Jukka Kekäläinen

Female ornaments revisited – are they correlated with offspring quality?

The evolution and signalling content of female ornamentation has remained an enduring challenge to evolutionary biologists, despite the fact that secondary sexual characters are widespread in females. While females usually invest significant amounts of their resources, including carotenoids, in offspring, all the resources allocated to elaborate ornamentation reduce resources available for other purposes. This may in turn constrain female fitness leading to dishonest female signalling. We review the literature for empirical studies on mutually ornamented species with conventional sex roles, by focusing on the association between female ornaments and quality of their offspring. We found 43 papers where 33 (77%) are bird-studies, nine (21%) are on fishes, and one (2%) is a lizard-study. Nine of these report negative, 14 non-existing, and 20 positive associations between female ornament and offspring quality. Eighteen of the bird studies (55%) show a positive association between the two traits investigated, whereas five (15%) of the studies report a negative association. The number of fish studies, although few, is skewed in the opposite direction with two (22%) and four (44%) studies supporting positive and negative association, respectively. A minority of studies on carotenoids-based ornaments reports a positive association (4 of 18 studies, or 22%) between the traits, which is low compared to studies on non-carotenoids-based ornaments (16 of 25 studies, or 64%). The above-mentioned relative large number of studies with negative association, especially common in studies on fishes and in carotenoids-based-ornaments, challenges the generality of the direct selection hypothesis to account for female fineries. This is important because this hypothesis seems to have strong support in recent literature on the topic. In the present paper, we also propose possible explanations for the observed differences between taxa and suggest directions and ideas for future research on the evolution of female ornamentation.

Do brain parasites alter host personality? — Experimental study in minnows

Despite that the existence of animal personalities is widely recognized, no consensus has been reached on the relative importance of different ecological factors behind their expression. Recently, it has been suggested that parasites may have a crucial role in shaping animal personalities, but only a very few studies have experimentally tested the idea. We infected Eurasian minnows (Phoxinus phoxinus) with the brain-encysted trematode parasite, Diplostomum phoxini, and studied whether infection could modify the personality of their hosts. Our results show that D. phoxini infection did not affect the mean levels of boldness, activity or exploration, but infected minnows showed higher repeatability in boldness and activity, and reduced repeatability in exploration. We also found that D. phoxini may be able to break the associations (behavioral syndromes) between behavioral traits, but that this effect may be dependent on parasite intensity. Furthermore, the effect of D. phoxini infection on personality of the hosts was found to be nonlinearly dependent on infection intensity. Taken together, our results suggest that D. phoxini parasites may shape the personality of their hosts, but that behavioral consequences of ecologically relevant infection levels may be rather subtle and easily remain undetected if only the mean trait expressions are compared.

Signals of major histocompatibility complex overdominance in a wild salmonid population

The major histocompatibility complex (MHC) contains the most variable genes in vertebrates, but despite extensive research, the mechanisms maintaining this polymorphism are still unresolved. One hypothesis is that MHC polymorphism is a result of balancing selection operating by overdominance, but convincing evidence for overdominant selection in natural populations has been lacking. We present strong evidence consistent with MHC-specific overdominance in a free-living population of Arctic charr (Salvelinus alpinus) in northernmost Europe. In this population, where just two MHC alleles were observed, MHC heterozygous fish had a lower parasite load, were in better condition (as estimated by a fatness indicator) and had higher survival under stress than either of the homozygotes. Conversely, there was no consistent association between these fitness measures and assumedly neutral microsatellite variability, indicating an MHC-specific effect. Our results provide convincing empirical evidence consistent with the notion that overdominance can be an important evolutionary mechanism contributing to MHC polymorphism in wild animal populations. They also support a recent simulation study indicating that the number of alleles expected to be maintained at an MHC loci can be low, even under strong heterozygote advantage.

Both male and female sexual ornaments reflect offspring performance in a fish.

Secondary sexual characters are often expressed in both sexes (mutual ornamentation), but are less often studied simultaneously. We studied the adaptive signaling function of male and female ornamentation in a mutually ornamented fish, the whitefish Coregonus lavaretus. In an experimental design in which nongenetic environmental effects were minimized, we found that highly ornamented females, males, and their parental combinations had offspring with better swimming performance and predator‐avoidance ability than less ornamented individuals or combinations. Furthermore, highly ornamented females had larger offspring that also had higher yolk volume than less ornamented individuals. Offspring swimming performance was not dependent on offspring size and was only weakly affected by yolk volume, which suggest that swimming performance and measured morphological traits are independent fitness measures. In conclusion, mutual ornamentation of whitefish may signal the quality of individuals in both sexes, which may indicate ongoing directional selection for these ornamental traits. However, offspring fitness traits were also dependent on parental combination, which suggests that genetic compatibility effects may weaken the directional selection and the indicator value of the ornamentation.

Experiments on the ectoparasitic deer ked that often attacks humans; preferences for body parts, colour and temperature.

The deer ked (Lipoptena cervi) can fail in its host search. Host search fails when an individual deer ked irreversibly accepts a host unsuitable for its reproduction (e.g. a human) and drops its wings. In northern Europe, the main host of the deer ked is the moose (Alces alces). The deer ked is increasingly causing serious problems for humans (for example, causing deer ked dermatitis) and is considered a threat for the recreational use of forests. The adult deer ked flies in early and mid-autumn to search for a host. Our aims were: (i) to study whether there are ways to avoid deer ked attacks by wearing particular clothing, and (ii) to evaluate deer ked host choice. Using human targets, we explored the cues the deer ked uses for host selection. We studied which part of the host body deer keds target and if body colour and temperature affect their choice. In our experiments, deer keds landed more on dark and red clothing than on white clothing. Moreover, deer keds mostly attacked the upper body parts and preferred the back side of the body over the front side. Finally, deer keds preferred the warmest areas of the host.

Individually assessed boldness predicts Perca fluviatilis behaviour in shoals, but is not associated with the capture order or angling method

Selectivity of recreational angling on fish behaviour was studied by examining whether capture order or lure type (natural v. artificial bait) in ice-fishing could explain behavioural variation among perch Perca fluviatilis individuals. It was also tested if individually assessed personality predicts fish behaviour in groups, in the presence of natural predators. Perca fluviatilis showed individually repeatable behaviour both in individual and in group tests. Capture order, capture method, condition factor or past growth rate did not explain variation in individual behaviour. Individually determined boldness as well as fish size, however, were positively associated with first entrance to the predator zone (i.e. initial risk taking) in group behaviour tests. Individually determined boldness also explained long-term activity and total time spent in the vicinity of predators in the group. These findings suggest that individual and laboratory-based boldness tests predict boldness of P. fluviatilis in also ecologically relevant conditions, i.e. in shoals and in the presence of natural predators. The present results, however, also indicate that the above-mentioned two angling methods may not be selective for certain behavioural types in comparison to each other.

Do highly ornamented and less parasitized males have high quality sperm? – an experimental test for parasite‐induced reproductive trade‐offs in European minnow (Phoxinus phoxinus)

Parasites take their resources from hosts and thus directly reduce available resources for hosts’ own body functions, such as growth and reproduction. Furthermore, parasite infections cause significant indirect costs to their hosts in terms of increased investments on immune defense. In this study, we investigated the impact of parasite infection on the sperm quality and expression of secondary sexual ornamentation (saturation of the red abdominal colouration and number of breeding tubercles) in the Eurasian minnow (Phoxinus phoxinus). We exposed minnows to a high and low dose of common nonspecific fish ectoparasite, the glochidia larvae of duck mussel (Anodonta anatina) and tested whether parasite infection leads to trade-off in sperm quality and/or ornamental expression. We found that glochidia infection reduces the curvature of the sperm swimming trajectory, number of breeding tubercles, and possibly male competitive ability, but does not affect expression of male color ornamentation. Furthermore, glochidia infection was found to reduce sperm motility, but only when all the noninfected individuals were excluded from the model. Supporting one of the predictions by phenotype-linked fertility hypothesis both in high-infection and low-infection group male breeding colouration was positively associated with sperm quality. Our results suggest that although glochidia infection may have negative impact on male reproductive success, parasite-induced costs may not create strong trade-off between breeding colouration and sperm quality or that such trade-off become detectable only in resource-limited conditions.

Male Facial Anthropometry and Attractiveness

The symmetry and masculinity of the face are often considered important elements of male facial attractiveness. However, facial preferences are rarely studied on natural faces. We studied the effect of these traits and facial metric parameters on facial attractiveness in Spanish and Colombian raters. In total, 13 metric and 11 asymmetry parameters from natural, unmanipulated frontal face photographs of 50 Spanish men were measured with the USIA semiautomatic anthropometric software. All raters (women and men) were asked to rank these images as potential long-term partners for females. In both sexes, facial attractiveness was negatively associated with facial masculinity, and preference was not associated with facial symmetry. In Spanish raters, both sexes preferred male traits that were larger in the right side of the face, which may reflect a human tendency to prefer a certain degree of facial asymmetry. We did not find such preference in Colombian raters, but they did show stronger preference for facial femininity than Spanish raters. Present results suggest that facial relative femininity, which is expected to signal, eg good parenting and cooperation skills, may be an important signal of mate quality when females seek long-term partners. Facial symmetry appears unimportant in such long-term mating preferences.

The information content of odour, colour and tactile cues in the mate choice of minnows

Sexual displays often involve many different signal components, which may give information about the same or different mate qualities. We studied the information content of different signals in male minnows (Phoxinus phoxinus) and tested whether females are able to discriminate between males when only olfactory cues are present. We found that females preferred the odour of males with a more saturated (i.e., redder) belly, but only when the females had been in physical contact with the males before the experiments. Instead, when unfamiliar males were used, females did not discriminate between male odours and also the overall swimming activity (mate choice intensity) of the females was significantly lower. More ornamented males had lower number of Philometra ovata parasites (indicated by belly saturation) and Neoechinorhynchus rutili parasites (indicated by belly hue) than their less ornamented counterparts. We did not find experimental evidence for female odour preference being linked to belly hue and breeding tubercle number, but in the nature these traits were associated with the condition factor of the males. Taken together, our results suggest that belly colouration and breeding tubercles give honest information on several aspects of male quality. In addition females may learn the association between male colouration and their olfactory signals and utilize this information when visual signals are not present.

Metabolic rate associates with, but does not generate covariation between, behaviours in western stutter-trilling crickets, Gryllus integer

The causes and consequences of among-individual variation and covariation in behaviours are of substantial interest to behavioural ecology, but the proximate mechanisms underpinning this (co)variation are still unclear. Previous research suggests metabolic rate as a potential proximate mechanism to explain behavioural covariation. We measured the resting metabolic rate (RMR), boldness and exploration in western stutter-trilling crickets, Gryllus integer, selected differentially for short and fast development over two generations. After applying mixed-effects models to reveal the sign of the covariation, we applied structural equation models to an individual-level covariance matrix to examine whether the RMR generates covariation between the measured behaviours. All traits showed among-individual variation and covariation: RMR and boldness were positively correlated, RMR and exploration were negatively correlated, and boldness and exploration were negatively correlated. However, the RMR was not a causal factor generating covariation between boldness and exploration. Instead, the covariation between all three traits was explained by another, unmeasured mechanism. The selection lines differed from each other in all measured traits and significantly affected the covariance matrix structure between the traits, suggesting that there is a genetic component in the trait integration. Our results emphasize that interpretations made solely from the correlation matrix might be misleading.