Because of their striking diversity, Lake Malawi cichlid fish have been well studied for male aggression, particularly among dominant males of closely related sister species within the framework of mate-choice and speciation. However, aggression in females has been largely ignored, and variation in aggressive behaviors between more distantly-related taxa is not well understood despite its potential impact in a complex community structure. To better understand variation in patterns of aggression between species, we presented males and females from five species of Lake Malawi cichlid with a non-predator intruder and recorded all movement and aggressive acts. Additionally, we measured excreted cortisol levels the day after the intruder assay to evaluate one physiological aspect of stress response. We identified species-specific patterns in both specific aggressive acts, and overall level of aggression. Additionally, we found that sexual dimorphism in aggressive acts varies by species and act, where the species with the most aggressive males also has aggressive females. Additionally, cortisol levels vary by taxa, and are associated with restless behavior in the intruder assay, but not levels of aggression. These findings have bearing on understanding sex differences in aggression and their impact on community structure in this important model of rapid evolution.fine-scale variation between sister species within the rock genus Metriaclima [M. aurora, M. lombardoi, M. callainos, and M. pyrsonotos], as well as the more distantly related, sand-dwelling species Aulonocara baenschi.
Results
Aggression differences by sex and speciesAt the broadest levels, behavioral patterns can be visualized with a time-line representation of the ethogram, or catalog of all the behaviors observed in the assay (figure S1), which allows for a non-quantitative overview of the patterns of behavior displayed by each individual fish. A comparison of mean summary values for each behavior (count or duration) by species, sex, and the species by sex interaction showed differences in the most aggressive behaviors. The aggressive behaviors lunging (F(9,61)=3.71, corrected p=0.0063) and chasing (F(9,61)=2.84, corrected p=0.044) have significant species main effects, as did the attentive fin display behavior (F(9,61)=4.69, corrected p=0.0008). Sex and the interaction term were left in the model despite non-significance because M. lombardoi shows aggressive behaviors in males but not females; while this may be an artifact, the pattern is clear for three behaviors, and thus deemed biologically relevant and may be detectable with a greater sample size (figure 1, top three panels). The quiver display was not significantly different by species in the model (F(9,61)=2.36, corrected p=0.117), but was notably only present in 4 of 8 M. aurora males, and a single M. lombardoi male, and thus the only male exclusive behavior recorded (figure 1, third panel from the top).The most aggressive species, M. aurora, and the least aggressive species, A. baenschi display ma...