Blue Tail?

A Potential Deimatic Display

Honors Research Proposal

Britney Free

December 2, 2021

Mentors: David A. Penning, K. Casey Cole

 Free 2

Abstract:

The aim for this study is to explore the possibility of the blue tail coloration in P.

fasciatus serving as a potential deimatic display and attempt to explain the adaptive value of

ontogenetic changes in tail color in P. fasciatus. A deimatic display is an anti-predator strategy

that revolves around startling the predator (Umbers et al. 2015), which at large is still in its

infancy of being explored; the mechanism and evolution behind its theory still far from being

fully understood (Badiane et al. 2018). P. fasciatus currently aren’t known to exhibit any

behaviors that are considered a deimatic display, but I hypothesize a tail-wagging behavior noted

in Cooper 1985 could serve two purposes one which being a deimatic display. I propose to test

this through examining the UV reflectiveness of the tails in P. fasciatus, as well as identifying if

a tail-wagging behavior is exhibited in response to a number of different simulated predatory

attacks.

Introduction:

Color often plays an important role as it can serve as an important signal in both intra-

and interspecific communication in several animal species. Apart from serving as cues for

recognition of an individual or species, color can cover a variety of other semantic meanings

(Williams and Rand 1977; Losos 1985; Macedonia et al. 2013; Luke 1989). Conspicuous colored

patches on animals often correlate with hierarchic status or serve as a sexually selected signal

that advertises male quality (Bergman et al. 2009; Stuart-Fox et al. 2006). This is common in

males of many fish, bird, or reptile species. As this often leads to a direct influence on their

mating success, these traits are often subject to sexual selection (Sinervo et al. 2000). However,
 Free 3

color can also be used to deter potential predators. Anti-predator strategies and responses are all-

pervading throughout our natural world taking many unique forms and mechanistic approaches,

from camouflage to aposematic coloration (Endler 1978; Humphries and Driver 1970; Stuart-

Fox et al. 2006). Camouflage is an adaption that plays off of the perception and cognitive

functions of another allowing an animal to pass as undetected when successful (Merilaita et al.

2017). An example of camouflage can be found in Rhinella margaritifera, a toad that closely

mimics the appearance of leaf litter in its shape and coloration (Barnett et al. 2021). Aposematic

coloration is an adaption that leaves the prey visually conspicuous to signal to predators that they

are an unfavorable prey item, this aversion by predators is often noted as a learned behavior by

predators (Prudic et al. 2007). Some species, however, rely on a completely different type of

anti-predator behavior such as deimatic displays, a strategy of startling the predator for a

moment, flashing suddenly a hidden warning sign, and thus gaining some time to escape

(Umbers et al. 2015). Color while not always core to deimatic displays, often times can play a

sizeable role in aiding to startle a predator. The mountain katydid (or bush cricket) would be a

great example of this as it is cryptic at rest, but when disturbed flashes spectacular abdominal

colors by lifting its wings (Umbers et al. 2015).

Recent debates regarding the deimatic display theory (Umbers et al. 2015, 2017;

Skelhorn 2016; Maldonado 1970) have helped establish and shape a useful theoretical

framework for the understanding, efficiency, and function of this type of display. The deimatic

display theory largely is based on the fundamental tenet, that the effectiveness of deimatic

displays depends on a sudden transition from an inconspicuous state (camouflage for example) to

a highly conspicuous display when a predatory attack is imminent. This transition in theory

would cause a reflexive recoil in the predator. Unlike other anti-predator strategies (aposematism
 Free 4

for example), deimatic displays do not require any learning from the predator to be effective

(Umbers et al. 2017; Prudic et al. 2007). Overall, deimatic displays are unique anti-predatory

behavior that, is still in its infancy of being explored; the mechanism and evolution behind its

theory still far from being fully understood (Badiane et al. 2018).

The chosen study species, the common five-lined skink (Plestiodon fasciatus) is small

and abundantly present lizard widely distributed over the eastern United States. Their body is

generally gray, brown, or black, in background color with five yellowish to white stripes along

their backs. Hatchlings and juveniles are characterized by bright blue tails that tend to fade

considerably with age (Fitch 2019). The adaptive value of ontogenetic changes in tail color in P.

fasciatus remains a poorly understood phenomenon that is often rarely addressed. A few clay

model studies conducted suggest that the tail is unlikely to be a form of aposematic coloration,

but rather a decoy to draw predators away from the body and towards their autonomous tails

(Watson et al. 2012; Batman et al. 2014). Yet, these clay model studies fail to account for

movement which has been shown to play an important role in clay model studies (Paluh et al.

2014). Clay models in many studies often don’t accurately represent the animal they were made

to mimic, which can lead to curious animals messing with brightly colored models that they’d

otherwise avoid if it were the real thing (Paluh et al. 2015). If the blue coloration is solely used

as a decoy mechanism in P. fasciatus that would put a considerable target on the backs of

juvenile skinks which leads me to believe that perhaps there is a secondary defensive mechanism

taking place before autonomy of the tail.

The aim for this study is to explore the possibility of the blue tail coloration in P.

fasciatus serving as a potential deimatic display. At face value this idea may be discarded or

overlooked as the tail of P. fasciatus doesn’t hold the same traditional property of surprise many
 Free 5

colors based deimatic displays often have. For example, the mountain katydid cited earlier flash

their spectacular abdominal colors by lifting its wings when under threat (Umbers et al. 2015).

However, I propose that P. fasciatus still be undergo a sudden transition from an inconspicuous

state to a highly conspicuous one through other means. One study documented an interesting tail-

wag behavior in response to snake predation in P. fasciatus (Cooper 1985). This behavior from

what I could find hasn’t really been studied empirically. Which leads me to hypothesize if

perhaps this tail-wag behavior could serve as two functions. First being perhaps the most obvious

based on current literature which is draw the predator’s attention away from the body the

autonomous tail as a decoy. My second proposed function would that perhaps this behavior is

serving as a deimatic display via the rapid movement of the bright blue coloration of the tail. The

rapid movement serving a transition state from somewhat inconspicuous to highly conspicuous.

Studies on blue tongued skinks have suggested that perhaps their blue tongues are a UV based

deimatic display (Badiane et al. 2018; Abramjan et al. 2015). Which further leads me to question

if we could perhaps, we’re seeing a similar effect in P. fasciatus as their tails also hold a blue

pigmentation. It’s a bit of stretch, but nonetheless an interesting exploration in field of study that

has still yet to be widely explored.

Specifically for this study I would like to test the UV reflectiveness of the tail in P.

fasciatus as well as identify if a tail-wagging behavior is exhibited in response to a number of

simulated predatory attacks. Additionally, I would note any other anti-predator behaviors in the

case some other interesting insights could be gleaned from the study. As this study in particular

would only looking at the behaviors P. fasciatus exhibits, if statistically significantly results are

found further studies would need to be conducted. Most notably, a study looking at predator’s

responses to the tail-wagging behavior of P. fasciatus.

 Free 6

Proposed Methodology:

Study species

Fifty common five-lined skink (P. fasciatus) are to be collected locally (25 adults and 25

juveniles with as close to an even male to female ratio as possible). Collection is to take place

during late spring / early summer as that’s when hatchling start to emerge. Collection is also to

be done at random and a considerable distance apart to avoid collecting clutch mates, this is done

to prevent a similar genetic makeup. The skinks are to be housed individually in 30-gallon tanks

with proper lighting, humidity, heating, and ground cover that were designed to mimic their

natural environment as closely as possible. To gain some baseline observations of the animals

there will be a weeklong adjustment period to captivity before testing occurs to reduce. During

this adjustment period contact minimal human exposure with the animals will be made.

UV measurements

A USB-2000 portable diode-array reflectance spectrophotometer with a R200-7-VIS-NIR

reading illumination probe and a PX2 strobe light source (Ocean Optics Inc.) will be used to

obtain spectral reflectance measurements of the tail, head, and mid body of the lizards.

Following a similar procedure to Badiane et al. 2018.

Behavioral experiments
 Free 7

Behavioral is to take place in an indoor 60 cm x 60 cm x 30 arena. Predator treatments will

consist of a bird (a stuffed American crow), a rodent (a stuffed shrew), a stuffed fox, a stuffed

domestic cat, and block of wood used as a control. Predator treatments were chosen based on

natural predators of P. fasciatus (Vanwormer 2002). Each lizard will participate in two

experimental trials a day during morning hours, which is the peak activity period for this species

(Fitch 1956). The order in which the lizards are to be tested is to be determined randomly. A

wooden stick with metal prongs to attached will be used to move model predators towards the

lizards during four simulated stages of predation in which predation risk is escalated. All trials

were filmed. In stage 1, we placed the lizard in the arena, beneath an opaque cup, and

subsequently introduced the static model predator 2 m away from the lizard. After 2 min, we will

remove the cup and recorded the lizards’ responses for 30 s. In stage 2, we slowly move the

predator 75 cm towards the lizard at a constant speed (25 cm every 10 s). In stage 3, we abruptly

moved the predator closer (approximately 20 cm from the lizard in) to simulate a single attack

and left the predator immobile for 2 min. In stage 4, we simulate intensive attacks by repeatedly

and rapidly presenting the predator to the lizard (no physical contact) for 15 s. From the recorded

footage, it will be recorded weather or not the lizard exhibited a tail shaking behavior. If the tail

shaking behavior occurred, then the time spent shaking will also be recorded. The distance of

predator until skink begins to exhibit some sort of response will also be noted. Any observations

of other defensive behaviors such as mouth gaping, body inflation, hissing, orientation of the

animal, and whether or not the lizard drops its tail will also be recorded, as well as any other

unexpected observations. The mass of each animal was also recorded after each day of testing.
 Free 8

Proposed Statistical Analysis:

To analyze the UV measurements a two-way ANOVA of the tail, head, and mid body

spectral readings of both juvenile and adult lizards will be run to determine if there’s a statistical

significance. If there a statistical significance an appropriate post hoc will be used.

To analyze the behavioral data a chi square test would be performed on our Yes/No data

set regarding whether a tail shaking behavior was displayed. An ANCOVA or ANOVA

preformed with the necessary post-hoc tests can be performed on additional data sets. An

example of an ANCOVA that would be potentially run would be the number of tail shakes as our

dependent variable, the mass of the lizard as the continuous independent variable, and the type of

predator as the categorical independent variable. For differentiating between juvenile and adult

skinks mass can be used as used in the example above, but this could alternatively be a

categorical variable (e.g. “Juvenile” “Adult”) characterizing the loss of blue pigmentation in the

tail. Effectively wither way the same story is told as there’s really no difference between what an

ANOVA and what a regression are doing. However, I think before I decided which route I’d like

to go for my experiment I’d have to look at the weight data of my skinks and make sure that

juveniles and adults aren’t within the same weight spread. This is highly unlikely generally as an

animal grows it gains mass up into adulthood, but it’s still better to go ahead and look at the data

trend when collected.

It’s hard to pinpoint for certain what tests should be used because data hasn’t been

collected and the proper assumption checks haven’t been run, but this is just a rough idea of how

the data would be analyzed.

 Free 9

Citations:

Abramjan, A., Bauerová, A., Somerová, B., & Frynta, D. (2015). Why is the tongue of blue-

tongued skinks blue? Reflectance of lingual surface and its consequences for visual

perception by conspecifics and predators. The Science of Nature, 102(7), 1-12.

Badiane, A., Carazo, P., Price-Rees, S. J., Ferrando-Bernal, M., & Whiting, M. J. (2018). Why

blue tongue? A potential UV-based deimatic display in a lizard. Behavioral Ecology and

Sociobiology, 72(7), 1-11.

Barnett, J. B., Michalis, C., Scott‐Samuel, N. E., & Cuthill, I. C. (2021). Colour pattern variation

forms local background matching camouflage in a leaf‐mimicking toad. Journal of

Evolutionary Biology, 34(10), 1531-1540.

Bateman, P. W., Fleming, P. A., & Rolek, B. (2014). Bite me: blue tails as a ‘risky-decoy’

defense tactic for lizards. Current Zoology, 60(3), 333-337.

Bergman, T. J., Ho, L., & Beehner, J. C. (2009). Chest color and social status in male geladas

(Theropithecus gelada). International Journal of Primatology, 30(6), 791-806.

 Free 10

Cooper Jr, W. E., & Vitt, L. J. (1985). Blue tails and autotomy: enhancement of predation

avoidance in juvenile skinks. Zeitschrift für Tierpsychologie, 70(4), 265-276.

Endler, J. A. (1978). A predator’s view of animal color patterns. In Evolutionary biology (pp.

319-364). Springer, Boston, MA.

Fitch, H. 1956. Life history and ecology of the five-lined skink, Eumeces fasciatus. Pp. 1-156 in

E Hall, A Leonard, R Wilson, eds. University of Kansas Museum of Natural History

Publications volume 8. Topeka, Kansas: University of Kansas

Fitch, H. S. (2019). Life history and ecology of the five-lined skink, Eumeces fasciatus. Good

Press.

Humphries, D. A., & Driver, P. M. (1970). Protean defence by prey animals. Oecologia, 5(4),

285-302.

Losos, Jonathan B. (1985). An Experimental Demonstration of the Species-Recognition Role of

Anolis Dewlap Color. Copeia, 1985(4), 905–910.

 Free 11

Luke, C. A. (1989). Color as a phenotypically plastic character in the side-blotched lizard, Uta

stansburiana (Doctoral dissertation, University of California, Berkeley).

Macedonia, J. M.; Clark, D. L.; Riley, R. G.; Kemp, D. J. (2013). Species recognition of color

and motion signals in Anolis grahami: evidence from responses to lizard robots.

Behavioral Ecology, 24(4), 846–852.

Maldonado, H. (1970). The deimatic reaction in the praying mantis Stagmatoptera biocellata.

Zeitschrift für vergleichende Physiologie, 68(1), 60-71.

Merilaita, S., Scott-Samuel, N. E., & Cuthill, I. C. (2017). How camouflage works.

Philosophical

Transactions of the Royal Society B: Biological Sciences, 372(1724), 20160341.

Paluh, D. J., Hantak, M. M., & Saporito, R. A. (2014). A test of aposematism in the dendrobatid

poison frog Oophaga pumilio: the importance of movement in clay model

experiments. Journal of Herpetology, 48(2), 249-254.

 Free 12

Paluh, D. J., Kenison, E. K., & Saporito, R. A. (2015). Frog or fruit? The importance of color and

shape to bird predators in clay model experiments. Copeia, 103(1), 58-63.

Prudic, K. L., Skemp, A. K., & Papaj, D. R. (2007). Aposematic coloration, luminance contrast,

and the benefits of conspicuousness. Behavioral Ecology, 18(1), 41-46.

Sinervo, B., Miles, D. B., Frankino, W. A., Klukowski, M., & DeNardo, D. F. (2000).

Testosterone, endurance, and Darwinian fitness: natural and sexual selection on the physiological

bases of alternative male behaviors in side-blotched lizards. Hormones and Behavior, 38(4), 222-

233.

Skelhorn, J., Holmes, G. G., & Rowe, C. (2016). Deimatic or aposematic?. Animal Behaviour.

Stuart-Fox, D., Whiting, M. J., & Moussalli, A. (2006). Camouflage and colour change:

antipredator responses to bird and snake predators across multiple populations in a dwarf

chameleon. Biological Journal of the Linnean Society, 88(3), 437-446.

Umbers, K. D. (2013). On the perception, production and function of blue colouration in

animals. Journal of Zoology, 289(4), 229-242.

 Free 13

Umbers, K. D., Lehtonen, J., & Mappes, J. (2015). Deimatic displays. Current Biology, 25(2),

R58-R59.

Umbers, K. D., De Bona, S., White, T. E., Lehtonen, J., Mappes, J., & Endler, J. A. (2017).

Deimatism: a neglected component of antipredator defence. Biology Letters, 13(4),

20160936.

Vanwormer, E. (2002). Plestiodon Fasciatus (Five-Lined Skink). Animal Diversity Web

https://animaldiversity.org/accounts/Plestiodon_fasciatus/

Watson, C. M., Roelke, C. E., Pasichnyk, P. N., & Cox, C. L. (2012). The fitness consequences

of the autotomous blue tail in lizards: an empirical test of predator response using clay

models. Zoology, 115(5), 339-344.

Williams, Ernest E.; Rand, A. Stanley (1977). Species Recognition, Dewlap Function and Faunal

Size. Integrative and Comparative Biology, 17(1), 261–270