Secondary contact in the European wall lizard

A critical mechanism underpinning current biological diversity is the extent to which one species mates with, or avoids mating with, another. However, little is known about the factors that mediate hybridisation, especially during the initial and rarely observed stages of secondary contact when interspecific interactions have not responded to selection. In particular, whilst hybridisation is ultimately a behavioural phenomenon, the role of behaviour in mediating hybridisation and how it is influenced by environmental and circumstantial factors is rarely investigated. Recently introduced species provide us with unequalled opportunities to study these factors. In this thesis I examine the role of behavioural mechanisms, in particular male-male competition and mate choice, in mediating mating patterns between two genetically and phenotypically distinct lineages of European wall lizard (Podarcis muralis) that have come into recent secondary contact through human introductions.
In Chapter Two, I investigated how sexual selection during allopatry is responsible for creating stark differences in phenotypic traits such as body size and weapon performance evident in the two lineages today, ultimately explaining the strong biases in dominance during territorial disputes between males. However, I also show that even given this asymmetry in male competitive ability, the extent to which it extrapolates into greater access to females in naturalistic, outdoor enclosures depends strongly on the spatial clustering of basking sites, a critically important resource for many ectotherms.
In contrast to initial predictions suggested by asymmetries in male competition outlined in the previous chapter, in Chapter Three I show that both paternity and courtship behaviour was strongly assortative in the outdoor enclosures. Further investigation through staged experiments on olfactory mate choice, mating trials and analyses on specific behavioural data obtained in an enclosure experiment, I show that lineage based dominance actually contributes to assortative mating patterns in conjunction with weak conspecific male choice. In contrast, female choice seems to play no role in mediating the mating patterns observed between the two lineages.
In Chapter Four I had the rare opportunity to examine the morphological and behavioural factors that predict why animals should hybridise in the first place, using the data obtained in the enclosure experiment above. I found that hybridisation was particularly common between small individuals of the larger lineage and large individuals of the smaller lineage; a result that corroborates the mechanisms determining the assortative patterns uncovered in Chapter Three. Additionally, hybridisation rates were particularly high in less dominant individuals, which I suggest is due to subordinate males having reduced opportunities for courting conspecific females due to male-male competition, requiring them to become less ‘choosy’ and therefore more likely to mate with heterospecifics.
Finally, secondary contact cannot occur without at least one lineage coming into a new environment, and yet relatively little attention is paid to how this environmental change can affect the signals involved in intraspecific communication and mate choice. In Chapter Five I show that a change in the amount of time male lizards spend thermoregulating (a likely consequence of arriving in a new environment) significantly changes the chemical composition of their scent marks. However, whilst female lizards were able to detect these effects, they did not seem to base their mating decisions on them. Nevertheless, this result raises interesting questions about the potential function and consequences of this plasticity, and highlights the importance of considering plasticity in chemical communication in heterogeneous environments.
Overall, this thesis shows the critically important role of behaviour in mediating intra- and interspecific mating patterns during recent secondary contact. In particular, it highlights how the direction and extent of hybridisation and competition are influenced by the degree to which differing morphological and behavioural phenotypes interact over a heterogeneous environment, particularly during the initial stage of secondary contact when mate choice has not had the chance to respond to the selective pressures of hybridisation.

Keywords: assortative mating
secondary contact
female choice
podarcis muralis
hybridisation
male-male competition
wall lizard
chemical communication
Advisor: Uller, T
Sheldon, B