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STUDENT GRANT PROGRAM

The Effect of Adaptive Evolution and Ecology on Thermoregulatory Behavior in Holbrookia maculata SGP: A
$1047.60
White Sands, New Mexico, is a site of recent evolutionary divergence in three lizard species. I will examine how previous
evolutionary adaptation of Holbrookia maculata (Lesser Earless Lizard) in White Sands has affected thermoregulatory
behavior. I hypothesize that there has been a directional change in phenotypic traits in Holbrookia maculata in White
Sands from their dark soils conspecifics that affects their thermoregulatory behavior. I will work with two undergraduates
and three doctoral students in White Sands, New Mexico, for approximately two one-month intervals. I will capture lizards
by hand and measure traits such as body size, body temperature, environmental temperature (surface and air), and dorsal
surface area. I will also record lizard features such as sex and age that will allow me to document variation in the wild. The
divergence of the two lizards has shown cascading affects on phenotype (Des Roches et al. 2011). The thermoregulatory
behavior of ectotherms is greatly affected by their environment. The changes in coloration among H. maculata in respect to
their surrounding environment must have influenced change in their thermoregulatory behavior. Habitats are changing due
to shifts in climate, understanding how ectotherms adapt relative to their environment will provide managers with foresight
on how to better protect similar species.
Isaiah Hoyer 100 hoye6378@vandals.uidaho.edu
081-34886 208-215-8841
102 South Lilly Street Moscow, Idaho 83843
College of Natural Resources
Dr. Luke Harmon LSS 347
Biological Sciences 208-885-0346

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Janet Rachlow jrachlow@uidaho.edu

Natural Resources
Tom Poorten tom.poorten@gmail.com
Biological Sciences
Mikki Brinkmeyer brin5845@vandals.uidaho.edu
Biological Sciences and Natural Resources

2010-48 10 May 2010

I will be capturing lizards in the eld and measuring their temperature and recording
morphological measurments. Lizards captured will also be transported back to the eld
lab to be photocopied for other measurements, only to be released the following day. No
direct harm is intended or anticipated from handiling the lizards.

STUDENT GRANT PROGRAM

Equipment {itemized)
Amount
1.
2.
3.
+.
5.

Equipment total:
Materials and Supplies {itemized)
1.
2.
3.
+.
5.
Naterials and supplies total:
Travel Costs {itemized)
1.
2.
3.
+.
5.
Travel total:
PersonnelJServices {itemized)
1.
2.
3.
+.
5.
Personnelfservices total:
Other Expenses {itemized)
1.
2.
3.
+.
5.
6.
7.
Other total:
Subtotal:
Facilities and Administrative Costs {S%):
Your Current Total:
Total must NOT EXCEED $5000.00 Total Request:
Name:
Student ID #:
Please hll out this form usning Adobe Acrobat
For all expenses listed under Equipment" indicate on page 2 of this PDF who will assume responsibility
for the equipment once the project has been completed.
All items listed under Other Expenses" must be accompanied by a justihcation statement on the 2 page of this PDF.
Proj ect Grant Budget Form
$970.00
$77.60
$1047.60
$1047.60
Isaiah Hoyer
081-34886
2 Carolina Cloacal Thermometers
2 Noose Pole (Cabela Crappie pole). 40.00
1 Pesola Light-Line Spring Scale
25.00
35.00
Thermocouplers 250.00
$350.00
2 Rite in the Rain eld note books 20.00
$20.00
Transport (Personal Auto from Almorgordo, NM to Moscow, ID)
Transport (ight from El Paso, NM to Spokane, WA)
250.00
350.00
$600.00
Just i f i cat i onlorallexpensesover$100andLhoselLemsllsLedunderCLherLxpenses",descrlbeLheexpensessuch
LhaLrevlewersandcommlLLeememberswlllbeableLolmmedlaLelyrecognlzeLherelevanceandnecesslLyofLhelLems.
lorallexpensesllsLedunder"LqulpmenL",lndlcaLewhowlllassumeresponslblllLyforLheequlpmenLonceLhepro[ecL
hasbeencompleLed.

Equi pment ( i t emi zed)

Number: Who will assume responsibility for the equipment once the project has been completed?


Justification for expense if over $100:

Number: Who will assume responsibility for the equipment once the project has been completed?


Justification for expense if over $100:


Number: Who will assume responsibility for the equipment once the project has been completed?

Justification for expense if over $100:

Number: Who will assume responsibility for the equipment once the project has been completed?

Justification for expense if over $100:

Number: Who will assume responsibility for the equipment once the project has been completed?

Justification for expense if over $100:





2
Dr. Luke Harmon and his lab members.
3
Dr. Luke Harmon and his lab members.
4 Dr. Luke Harmon and his lab members.
Thermocouplers include a digital thermometer and attachments that will allow me
measure lizards and model lizards (clay/copper) temperatures. Multiple attachments
will able efcient data collection by leaving the attachments with the model lizards.
1
Dr. Luke Harmon and his lab members.
Mat er i al s and Suppl i es ( i t emi zed) lorallexpensesover$100.
Number: Just ificat ion for expense:

Number: Just ificat ion for expense:

Number: Just ificat ion for expense:

Number: Just ificat ion for expense:

Number: Just ificat ion for expense:


Tr avel Cost s ( i t emi zed) lorallexpensesover$100.
Number: Just ificat ion for expense:

Number: Just ificat ion for expense:

Number: Just ificat ion for expense:

Number: Just ificat ion for expense:

Number: Just ificat ion for expense:

1
Part of Dr. Rosenblum's and Simone Des Roches Grant will cover travels costs
associated with driving. This amount will roughly cover my portion for Gas and
Hotels.
2
Part of Dr. Rosenblum's and Simone Des Roches Grant will cover travels costs
associated with ying. This is the amount necessary to purchae a one-way ticket
after the eld season is over.

To enrich education through diversity the University of Idaho is an equal opportunity/affirmative action employer


Apiil 12, 2u12

Beai SuP Committee,

I am wiiting with my full suppoit foi Isaiah Boyei's application foi the 0I Stuuent uiant
Piogiam. I am an Assistant Piofessoi in the Bepaitment of Biological Sciences at the
0niveisity of Iuaho anu Isaiah's co-supeivisoi (with Bi. Eiica Rosenblum at 0C Beikeley).

I have no uoubts at all about Isaiah's ability to caiiy out the pioposeu ieseaich. Be is
alieauy veiy involveu with the White Sanus ieseaich team, anu iegulaily attenus lab
meetings anu uiscussions. Bis pioposeu woik will fill a much-neeueu gap in the oveiall
ieseaich piogiam at White Sanus.

Bi. Rosenblum has obtaineu extia-muial funuing fiom NSF foi components of hei lab's
ieseaich at White Sanus. Bowevei, we uo not have auequate funus foi Isaiah's SuP
pioposal, which uesciibes woik outsiue of the scope of the ieseaich. All necessaiy
iegulatoiy committee appioval has been gianteu foi Isaiah's ieseaich (Animal Caie anu
0se Piotocol #2u1u-48). Please contact me if you have any questions.

Sinceiely,



Luke }. Baimon
Assistant Piofessoi
Biological Sciences
lukehuiuaho.euu
2u8-88S-uS46

Student Grant Proposal Committee
University of Idaho

Dear SGP Committee,
White Sands, New Mexico is a site of recent evolutionary divergence in three lizard
species. The emergence of the gypsum dunes was followed by rapid rates of colonization of
organisms from the surrounding dark soil habitat. The unique geology of the White Sands gives
scientists a remarkable chance to study evolutionary ecology. I am applying for the student grant
to fund my senior thesis research on the affect of thermal regulatory behavior on the divergence
of Holbrookia maculata (Lesser-earless lizard).
The selective forces acting upon cryptic coloration may affect the thermal regulatory
behavior on a poikilotherm. Thermoregulation is the ability for an organism to maintain its body
temperature with in certain limits. Poikilotherms allow their body temperature to fluctuate
relative to their environment, e.g. alligators, frogs and lizards.
This year I have worked closely with many doctoral students in the Harmon and
Rosenblum labs. I have been positively influenced by the doctorial students Simone Des Roches,
Tyler Hether and Kayla Hardwick, who have helped me develop my proposed research in White
Sands this summer. I hypothesize that there has been a directional change in phenotypic
traits in Holbrookia maculata in White Sands from their dark soils conspecifics that affects
their thermoregulatory behavior. Specifically, I predict: a) selection for lizards with substrate
matching has concordantly affected their thermoregulatory behavior, b) the overall body
temperature is lower in White Sand H. maculata due to divergent evolution, c) the white morph
H. maculata has evolved a larger dorsal surface area to maximize heat gain to compensate for
their white coloration compared to their dark soil conspecifics, and d) the white morph H.
maculata differs in habitat use and are able to travel further from cover because they are less
likely to reach their thermal maximum than their dark soil conspecifics. I will determine the
relationships among morphology, habitat use, and thermal behavior to elucidate differences in
evolutionary ecology between White Sands and dark soils lizards.
I hope to make my own contribution to ecological evolution on the lizards of White
Sands with my research. The Rosenblum lab has contributed in-depth research on the White
Sands system. Research focuses on the characteristics of recent ecological adaptation. My
research on how recent evolution influences thermoregulatory behavior will demonstrate other
interactions of features that are affected by natural selection.
Requested funding will help cover costs of travel and equipment. At this time, existing
funding does not completely cover my thermoregulatory study. Necessary equipment for my
research includes cloacal and general use thermometers, replacement noose poles for capturing
the lizards, and a pesola spring scale for weighing the lizards. Further funding is necessary to
efficiently allocate my effort on data collection.
I have already gained invaluable experience in researching my subject and writing this
grant proposal. The funding received will blanket my concerns of inadequate data collection
potential and fuel my experience gained working in the field, analysis and the underlying goal of
a published paper. This senior project is the glaze over my undergraduate degree and will shine
bright promises for a prospering future towards the work force and graduate school.

Sincerely,
Isaiah Hoyer

Student Grant roposa| 2012
1he LffecL of AdapLlve LvoluLlon and Lcology on 1hermoregulaLory
8ehavlor ln nolbtooklo mocoloto (Lesser Larless Llzard)
lsalah Poyer
081-34886
hoye6378[vandals.uldaho.edu
208-213-8841
Adaptive Evolutionary Ecology Isaiah Hoyer

1
Narrative
Objective
White Sands National Monument, New Mexico, is a site of recent evolutionary
divergence in three lizard species. The unique geology of the White Sands gives scientists a
remarkable chance to study evolutionary ecology. Over thousands of years, the inland of Lake
Otero receded to a temporal shallow basin that is now called Lake Lucero, reveling
approximately 275 square miles of white gypsum. The dunes are a dynamic and growing habitat
that is largely shaped by the consistent eastwardly blowing winds (Kocurek 2007). The gypsum
dune field is the largest of its kind on Earth! In the last 6000 years, several of the plants and
animals in the neighboring dark soiled habitat have had the opportunity to colonize White Sands
and adapt to its dynamic environment.
There is potential for adaptation by organisms inhabiting any newly formed, recently
devastated or isolated habitats. The exploitation of White Sands by the lizard species is shaped
by evolutionary driving forces. In comparison to the surrounding dark soils, White sands has
reduced species richness and species abundance for potential competitors and predators (Des
Roches et al. 2010). The available habitat of the newly formed gypsum dunes resulted in feats of
adaptation in both plants and animals. The plants that persist in White Sands must be hardy
enough to withstand being buried and survive in the prevailing winds that shape the dynamic
habitat. Much of the wildlife in the White Sands environment has evolved cryptic coloration to
match their surroundings (Harmon and Rosenblum 2010). It is likely that preferential predation
on mismatched lizards has selected for this cryptic coloration (Rosenblum 2005).
The selective forces acting upon cryptic coloration may affect the thermal regulatory
behavior of ectotherms, organisms that allow their body temperature to fluctuate relative to their
environment, e.g. alligators, frogs and lizards (Tucker et al. 1963). Thermoregulation is the
ability of an organism to maintain its body temperature within certain limits. Background color
matching and color change affect rates of bodily warming in lizards (Norris 1967). A lizard will
regulate its body temperature behaviorally (Stevenson 1979). Many lizards can increase or
decrease heat absorption by changing their color (Cole 1943); however, color also has a genetic
basis. Most lizards, including individuals that persist in White Sands, are unable to change their
color entirely to match their neighboring habitat (i.e., the white lizards cannot be as darkly
colored as their dark conspecifics and the dark lizards cannot be as white as their white
conspecific; Rosenblum 2005). Norris observed that in early mornings some lizards were darker
in color, presumptively facilitating warming, than throughout the day when they were more
faithfully matching the substrate.
Three species of lizards have a white and dark morphs associated to the color of their
habitat; Holbrookia maculata, Aspidoscelis inornata, and Sceloporus undulata (Harmon and
Rosenblum 2010). The lesser earless lizard, H. maculata, is a species of interest for the
thermoregulatory study because it is more conspicuous than the other lizards, and it shows the
most dramatic change in color (Harmon and Rosenblum 2010). There is little to no gene flow
between the populations in White Sands and dark soils habitats (Rosenblum et al. 2007). The
isolation of H. maculata gives the greatest expected difference in thermoregulatory behavior.
The other lizard species show more interaction among color morphs due to overlapping ranges.
Also, H. maculata evolved its skin color differently than A. inornata and S. undulatus. The
mechanisms for the blanched phenotype in H. maculata are unknown although it is speculated
that a mutation in the Melancortin-1 receptor gene (Mc1r) is responsible as documented in A.
inornata and S. undulates (Rosenblum 2005).
Adaptive Evolutionary Ecology Isaiah Hoyer

2
There are many potential differences among the color morphs of H. maculata that I might
find. Due to the reduced efficiency of solar radiated heat absorbance, the white morph may have
evolved a larger dorsal surface area to maximize its heat gain and compensate for its color loss.
Lizards receive their energy in the form of direct sunlight, reflected sunlight, erratically scattered
skylight, thermal radiation from the atmosphere, conduction from the ground, and surrounding
vegetation. The amount of energy absorbed depends on the lizards surface area exposed, the
angles of the surfaces, and the absorptivity of the surface to the quality of incident radiation
(Bartlette and Gates 1966). Interestingly the gypsum that makes up White Sands has less
conductive properties (retains less heat) than the darker surroundings soils (Scheidt et al. 2010).
Therefore, it is possible that the White Sand morphs could have evolved an overall lower body
temperature.
Another interesting difference between the color morphs could be distance from cover. It
is possible that the white color of a lizard will allow it to spend more time in the open due to its
reflectivity and inferior sun absorption capability. A white object is less efficient at sun
absorption than a darker object. As a result from the evolved color change, the white morph H.
maculata is potentially able to spend more time foraging and exhibit increased activity further
from cover. It takes the white morph H. maculata longer to reach their thermal maximum than
dark morph H. maculata.
Examples of lizard behaviors that affect body temperature are perch choice, time in
shade, and the time in sun. Morphological examples in lizards that affect body temperature are
dorsal skin brightness, proximity of blood vessels to the surface of the skin, body size and body
surface area. With my research, I will examine how previous evolutionary adaptation of
Holbrookia maculata to White Sands has affected thermoregulatory behavior. I hypothesize
that there has been a directional change in phenotypic traits in Holbrookia maculata in
White Sands from their dark soils conspecifics that affects their thermoregulatory
behavior. Specifically, I predict: a) selection for lizards with substrate matching has
concordantly affected their thermoregulatory behavior, b) the overall body temperature is lower
in White Sand H. maculata due to divergent evolution, c) the white morph H. maculata has
evolved a larger dorsal surface area to maximize heat gain to compensate for their white
coloration compared to their dark soil conspecifics, and d) the white morph H. maculata differs
in habitat use and are able to travel further from cover because they are less likely to reach their
thermal maximum than their dark soil conspecifics. I will determine the relationships among
morphology, habitat use, and thermal behavior to elucidate differences in evolutionary ecology
between White Sands and dark soils lizards.
Predictions b and c are both viable because dorsal surface area is very plastic. Lizards are
able to contort their body to increase surface area and increase heat absorption. When a lizard is
looking to shed heat they will minimize their surface area in addition to seeking cooler
microhabitat. Therefore White Sands H. maculata is able to have an overall lower body
temperature and a larger dorsal surface area than dark soils H. maculata.
For two months during the summer 2012, I will travel to White Sands, New Mexico, to
study how past adaptation and current ecological surroundings affect thermoregulatory behavior
of the two color morphs of Holbrookia maculata. Furthermore, I will compare habitat use (perch
selection, temperature selection) by the two color morphs in their separate habitats to determine
divergence in thermoregulatory behavior. The research will contribute to a larger pool of
evolutionary ecology work of the White Sand lizards. Dr. Erica Bree Rosenblums lab has
performed research on colonization, genetics of adaptation, and ecologically relevant
Adaptive Evolutionary Ecology Isaiah Hoyer

3
morphological traits, sexual communication and functional morphology. Ongoing research
includes several experiments and studies exploring ongoing natural selection on color in White
Sands. My own research will help determine whether this selection on color change also
influences thermoregulatory behavior.

Importance
Investigator
As an undergraduate, I will learn firsthand what is involved in ecological field research. I
will gain experience in experimental design, collecting and analyzing data, as well as working
cooperatively towards a common goal with other scientists. During my time in White Sands, I
will be involved with an outreach project with local students. In past field seasons, the lab team
designated a full day field course open to all local residents and students in the area. I will gain
qualification in explaining my research and involvement in White Sands to many different age
groups and backgrounds. Working with two other undergraduates and two graduate doctorial
students will prepare me for graduate school, pursuing a career as a field technician and/or
researching positions. The interactions with the community will give me experience in
understanding the publics desires and curiosities.
The research I will conduct this summer and the analysis afterwards will fulfill my senior
thesis requirement for my major in Ecology and Conservation Biology. The knowledge I have
gained over my undergraduate career will be greatly exercised during this project.

Ecology and Biology
White Sands, New Mexico gives visitors and students a great opportunity to experience
the recent evolutionary ecology of its inhabitants. As a federally protected national monument it
is protected from human disturbance. The interactions among the wildlife are greatly apparent.
As in many desert environments there is low relative diversity in plants and animals when
compared the tropic regions of the world. The ecotone, transition areas from White Sands to dark
soils, is a very busy place for natural selection to act on the lizards. There is high selection of
mismatch prey from avian predators (Rosenblum 2005).
Studying the thermal ecology of the H. maculata in White Sands will add knowledge of
what traits have been affected by their evolution. The unique habitat of White Sands can clearly
show what major forces act upon adaptive evolution. The White Sand system is rare window in
time of a stage in speciation. Understanding how selection leads to evolution in new
environments is key to understanding biological diversity.
Conservation implications can be addressed from better understanding of how traits are
affected by colonization. Studying the influences that the environment poses on a species
provides more accurate predictions for the future health of colonization populations.
Understanding the constraints and benefits for species divergence will support conservation
efforts of endangered/threatened species of similar habitats.
White Sands is one of the most recent sights to see vertebrate adaptive evolution in North
America. White Sands gives Biologists a rare opportunity to study real time evolution on traits of
known functional significance in the field.




Adaptive Evolutionary Ecology Isaiah Hoyer

4
University of Idaho
Dr. Rosenblum and Dr. Harmon have had many publications and study opportunities that
have risen from the evolutionary work in the White Sands. Many papers have been published
since, as well as funding. The outreach to the worlds biggest gypsum dune holds fast to
University of Idahos spirit in leadership and discovery. Research in White Sands, New Mexico,
adds to the diversity and elasticity of this great University. Supporting my research sheds light on
the fact that Idaho has interest outside the Northwest.

Methods and Analysis
I will work with two undergraduates and three doctoral students in White Sands, New
Mexico, for approximately two one-month intervals. The necessary Institutional Animal Care
and Use Committee (IACUC) protocol forms have been submitted and approved for University
of Idaho and University of California Berkeley. I will capture lizards by hand and measure traits
that will allow me to document variation in the wild: body size, weight, shape, body temperature,
environmental temperature (surface and air), dorsal surface area, and brightness, hue and chroma
of the dorsal color. I will record the following lizards features: sex (male or female), age
(juvenile or adult), and whether or not the female is gravid (pregnant). Gravid females lizards
have been known to have overall lower body temperatures than males or non-gravid females
(Andrews 2005). I will also measure distance from cover, percent vegetation, and perch choice. I
will measure distance from cover to the nearest ten centimeters. Percent vegetation will be
calculated using a visual index of vegetation in the conical area forming above the individual to
the nearest five percent. Perch choice will be identified as ground (not associated with
vegetation), tree/shrub. Perch height and diameter will be estimated by eye to nearest centimeter.
Thermoregulation has been correlated to
fitness through costs and benefits associated to
environmental factors (Slatkin 1976). Lizards
become thermoconformers when costs of
thermoregulation outweigh the potential benefits.
There is an opportunity cost to a lizard spending time
thermoregulating. A lizard that spends less time
thermoregulating is able to engage in other important
activities such as foraging or mate searching (Nadeau
2005). The thermal data and morphological
measurements combined with a regression analysis
will allow me to statistically test for differences
among the individuals and color morphs.
I will quantify multiple characters
simultaneously including brightness and body
temperature that directly affect thermoregulation. I
will test the effect of species age class (juveniles or
adults), sex (male or female) and morphology
(snout-vent, forelimb, hind limb, tail length, body
temperature, and brightness etc.) on distance from
cover, perch height/diameter, and percent
vegetation using ANOVA or ANCOVA (R
Development Core Team 2012) with an alpha
Figure 1. Example of measuring
morphological traits from a lizard scan
(Holbrookia maculata). (SVC=Snout-vent-
length, T=tail-length, H=hind-limp, F=fore-
limb).
Adaptive Evolutionary Ecology Isaiah Hoyer

5
value of P=0.05. At each field site area I will capture approximately ten lizards. The H. maculata
home range size has not been well evaluated in White Sands, nevertheless this species generally
has a small home range (Hulse 1985). H. maculata is a conspicuous species and individuals will
usually bask on the same perch throughout a field season (Des Roches, personal observation).
Based on these observations, I plan to conduct sampling within in several small (1 km
2
)
interdunes (between dune areas that contain most of the vegetation and animal life). I hope to
find at least four interdune areas (two dark soils and two white sand soils) that harbor H.
maculata. We already have some evidence of selection for brightness, hue and chroma (Des
Roches et al. In prep), and my study will test whether thermoregulation has also been affected.
I will mark each individual with a sharpie streak on its tail, which will avoid any
duplicate data. The sharpie will not harm the lizard and will wash off in a few days. My total
target is to mark 40 individuals in four interdunes (about 10 lizards per interdune). In about five
days, I will sample, mark, measure and return all the individuals in an interdune. Therefore, it
will take me about 25 days to extensively sample 40 individuals at each capture event. The
lizards will be captured by noose and by hand. Upon capture I will first measure the lizards body
temperature with a Carolina cloacal thermometer and then subsequently measure the air,
substrate or perch temperature with a general purpose thermometer. The lizards will then be
transported to the field lab where the following measurements will take place: snout-vent-length,
total mass, head shape, fore and hind limb length including toe length, inter-limb length, pelvic
width, and tail length.
To ensure correct measurements of the lizards they will each be scanned dorsally and
ventrally on a standard photocopier (Figure 1). The dorsal scans will allow me to calculate
relative surface area, the area that absorbs the majority of solar radiation. Also, the scans will
allow me to re-measure if necessary. Also, at our field lab, the brightness, hue and chroma will
be measured using a spectrometer. At my leisure, after recording the trait measurements and
photographing individuals, they will be returned to their place of origin the following day.

Discussion
The White Sands system has much to offer to evolutionary biologists. The unique
adjacent environments of White Sands and the dark soils give implications on divergence of the
lizards. Holbrookia maculata color morphs are relatively isolated from each other suggesting the
most extreme differences in morphology compared to A. inornata and S. undulates which exhibit
overlapping home ranges. Climate and phenotype affect associative costs of maintenance and
activity i.e. foraging, time basking, growth, reproduction and space utilization (Tracy et al.
1983). The thermal environment has potential to influence the physiology and behavior of lizards
(Hager 2000). The evolution of H. maculata has many factors that could affect their
thermoregulatory behavior.
In all organisms there are many environmental factors that affect body temperature.
Poikilotherms generally depend on absorption of solar radiation for their heat gain, and
conduction from warmed surroundings. Interestingly the gypsum that makes up White Sands has
less conductive properties (retains less heat) than the darker surroundings soils (Scheidt et al.
2010). Therefore, it is possible that the White Sand morphs could have evolved an overall lower
body temperature. Another interesting difference between the color morphs could be distance
from cover. It is possible that the white color of a lizard will allow it to spend more time in the
open due to its reflectivity and inferior sun absorption capability. A white object is less efficient
at sun absorption than a darker object. As a result from the evolved color change, the white
Adaptive Evolutionary Ecology Isaiah Hoyer

6
morph H. maculata is potentially able to spend more time foraging and increased activity further
from cover. It takes the white morph H. maculata longer to reach their thermal maximum than
dark morph H. maculata.
There are many potential differences in the color morphs of H. maculata that I might
find. Due to the reduced efficiency of solar radiated heat absorbance, the white morph may have
evolved a larger dorsal surface area to maximize its heat gain and compensate for its color loss.
The divergence of H. maculata has led to many changes in traits and adaptations (Des Roches et
al. 2010).
My research will contribute to past studies on thermoregulation, physiology, and
evolution. If thermal differences in the conspecific lizards are significant, than morphological
studies on their absorption of heat would prove interesting. Do they absorb solar radiation or
conduction heat differently? Are there differences in the proximity of blood vessels to the surface
of their skin? Are White Sand H. maculata more efficient at oxygen consumption than the dark
morphs i.e. estimation by heart rate during heating (Tucker 1964). A more thorough study on
thermal preference could determine if the lizards vary in their thermal maximum and thermal
minimum. We could test whether a white lizard prefers a lower temperature on dark soils and
visa versa for the dark lizard on white sands. Dr. Rosenblum also suggested that we test whether
a lizard is able to distinguish color. Do the lizards know when they are matching or mismatching
their surroundings? Has the White Sand H. maculata evolved a larger in body size to ensure less
fluctuation in their body temperature (Stevenson 1985), allowing them to increase their activity
further from cover?
As in many research projects, the questions and correlations are endless. The recent
colonization of the White Sands gives biologist a rare opportunity to study the effects of
evolution. The University of Idaho, the field of evolutionary ecology and biology, and me will
benefit from the research conducted in White Sands, New Mexico. After all Theodosius
Dobzhansky says it best, nothing in biology makes sense except in the light of evolution.

Timeline
May 2012: Meet with other student field assistants; learn on field protocols.
May 15 2012: Depart for NM by car.
May-June 2012: Collect, mark and measure 40 Holbrookia maculata (20 in the White Sands and
20 in adjacent dark soils habitat).
June 20 2012: Depart from NM by plane.
August 1 2012: Depart for NM by plane.
August 2012: Collect more Holbrookia maculata in the dark soils and White Sands.
August 20 2012: Depart from NM by car.
September 2012-April 2013: Analyze behavior and morphology over one field season.
April 2013: Present results of ECB Senior Thesis to the College of Natural Resources







Adaptive Evolutionary Ecology Isaiah Hoyer

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Bibliography
Andrews, Robin M. "Geographic Variation in Field Body
Temperature of Sceloporus Lizards." Thermal Biology 23.6 (1998): 329. Print.
Barlett, Peter N., and David M. Gates. "The Energy Budget of a Lizard on a Tree Trunk."
Ecology 48.2 (1967): 315-22. Print.
Cole, LaMont C. "Experiments on Toleration of High Temperature in Lizards with Reference to
Adaptive Coloration." Ecology 24.1 (1943): 94-108. Print.
Des Roches, S., Robertson, J. M., Harmon, L. J. & Rosenblum, E. B. 2010 Evidence for
ecological release in White Sands lizards. For Proc. Roy. Soc. B.
Hager, Stephen B. "Variation in Body Temperature and Thermoregulatory Behavior between
Two Populations of the Lesser Earless Lizard, Holbrookia Maculata." Contemporary
Herpetology (2000). Print.
Hager, S. B. 2001 "Microhabitat Use and Activity Patterns of Holbrookia Maculata and
Sceloporus
Undulatus at White Sands National Monument, New Mexico." Journal of Herpetology
35.2 (2001): 326-30. Print.
Hulse, Arthur C. "Home Range Size in Holbrookia Maculata (Iguanidae) from Southeastern
Arizona." The Southwestern Naturalist 30.4 (1985): 608-10. Print.
Kocurek, G., M. Carr, R. Ewing, K. G. Havholm, Y. C. Nagar, and A. K. Singhvi. "White Sands
Dune Field, New Mexico: Age, Dune Dynamics and Recent Accumulations."
Sedimentary Geology 197 (2007): 313-31. Print.
Nadeau, Gabriel Blouin-Demers And Patrick. "The Cost-Benefit Model of Thermoregulation
Does Not Predict Lizard Thermoregulatory Behavior." Ecology 86.3 (2005): 560-66.
Print.
Norris, K. S. "Color Adaptation in Desert Reptiles and Its Thermal Relationships." Ecology
(1967): 162-229. Print.
Rosenblum, Erica Bree. "The Role of Phenotypic Plasticity in Color Variation
of Tularosa Basin Lizards." Copeia 3 (2005): 586-96. Print.
Rosenblum, E. B., Hickerson, M. J. & Moritz, C. 2007 A multilocus perspective on colonization
accompanied by selection and gene flow. Evolution 61, 2971-2985.
Rosenblum, Erica Bree and Harmon, Luke J. "Same Same but Different: Replicated Ecological
Speciation at White Sands." Evolution (2010). Print.
Slatkin, Raymond B. Huey and Montgomery. "Cost and Benefit of Lizard Thermoregulation."
The Quarterly Review of Biology 51.3 (1976): 363-84. Print.
Stephen Scheidt, Michael Ramsey, and Nicholas Lancaster. "Determining Soil Moisture and
Sediment Availability
at White Sands Dune Field, New Mexico, from Apparent Thermal
Inertia Data." Journal of Geophysical Research 115.F02019 (2010): 1-23. Print.
Stevenson, Raymond B. Huey and R. D. "Integrating Thermal Physiology and Ecology of
Ectotherms: A Discussion of Approaches." American Zoologist 19.1 (1979): 357-66.
Print.
Stevenson, R. D. "Body Size and Limits to the Daily Range of Body Temperature in Terrestrial
Ectotherms." The American Naturalist 125.1 (1985): 102-17. Print.
Tracy, Warren P. Porter and C. Richard. "Biophysical Analyses of Energetic, Time-Space
Utilization, and Distributional Limits." Lizard Ecology. Ed. Raymond B. Huey, Eric R.
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Pianka, and Thomas W. Shoener. Cambridge and London: Harvard University Press,
1983. 55-83. Print.
Tucker, George A. Bartholomew and Vance A. "Control of Changes in Body Temperature,
Metabolism, and Circulation by the Agamid Lizard, Amphibolurus Barbatus."
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Tucker, George A. Bartholomew and Vance A. "Size, Body Temperature, Thermal Conductance,
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Additional Information
Institutional resources available-
Dr. Rosenblum and Dr. Harmon labs have accumulated equipment from grants over the
past two field seasons. Equipment they have accumulated includes noose poles, spectrometer,
and general-use thermometers. Many noose poles are damaged or of little effective use. New
noose poles are necessary to ensure safe capture of the lizards. The specific cloacal
thermometers, spring scale and other equipment is required for accurate fulfillment of my
research. Using damaged or general equipment will lessen the comparability and significance of
my data.

Other Sources of funding-
In addition to the SGP I have applied for the Biological Science Research Grant and
Berklund Undergraduate Research Grant. The Biological and Berklund Research Grants are
intended to pay my salary for my data collection over the summer. My data collection will take
up the majority of my summer and work opportunities. I will need a salary to pay for my cost of
living for the next school year.

Plan to disseminate to the university-
This research fulfills the requirements for my senior thesis project under the Ecology:
Conservation Biology degree program. Not only will I present my proposal to board of
professors and peer, I will also present my finding in Spring 2013 along with a final paper. My
results will be accessible upon submission and my final poster will be on display either at the
Biological science building or at the college of Natural Resources.

Regulatory committee approvals-
As a field assistant to the doctorial students (Simone des Roches, Kayla Hardwick and
Tyler Hether) I have already completed the necessary liable Institutional Animal Care and Use
paper work (IACUC). Therefore I am covered under their research and able to appropriately
handle the animals.

Other-
As a senior in Ecology: Conservation Biology I am excited to put my knowledge that I
have accumulated a the University of Idaho to use. I have field experience in Dalby, Australia
where I conducted a intense four day ecological assessment on a long-term ecological field site. I
shadowed a masters student at small mammalian facility at Washington State University. During
the my six months shadow program I accumulated a journal of her various protocols and process
of conducting research and experience in animal care. Beginning fall 2011 to present I have been
working in Dr. Rosenblums lab with Tom Poorten a doctoral student studying the chytrid fungus
batrachochytrium dendrobatidis. I have also taken one upper division statistic course that will
benefit me during the analysis portion of my project. This spring I have been working in a
advance evolutionary ecology research team taught by Simone Des Roches. The team are post-
White Sand field assistants. The team and I are compiling last years data and researching other
papers to compile a hopeful published paper.