Research     

Brief Overview
  

I am currently researching a hypothesis of trade-offs between mimicry and thermoregulation efficiency in the California Mountain Kingsnake (Lampropeltis zonata). I began collecting data for my graduate work as an undergraduate at U.C.Santa Cruz and have been compiling data from breeding experiments ever since. The aim of the original breeding experiments was to explore the degree of phenotypic plasticity in pattern development of L. zonata by experimentally adjusting incubation temperatures. The data from those initial years of captive propagation have since been used to develop and defend my hypothesis of the aforementioned trade-offs. The California mountain kingsnake is a member of a diverse group of tri-colored (white, black and red banded) North American colubrid snakes. Though non-venomous, they are thought to be Batesian mimics of dangerous tropical coral snakes in the genera Micrurus and Micruroides. Batesian mimicry involves a potential predator, an unpalatable model species, and a palatable mimetic species.

Swainson's Hawk Buteo swainsoni Photo by Carl Schwartz (USFWS) Possible prey

        Arizona Coral Snake                      California Mountain Kingsnake
Micruroides euryxanthus                              Lampropeltis zonata
"Model"                                                  "Mimic"
Photo by Tom Brennan                           Photo by Gerold Merker


     The mimic is afforded the greatest amount of protection by possible 
predators if its pattern closely resembles the model.  Therefore, selection 
favors the mimetic pattern that most closely resembles the model species.  However, the 
more the mimic and unpalatable model resemble each other the greater the chance of 
possible predation upon the model.  This is a result from predators learning that the 
aposematic signal is not always associated with lethal consequences.  Concurrently,
 selection towards differentiation from the pattern of the most abundant mimic is 
occurring in the model species – an antagonistic effect of Batesian mimicry.

     Sympatry between model and mimic is typically considered a critical 
condition of the Batesian mimicry system that, if absent, should decrease the 
protection afforded the mimic.  Many kingsnake species, however, including
L. zonata, are distributed outside the range of coral snakes.

Avian migration could be the key mechanism that perpetuates Batesian mimicry
where the model and mimic are allopatric.  Several predatory bird species such
as kites, eagles and hawks, annually engage in latitudinal migration routes
from Mexico north through California's coastline, a route beginning within coral
snake range and ending within the range of L. zonata.  

Below is the geographic distribution of L. zonata and all of its previously 
recognized subspecies in different colors.  A phylogeographic study using mtDNA
showed that the subspecific taxa of L. zonata are not genetically distinct and invalid 
(Rodriquez-Robles, 1999).  As a result of the study the seven recognized subspecies of
 L. zonata were dissolved, resulting in a single species with no subspecies.  However, I have provided 
the historical range map of the seven subspecies which depicts the overall geographic range 
of L. zonata in its current monophyletic distribution.

Whether avoidance of coral snake patterns stems from innate recognition of aposematic signals or is a learned behavior, avian predators should avoid the signal as a direct result of spending approximately six months each year within the range of a lethal model. Failure to distinguish between a model and mimic could be lethal. However, not all populations of L. zonata display a highly mimetic pattern. A trade-off may exist between mimetic resemblance and physiological functions; populations that inhabit high elevations have patterns severely reduced in red, and instead have expanded black bands that may increase thermoregulatory efficiency. Below are three examples of pattern that demonstrate the variation between populations of L. zonata.

Highly confluent (L. zonata – average coast variant)

Poor confluency (L. zonata – average Sierra variant)

Lacking all red pigmentation (L. zonata – atypical Sierra variant)

From field collected and in-lab breeding data gathered over the last five years there are significant differences in the average amount of triad confluency (i.e., where a black ring is completely interrupted by a red band dorsally) and the amount of black pigmentation between regionalized populations of L. zonata. The Coast populations have the least amount of black pigmentation and are highly confluent, while the Sierra populations, high elevation populations, have the greatest amount of black pigmentation (63.67% average) of all L. zonata I have examined. This high elevation pattern may be necessary for increasing thermoregulatory efficiency because of shorter seasons and overall cooler temperatures. However, as the amount of black pigmentation increases, the strength of the mimetic signal is likely compromised. Thus, the anti-predatory advantage arising from mimicry, and in thermoregulatory efficiency arising from a darker-colored dorsal pattern, may create an evolutionary trade-off. Hypothetically then, coast populations are mimetic, whereas high altitude populations of L. zonata, thermoregulation has maybe become the dominant pressure, outweighing mimicry. Different species and subspecies of tricolor snakes (the majority being from the genus Lampropeltis) range throughout many of the lower forty eight states. Many do not share the same geographical distribution or overlap in range with any of the three coral snake species native to the United States. Therefore, these snakes are phenotypically maladaptive – they are highly conspicuous, brandishing warning coloration and having no type of secondary defense to back it up (i.e. venom). The goal of my research is to explain the distribution of North American tricolored colubrids with ranges outside the range of coral snakes.