account of the study area, amphibians, and sampling protocol was described in Chapters
II and III.
I used total daily capture per species (i.e., summed across playas within landuses
and years) as an estimate of daily species-specific abundance (Dodd and Scott 1994).
Summing across playas (i.e., local populations) was a reasonable estimate of relative
population size within landuses each year, assuming playas were mutually independent
(Hanski 1997). Species-specific abundance was standardized (hereafter percent
abundance) by dividing species-specific daily capture {N\t) by species-specific total
capture (i.e., Aî, sum of captures over t days) within landuses each year (Hurlbert 1978).
Thus, species-specific percent abundance represented the proportion of the population of
the /• species present on the t day. Standardization was necessary to remove abundance
effects and compare niche overlap of species equally among time periods and between
landuses (Hurlbert 1978, Paton and Crouch 2002). Time periods (i.e., 1, 2, and 3)
represented 3 equal sequential time series in each year (i.e., 22 and 17 days per period in
1999 and 2000). I chose to divide each yearly time series into 3 periods, because
previous studies in other regions of North America suggested amphibians temporally
partition resources into ca. 2-3 within-year time periods (e.g.. Murphy 1963, Wemer and
McCune 1979, Wilbur 1980). A different temporal division (e.g., 2 or 4) of the within-
year time series may have revealed different results.
Temporal niche partitioning was determined by inspecting time series plots and
comparing frequency distributions of percent abundance for pairs of species among the 3
time periods within years. Frequency distributions also were compared between cropland
84
and grassland landscapes. Frequency distributions were constmcted by tallying number
of days within time periods, landuses and years that percent abundance of pairs of species
exceeded each other. Thus, each frequency distribution represented the pairwise species
comparison of standardized temporal abundance. I considered differences between
species as a reasonable measure of temporal niche partitioning, because standardized
abundance of species pairs was compared to the null distribution of equal relative
frequency among time periods and between years (see below). This approach to
quantifying niche partitioning is similar to Hurlbert (1978).
I tested for differences in frequency distributions (a = 0.05) among time periods
and between landuses by each year using logistic regression with a generalized logit
model (Agresti 1990:307, Stokes et al. 2000:257- 259). Nonaddivity of main effects
(i.e., time and landuse) was tested by including an interaction term in the model (Stokes
et al. 2000:259). Normalized chi-square tests of homogeneity (i.e., Z-tests) Bonferroni
corrected were used to test pairwise differences in frequency distributions when the main
effect logit model was significant (Zar 1984:395-396, Milton and Amold
1995:667-670). If main effects were not additive, analyses were separated by periods
and landuses, and pairwise Z-tests performed. I asstimed the categorical response of t
days followed a multinomial distribution per effect and frequency distributions were
independent (Agresti 1990:306).
Results
Time series plots of pairwise species-specific percent abundance suggested that
amphibian species temporally segregated resources on the Southem High Plains (Figure
85
5.1). The main effect logistic regression indicated that frequency distributions were
different among time periods both years between GPT and NSF (x 2=14.9, P<0.001
[1999]; 5cV9.6, P=0.008 [2000]), PSF and NSF (xV29.9, P<0.001 [1999]; xVl8.4,
P<0.001 [2000]), BTS and NSF (xV40.4,P<0.001 [1999]; x V 13.3,/'<0.001 [2000]),
BTS and PSF (xV31.2, P<0.001 [1999]; xV8.6, P=0.014 [2000]), and GPT and BTS
(xV33.5, P<0.001 [1999]; x V 9 . 1 , P=0.0\ 1 [2000]), and GPT and PSF in 1999
( x V l 6 . 3 , P<0.001 [1999]; x^2=18.4, 7^=0.099 [2000]). Temporal niche partitioning did
not differ between landuses for all species pairs both years (xî=0.001-1.75,
P=0.19-0.98).
Nonadditivity was violated for GPT and NSF in 1999 (xVl4.1, P<0.001), BTS
and NSF in 2000 ( x V l 1-9, P<0.001), GPT and PSF in 1999 (xV9.4, P=0.009), BTS
and PSF in 2000 (xVl4.4, P<0.001), and GPT and BTS in 2000 ( x V l 1.2, P=0.004);
therefore, these analyses were separated by landuse. Frequency distributions were
different among time periods between GPT and NSF in grassland in 1999 (x^2=15.9,
P<0.001), between GPT and PSF in cropland in 1999 (xVl6.8, P<0.001), and between
BTS and NSF (x^2=15.4, P<0.00\), BTS and PSF (xVl6.8, P<0.001), and GPT and BTS
(X^2=12.9, P=0.002) in cropland in 2000. Differences were not detected in frequency
distributions among time periods for GPT and NSF in cropland in 1999 (x 2=4.3,
P=0.114), GPT and PSF in grassland in 1999 (xV4.6, P=0.099), and for BTS and NSF
(X^2=1.5, P=0.482), BTS and PSF (xVo.65, P=0.723), and GPT and BTS (xVo.76,
P=0.684) in grassland in 2000.
86
87
Figure 5.1. Time series plots for pairs of amphibian species residing at 16 playa wetlands
in 2 landuses and years on the Southem High Plains of Texas. Linear trajectories
represent total daily capture per species summed across playas (i.e., 4 per landuse per
year). Daily capture per species was standardized using species-specific capture for the
entire time series (Hurlbert 1978), thus each node on the trajectory represents the
proportion of the population present during that day (i.e., percent abundance). The solid
line is the first species listed for the pair in the upper right comer of the graph. NSF =
New Mexico spadefoot {Spea multiplicata), PSF = plains spadefoot {S. bombifrons), BTS
= barred tiger salamander {Ambystoma tigrinum mavortium), and GPT = Great Plains
toad {Bufo cognatus).
Cropland, 1999 GPT vs. NSF Grassland, 199£
GPT vs. NSF
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10 20 30 40 50 60 30 40
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Cropland, 2000 GPT vs. NSF Grassland, 2000

GPT vs. NSF
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Cropland, 2000 PSF vs. NSF Grassland, 2000 PSF vs. NSF
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88
Cropland, 1999 Grassland, 1999p^ B T S v s . NSF
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BTS vs. NSF BTS vs. NSF
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GPT vs. PSF GPT vs PSF
c ra
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G P T vs. PSF
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Figure 5.1. Continued.
89
5 - 5 -'
Cropland, 1999 i
BTSvs PSF
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G P T vs. BTS G P T vs. BTS
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Figure 5.1. Continued.
90
In general, pairwise Z-tests indicated that a larger proportion of the GPT
population was present during time 2 than the NSF populafion (Figure 5.2). Also, a
larger proportion of the PSF population was present during time 2 than NSF and GPT
populations. Barred tiger salamander was most active during time 3 compared to NSF,
PSF, and GPT in 1999. However, a larger proportion of the BTS population was present
in time 1 than NSF, PSF, and GPT populations in cropland in 2000 (Figure 5.2).
Discussion
This represents the first documentation of temporal niche partitioning in semi-arid
Great Plains amphibian communities. Even though amphibians on the Southem High
Plains generally emerge and breed simultaneously and clutch times are similar (see
Chapter II life history review. Rose and Armentrout 1974, 1976, Degenhardt 1996:38, 40.
51, Sullivan and Femandez 1999), plains spadefoot was more active during the middle of
both years compared to New Mexico spadefoot and Great Plains toad. Creusere and
Whitford (1976) also observed this temporal relationship with plains spadefoot and Great
Plains toad and a different spadefoot {Scaphiopus couchii) in New Mexico. Plains
spadefoot may be more active after New Mexico spadefoot and Great Plains toad because
of exploitative competition for food resources (Schoener 1974, 1983). Anderson et al.
(1999^) documented some diet overlap among adults of these 3 species. Moreover,
plains and New Mexico spadefoots use similar habitats for breeding and can breed
simultaneously (Bragg 1965, Anderson et al. 1999a, M. J. Gray, unpublished call survey
data); thus, temporal niche partitioning between spadefoots may be a consequence of
interference competition for breeding sites (Wilbur 1980, Toft 1985). Similarly, Great
91
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Plains toad was more acti\e during the second time period than New Mexico spadefoot.
which may ha\e been a consequence of diet o\erlap, because these species use different
microhabitats for breeding (Anderson et al \999a,b).
Barred tiger salamander was most acti\e later in 1999 than all other species. I
believe activity pattems of barred tiger salamander probably were related to
developmental time or physiological constraints associated with probability of
desiccation not competition (Toft 1985). Although barred tiger salamander can develop
as rapidly as anurans on the Southem High Plains. the> may prolong development or
become neotenic during >ears of favorable rainfall (Petranka 1998). Rainfall and
hydroperiod in playas was greater in 1999 than 2000 (Chapter II): thus, barred tiger
salamander may have been more active later in 1999 than other species because
favorable conditions in the aquatic en\ ironment prolong development hence metamorph
emergence. Rainfall also was greater during the last time period (17.5 cm) than the 2
previous periods (14.5 cm and 10.5 cm for 1 and 2, respectively) in 1999. possibly
facilitating dispersal of all age classes (Petranka 1998). If temporal pattems were a
consequence of evolutionary competition (i.e.. ghost of competition past, Connell 1980),
barred tiger salamander should have been more active later in both years; this was not
observed.
Barred tiger salamanders also were most active earlier than all other species in
2000 at cropland playas. Similarly, I suspect this ma>' be a consequence of differential
hydroperiods between landuses (Chapter II). Two of 4 cropland playas were dry prior to
time period 2, whereas all grassland playas retained water through time period 3 in 2000.
101
Perhaps, early playa drying in cropland landscapes caused barred tiger salamanders to
emigrate in search of altemate water sources, resulting in increased levels of activity
(Semlitsch 19876). I speculate that anurans remained near playa bottoms and aestivated
in subterranean burrows following playa drying (McClanahan et al. 1994).
Temporal niche partitioning implies niche overlap and competitive exclusion in at
least one other niche dimension (Schoener 1974. Huey and Pianka 1983, Pianka 1994).
thus seemingly a competitive dominant exists among Southem High Plains anuran
species. Data from Chapter III suggested that New Mexico spadefoots can become
competitively dominant in the absence of larval and neotenic barred tiger salamanders.
Laboratory studies (e.g., Morin 1983, Wilbur 1987, Dayton and Fitzgerald 2001) indicate
that certain spadefoot tadpoles are superior competitors for food resources compared to
other anurans (e.g., Bufo, Rana). Therefore, New Mexico spadefoots may have displaced
the other 2 anurans from earlier activity over evolutionary time, resulting in plains
spadefoot and Great Plains toad being more active during the middle portion of the
breeding season. The time series plots (Figure 5.1) support this inference, because
species-specific percent abundance was > 0 for most days. However, no experiments
exist comparing the relative competitive ability among these species for food or other
resources in terrestrial landscapes. To test the above hypothesis, a shared limiting
resotirce (e.g., food, cover) would need to be identified among these species (Pianka
1976). Then, an experiment could be conducted where individuals were allowed to
compete for the limiting resource in allopatric and sympatric scenarios (Pianka 1976).
102
Numerical or fitness response could be compared subsequently among species (Pianka
1976).
It also has been hypothesized that interspecific competition in harsh environments
is low or nonexistent compared to more stable habitats (Wiens 1977, Huston 1979,
Strong 1983). This idea was stimulated from the observation that biodiversity is greater
in stable climates (e.g., tropics) than in variable ones (e.g., temperate regions, Pianka
1966). Theoretical studies since then have shown that competition and competitive
exclusion operate independently of environmental stability (May and MacArthur 1972,
Holt 1985, Chesson and Huntly 1997); however, few empirical examples exist. Because
climate on the Southem High Plains is semi-arid and hydroperiod in playa wetlands is
variable (Haukos and Smith 1994), these results seem to support the prediction that
interspecific competition and niche partitioning exist in variable environments.
Moreover, it has been suggested that temporal niche partitioning exists most often with
species, such as insects and larval amphibians, that exploit transient resources (Schoener
1974, Toft 1985). Considering the environmental variability in playas and the explosive
breeding strategy of Southem High Plains amphibians, these results also seem to support
this hypothesis.
Although anthropogenic landuse affected dynamics and demographics of
Southem High Plains amphibians (Chapters III and VI), it did not affect temporal niche
partitioning. Perhaps, small sample size and variability in multinomial distribution
pattems prevented detection of biologically meaningful differences. Altematively,
temporal resource use may not be as phenotypically plastic as polymorphic characters or
103
developmental time in amphibians. For example, spadefoots may develop carnivorous
feeding apparatuses in wetlands with short hydroperiods to capitalize abundant transient
invertebrate food resources (Pfennig 1992). Also, many amphibian species may
accelerate growth and undergo metamorphosis at a smaller body size in wetlands with
shorter hydroperiods (Newman 1992). In Chapter II, I suggested the capability of
phenotypic plasticity in development might explain differences in postmetamorphic body
size among playas with differing hydroperiods.
Because amphibians behaviorally hydroregulate (Duellman and Tmeb 1994), it is
reasonable to hypothesize that activity pattems would follow water availability, hence
temporal resource use be shortened in wetlands with rapid hydroperiods (i.e., cropland
playas, Luo et al. 1997). Thus, temporal niche partitioning should have been affected by
landuse, but this was not observed. Accordingly, I hypothesize that change in temporal
resources use may be beyond the genotypic norm of reaction for Southem High Plains
amphibian species (Via and Lande 1985, Gomulkiewicz and Kirkpatrick 1992).
Conservation implications of possible genotypic rigidity in temporal niche
partitioning are important. Paton and Crouch (2002) and Snodgrass et al. (2000) suggest
that duration and timing of wetland hydroperiods are critical to survival and reproduction
of pond-breeding amphibians because of specific-specific phenology in temporal use; my
results support their suggestions. Species, such as plains spadefoot and Great Plains toad,
that use resources later than other species (e.g., New Mexico spadefoot) may have
reduced fitness in anthropogenically modified wetlands, because shortened hydroperiods
may decrease probability of survival and reproduction. Consequently, these species may
104
have a greater probability of local extinction. I recommend retention and restoration of
grasslands around playa wetlands, because hydroperiods generally are shorter in cropland
landscapes from sedimentation (Luo et al. 1997).
105
CHAPTER VI
CHAOS AND REGULATING MECHANISMS
IN A SOUTHERN HIGH PLAINS AMPHIBIAN ASSEMBLAGE
Abstract
Biological chaos is a dynamic condition of nonlinear ecological time series that
exhibits sensitive dependence on initial conditions. Chaos has been demonstrated in
various discrete biological models; however, it rarely has been documented in natural
ecosystems. Simulations suggest that chaos may be a fundamental characteristic of
spatially structured individuals, and its presence can reduce probability of metapopulation
extinction. It also has been hypothesized that anthropogenic disturbance may influence
occurrence of chaos. Amphibians exist in spatially structured playa wetlands embedded
in natural and anthropogenically disturbed landscapes on the Southern High Plains
(SHP). Thus, I tested the difference in occurrence of biological chaos in a SHP
amphibian assemblage between landuses (i.e., cultivation vs. grassland) at 16 playa
wetlands during 1999 and 2000 (4 playas per landuse per year). Also, deterministic and
stochastic difference equations can be used to model assemblages and determine
regulating mechanisms. Thus, I compared occurrence of chaos in a SHP amphibian
assemblage predicted by 5 common difference equations with actual dynamics. For 3
deterministic equations, I also tested for differences in parameter estimates between
iteration and least squares techniques. The amphibian assemblage time series was
constructed from total daily captures in pitfall traps summed across species at each playa.
Existence of chaos was determined via estimating Liapunov exponents per playa and
106
using published stability estimates. Occurrence of chaos was greater at grassland (7/8)
than at cropland (1/8) playas, indicating that biological chaos was a fundamental
characteristic of the SHP amphibian assemblage and its occurrence was altered by
anthropogenic disturbance. The Ricker logistic function with a stochastic parameter
(M+i=Mexp[r(l- N[IK) + ex\) predicted occurrence of chaos similar to actual dynamics,
suggesting that density dependence and environmental stochasticity may be important
regulating mechanisms in this amphibian assemblage. The stochastic Ricker function
predicted that intrinsic rate of increase at grassland playas was greater than at cropland
playas. Intrinsic rates of increase also were greater using an iterative method rather than
the least squares method for estimation in all difference equations. Anthropogenic
disturbance of landscapes surrounding wetlands altered the dynamics of the spatially
stmctured amphibian assemblage on the Southem High Plains, and perhaps, increased
probability of metapopulation extinction via reducing natural occurrence of biological
chaos. Parameter estimates for the stochastic Ricker function are presented for future
simulation of amphibian assemblage dynamics in disturbed and undisturbed wetland
systems.
Introduction
Examining the dynamical behavior of ecological time series is a critical prelude to
understanding regulating mechanisms of assemblages and populations (Edelstein-Keshet
1988) and initiating conservation endeavors. Most biological theories assume abundance
trajectories approach equilibrium and the population or assemblage is inherently stable
(DeAngelis and Waterhouse 1987). However, complex dynamics, such as chaos, have
107
been demonstrated in simple discrete deterministic models (May 1974. 1975fl. 1976a).
Interestingly though, chaos has been detected in few natural assemblages (Turchin and
Taylor 1992). Assemblages that exhibit chaos may have a greater probability of
extinction than those near positive stable equilibria (Hastings et al. 1993, Shulenburger et
al. 1999). Altematively, chaos may be a critical component of spatially stmctured
assemblages, because it can reduce the probability of metapopulation extinction by
increasing local abundance noise (Allen et al. 1993). Anthropogenic modifications of the
landscape also may induce transition from stable equilibria to chaos or vice versa
(Berryman and Millstein 1989, Berryman 1991). This may be especially critical for
assemblages in decline or near extinction thresholds, such as amphibians (Houlahan et al.
2000).
Populations, and possibly assemblages, are regulated via density dependent or
independent mechanisms (Lotka 1925, Andrewartha and Birch 1954). Densit}
dependence implies that dynamics are a consequence of positive or negative feedback
among individuals (Roughgarden 1998). Competition and predation have been
considered important negative feedback processes in most natural populations and
assemblages (May 1976b). Competition and predation can be modeled over discrete time
intervals using difference equations (Edelstein-Keshet 1988). The discrete logistic
growth and Ricker functions are commonly used to model competition (Hastings 1996).
A negative quadratic term can be added to these equations to incorporate a predation
effect (Edelstein-Keshet 1988). Additionally, a stochastic parameter {st) can be included
to model density independent factors (i.e., environmental stochasticity, Turchin and
108
EUner 2000). Parameters of these models can be estimated using observed data and least-
squares estimation or by solving sequential iterations of the difference equations
(Edelstein-Keshet 1988). Parameter estimates subsequently can be used to predict
population or assemblage dynamics. If predictions are similar to actual dynamics, the
regulating mechanism mathematically associated with the function (e.g., densit}
dependence) may be an important process in the population or assemblage (Turchin and
Ellner 2000). Thus, regulating mechanisms in populations and assemblages can be
discemed using observed data and difference equations (Elkinton 2000).
Amphibians exist in spatially stmctured wetlands called playas on the Southem
High Plains (SHP, Bolen et al. 1989). Anthropogenic landuse (i.e., grassland and
cultivation) affects postmetamorphic body size (a fitness correlate), demographics, and
community stmcture of amphibians (Chapters II and III). Because amphibians are
spatially structured on the SHP, the existence of chaotic dynamics may be critical.
Anthropogenic disturbance of the landscape surrounding playas may dismpt natural
dynamics. Thus, my first objective was to determine the presence of chaos in a SHP
amphibian assemblage at 16 playa wetlands from an existing time series and test for
differences in occurrence of chaos between landuses. I examined assemblage instead of
population dynamics because species-specific analyses may have lower probability of
chaos detection (Turchin 1991, 1993, Turchin and Taylor 1992). My second objective
was to determine regulating mechanisms in the SHP amphibian assemblage via
estimating parameters for deterministic and stochastic versions of the discrete logistic and
Ricker functions and comparing their predictions to actual dynamics. I assumed
109
regulating mechanisms (e.g., density dependence) in populations also operated in
assemblages. Turchin (1991) also made this assumption in his chaotic analysis of multi-
species vole data. Lastly. I quantified differences in parameters between iteration and
least-squares estimation techniques. This study provides insight into determining if
biological chaos was a fundamental characteristic of a spatially structured amphibian
assemblage, and whether density dependence, predation, or environmental stochasticity
were important regulating mechanisms.
Methods
This study was conducted at 16 playa wetlands (8 grassland, 8 cropland) on the
SHP of Texas during 1999 and 2000. Playas were considered to be in grassland if >75%
of the surrounding landscape (i.e., <3 km from the playa center) was undisturbed and
vegetated, and cropland if >75% of the surrounding landscape was cultivated (Chapter
III). Playas were partially enclosed (i.e., 25% of circumference) with 60-cm high drift
fence and 19-L pitfall traps (Dodd and Scott 1994). Pitfall traps were checked altemate
days for captures from 16 May-17 October 1999 and 19 April-18 August 2000. Thus,
the amphibian time series represented altemate equally spaced days nested within 2
years. Captured amphibians were enumerated, marked uniquely, and released. A more
detailed account of the study area, amphibians, and sampling protocol has been described
previously (Chapters II and III).
I used total daily capture per playa (natural-log transformed) as an estimate of
relative assemblage size (i.e., Nt, Dodd and Scott 1994). As in Chapter III, I assumed if
any bias existed by using mean daily capture to estimate relative daily abimdance of the
110
assemblage, it was constant between landuses and years. Daily capture was summed
across species for Nt to incorporate positive and negative feedbacks into dynamics
(Schaffer and Kot 1986). As mentioned, Turchin (1991, 1993) summed across vole
species using data of Henttonen et al. (1984) for chaos analyses. Turchin (1991) also
suggested and later demonstrated (Turchin and Taylor 1992) that modeling a nonlinear
time series at lower dimensionality (e.g., number of species, time lags) than the tme
overall dimension of the system can decrease detection of real chaos. I did not sum
captures across playas (i.e., local assemblages), because simulations have demonstrated
that it can reduce probability of detecting actual chaos (Rohde and Rohde 2001). I also
used da>s as time units instead of years or generations. Thus, I assumed regulating
mechanisms, such as density dependence, operated within years for my amphibian
assemblage (Wilbur 1980).
Time series plots were constmcted to graphically illustrate actual dynamics at
each playa. Then, I assumed the amphibian assemblage was govemed by an underlying
difference equation of the form Nt, Nt+i=F(N(). The presence of chaos was determined
via computing local Liapunov exponents per playa as,
Am{t) = -HAt + rn-\}-J{t + l)J{t)\\ , (6.1)

m
where J(t) was the first derivative of F(N) along the assemblage trajectory and m was the
playa-specific length of the time series (Ellner 2000). Liapunov exponents measure the
average deviation in slope of F(N) over time (Abarbanel 1996). They also measure the
rate of trajectory divergence of F(N), commonly called sensitive dependence on initial
conditions, which is tantamount to chaos (Ellner 2000). Positive ^^{t) indicates the
111
presence of chaos; negative exponents suggest convergence to some stable or periodic
equilibrium (Ellner 2000). Inasmuch as equation (6.1) is the natural-log geometric mean
of DF(N), the presence of chaos merely implied the average slope of the trajectory of
F(N) > 1. For detecting chaos in the SHP amphibian assemblage, I initially assumed no
specific form of F(N) (i.e., Nt+i=F(Nt)), because regulating mechanisms (i.e., the
underlying ftanction) were unknown. Thus, Liapunov exponents were estimated directly
by computing the slope of the linear trajectory between time periods (i.e., equally spaced
days) per playa (Olsen and Degn 1985),
N{t + m-\) N{t + \) N{t) (6.2)

m dN dN dN
I assumed the slope of each linear trajectory was a reasonable estimate of the derivative
of A^,. Differences in occurrence of chaos (i.e., positive 'km{t)s) between landuses was
tested (a=0.05) using a Fisher's exact test (Conover 1980:167).
Subsequently, I assumed that density dependence was the primary regulating
mechanism in the SHP amphibian assemblage and followed discrete logistic growth.
A^/+1 = A^^ 1 + r 1- (6.3)
where, r = intrinsic rate of increase and A'= carrying capacity (May 19756). Equation
(6.3) simplifies to
(6.4)
Nt K
Finally, making the following change of variables {a = \ + r, b = r/K), equation (6.4)
simplifies to
^ ^ = a-bNr, (6.5)
Nt
112
which can be solved using 2 successive iterates (Edelstein-Keshet 1988). For example, if
Aî, N2, and N3 = 4.9, 3.6, and 3.7, it follows that NiINx ~ 0.735 and N^ÎNi = 1.028,
resulting in 2 equations of the form (6.5) with 2 unknowns (i.e., 0.735 = a-4.9b , 1.028 =
a-3.6b). These equations can be solved for a (0.622) and b (-0.113) and subsequently r
(-0.378) and K (3.36). Using daily capture per playa and equation (6.5), I estimated r
and K for each successive pair of iterates in the amphibian assemblage time series, and
computed the arithmetic mean for each parameter per playa. It should be noted that
because my time series was within years and I used difference instead of differential
equations, r and K may not represent the true intrinsic rate of increase and carrying
capacity of the assemblage; they were merely control parameters of these functions.
Because Mean become negative using equation (6.3) if M > K{\+r)lr, which is
biologically unrealistic, I then assumed the amphibian assemblage followed Ricker
logistic growth,
Nt+\ = Nt^^V
'I'-i (6.6)
which is positive for all positive initial conditions (May 19756). Equation (6.6)
simplifies to
In Nt+\ = '-^Nt • (6.7)
\ Nt J
Finally, making the following change of variables {a = r,b = r/K), equation (6.7)
simplifies to
In ^Nt.i^ = a-bNt , (6.8)
V Nt )
which can be solved using successive iterates as described previously
113
Previous research (e.g., Morin 1981, Chapters II and III) also suggests that
predators may be an important regulating mechanism in amphibian assemblages.
Because predation generally has a negative effect on M , I assumed Ricker logistic
growth then added a negative quadratic term for predation to equation (6.6),
A^/+l = A^/exp ,fl_A^l_,^2 (6.9)

K t
Equation (6.9) simplifies to
In {EI±I\ = f- — Nt-cNf (6.10)

\ Nt J
Finally, making the following change of variables {a = r, b = r/K, and c=c), equation
(6.10) simplifies to
In Nt+\ = a-bNt-cNJ (6.11)

\ Nt )
which can be solved using 3 successive iterates similar to previously described.
Parameters in equations (6.5), (6.8) and (6.11) also can be approximated using
least-squares estimation (e.g., Turchin 1990). Letting, yt = M+i/M for equation (6.5), yt =
ln(M+i/M) for equation (6.8), and x, = M for both, equations (6.5) and (6.8) reduce to
y- = a + bxi . (6.12)
which is a linear function. Parameters a and b can be solved by fitting a line to the entire
data set and using the least-squares algorithm (Milton and Amold 1995:386). Similarly,
letting >^, = \n{Nt+i/Nt) and x, = M, equation (6.11) reduces to
y. = a + bxi-cxj , (6.13)
114
which is a quadratic function. Parameters a, b, and c can be solved similarly by fitting a
quadratic curve to the data set and using least-squares estimation. Thus, I estimated
parameters in equations (6.3), (6.6), and (6.9) per playa using an iterative method and
least-squares estimation, and tested for differences between estimation techniques by
computing the difference {D) between estimates per playa then performing a one-sample
Mest on D (HQ: D=0). The aforementioned was necessary because estimation samples
per playa were not independent (Milton and Arnold 1995:353).
Estimating assemblage dynamics in natural systems using deterministic functions
may not be completely realistic because of inherent environmental stochasticity (Ellner
and Turchin 1995). This might be especially tme for amphibian assemblages inhabiting
semi-arid environments, such as on the SHP, because their aboveground dynamics can be
strongly influenced by ambient conditions. Accordingly, Turchin and Ellner (2000)
suggested a stochastic parameter (ct) be added to deterministic functions to increase
realism and account for variability in dynamics associated with the environment. Thus, I
added St to equations (6.6) and (6.9) to create stochastic versions of the Ricker logistic
and Ricker logistic with predator term, respectively:
Nt+\ = Nt^^P + £t (6.14)

V KJ
and.
Nt+\ = Nt^^P ,i-f)-c;v?.. (6.15)
Parameters in equations (6.14) and (6.15) were approximated by fitting the functions to
the observed data and using least-squares estimation (Turchin and Ellner 2000), which
115
was done using PROC NLIN in SAS® (Freund and Littell 2000). A reasonable estimate
of ft for the function is mean-squared error of the nonlinear regression (Turchin and
Ellner 2000). Residuals can be used as time-specific CtS, which were needed to classify
dynamics (discussed later).
Turchin and Ellner (2000) also suggested density-dependent difference equations
(e.g., equations 6.3, 6.6, 6.9, 6.14, and 6.15) should be constmcted with lags to increase
prediction realism. Lags should correspond to duration of delayed density dependence,
which is the embedding dimension of the time series. Because no information exists on
the presence of delayed density dependence in the amphibian assemblage on the SHP, I
assumed embedding dimension = 1 for all aforementioned difference equations. This is
reasonable for many vertebrate assemblages because density dependence and predation
usually most strongly affects individuals in the adjacent time period (Turchin and Ellner
2000). Moreover, graphs of autocorrelation functions from my time series (M. J. Gray.
unpublished data) suggested lack of correlation after 1 time lag (Turchin 1990, 1996).
Thus, I did not incorporate lags into presented functions.
Predicted dynamics from difference equations were classified as chaotic or not
chaotic using published stability estimates from May (19756) for equations (6.3) and
(6.6), and estimating Liapunov exponents for equations (6.9), (6.14), and (6.15). If r >
2.59 and 2.69, dynamics were classified as chaotic for equations (6.3) and (6.6),
respectively (May 19756). Estimation of Liapunov exponents for the other difference
equations was straightforward using equation (6.1), because F{N^ (i.e., the right hand
side of the difference equations) was a constant (c) after solving using parameter
116
estimates. £t and M (Ellner 2000). It follows. DF(M) was mereh' e' for equations (6.9).
(6.14), and (6.15), because the time-specific derivative {J(t)) of M+i= e'N^ was e' (Ellner
2000). Posifive Liapunov exponents indicated chaotic dynamics (Olsen and Degn 1985).
Predicted occurrence of chaos for each difference equation was tested between
landuses and actual dynamics (i.e., chaos occurrence estimated assuming no specific
F(N)) using a main effect logistic regression with a generalized logit model (Stokes et al.
2000:257). Parameter estimates per playa were presented for functions that predicted
chaotic frequency similar to the actual dynamics. Lastly, estimates of parameters for all
difference equations were tested between landuses using a 2-sample /-test (Milton and
Amold 1995:347).
Results
Time series plots of relative abundance at each playa clearly suggested that
within-year dynamics of the SHP amphibian assemblage were not linear and may not
approach a single positive stable equilibrium (Figures 6.1-6.2). Measures of stability for
actual and predicted dynamics from difference equations indicated that chaos was a
fundamental characteristic of the SHP amphibian assemblage (Table 6.1). Actual
occurrence of chaos was greater at grassland (7/8) than cropland (1/8) playas (Table 6.2).
The stochastic Ricker function predicted occurrence of chaos similar to actual dynamics
(i.e., significant and non-significant landuse and model effect, respectively, Table 6.2);
least-squares estimates of its parameters per playa are presented (Table 6.3). The
deterministic and stochastic Ricker fiinctions with a predator term also predicted
dynamics similar to actual dynamics (i.e., no model effect); however, no landuse effect
117
118
Figure 6.1. Time series plots of amphibian assemblage dynamics at 8 playa wetlands
during 16 May-17 October 1999 on the Southem High Plains, Texas. Relative
abundance was total daily capture (natural-log transformed) of all species in pitfall traps;
days were altemate equally spaced days within the time series. Time series for playas 1
and 4 were delayed because rain caused flooding over pitfalls. Playas 1-4 and 5-8 were
located in cropland and grassland, respectively.
6.5
5.5
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10 20 30 40 50 60
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119
120
Figure 6.2. Time series plots of amphibian assemblage dynamics at 8 playa wetlands
during 19 April-18 August 2000 on the Southem High Plains, Texas. Relative
abundance was total daily capture (natural-log transformed) of all species in pitfall traps;
days were altemate equally spaced days within the time series. Playas 9-12 and 13-16
were located in cropland and grassland, respectively.
6 -f
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121
Table 6.1. Stability of amphibian assemblage trajectories at 16 playa wetlands during 16
May-17 October 1999 and 19 April-18 August 2000 on the Southem High Plains, Texas.
Stability
F(Ny Year Landuse Playa Parameter Classification'^
NSF 1999 Cropland 1 -0.004 Not Chaotic
2 -0.015 Not Chaotic
4 0.001 Chaotic
Grassland 5 0.003 Chaotic
6 0.001 Chaotic
7 0.001 Chaotic
8 0.008 Chaotic
2000 Cropland 9 -0.007 Not Chaotic
14 0.015 Chaotic
15 0.015 Chaotic
DDL 1999 Cropland 1 4.465 Chaotic
2 3.691 Chaotic
3 2.693 Chaotic
4 3.931 Chaotic
6 3.242 Chaotic
7 2.791 Chaotic
8 2.841 Chaotic
2000 Cropland 9 2.516 Not Chaotic
10 2.842 Chaotic
11 3.087 Chaotic
12 2.271 Not Chaotic
16 2.861 Chaotic
122
Stability
F(Nf Year Landuse Playa Parameter'' Classification'^
DDR 1999 Cropland 1 3.881 Chaotic
2 3.105 Chaotic
3 3.029 Chaotic
4 3.706 Chaotic
Grassland 5 2.615 Not Chaotic
6 2.922 Chaotic
7 2.665 Not Chaotic
8 3.021 Chaotic
2000 Cropland 9 2.305 Not Chaotic
10 2.706 Chaotic
11 2.817 Chaotic
DSR 1999 Cropland 1 0.006 Chaotic
2 0.004 Chaotic
6 0.009 Chaotic
8 0.008 Chaotic
14 0.024 Chaotic
15 0.015 Chaotic
16 0.011 Chaotic
123
Stability
F(nf Year Landuse Playa Parameter' Classification'
DRP 1999 Cropland 1 -0.036 Not Chaotic
2 0.023 Chaotic
Grassland 5 -0.093 Not Chaotic
7 0.002 Chaotic
8 0.087 Chaotic
15 0.101 Chaotic
16 0.252 Chaotic
SRP 1999 Cropland 1 -0.029 Not Chaotic
2 0.021 Chaotic
4 -0.423 Not Chaofic
7 0.006 Chaotic
8 0.098 Chaotic
10 -0.015 Not Chaofic
15 0.099 Chaotic
16 0.248 Chaotic
124
^ S F - no specific form assumed (i.e., Nt+i=F(Ni), tme population dynamics), DDL =

discrete deterministic logistic (M+i=M + rNt[\- Nt/K]), DDR = discrete determinisfic
Ricker (M+i=Mexp[r(l- N^/K)]), DSR = discrete stochastic Ricker (M+i=Mexp[r(l- M / ^
+ etl), DRP = discrete determinisfic Ricker with predator term (M+i=Mexp[r(l- Nt/K) -
cNt ]), and SRP = discrete stochastic Ricker with predator term (M+i=Mexp[r(l- Nt/K) -
cNt^ + et]) {May 1975b).
''Stability parameter was X (Liapunov exponent) for NSF, DSR, DRP, and SRP and r
(intrinsic rate of increase) for DDL and DDR (May 19756, Olsen and Degn 1985).
"Dynamics classified as chaotic when A > 0, and r > 2.59 and r > 2.69 for DDL and DDR,
respecfively (May 19756, Olsen and Degn 1985).
125
Table 6.2. Occurrence of chaos between landuses and models in amphibian assemblages
at 16 playa wefiands during 16 May-17 October 1999 and 19 April-18 August 2000 on
the Southem High Plains, Texas.
Percent Chaos Landuse Effect Model Effect"
F(N/ Cropland Grassland X'^ P X^ P
NSF 12.5 87.5 9.0 0.003 NT NT
DDL 75.0 75.0 0.0 1.0 4.26 0.039
DDR 75.0 37.5 2.29 0.131 6.35 0.012
DSR 25.0 87.5 6.35 0.012 0.18 0.675
DRP 12.5 50.0 2.62 0.106 0.96 0.326
DSP 12.5 50.0 2.62 0.106 0.96 0.326
^ S F = no specific form assumed (i.e., Nt+\=F(Nt). true population dynamics), DDL
discrete deterministic logistic (M+i=M + rNt[\- Nt/K]), DDR = discrete deterministic
Ricker {Nt+i=NtQxp[r{l- Nt/K)]), DSR = discrete stochastic Ricker {Nt+i=Ntexp[r{\- Nt/K)
+ Et]), DRP = discrete deterministic Ricker with predator term {Nt+\=Ntexp[r{\- Nt/K) -
cNt ]), and SRP = discrete stochastic Ricker with predator term {Nt+\^Nttxp\r{\-Nt/K) -
cM^ + et]) (May 19756).
''Model effect was pairwise Fisher's exact tests in percent chaos between NSF (i.e., tme
dynamics) and other functions; no test (NT) was performed on NSF and itself
126
Table 6.3. Least-squares esfimates of parameters for the discrete stochastic Ricker
equation^ from amphibian assemblages at 16 playa wetlands during 16 May-17 October
1999 and 19 April-18 August 2000 on the Southem High Plains, Texas.
Parameter Estimate
Year Landuse Playa r K Et MSreg" F" P
1999 Cropland 1 0.273 4.493 0.057 0.168 5.65 0.022
2 0.292 3.551 0.174 0.798 9.14 0.004
3 0.308 3.795 0.113 0.392 6.60 0.013
4 0.118 3.407 0.116 0.189 2.46 0.123
Grassland 5 0.593 4.064 0.247 1.578 12.71 <0.001
6 0.673 3.187 0.211 2.561 24.27 <0.001
7 0.813 3.924 0.175 2.394 27.35 <0.001
8 0.486 3.251 0.209 1.568 15.02 <0.001
2000 Cropland 9 0.491 3.041 0.228 1.277 11.03 0.002
10 0.346 3.807 0.131 0.421 6.07 0.018
11 0.457 3.996 0.111 0.643 11.04 0.002
12 0.677 2.992 0.317 2.955 18.55 <0.001
Grassland 13 0.525 2.113 0.189 1.186 12.54 0.001
14 0.467 2.731 0.174 1.059 12.19 0.001
15 0.654 3.679 0.189 1.571 16.55 <0.001
16 0.511 3.509 0.159 1.025 12.83 <0.001
^ h e discrete stochastic Ricker equation is A^t+i=Mexp(r[l-A^t/-^ + ^t); parameter
estimates can be used for predicting playa- and time-specific assemblage dynamics.
''Mean-square regression (MSreg) is variation in assemblage dynamics explained by the
discrete stochastic Ricker equation (Freund and Littell 2000:185-189).
'^Numerator, denominator degrees of freedom of F were 2,65 and 2,50 for 1999 and 2000,
respectively.
127
was detected, which was inconsistent with observed dynamics (Table 6.2). The
stochastic Ricker function predicted mean intrinsic rate of increase was greater at
grassland than cropland playas; carrying capacities were similar (Table 6.4). No
differences in parameters between landuses were detected for other functions (Table 6.4).
Lastly, estimates of parameters for the deterministic logistic, Ricker, and Ricker with
predator term functions were different the between iterative method and least-squares
estimation (Table 6.5). Estimates of intrinsic rate of increase were greater using the
iterative method than by least squares (Table 6.5). Parametric assumptions of residual
normality (P>0.79) and homoscedasticity {P>0.23) were satisfied for all ^tests.
Discussion
Occurrence of chaos in the SHP amphibian assemblage was greater at grassland
than cropland playas. These results represent the first empirical evidence of chaos in
spatially stmctured amphibian assemblages, and more importantly, that its frequency can
be affected by anthropogenic disturbance. Occurrence of chaos may have been lower at
cropland playas, because anthropogenic disturbance might have affected normal
assemblage processes, such as trophic stmcturing or interdemic exchange of individuals
among spatially stmctured populations. Agricultural landscape use and stmcture can
affect local population demographics and body size of amphibians (Chapters II-IV). In
particular, mean daily abundance of some species was greater at cropland than grassland
playas, perhaps because of the anthropogenic reduction of an intraguild predator (i.e.,
barred tiger salamander, Ambystoma tigrinum mavortium) in cropland playas. Also,
differential viscosity of anthropogenic crop types may have influenced movements of
128
Table 6.4. Iterafive and least-squares estimates of parameters for deterministic difference
equafions used to model amphibian assemblages at 16 playa wetlands during 16 May-17
October 1999 and 19 April-18 August 2000 on the Southem High Plains, Texas.
Estimation Technique Paired
F(N/ Parameter Iteration Least-Squares Difference'^
X SE X SE X SE /15 P
DDL r 3.17 0.15 0.78 0.09 2.39 0.21 11.42 <0.001
K 3.94 0.16 4.12 0.16 0.17 0.07 2.49 0.025
DDR r 2.74 0.15 0.48 0.05 2.26 0.18 12.57 <0.001
K 3.89 0.15 3.47 0.15 0.43 0.08 5.27 <0.001
DRP r -1.52 4.10 0.89 0.13 11.33 3.02 3.74 0.002
K 2.40 0.26 2.89 0.60 1.48 0.61 2.42 0.029
c 0.10 0.22 -0.03 0.008 0.61 0.17 3.66 0.002
"^DL = discrete determinisfic logisfic (M+i=M + ^ ^ [ 1 - Nt/K]), DDR = discrete
deterministic Ricker {Nt+i=NtQxp[r{\- Nt/K)]), and DRP = discrete deterministic Ricker
with predator term (M+i=Mexp[r(l- Nt/K) -cM^]) (May 19756).
Iterative estimates were computed by making a change of variable {a=\+r and b=r/K for
DDL, a=r and b=r/K for DDR, and a=r, b=r/k, and c=c for DRP), solving 2-3 iterates
using daily capture per playa for M+i and Nt, and computing the arithmetic mean within
then across playas; least-squares estimates were ordinary least-squares computed by
fitting the deterministic function to the entire data set and estimating parameters that
allow the best available fit (Milton and Amold 1995:386).
'^Mean paired difference {D) was the absolute difference between iterative and least-
square estimates averaged across playas; a one-sample /-test (HQ: D = 0 ) was performed on
arithmetic mean of D to test for difference between parameter estimates because samples
were not independent (Milton and Amold 1995:353).
129
Table 6.5. Esfimates of parameters between landuses for difference equations used to
model amphibian assemblages at 16 playa wefiands during 16 May-17 October 1999 and
19 April-18 August 2000 on the Southem High Plains, Texas.
Cropland Grassland
F(nr Parameter X SE X SE tu P
DDL r 3.19 0.27 2.74 0.14 1.48 0.161
k 4.19 0.18 3.75 0.27 1.36 0.196
DDR r 2.92 0.24 2.56 0.17 1.27 0.225
k 4.09 0.19 3.69 0.22 1.35 0.199
DSR r 0.37 0.06 0.59 0.04 3.02 0.009
k 3.64 0.18 2.77 0.23 1.14 0.275
DRP r 4.07 6.78 -7.02 4.21 1.39 0.186
k 2.52 0.46 2.28 0.26 0.46 0.651
c -0.17 0.39 0.37 0.21 1.23 0.239
DSP r 0.62 0.18 1.02 0.16 1.64 0.123
k 1.85 0.32 2.35 0.23 1.25 0.231
c -0.02 0.01 -0.05 0.02 2.06 0.361
"^DL = discrete deterministic logisfic (M+i=M + rNt[\- NJK]), DDR = discrete
deterministic Ricker {Nt+\=NtQxp[r{\- Nt/K)\), DSR = discrete stochastic Ricker
(M+i'^Mexp[r(l- Nt/K) + et]), DRP = discrete deterministic Ricker with predator term
{Nt+\=Nttxp[r{\- Nt/K) -cN^]), and SRP = discrete stochastic Ricker with predator term
(M+i=Mexp[r(l- Nt/K) -cN^ + Et]) (May 19756).
130
individuals among local assemblages. Existence of predafion and interdemic movement
of individuals among grassland playas may act as natural destabilizing mechanisms,
owing to the increased occurrence of chaos in undisturbed grassland playas. Allen et al.
(1993) demonstrated that chaos was an important characteristic of simulated spatially
structured populations, because it can reduce probability of metapopulation extinction by
amplifying local population noise. Thus, a spatially structured amphibian assemblage
surrounded by cultivation may have a greater probability of extinction than one
surrounded by an undisturbed landscape.
Differences in occurrence of chaos in the SHP amphibian assemblage also may
have been related to density dependence and environmental stochasticity. The stochastic
Ricker function predicted occurrence of chaos most similar to actual dynamics,
suggesting that density dependence is an important regulating mechanism in the
assemblage. Density dependence has been hypothesized and shown to regulate some
amphibians (Wilbur 1980, 1984). Density dependence probably is a regulating
mechanism in the SHP amphibian assemblage, because most species are explosive
breeders that emerge simultaneously and produce numerous offspring (see species review
in Chapter II). Consequently, relative abundance of individuals can increase rapidly. For
example, total daily capture increased from 30 to 14,623 individuals in 1 week at 1 playa
in 1999. I suspect competition for food, cover, and mating resources may become
limiting under these and less extreme dynamics, inducing density dependent regulation.
Density independence also seems to be important in regulating the SHP amphibian
assemblage, because the deterministic Ricker model did not predict occurrence of chaos
131
similar to actual dynamics. Thus, these results support Ellner and Turchin (1995)
hypothesis that including a stochastic parameter in completely deterministic models
makes them more realistic.
The stochastic Ricker function also predicted that intrinsic rate of increase was
greater in grassland than cropland playas. Elementary stability analyses demonstrate that
as the control parameter of an underlying function (such as r) increases, probability of
progression to a chaotic state also increases (e.g.. May 1975). The aforementioned may
be the functional response to suspected differential predation, interdemic movement, and
density dependence between landuses, and the mathematical reason for differences in
chaotic occurrence between landuses.
Although no differences were detected, the stochastic Ricker function also
predicted mean carrying capacity was greater at cropland (x=38.1 captures/day, SE=1.19)
than grassland (x=15.9 captures/day, SE=1.26) playas. As I hypothesized in Chapters II
and III, carrying capacity may be lower in grassland playas because larval and neotenic
barred tiger salamanders may be exhibiting top-down control. Indeed, deterministic and
stochastic Ricker functions with predator terms predicted occurrence identical and similar
A
to actual dynamics in cropland and grassland, respectively. Nutrient loading also may be
greater in cropland than grassland playas due to agriculture (Freemark and Boutin 1995),
facilitating greater numbers of individuals in cropland playas via increasing food
abundance (Chapter III). Although abundance and carrying capacity may be higher in
cropland playas, the chaos results herein indicate that dynamics of an amphibian
assemblage in disturbed wetlands are altered from their natural state in grasslands.
132
Estimates of intrinsic rate of increase were greater using the iterati\ e method than
least squares for all difference equations. This probably was a consequence of
methodological differences in their algorithms. As described previously, sets of
difference equations were solved and parameters estimated for 2-3 sequential time
iterates for the iterative technique. Thus, parameters per playa were approximated from
m/2 or m/3 samples (where m is the duration of the time series) for non-predator and
predator difference functions, respectively. The arithmetic average then was calculated
per playa for each parameter estimate. In contrast, the least-squares technique fit the
difference function to the entire data such that residual sum-of-squares v\ as minimized,
and obtained one sample estimate per parameter per playa. Consequently, iteration may
capture the true dynamics of a chaotic time series better than least-squares estimation
because it is based on more samples. However, limited simulations (J. Surles, Texas
Tech University, unpublished data) suggest that least-squares estimates are less \ ariable
and more robust than those from the iterative method.
Ecological and Mathematical Implications
Chaos was a ftjndamental characteristic of a spatially stmctured amphibian
assemblage on the SHP, and anthropogenic disturbance around playa wetlands can
dismpt its occurrence. Chaos may exist naturally in this assemblage because of densit>
dependent and independent regulation. Dynamics also may be a consequence of
predation and interdemic movement (Chapters III and IV). Additional research is needed
to test these hypotheses. Regardless, dynamics of amphibians in cropland wetlands are
altered from their natural state in grasslands. These results and those of Allen et al.
133
(1993) on simulated metapopulations suggest that spatially stmctured amphibians
positioned in cropland may have a greater probability of extinction, because of reduced
natural occurrence of chaos. Future research needs to discem differences in the
mechanisms regulating amphibian assemblages and populations between these
anthropogenic landuse types. Subsequently, we need to evaluate conservation techniques
(e.g., wetland buffer restoration, corridor construction) that restore dynamics and
demographics similar to an undisturbed state. Amphibian dynamics at undisturbed and
disturbed wetlands can be predicted initially by using the stochastic Ricker model and
parameter estimates presented herein (Table 6.3). The true underlying function
describing dynamics and the relationship of regulating mechanisms in the SHP
amphibian assemblage likely is more complex than the stochastic Ricker function. Thus,
future modeling endeavors should incorporate additional parameters, such as immigration
and emigration, and investigate other relationships of density dependence (e.g.,
N,+]âNt/[l+{bNty], Maynard Smith 1974) and predation. Future models also should
include a stochastic parameter (ct) to improve predictive ability. Additional density
independent state variables (e.g., rainfall, water chemistry) could be added to refine
models. Cross validation techniques and information statistics (e.g., Akaike information
criterion) can be used to compare predictability among difference equations (Turchin and
Ellner 2000). Lastly, species-specific analyses would be useful to discem if chaos is a
ftjndamental characteristic of SHP amphibian populations. Reduced detection of chaos at
the population-level may give insight into the embedding dimension of the amphibian
assemblage, and provide support for combined assemblage analyses as presented here.
134
CHAPTER VII
CONCLUSION
This study provides evidence that anthropogenic disturbance surrounding
wetlands affects body size, demographics, and chaotic dynamics of amphibians on the
Southern High Plains. In particular, postmetamorphic body size of amphibians was
greater at grassland playa wetlands than at playas surrounded by agricultural cultivation
(Chapter II). The presence of biological chaos also was altered by anthropogenic
disturbance (Chapter VI). Amphibian assemblages surrounded by grassland exhibited
chaotic dynamics more often than assemblages in cropland landscapes. Abundance of
New Mexico and plains spadefoots {Spea multiplicata, S. bombifrons) was positively
affected by cultivation; abundance of other species and diversity of the amphibian
assemblage was not affected (Chapter III). Grassland playas exhibited source dynamics
most often, while playas in cultivated landscapes generally were sinks (Chapter III).
Landuse did not affect temporal niche partitioning among amphibian species (Chapter V).
Geometric indices representing spatial positioning of playas and landscape complexity
were positively related with amphibian abundance and affected community composition
(Chapter IV). A stochastic Ricker equation predicted that density dependence and
environmental stochasticity were possible regulating mechanisms in the Southem High
Plains amphibian assemblage (Chapter VI).
Various abiotic and biotic factors probably infiuenced amphibian responses to
anthropogenic disturbance treatments (cultivation vs. grassland). Differences in body
size between cultivated and grassland landscapes may have been related to wetland
135
hydroperiod, agricultural chemicals, availability of food resources, or density of
conspecifics (Chapter II). Occurrence of biological chaos probably was greater in
grassland landscapes, because intrinsic rate of increase was greater at grassland than
cropland playas (Chapter VI). Differences in chaotic dynamics between anthropogenic
landuses may have been related to competition, predation, interdemic movement, or
environmental stochasticity (Chapter VI). Species-specific vagility and anthropogenic
reduction of barred tiger salamanders {Ambystoma tigrinum mavortium) in the aquatic
environment may have positively influenced spadefoot abundance at cropland playas
(Chapter III). Rigidity in the genie norm of reaction for temporal resource use may have
resulted in no differences between landuses in temporal niche partitioning (Chapter V).
Amphibian abundance was positively related to playa juxtaposition, probably because
probability of interdemic movement increases with decreasing dispersal distance
(Chapter IV). Abundance might be positively related to landscape complexity, because
amphibians may perceive complex agricultural landscapes as impervious, resulting in
increased abundance near natal wetlands (Chapter IV). Most of these inferences are
based on results in lab and field experiments on amphibians elsewhere or from
predictions by difference equations; thus, they are speculative. More research is needed
(discussed later). Regardless, results herein strongly suggest that agricultural cultivation
around wetlands affects amphibian populations, thus the relative importance of various
regulating mechanisms (e.g., competition, predation, interdemic movement).
Although spadefoot abundance was positively influenced by cultivation, I
recommend retention and restoration of native grasslands surrotmding playa wetlands.
136
Cropland playas exhibited sink dynamics whereas playas in grassland landscapes were
most often sources of individuals (Chapter III). Inasmuch as body size is positively
related to survival and reproduction (thus evolutionary fitness) in amphibians,
populations surrounded by cropland also may have a greater probability of extinction
(Chapter II). Presence of biological chaos may increase persistence of spatially
structured populations by increasing local population noise; thus, an amphibian
metapopulation in a cropland landscape may have greater probability of exfinction than
one in grassland (Chapter VI). Incidence of disease, parasite infestation and
malformations also can be positively related to amphibian abundance thus increase in
cropland landscapes (Carey et al. 1999). Lastly, I suspect that interdemic movement is
reduced in cropland landscapes, which can have negative demographic and genetic
consequences on amphibian populations (Brown and Kodric-Brown 1977, Hastings and
Harrison 1994).
Additional research is needed on the effect of anthropogenic disturbance on
Southem High Plains (SHP) amphibians. I recommend the following studies: (1)
determine the effect of landscape use and structure on interdemic genetic variability of
SHP amphibians, (2) estimate species-specific vagility of SHP amphibians in natural and
anthropogenic cover types, (3) estimate home ranges, dispersal probabilities, and micro-
habitat use of SHP amphibians, (4) simultaneously examine the effect of anthropogenic
landuse on sedimentation rate, water chemistry, aquatic and terrestrial food resources,
and vegetation characteristics in playa wetlands, (5) investigate the effect of landuse on
body size, demographics, and dynamics of SHP amphibian larvae populations, (6)
137
examine pathogenic incidence between landuses in SHP amphibians, (7) determine the
effect of UV-B radiation on hatching and survival of SHP amphibian larvae, and (8)
determine the effect of landuse on the relative importance of regulafing mechanisms (e.g..
competition and predation) in aquatic and terrestrial amphibian populations. These
studies could be accomplished through multiple concurrent lab and field experiments. A
manipulative study in which populations were monitored in a cropland landscape before
and after grassland restoration, or alternatively in a grassland landscape before and after
disturbance, would be novel and provide strong inferential insight. Results from the
aforementioned research could be used to predict population dynamics using difference
equations or discrete Markov chains. Moreover, landscape changes could be simulated
using the geographic information system to predict the effect of restoration techniques
(e.g., entire grassland restoration, corridor construction) or disturbance on amphibian
populations. Lastly, a long-term amphibian population monitoring program needs to be
established on the SHP, and demographic information of playa amphibians tied to global
declines.
The Southem High Plains is an incredible region to examine the effect of
anthropogenic disturbance on amphibian populations. Thousands of playa wetlands in
cultivated and grassland landscapes allow comparative examination of human impacts on
wildlife populations with substantial statistical replication. Playas also are spatially
stmctured; thus, they are ideal systems to test various hypotheses of metapopulation
theory. Therefore, an experimenter can investigate progressive concepts in conservation
biology and landscape ecology simultaneously in nature—This is a rare scientific
138
opportunity. I encourage continued amphibian research on the Southem High Plains.
Indeed, Ann Anderson's (1997) research and this study only have scratched the surface of
our understanding of basic life history for SHP amphibians and the effect of human
disturbance on their population processes. I am confident the stellar researchers at Texas
Tech University will guarantee the progression of science and our knowledge. Buena
suerte y que la ciencia provea la luz\
139
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York, New York, USA.
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166
APPENDIX A
PERCENT COVERAGE OF COVER TYPES IN LANDSCAPES
SURROUNDING PLAYA WETLANDS
167
Table A.l. Percent coverage of cover types in landscapes^ surrounding 16 playa wetlands
between landuses on the Southem High Plains, Texas, 1999 and 2000.
Landuse"
Cover t> pe*^ Cropland Grassland
X SE X oh
Alfalfa Medicago sativa 0.12 0.08 0.02 0.01
Barley Hordeum spp. 0 0 0.04 0.04
Com Zea mays 2.11 1.31 1.07 0.47
Cotton Gossypium hirsutum 60.3 5.37 2.75 1.27
Ha> Phalaris arundinacea 0.43 0.19 1.17 0.67
Kleingrass Panicum coloratum 0.45 0.19 0.34 0.17
Millet Pennisetum glaucum 0.15 0.15 0 0
Oats Arena sativa 0.21 0.11 0.09 0.08
Rye Secale cereale 0.17 0.17 0 0
Sorghum Sorghum vulgare 4.78 1.39 3.21 0.64
Soybean Glycine max 1.14 0.81 0 0
Sunflower Helianthus annuus 0.79 0.24 0 0
Vegetables (various spp.) 0.04 0.02 0 0
Wheat Triticum aestivum 6.95 2.28 10.9 2.96
Fallow 2.55 0.66 1.65 0.52
Buildings 0.96 0.24 0.38 0.12
Maintained county roads 1.67 0.09 1.05 0.09
Interstate 0.32 0.32 0 0
Other anthropogenic structures 0.02 0.02 0.01 0.01
Canyon 0.96 0.96 0 0
Native grass 3.19 0.67 28.8 4.34
Replanted grass 8.47 1.29 46.1 6.78
Playa wetland 3.86 0.32 2.35 0.23
Water 0.29 0.14 0.02 0.02
^Landscapes were 2830-ha circular plots (i.e., 3-km radius) with their origins positioned
at the center of each study playa.
''Percent cover of cotton, county roads, and native and replanted grass differed (P<0.006)
between landuses by Wilcoxon 2-sample nonparametric tests (Conover 1980:216); no test
was performed between landuses when x and SE=0; « = 8 playas/landuse.
'Ôther anthropogenic stmctures included cemeteries, quarries, and airstrips; replanted
grass was cultivated previously then revegetated by the Conservation Reserve Program.
168
APPENDIX B
ANOVA TABLES FOR CHAPTER III
169
Table B.l. Statistics for ANOVAs testing differences in relative daily abundance and
diversity of amphibians between landuse t>pes (cropland \s. grassland) and years and
among species at 16 playa wetlands on the Southem High Plains, Texas. May-October
1999 and April-August 2000.
Variable Tests and Effects^'' df^ F P
Abundance 3-wa> ANOVA
Landuse 1 11.97 <0.001
Year 1 10.54 0.002
Species 6 55.67 <0.001
Landuse x Year 1 0.80 0.375
Landuse x Species 6 8.36 <0.001
Year >: Species 6 2.52 0.051
Landuse x Year x Species 6 1.40 0.223
2-wa> ANOVA
NSF
Landuse 20.33 <0.001
Year 2.63 0.131
Landuse x Year 2.34 0.152
BTS
Landuse 1.26 0.283
Year 8.08 0.015
GPT
Landuse 0.01 0.913
Year 2.35 0.152
PSF
Landuse 7.94 0.016
Year 0.51 0489
SCF
Landuse 0.23 0.639
Year 1.99 0.184
PLF
Landuse 0.156 0.156
Year 0.289 0.289
WHT
Landuse 0.33 0.576
Year 1.00 0.338
170
Table B.l. Continued.
Variable Tests and Effects*^^ df F
Abundance 1-way ANOVA
Cropland
Species 6 34.20 <0.001
Grassland
Species 6 18.74 <0.001
1999
Species 6 20.06 <0.001
2000
Species 6 15.56 <0.001
Diversity' 2-way ANOVA
Landuse 1 0.91 0.359
Year 1 0.29 0.602
Overall 3-way ANOVA on abundance was separated b\ species to test for differences
between landuses and years (i.e., 2-way analyses) then by landuse and year (i.e.. l-wa>
analyses) to test the species effect, because species interacted with landuse and year in
the 3-wa>' analysis.
''NSF = New Mexico spadefoot {Spea multiplicata), BTS = barred tiger salamander
{Ambystoma tigrinum mavortium). GPT = Great Plains toad {Bufo cognatus), PSF =
plains spadefoot {S. bombifrons), SCF = spotted chorus frog {Pseudacris clarkii). PLF =
plains leopard frog {Rana blairi), and WHT = Woodhouse's toad {B. woodhousii).
'Êrror (denominator) df for 3-way ANOVA, 2-way ANOVAs. and 1-way ANOVAs was
84 (2 landuses x 2 years x 7 species x 3 playas), 12 (2 landuses x 2 >ears x 3 playas). and
49 (7 species x 7 playas), respectively.
'^Diversity was calculated using the Shannon-Weaver algorithm (Hair 1980:273); no
species effect was included in analyses because all species were used in estimation of the
index.
171
APPENDIX C
AGE CLASS .\NOVA TABLES FOR CHAPTER III
172
Table C.l Stafistics for ANOVAs for species (by age class) that differed significantly in
relati\ e dail>' abundance between landuse types (cropland vs. grassland) or years (see
Table B.l) at 16 playa wetlands on the Southern High Plains, Texas, May-October 1999
and April-August 2000.
Species Tests and Effects df
NSF 2-way ANOVA
Metamorph
Landuse 12 6.79 0.023
Year 12 13.99 0.003
Subadult
Landuse 12 17.18 0.001
Year 12 50.06 <0.001
Landuse x Year 12 21.26 <0.001
Adult
Landuse 12 14.02 0.002
Year 12 0.10 0.758
Landuse x Year 12 048 0.499
PSF Metamorph
Landuse 12 1.93 0.189
Year 12 12.1 0.005
Subadult
Landuse 12 6.74 0.023
Year 12 11.69 0.005
Adult
Landuse 12 3.71 0.078
Year 12 7.88 0.016
BTS Metamorph
Landuse 12 2.07 0.176
Year 12 10.50 0.007
Adult
Landuse 12 0.22 0.648
Year 12 0.76 0.399
NSF 1-way ANOVA
Metamorph
Landuse in 1999 1,6 17.46 0.006
Landuse in 2000 1,6 0.17 0.698
Year in Cropland 1.6 34.84 0.001
Year in Grassland 1,6 0.11 0.747
173
T a b l e d . Continued.
Species Tests and Effects df F P
NSF 1-way ANOVA
Subadult
Landuse in 1999 1,6 0.36 0.570
Landuse in 2000 1,6 22.55 0.003
Year in Cropland 1,6 43.66 <0.001
PSF 1-way ANOVA
Subadult
Landuse in 1999 1,6 1.96 0.211
Landuse in 2000 1,6 7.79 0.032
Year in Cropland 1,6 10.59 0.017
"^NSF = New Mexico spadefoot {Spea multiplicata), PSF = plains spadefoot {S.
bombifrons), and BTS = barred tiger salamander {Ambystoma tigrinum mavortium).
Overall 2-way ANOVA on abundance of significant species was separated by landuse
and year for metamorph NSF and subadult NSF and PSF (i.e., 1-way analyses), because
landuse and year main effects interacted.
174
APPENDIX D
CROPLAND AND GRASSLAND LANDSCAPES
175
176
Figure D.l. Anthropogenic and natural cover types in landscapes (i.e., 2,830-ha [3-km
radius] circular plots) surrounding 16 playa wetlands on the Southem High Plains, 1999
and 2000. Each study playa is positioned at the landscape plot origin. Crop types
included alfalfa {Medicago sativa), barley {Hordeum spp.), com {Zea mays), cotton
{Gossypium hirsutum), domestic grass {Panicum coloratum), hay {Phalaris
arundinacea), millet {Pennisetum glaucum), oats {Avena sativa), sorghum {Sorghum
vulgare), soybean {Glycine max), sunflower {Helianthus annuus), various vegetable
species, and wheat {Triticum aestivum). Fallow was arable but unplanted land. Non-
playa water was permanent water resulting from anthropogenic modifications of the
landscape (e.g., tailwater pit, effluent treatment pond). Conservation Reserve Program
(CRP) grasslands were cultivated previously and replanted with native and exotic grasses
for erosion control and wildlife habitat.
Cropland Landscapes, 1999
0 1,000 2,000 4,000 6,000 8,000

I Meters
_ _ _ Alfalfa Sorghum Airstrip
I I Com Soybean Buildings
1111111 Cotton Sunflower Maintained Roads
I I Domestic Grass Vegetables Playa Wetland
Hay Wheat Non-playa Water
i l i i Milet Fallow Native Grassland

f^^m
I Oats %>mi 1-27 CRP Grassland
Figure D. 1. Continued.
177
Grassland Landscapes, 1999
N
0 1,000 2,000 44K)a 6.000 8.000

(Meters
Com Hwy385
Cotton Buildings
^?^
Domestic Grass Maintained Roads
I Hay Playa Wetland
Sorghum Native Grassland
Wheat CRP Grassland
Fallow
Figure D.l. Continued.
178
Cropland Landscapes, 2000
0 1,000 2,000 4J0OO 6,000 8,000

I Meters
^ ^ 1 Vegetables Raya Wetland
n u l l Cotton Norvplaya Water
Kss Domestic Grass p• -• "•• j FaHow Native Grassland
r n n a y I j Cemetery CRP Grassland
J H J H Sorghum ^ H H Buildings V\^rte River Canyon
Sunflower ^ ^ H Maintained Roads
179
Grassland Landscapes, 2000
0 1,000 2,000 4,{KK) 6,000 8,000

[Meters
Alfalfa Sofghum Playa W ^ b n d
Barley \/\*ieat Non-playa Water
Com Fallow Native Grassland
Cotton Buildings CRP Grassland

Domestic Grass Gravel Pit Bumt Grassland
J Hay Maintained Roads
t Oats
180

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EFFECT OF ANTHROPOGENIC DISTURBANCE AND

LANDSCAPE STRUCTURE ON BODY SIZE, DEMOGRAPHICS, AND

CHAOTIC DYNAMICS OF SOUTHERN HIGH PLAINS AMPHIBIANS

MATTHEW JAMES GRAY, B.S., M.S.

Submitted to the Graduate Faculty

opportunity to continue my education and professional development at Texas Tech

and conducting research independently. Dedicating my dissertation to him is not enough

Next, I thank my minor professor. Dr. Linda J. S. Allen of the Department of

burgeoned my interest in biomathematics. I now understand and appreciate the

usefulness of mathematics in explaining ecological phenomena. Indeed, her tutelage

Each of these individuals helped me develop important quantitative skills.

my four years at TTU. My dissertation is a testament of their devotion and support.

numerous stimulating discussions on experimental design and statistics. I am grateful to

Biological Sciences) for editorial comments on my dissertation. I also thank the

Raquel Leyva for instructing me in ERDAS®, ESRI®, and FRAGSTATS® software.

I am deeply indebted to the landowners of my study playas: A. Arajo, L. Crum, K.

Gabel, B. Harrell, R. Hunter, B. Kimbrough, M. Marley, S. McPhearson, J. Merritt, G.

Olson, S. Olson, D. Schulte, E. Schulte, C. Sellers, E. Verett, and M. Wilmeth. Their

many individuals. I gratefully thank: A. Anderson, J. Anderson, J. Avey, R. Brenes, W.

Dayawansa, E. Edwards, J. Emrich, D. Garza, D. Ghioca, W. Huskisson, A. Kauffman,

R. Kostecke, P. Lemons, A. Marshall, R. McCaffrey, D. Miller, K. Offill, J. Pautz, C.

Perchellet, R. Phillips, A. Quarels, D. Ray, W. Redell, S. Sebring, C. Shavlik, and C.

him equally. Muchisimo gracias y pura vida, Sementall

Finally, I am grateful for the incredible support of my family: James S. Gray

patience gave me the strength to succeed.

interaction: Drs. Harold H. Prince (Michigan State Universit>). Richard M. Kaminski

(Mississippi State Universit>'). Francisco J. Vilella (Mississippi Cooperati\ e Fish and

Thank \ ou so much gentlemen!

LIST OF FIGURES xiii

II. EFFECT OF LANDUSE AND RAINFALL ON POSTMETAMORPHIC

III. EFFECT OF LANDUSE AND YEAR ON POPULATION

IV. INFLUENCE OF LANDSCAPE STRUCTURE ON COMMUNITY

V. TEMPORAL NICHE PARTITIONLNG IN SOUTHERN HIGH PLAINS

VI. CHAOS AND REGULATING MECHANISMS IN A SOUTHERN HIGH

VII. CONCLUSION 135

LITERATURE CITED 140

A. PERCENT COVERAGE OF COVER TYPES IN LANDSCAPES

B. ANOVA TABLES FOR CHAPTER III 169

C. AGE CLASS ANOVA TABLES FOR CHAPTER III 172

D. CROPLAND AND GRASSLAND LANDSCAPES 175

Amphibian populations are declining globally. Anthropogenic disturbance of

landscapes surrounding wetlands may affect fitness, demographics, and dynamics of

amphibian populations. Spatial positioning and relati\'e connectedness of wetlands also

may influence population demographics. Thus, I examined the effect of anthropogenic

fitness correlate), demographics, and dynamics of amphibians at 16 playa wetlands on the

salamanders {Ambystoma tigrinum mavortium) in 1999 than 2000. Mean daily

abundance of amphibians also was positively related to landscape structure indices

representing geometric complexity and spatial positioning of wetlands. In general, as

distance decreased, mean daily abundance of amphibians increased. Additional

amphibian populations; however, no differences existed between landuses. Lastly,

biological chaos in the amphibian assemblage existed at 1 of 8 cropland and 7 of 8

dynamics most accurately.

Anthropogenic disturbance surrounding wetlands affects body size,

demographics, and dynamics of SHP amphibians. Spatial positioning of wetlands and

landscape complexity may be as or more important than general landuse in affecting

amphibian demographics. Armual differences in body size and abimdance suggest

rainfall may be important in influencing amphibian populations. Although spadefoot

abundance was positively influenced by anthropogenic disturbance, I recommend

retention and restoration of grasslands surrounding playa wetlands because landscape

an undisturbed state. These results have important implications in conservation biology,

landscape ecology, and basic ecological and mathematical theory.

4.1. Pearson coefficients of correlations and p-values (located beneath coefficients)

6.1 Stability of amphibian assemblage trajectories at 16 playa wetlands during 16