Guy Sion PhD proposal 18 May 2011

The effects of body state on decision making in the gecko

Ptyodactylus guttatus

Proposal outline
1. Introduction
1.1. Background
1.1.1 Morphological characters
1.1.2. Energy reserves and decision making
1.1.3. Behavior
1.2. The goal
1.3. Hypotheses & ensuing predictions
2. Methods
2.1. Study organism
2.1.2. Study Area
2.1.3 Abiotic parameters
2.1.4. Sede Boqer study site
2.2. Marking
2.3. Body Measurements
2.4. Behavioral parameters
2.5. Body state
2.6. The experimental design
2.7. Predictions in tables
3. Timetable
4. Discussion
References
 1. Introduction:

Behavioral ecology studies the ecological and evolutionary basis for animal behavior.

The role of behavior is in enabling an animal to adapt to its environment. To achieve this goal to adapt,

the individual may use behavioral functions, such as reptiles basking or hiding under a rock to regulate

temperature behaviorally instead of physiologically (Huey et al., 1977). However, behavior may

change the course of evolution of an animal. A change in the environment may induce a response in the

behavior, that may in turn, impose a genetic change over time due to a new selective pressure. (Avital

and Jablonka, 2000). The behavior of an animal is a very important component in the understanding of

the evolutionary and ecological context. However, studying animal behavior takes time and allegedly

cannot be recorded from dead animals e.g. in museum collections. I claim I can learn on population

behavior from carcasses, by morphometric characters e.g. directional asymmetry (DA) can inform me

on the general risk taking and social status, of the population, and so is tail injury for social status. I

suggest the museum as an important resource to learn on the behavior of past and present populations.

In this study, I wish to predict an animal behavior, using the morphological component among the

factors that may affect the decision making of foraging strategy and social hierarchy. The behavior I

want to study is decision making and social status. The decision I will study is foraging strategy: how

far from cover will the lizard ambush for prey. The social status will be studied by proximity to a

covered perch, night lamps. I am interested especially in the effects of body state in terms of fat store

and body asymmetry on the decision making. Therefore, lizards that can shed their tail are ideal subject

for such experiments. The tail contains valuable fat deposition. Losing a tail is losing a fat storage. The

lizard I chose is Ptyodactylus guttatus, which was studied for DA (Werner et al., 1991). I suspect that

DA interact with body state, in that DA may reflect developmental instability and thus, affect behavior

to adapt to its environment; alternatively, that the DA may reflect an evolutionarily functional

importance or new adapted function. Thus, left biased DA individuals may adopt different foraging
strategy from right biased DA individuals in order to adapt to the environment. The effect may be on

the decision or on the procedure of reaching it, or both. The parameters of body state I will study, relate

to fat store such as tail base width, and DA and its possible link to body fat percentage. Although these

connections are not known, one may argue it implies from the connection between DA and stress

(Kark, 2001) Thus, these morphometric characters may affect the individual decision making and risk

management.

1.1.1. Morphological characters:

1. Sex: The sex has a tremendous effect, on the use of energy and fat store. Energy loss of fat stored in

the tail has a different effect on females and males. Females reduce their clutch, while only males'

speed is reduced e.g. in Eulamprus tympanum a skink (Doughty and Shine, 1998). Females regenerate

their tails significantly faster than males (e.g., in Skink, Chapple and Swain, 2002). Females and males

may also differ in prey preferences (Castilla et al., 2008). Therefore, I expect possible different

behavior interests in the sexes, such as different prey fat content that yield a different fat percentage

level and different risk taking strategy. Thus, I treat the sexes separately to be compared in the end.

Doing that, in terms of morphology traits is not a precedent (Ligon, 1968; Perry, 1996; Lachman et al.,

2006; Werner et al., 2006).

2. Asymmetry:

A study on three species of Ptyodactylus geckos showed significant minor directional

asymmetry (DA). The study raises the question of the possible evolutionary role of minor DA and

suggests it as a potential base for evolutionarily functional DA. Moreover, DA side-dominance may

differ between related taxa and possibly between sexes (Werner et al., 1991).

Directional asymmetry, defined as a greater development of a character on one side of the plane

or planes of symmetry than on the other. The ecological meaning of minor directional asymmetry is an

open question. Unlike fluctuating asymmetry (FA), that results from the inability of organisms to
develop in precisely determined paths, thus, indicates developmental instability (Van Valen, 1962), DA

is considered by some as a potentially functional basis for evolutionary processes. Recent studies link

DA with fitness with two different approaches: One approach is to see DA as another phenotypic

manifestation of developmental instability (Kark, 2001; Velickovic and Jakovsev-Todorovic, 2006).

Another approach is to search for functional importance of DA in the frame of evolution (Seligmann,

1998). Here I test possible relationships between DA and behavior, such as dominance and risk-taking

foraging strategy, using the gecko’s eye-size asymmetry as indicator of its social status and/or decision

making. The basis for using the animal eye size, derived from the brain-eye link, as optic nerve number

two, is actually an extension of the brain. Thus, possible link between risk taking and eye size could be

point to the brain directional asymmetry as it is explained in human morphometry difference of digit 2

and 4 from directional asymmetry of right and left hemisphere that induce aggression hormones and

causing digit DA (Coyne et al., 2007). Previous morphometric studies (Werner et al., 1991; Seligmann,

1998; Seligmann, 2003; Almog et al., 2005; Lachman et al., 2006) have used museum specimens, to

link assorted characters and taxa of Lepidosauria to tail injury. One important result was the useful use

of "Seligmann effect" to differentiate different populations to different species or sub-species by

excluding tail-injured individuals (Seligmann, 1998; Almog et al., 2005; Lachman et al., 2006).

This study observed live individuals in their natural habitat, to test the hypothesis that DA

correlates with risk-taking behavior. This connection of morphometric measurements of live

individuals with the behavioral field observations can give the lab measurements of dead animals, new

context and may demonstrate that museum specimens can inform about behavior. The prediction is

based on Seligmann (1998) finding that left-biased individuals have more tail injuries. A possible

explanation I raise is that left-biased individuals will tend toward riskier foraging strategies. According

to Seligmann (1998), minor directional asymmetry may occur, if there is small systematic difference,

tendency toward right or left, such as in the "Seligmann effect", with a possible evolutionary functional
role. The "Seligmann effect", is an inherent different meristic biometry (inborn) difference between

those with tails and those with damaged tails. The effect predicts better statistical distinction for

taxonomic comparison of close taxa, (only the full-tail specimens are used, despite the smaller sample),

this due to the smaller variation, when you 'clean' the statistical noise of the tail injured individuals.

Obviously the morphologically extreme, deviating, specimens are expected to be accident prone.

(Seligmann, 1998; Almog, et al., 2005; Lachman et al., 2006). Hence, if one compares behavior of tail

retaining and tail losing of lizards, possibly some behavior difference is inborn, not due to the tail.

However, this next step of observing lizards in the wild and compare the behavior and risk-taking

strategy of left and right biased DA individuals, was never done before. Moreover, it also may function

as an external honest signal that may be disclosed to potential females or rival males and predators.

Studies on the effects of asymmetry on mate preferences shows preferences toward Symmetry

over FA (Fluctuating Asymmetry) in lizards (Lopez et al., 2002), and other taxa such as birds (Møller,

1992) or humans (Møller et al., 1995; Simmons et al., 2004). Studies on the effects of stress on

asymmetry (Leamy et al., 1999), shows that stress or exposure to toxic substances can cause FA and

DA.

Although DA was not considered as reliable indicator of developmental instability like the FA,

increasing evidence suggests that DA in addition to FA, may often reflect developmental instability of

bilateral traits, expressing stress (Graham et al., 1993; 1998; Kark et al., 2001).

3. Tail condition:

Since a substantial amount of fat is stored in the tail, it is an important energy storage organ in some

species of lizards (Bustard, 1967; Daniels, 1984). Thus, the state of the tail: Amputated tail, tailless or a

full intact tail may affect and disclose the body state of the individual, as the tailless individuals may

reduce energy stores and delay the animal’s next reproductive episode or reduce size and number of
young (Smyth, 1974; Dial and Fitzpatrick, 1981; Wilson and Booth, 1998); Possibly, that a regenerated

tail may signal a lipid loss, regardless of the true lipid content. The information on tail condition is

disclosed by coloration: Intact tail has dark rings along the tail and a black tip. Regenerated tail is

unicolor.

4. Tail Base Width (TBW), may be used as an indicator of fat store and hence of body condition

(Bauwens, 1985; Donoghue, 1998; Doughty and Shine, 2003; Vervuast et al., 2008). Moreover, using

the tail-base width as the independent variable in a linear regression model may yield an accurate

prediction of tail fat content mass (g). This has newly been demonstrated by Doughty and Shine, (2003)

in lizards lacking abdominal fat bodies, possessing caudal fat only (Doughty and Shine, 2003). If tail

fat is proportional to body fat, thus, body fat may be as well be predicted by TBW, possibly in lizards

with abdominal fat as well. Predicting total body fat percentage may be obtained by the use of TBW

only, or with other morphometric characters such as mass and RA. A statistical model may be

constructed, to predict body fat percentage, by comparing results of body fat percentage, with

morphometric characters that may indicate fat percentage such as tail base width.

1.1.2. Energy reserves and decision making:

Energy requirement has great effect on risk taking. In dire straight even safety gets lower value.

Thus, decision-making procedure supposed to be effected by energy requirement in the form of body

state (Clark, 1994; Kotler et al., 2004; Simmonds et al., 2010). I predict that status and risk interact and

maybe rely on the body state. Body Index is an index for body state. Dual energy X-ray absorptiometry

(DEXA) enables to measure directly the body state, by measuring the body fat percentage. Instead of

evaluate it indirectly by a body index. Another advantage is in multiple measurements in-situ, during

the life time of the animal (Secor and Nagy, 2003). The PIXImus is such a machine that uses DEXA to

measure bone densitometry and body composition giving values for fat mass (g) lean mass (g) and fat

percentage. The process requires careful sedation of the animal. I will use Isufluran for the sedation.

The sedation might endanger the animal, and it is possible that some individuals will not survive the
sedation. The use of Isofluran diminishes these dangers. It is relatively expensive, but safer to use. In

comparison to ether e.g., that may cause pulmonary edema. An example to a body condition index is

the tail base width (TBW). It may serve as a predictor of fat store in the tail. In combine with other

morphometric characters, such as mass/rastrum-anus (RA) (Werenr, 1971) with compare to DEXA

measurement results for body fat, a statistical model may be constructed for predicting body fat

percentage.

1.1.2.1. DEXA storage: The piximus store the DEXA data in the computer and allows

manipulating the scanned areas, so it is possible to compare the fat content of a specific zone

such as the tail, even if we scanned the whole animal. This way allows comparing the fat

content stored in a renewed tail and an intact tail.

1.1.3. Behavior:

Measurable expressions of risk, asset protection and energy reserves:

Social status by proximity to night lamps: The night lamps are preferable locations, occupied by

territorial males and their females who are in better body condition relative to non-territorial males

(Werner, 1972; Johnston and Bouskila, 2007).

Risk by foraging distance away from cover: Perch closer to the ground, riskier but more rewarding.

Perch on the high third of wall length will be considered as a high perch, as oppose to low perch or

ground perch

1.2. The goal:

To study what affects behavior such as decision making in general, and risk management in

particular, the possible interact with social status and to study possible relation of these behavior

with each other and possible relation of behavior to morphological traits. For that I use body state

and morphological components, as suspects that may affect decisions. A preferable result could lead

to recovering behavioral information from morphometric data from carcasses in museum

 collections.

To reach this goal, I intend to test the hypothesis that DA correlates with risk-taking behavior, as

predicted by Seligmann (1998). DA may function as an honest signal that may be perceived by females

(potential mates) or rival males and predators. Thus, this research should also explore the evolutionary

role of minor directional asymmetry, and mechanism that explain the relationships between DA, tail

injury, social hierarchy status and foraging strategy. To test the hypothesis that fat deposit in the tail is

proportional to whole body fat, experiments will use X-Ray to measure fat content, so body fat could

be predicted easily from morphometric data such as the tail base width, if hypothesis will be supported.

1.3. Hypotheses & ensuing predictions:

Predictions and experimental design:

1.3.1. Predictions based on the hypothesis that:

Morphometric characters (DA, Sex, tail condition, and fat%) affect decision making and risk

management:

(1). Behavior will be affected by DA. Left-biased individuals will tend toward riskier foraging

strategies.

(2). DA will be correlated with tail injury (Seligmann, 1998).

(3). Individuals with a deficit in their energy balance will be out active both in day and night to

obtain needed resources.

1.3.2. Predictions based on the hypothesis that:

Directional asymmetry (DA) is correlated with dominance:

Individuals with a right biased DA will:

1. Be in better body condition (Higher body fat percentage).

2. Perch higher.

3. Tend to win over left biased individuals at male-male conflicts on territory.

4. Tend to win in competition for females.

5. Left biased individuals are preferred by predators, when given a choice.

6. Tend to be territorial, in proximity to night lamps.

1.3.3. Predictions based on the hypothesis that:

Tail injury (regenerated or amputated) bears a vulnerability honest signal:

Males with a regenerated tail will:

1. Tend to win at conflicts in captivity if tail is colored with rings to mimic intact tail. If tail is not

colored, tend to lose.

2. Males with regenerated tail will be preferred by females if tail is colored with rings to mimic

intact tail, against individuals whose intact tail is colored to mimic a regenerated tail. If tail is not

colored, they will tend to lose.

3. Preferred by predators, when given a choice.

4. Tend not to be present at lighted ambush spots, (under lamp).

5. If active at moonlight night, will not be exposed to moonlight, only shaded ambush spots.

Daylight activity predicted to be mostly of tailless individuals.

1.3.4. Predictions based on the hypothesis that:

Fat deposited in the base of the tail is proportional to the total body fat content

Predictions:

1. Tail base width may serve as predictor of body fat percentage.

2. Constructing a statistical model for predicting body fat percentage, based on DEXA results and

morphometric parameters such as tail base width is possible.

1.3.5. The new model of predicting body fat percentage will be applied for geckos, and compared

against orthodox body condition indices (e.g. Mass/RA). The best method to explain the relation

between body index and risk taking: proximity to night lamp, longer activity, forage far from cover,

will be use.

Predictions:

1. Body fat percentage (as measured by DEXA) and predicting the body fat percentage, are

more reliable methods with a better (positive) correlation to risk-taking decision and risk

management, than the orthodox methods for body condition (e.g. Mass/RA). (Low body

index=Riskier foraging).

2. Individuals with low fat% (predicted or measured) will choose a high risk strategy:

1.3.5.1. Forage far from cover, closer to the ground.

1.3.5.2. Active in moon light nights.

1.3.5.3. Time out of cover will be longer.

1.3.5.4. Shorter inactivity period during winter

1.3.6. Predictions based on the hypothesis that:

Sub-adults present foraging strategy that considers possible future competition on social status.

Sub-adults will:

1. Tend to condition the decision, whether to go out foraging, on the body state, status and

activity of other sub-adults, and not only on their own body condition.

2. Sub-adults winter inactivity period is predicted to be shorter than their physiological

ability that can allow them to stay under cover much longer, during winter. So in captivity the

sub-adults demonstrate endurance to famine, which is much longer than the winter inactivity

observed in the field.

1.3.7. Predictions based on both hypotheses that:

a. Dominancy is correlated with directional asymmetry (DA).

b. Sub-adults present foraging strategy that considers possible future competition on social status.

Sub-adults will:

1. Not tend to condition the decision, whether to forage, on the body state, status and

activity of other sub-adults, if they are left biased (DA).

2. Demonstrate shorter winter inactivity, if they are left biased (DA) compared with right

biased individuals (DA).

1.3.8. Predictions based on the hypothesis that:

Directional asymmetry of eye diameter is derived from directional asymmetry of the brain (right

hemisphere vs. left):

 1. Detour responses will be correlated with DA in that detour around a barrier will be to the left for

left biased geckos; and to the right, for right biased geckos.

(Modified from Vallortigara, Regolin, and Pagni, 1999; and Vallortigara, 2000).

2. Approaching food will be correlated with DA (opposite to left and right-biased). So one eye

detects food better and the other eye detects predator better- will be correlated with DA bias

(Modified from Rogers and Anson, 1979).

3. Brain morphology will differ according to DA of eye diameter; in that left biased geckos will

have a larger right brain hemisphere and right biased individuals will have a larger left brain.

4. Brain morphology will differ according to DA of eye diameter, in that left biased geckos will

have a higher neuron density in right brain and right biased individuals will have a higher

neuron density in left brain.

{ How on earth will you do all this, including the ontogenetic change? Maybe

you should sell it to Anat Barnea and she should seek a grant for your post-

doc. }

1.3.9. Predictions based on the hypothesis that:

Directional asymmetry of eye diameter is derived from directional asymmetry of the brain (right

hemisphere vs. left) and mediated by hormones:

 1. Stress hormones will differ with DA of eye diameter, in that left biased gecko will have a higher

stress hormone level and right biased individuals will have a lower stress hormone level.

2. Aggression hormones will differ with DA of eye diameter, in that left biased geckos will have a

higher hormone level and right biased individuals will have a lower hormone level, as seen in

human digits DA (Coyne et al., 2007).

1.3.9.1. Predictions based on the hypothesis that:

By controlling the environmental stress, population composition of DA is controlled:

1. Geckos that will be raised under constant conditions of temperature, excess food and low

exposure to predators, will have a higher rate of right biased DA individual than a group of

geckos that will be held under higher density, lower food regime and under stress induced by

exposure to predators.

2. Methods

2.1. Study organism

I chose the Israeli Fan-toed gecko lizard (Ptyodactylus guttatus), as a study organism model,

since it is not endangered, a common lizard that show DA (Werner et al., 1991). Moreover, it

demonstrates behaviors related to social dominance. Some behavior such as status may be predicted

successfully by a morphological sign of tail loss (Sion, 2006) and observe behaviorally by the height of

the perch, as these rupicolous geckos are magnificent scansorial. The gecko P. guttatus uses caudal

autotomy to escape predators, followed by tail regeneration. Body size (adults are 60-90 mm) in this

species has a substantial effect on the social status of individuals (Frankenberg, 1978; D. Werner, 1972;

Werner, 1965, 1986, 1991; Johnston and Bouskila, 2007). Competitive exclusion based on body size
was demonstrated in captivity (D. Werner, 1972), and is expected in the wild.

The dominant males are territorial; occupy the best perches for foraging (Werner, 1972;

Frankenberg, 1978), such as under night lamps. The females of a dominant male lay their eggs at the

same site (Frankenberg, 1978; Werner, 1965, 1986, 1991). The dominant males mate and prevent other

males to mate on their territory, threaten other males and thus affect their behavior (Johnston and

Bouskila, 2007). They are more active and aggressive and perch higher (Werner, 1972; Frankenberg,

1978);mostly in best body condition (Frankenberg, 1978; Johnston and Bouskila, 2007).

2.1.2. Study Area: Jerusalem (Hebrew University) for captivity behavior experiments. The

Shomron region, and Sede Boqer (Midrasha) in the Negev region, for collecting geckos, and

most outdoor observations in Sede Boqer.

2.1.3. Abiotic parameters: The parameters recorded will be: date, hour, and distance from cover,

wall temperature (º C), moon condition, and proximity to night lamp.

2.1.4. Sede Boqer study site:

The outdoor behavioral study was conducted at Midreshet Ben-Gurion in the northern

Negev desert. The study site was a complex of guest rooms surrounded by a two-meter high

wall, 13x150 meters. P. guttatus inhabited densely the southern part of the premises, and

geckos utilized many of the lamps erected on some of the buildings and on the southern

wall.

2.2. Marking: The study will take place both in the lab and in the field. Geckos will be collected and

marked under permits from the Nature Reserves Authority, Israel.

Short term mark will be made by non-poisonous (child-safe) felt tip pen, long term mark by Destron

and Trovan microchips (Pit Tags). The microchips will be injected subcutaneously with a needle

through a cut made by scalpel, closed by super glue. The incision is made alongside the abdomen, in

between the hind and fore leg. The cut is made closer to the hind leg. The chip is inserted gently by

hand toward the fore leg through the cut, while the chip advances subcutaneously toward the fore leg.
When the chip does not touch the cut, the cut in the skin is closed with superglue. So it is between skin

and muscle.

2.3. Body Measurements: Sex, tail type and scars, weight by scale, while Rastrum-Anus (RA),

width of tail base and tail length, by caliper. Directional asymmetry will be studied and determined,

using comparison of left and right factors of body measurements: eye size and head size (by

caliper) while DA as shown in Van Valen (1962). If not caught, data will be collected such as, tail

condition and estimated body size.

2.4. Behavioral parameters: Foraging proximity to night lamps, foraging distance from cover, inactivity

period during winter (as the minimum number of consecutive days with no individuals of each group;

e.g. DA male adults), and determination of sex for possible sexual effect on behavior.

2.5. Body state: Body Index: an index for body state; an attempt to predict body fat percentage. Tail

base width and Mass/RA, Mass/RA ², Mass/RA ³, and residuals of body mass vs. RA (in linear

regression) will be compared, using DEXA measurement for body fat mass.

2.6. The Experimental design:

Calculating directional asymmetry: using comparison of left and right eye diameter. To neutralize

allometric effects of different body sizes, the subtraction of right eye diameter minus left eye

diameter will be divided by the mean value of eye diameter (Van Valen, 1962), and by percentage

of (RA) body size. This is called PERCRA, as suggested by Hoofien as an international word

(Werner, 1971). The data will be tested for possible links to behavior, such as social status, risk-

taking, and morphometric, such as sex, tail condition and body size and state.
2.7. Predictions based on the hypothesis that:

Morphometric characters (DA, Sex, tail condition, and body index) affect decision making by

risk management:

Predictions:

1. Left biased DA will not forage in proximity to night lamps.

2. Left biased DA will have more tail injuries.

3. Left biased DA will forage far from cover.

4. Left biased DA will have lower fat percentage.

5. Right biased DA will forage in proximity to night lamps.

6. Right biased DA will have less tail injuries.

7. Right biased DA will forage closer to cover.

8. Right biased DA will have higher fat percentage.

9. Males will have more tail injuries than females.

Predictions/O Proximity More tail Forage far Fat%

bservations to night injuries from cover

lamps
Left biased

DA - + + -

Right biased

DA + - - +
Males - +

2.9.1. Museum collection: The prediction that DA has a correlation with tail injury will be tested

by examining preserved specimens from the collection and live individuals from the field. A

secondary outcome is tackling the argument that a morphometric measurement, from

preserved cadavers, is problematic due to possible deformations during preservation:

Predictions:

1. Live geckos, left biased DA will have more tail injuries.

2. Preserved geckos, left biased DA will have more tail injuries.

Predictions More tail injuries

Left biased DA (Live geckos)

Left biased DA (Preserved geckos)

Thus a similar result with live and preserved specimens disproves preservation effects.

2.9.2. Lab behavioral manipulation:

The prediction that left biased DA is correlated with low status will be tested in the lab (in

addition to the field) by lamp proximity. In a small arena two males of the same size group could

compete for food, territory (night lamp) and on access to females. Left biased individuals will be

confronted with right biased individuals of the same body size, and individuals with no significant

asymmetry, with the same feeding regime.

The prediction that, directional asymmetry individuals are preferred by predators when given a choice,

will be tested in the lab. Three different taxa representative: 1) snake 2) bird 3) cat; will be used as

predators, with two individuals in each trial. One with a DA of eye size (left or right) and the other

without DA of eye size, each encounter will be recorded to examine the preference if any, of the

predator (the geckos will not actually be eaten):

Predictions/T Left Right No DA

reatments biased biased

DA DA
Night lamp

- + +
male-male

arena - + +
Diurno-

nocturnal - + +
foraging
Predators

preference + - -
Females

preference - + +

2.9.4. Male-male Lab manipulation: In the same arena, the tendency to win at conflicts with a fake

tail will be tested. First the regenerated tail is colored with rings to mimic intact tail. Another

manipulation would be to mimic a regenerated tail, by coloring an intact original tail with a uniform

color to hide the dark rings.

{First you will have to investigate how minor size and fat differences , and maybe last ,meal,

affect the outcome of contests between “equal” geckos. To be really exact and dependable, you

should first test the individual optimum temperature, to avoid a situation where one contestant

enjoys his optimum but the other not. Meaning, to apply the principle of correct comparative

physiology, to intra-specific work, especially where there is a reason to suspect that it matters

(Arad’s bimodal result)}.

The prediction that, individuals with tail injury (regenerated or not) are preferred by predators,

when given a choice, will be tested in the lab. Three different taxa representative: 1) snake 2) bird 3)

cat; will be used as predators, with two individuals at each trial. One with a tail injury and the other

with no tail injury (intact tail) encounter will be recorded to examine the preference if any, of the

predator.

The tail manipulation: In the same conditions a fake tail will be tested for predators' preference of

tail condition. First the tail is colored with rings to mimic intact tail. Another manipulation would be to

mimic a regenerated tail, by coloring an intact original tail with a uniform color to hide the dark rings.

2.9.3. Predictions based on the hypothesis that:

Tail injury (regenerated or not) bears a vulnerability honest signaling:

Origina Tail injury Fake Fake

l tail regenerated Reg. Orig.

or not tail tail

Night lamp + - + -
male-male

arena + - - +
Diurno-

nocturnal
foraging - + - +
Predators

preference - + + -
Females

preference + - - +
Moonlight

activity + - + -

The male-male interaction experiments set will test those predictions; while behavior will be recorded

in the lab in a small arena where two adult males could compete on food, access to females and on

warm source as a night lamp.

2.9.5. Predictions based on the hypothesis that:

Fat deposited in the base of the tail is proportional to the total body fat content

Tail base Total body Tail Body

fat fat content fat% fat%

Is the
content
column

Proportional

to the row?
Tail base + + + +

width
Predicted tail + + + +

fat%
Predicted + + + +

body fat%
2.9.6. Predicting fat percentage and risk managment:

Predictions based on the hypothesis that:

A reliable body index is strongly related to decision making and risk management.

Predicted Low fat% Low fat% Body condition

activity (Predicted) indices

(e.g. mass/RA)
Forage far

from cover + + -
Moon nights

activity + + -
Time out of

cover + + -
Short winter

inactivity + + -
Diurno-

nocturnal + + -
activity

2.97. Predictions based on the hypothesis that:

Regenerated tail has a different ability of fat deposition, thus a different risk predation strategy

is imposed.
 Regenerated Original

tail tail
Proportional
High fat% + -
moon light - +
Time out of - +

cover
Forage far - +

from cover
• Comparison between tails of the same tail length and body size.

Experimental design:

The piximus store the DEXA data in the computer and allows manipulating the scanned areas,

so it is possible to compare the fat content of a specific zone such as the tail, even if we scanned the

whole animal. This way allows comparing the fat content stored in a renewed tail and an intact tail.

The results of comparing the fat content stored in the two tail type will be tested for possible correlation

with behavioral parameters: Active in moon light nights, time out of cover, foraging distance of cover,

proximity to night lamp.

2.9.8. Predictions based on the hypothesis that:

Sub-adults presents foraging strategy, considers possible future competition on social status.

Sub-adults Sub-adults Adults Juveniles

Condition Right DA Left

foraging activity DA
on:
Self fat% + + + +
Others fat% + - - -
Forage far from

cover - + - +
physiological

ability to prolong

winter inactivity + + + -
Long winter

inactivity - + + -

Experimental design: The experiment will test the hypothesis that sub-adults present a foraging

strategy that reflects future competition on social status. Based on that hypothesis, I assume a certain

mechanism, that sub-adults winter inactivity period is shorter than their physiological ability that can

allow them to stay inactive much longer.

To test this prediction, I will combine field and lab observations and manipulations. The

winter inactivity length will be recorded in the field, and compare with the max. number of days

that sub adults can endure with no food or water. The prediction is, that the ability of the adults to

do so is much longer than the inactivity length, they demonstrate in the field. While the juveniles'

ability in the lab to survive long starvation will be close to that they demonstrate in the field. The

starvation will be monitored, and with the first sign of distress or apathy the individual will be fed,

or force fed if necessary.

2.9.9. Predictions based on the hypothesis that:

Directional asymmetry of eye diameter is derived from directional asymmetry of the brain (right

hemisphere vs. left):

Eye diameter DA Right Left

DA DA
Detour response (R) + -
Detour response (L) - +
Larger right brain - +
Larger left brain + -
Higher neural

density in right brain - +

Lower neural density

in right brain + -

Approaching threat,

predator (or model)

on right side (R eye) + -

Approaching food

on right side (R eye) + -

Stress hormone level - +

Aggression hormone

level + -
Stressful
 environment - +

3. Timetable:

Complete chapter on tail base as non-invasive body index, by September, 2010.

Done, awaits small calibration experiments for tail fat content- Will be done by May.

Complete chapter on asymmetry by December, 2010. Done, awaits the small experiment mention

above.

Complete fieldwork by June, 2011.

Complete lab experiments by September, 2011.

Complete the chapter on endocrinology, by September, 2011.

Complete museum measures by November, 2011.

Complete analysis by February, 2012.

Complete final report by June, 2012.

4. References:

Almog, A., Bonen, H., Herman, K. and Werner, Y. L. 2005. Subspeciation or none? The hardun in the

Aegean (Reptilia:Sauria: Agamidae: Laudakia stellio). Journal of Natural History 39:567–586.

Avital, E. and Jablonka, E. 2000. Animal Traditions: Behavioural Inheritance in

Evolution. Cambridge University Press, Cambridge.

Bustard, H. R. 1967. Gekkonid lizards adapt fat storage to desert environments. Science 158:1197–

1198.

Bauwens, D. 1985. Demografische kenmerken en aantalsdynamiek in een populatie van de

levendbarende hagedis (Lacerta vivipara). Doctoraatsthesis, U.I.A., Wilrijk.

Castilla, A. M., Herrel, A. and Gosa, A. 2008. Mainland versus island differences in behaviour of

Podarcis lizards confronted with dangerous prey: the scorpion Buthus occitanus. Journal of Natural

History 42:2331–2342.

Chapple, D. G. and Swain, R. 2002. Effect of caudal autotomy on locomotor performance in a

viviparous skink (Niveoscincus metallicus). Functional Ecology 16:817-825.

Clark, C. W. 1994. Antipredator behavior and the asset-protection principle. Behavioral Ecology

5:159–170.

Coyne, S. M., Manning, J. T., Ringer, L. and Bailey, L. 2007. Directional asymmetry

(right-left differences) in digit ratio (2D:4D) predict indirect aggression

in women. Personality and Individual Differences 43:865–872.

Daniels, C.B., 1984. The importance of caudal lipid in the gecko Phyllodactylus marmoratus.

Herpetologica 40:337–344.
Dial, B. E. and Fitzpatrick, L. C. 1981. The energetic costs of tail autotomy to reproduction in the lizard

Coleonyx brevis (Sauria: Gekkonidae). Oecologia 51:310–317.

Donoghue, S., Vidal, J., Kronfeld, D., 1998. Growth and morphometrics of Green Iguanas (Iguana

iguana) fed four levels of dietary protein. Journal of Nutrition 128:2587–2589.

Doughty, P. and Shine, R. 1998. Reproductive energy allocation and long-term energy stores in a

viviparous lizard (Eulamprus tympanum). Ecology 79:1073–1083.

Doughty, P., Shine, R. and Lee, M. S. Y. 2003. Energetic costs of tail loss in a montane scincid lizard.

Comparative Biochemistry and Physiology - Part A: Molecular & Integrative Physiology 135:215-219.

Frankenberg, E. 1978. Pupillary response and visual behaviour with relation to activity times in geckos

of various species. Dissertation for PhD. Hebrew University, Israel.

Frankenberg, E. 1978. Interspecific and seasonal variations of daily activity times in gekkonid lizards.

Journal of Herpetology 12:505-509.

Graham, J. H., Emlen, J. M., Freeman, D. C. and Leamy, L. J. 1998. Directional asymmetry and the

measurement of developmental instability. Biological Journal of the Linnean Society 64:1-16.

Graham, J. H., Freeman, D. C. and Emlen, J. M. 1993. Antisymmetry, directional asymmetry, and

dynamic morphogenesis. Genetica 89:121-137.

Huey, R. B., Pianka, E. R. and Hoffman, J. A. 1977. Seasonal variation in thermoregulatory behavior

and body temperature of diurnal Kalahari lizards. Ecology 58:1066-1075.

Johnston, G. and Bouskila, A. 2007. Sexual dimorphism and ecology of the gecko, Ptyodactylus

guttatus. Journal of Herpetology 41:506–513.

Kark, S. 2001. Shifts in Bilateral Asymmetry within a Distribution Range: The Case of the Chukar

Partridge. Evolution 55:2088-2096.

Kotler, B. P., Brown, J. S. and Bouskila, A. 2004. Apprehension and time allocation in gerbils: The

effect of predatory risk and energetic state. Ecology 85:917–922.

Leamy, L. J., Doster, M. J. and Huet-Hudson, Y. M. 1999. Effects of methoxychlor on directional and

fluctuating asymmetry of mandible characters in mice. Ecotoxicology 8:63-71.

Lachman, E., Carmely, H. and Werner, Y. L. 2006. Subspeciation befogged by the “Seligmann effect”:

the case of Laudakia stellio (Reptilia: Sauria: Agamidae) in southern Sinai, Egypt. Journal of Natural

History 40:1259–1284.

Leamy L. 1999. Heritability of directional and fluctuating asymmetry for mandibular char- acters in

random-bred mice. J. Evol. Biol 12:146-55.

Ligon J. D. 1968. Sexual Differences in Foraging Behavior in Two Species of Dendrocopos

Woodpeckers. The Auk 85: 2. 203-215.

Lopez, P., Munoz, A. and Martin, J. 2002. Symmetry, male dominance and female mate preferences in

the Iberian rock lizard, Lacerta monticola. Behavioral ecology and sociobiology 52:342-347.

Møller, A. P. 1992. Female swallow preference for symmetrical male sexual ornaments. Nature

357:238-240.

Møller, A. P., Soler, M. and Thornhill, R. 1995. Breast asymmetry, sexual selection, and human

reproductive success. Ethology and Sociobiology 16:207-219.

Perry, G. 1996. The evolution of sexual dimorphism in the lizard Anolis polylepis (Iguania): evidence

from intraspecific variation in foraging behavior and diet. Canadian Journal of Zoology 74:1238–1245.

Rand, A. L. 1952. Secondary sexual characters and ecological competition. Fieldiana Zoology 34:65-

70.

Rogers, L. J., and Anson, J. M. 1979. Lateralization of function in the chicken forebrain.

Pharmacology Biochemistry and Behavior 10:679–686.

Secor, S. M. and Nagy, T. R. 2003. Non-invasive measure of body composition of snakes using dual

energy X-ray absorptiometry. Comparative biochemistry and physiology 136:379–389.

Seligmann, H. 1998. Evidence that minor directional asymmetry is functional in lizard hindlimbs.

Journal of Zoology 245: 205–208.

Seligmann, H., Beiles, A. and Werner, Y. L. 2003. More injuries in left-footed individual lizards and
Sphenodon. J. Zool. (Lond.) 260:129–144.

Simmons, L. W., Rhodes, G., Peters, M. and Koehler, N. 2004. Are human preferences for facial

symmetry focused on signals of developmental instability? Behavioral Ecology 15:864-871;

Simmonds, M., Emmanuel, J. J., Drew, M. E., Batterham, R. L. and Dolan, R. J. 2010. Metabolic state

alters economic decision making under risk in humans. PLoS ONE 5(6): e11090.

doi:10.1371/journal.pone.0011090

Sion, G. 2006. Foraging Strategy & Social Hierarchy are State Dependent in the Gecko Ptyodactylus

guttatus. Dissertation for M.Sc. Ben-Gurion University of the Negev, Israel.

Smyth, M. 1974. Changes in the fat stores of the skinks Morethia boulengeri and Hemiergis peronii

(Lacertilia). Aust. J. Zool 22:135–145.

Vallortigara, G., Regolin, L., and Pagni, P. 1999. Detour behavior, imprinting, and visual lateralization

in the domestic chick. Cognitive Brain Research 7:307–320.

Vallortigara, G. 2000. Comparative neuropsychology of the dual brain: A stroll

through left and right animals’ perceptual worlds. Brain Lang 73:189-219.

Van Valen, L. 1962. A study of fluctuating asymmetry. Evolution 16:125–142.

Velickovic, M. V. and Jakovcev-Todorovic, D. 2006. Directional asymmetry estimates developmental

instability in plants. Genetika 38(2):169-17.

Vervuast, B., Lailvaux, S. P., Grbac, I. and Van Damme, R. 2008. Do morphological condition indices

predict locomotor performance in the lizard Podarcis sicula? Acta Oecologica 34(2):244-251.

Werner D. 1972. Beobachtungen an Pt.vodacr.vlus husselquistii guttatus (Geckonidae). Verhandl.

Noturf. Ges.Base1 82, 54-87.

Werner, Y. L. 1965. Uber die israelischen geckos der gattung Ptyodactylus und ihre biologie.

Salamandra 1:15–25.

Werner, Y. L. 1971. Some suggestions on the standard expression of measurements. Syst. Zool.

20:249-252.

Werner, Y. L. 1986. Ecology of eggs and laying sites of Ptyodactylus geckos. In Rocek, Z., editor. ed.

Studies in Herpetology 441–444.

Werner, Y. L., Rothenstein, D. and Sivan N. 1991. Directional asymmetry in reptiles (Sauria:

Gekkonidae: Ptyodactylus) and its possible evolutionary role, with implications for biometrical

methodology. Journal of Zoology 225:647-658.

Werner, Y. L., Takahashi, H., Yasukawa, Y., and Ota, H. 2006. Factors affecting foraging behaviour, as

seen in a nocturnal ground lizard, Goniurosaurus kuroiwae kuroiwae. Journal of Natural History 40:

439–459.
Wilson, R. S. and Booth, D. T. 1998. Effect of tail loss on reproductive output and its ecological

significance in the skink Eulamprus quoyii. J. Herpetol. 32:128–131.