Male Red Grouse Research Articles

Intra-sexual competition modulates calling behavior and its association with secondary sexual traits

The expression of elaborate sexual displays is associated with individual quality ensuring reliable information about the bearers. However, the associated cost of expressing enhanced sexual traits is expected to change according to environmental circumstances. Specifically, the cost of maintaining or producing a signal is predicted to increase when environmental conditions are unfavorable, which may lead to a reduction in signal expression as shown in several species. Here, we compared the calling behavior of male red grouse Lagopus lagopus scoticus living in an area of experimentally increased intra-sexual competition to that of males living in a control area. Levels of intra-sexual competition were experimentally manipulated by testosterone implants in a subset of captured males. In addition, we compared the association between two sexual traits, calling behavior and comb size, of males living in these two areas. Although call frequency was not affected by different levels of intra-sexual competition, males from the control area performed shorter calls than individuals from the treatment area. Additionally, a positive association between comb size and call duration was found only for males in the area of lower aggressiveness. We suggest that environmental conditions influence the expression of multiple plastic sexual traits, depending on the costs and the information conveyed about different individual qualities. Although sexual signals are considered reliable indicators of individual quality, environmental heterogeneity may modulate their expression and reliability. We experimentally manipulated levels of intra-sexual competition in a wild population of red grouse by increasing testosterone levels using implants in a subset of males. We observed that the social context shapes the expression of sexual traits. In less competitive conditions, males performed shorter calls and call length was positively related to the size of the supra-orbital comb, a relevant secondary sexual trait. Under higher intra-sexual competition, this relationship was decoupled and investment was directed at increasing call length.

Intra-sexual competition alters the relationship between testosterone and ornament expression in a wild territorial bird

In a reliable signalling system, individual quality is expected to mediate the costs associated with ornamental displays, with relatively lower costs being paid by individuals of higher quality. These relative costs should depend not only on individual quality, but also on levels of intra-sexual competition. We explored the current and delayed effects that testosterone implants have on bird ornamentation in populations with contrasted population densities, as a proxy for intra-sexual competition. In a replicated experiment, we manipulated testosterone in 196 yearling male red grouse Lagopus lagopus scoticus in autumn in populations of high and low levels of intra-sexual competition. Males were assigned to one of three exogenous testosterone (T) treatments: empty implants (T0), small T implants (T1) or larger T implants (T2). We monitored subsequent changes in testosterone levels, ornament size and carotenoid-based colouration, carotenoid levels and body condition from autumn to spring. Testosterone implants increased testosterone levels, comb redness and comb size, and decreased body condition but these effects depended on levels of intra-sexual competition. Specifically, T2-implanted birds increased testosterone levels and comb size more, and reduced body condition more, in populations where intra-sexual competition was low. In the following spring, testosterone levels of T2-treated birds kept increasing in populations where intra-sexual competition was high but not in populations where intra-sexual competition was low. Our results highlight that levels of intra-sexual competition alter the relationship between testosterone levels and ornament expression, influencing their condition-dependence; they also indicate that the outcome of standard hormone manipulation conducted in free-living animals vary depending on the population context.

Social context decouples the relationship between a sexual ornament and testosterone levels in a male wild bird

In order to maximise fitness individuals should adjust their level of signalling according to their surrounding social environment. However, field experiments showing such adjustment of current signalling associated to changes in social context are lacking. Here, we manipulated levels of male aggressive- and dominance-related displays in a wild bird in our treated area by increasing testosterone levels using implants in a subset of males. We then compared the expression of sexual signals (i.e. comb size) between non-treated red grouse Lagopus lagopus scoticus males from control and treatment areas. We further explored the potential endocrinological mechanism linking social environment and signal expression by analysing testosterone levels in all males. Our treatment successfully increased overall aggressive- and dominance-related behaviours in the treatment area. Furthermore, testosterone-implanted birds increased their comb size as repeatedly shown in previous studies in male red grouse. Interestingly, untreated males living in the treatment area decreased their comb size, whilst increasing testosterone levels. Since comb size is a signal of dominance, untreated males from the treatment area may have perceived themselves as subordinate individuals and decreased their signalling levels to avoid confrontations with testosterone-treated, dominant individuals. In conclusion, our findings show that social context has the potential to regulate sexual signalling and testosterone levels. Our results highlight the role of social context when exploring the link between testosterone and behaviour, as it may reverse the relationship between both traits. Our results suggest that social context affects signalling and testosterone independently.

Transcriptomic response of red grouse to gastro‐intestinal nematode parasites and testosterone: implications for population dynamics

A central issue in ecology is in understanding the relative influences of intrinsic and extrinsic effects on population regulation. Previous studies on the cyclic population dynamics of red grouse (Lagopus lagopus scoticus) have emphasized the destabilizing effects of either nematode parasites or territorial behaviour and aggression. The potential interacting effects of these processes, mediated through density-dependent, environmentally induced alterations of host immunocompetence influencing susceptibility to parasites have not been considered. Male red grouse at high density are more aggressive, associated with increased testosterone, which potentially could lead to reduced immunocompetence at a stage when parasites are most prevalent. This could depress individual condition, breeding performance and survival and thus drive or contribute to overall reductions in population size. Here, we characterize the transcriptomic response of grouse to nematode parasite infection and investigate how this is subsequently affected by testosterone, using a microarray approach contrasting red grouse with high and low parasite load at both high and low testosterone titre. A suite of 52 transcripts showed a significant level of up-regulation to either chronic parasite load or experimental parasite infection. Of these, 51 (98%) showed a reduced level of expression under conditions of high parasite load and high testosterone. The genes up-regulated by parasites and then down-regulated at high testosterone titre were not necessarily associated with immune response, as might be intuitively expected. The results are discussed in relation to the fitness and condition of individual red grouse and factors influencing the regulation of abundance in natural populations.

GROUSE COMMUNICATE STATE OF HEALTH WITH RED COMBS

![Figure][1] Parents of teenagers know just how much preening goes on before a date, and animals are no different: male peacocks display their impressive tails, blue footed boobies flash their feet and male red grouse get out an impressive pair of red combs above their eyes. Francois Mougeot

Ornamental comb colour predicts T-cell-mediated immunity in male red grouse Lagopus lagopus scoticus

Sexual ornaments might reliably indicate the ability to cope with parasites and diseases, and a better ability to mount a primary inflammatory response to a novel challenge. Carotenoid-based ornaments are amongst the commonest sexual signals of birds and often influence mate choice. Because carotenoids are immuno-stimulants, signallers may trade-off allocating these to ornamental colouration or using them for immune responses, so carotenoid-based ornaments might be particularly useful as honest indicators of immuno-compentence. Tetraonid birds, such as the red grouse Lagopus lagopus scoticus, exhibit supra-orbital yellow-red combs, a conspicuous ornament which functions in intra- and inter-sexual selection. The colour of combs is due to epidermal pigmentation by carotenoids, while their size is testosterone-dependent. In this study, I investigated whether comb characteristics, and in particular, comb colour, indicated immuno-competence in free-living male red grouse. I assessed T-cell-mediated immunity using a standardised challenge with phytohaemagglutinin. Red grouse combs reflect in the red and in the ultraviolet spectrum of light, which is not visible to humans but that grouse most likely see, so I measured comb colour across the whole bird visible spectrum (300-700 nm) using a reflectance spectrometer. I found that males with bigger and redder combs, but with less ultraviolet reflectance, had greater T-cell-mediated immune response. Comb colour predicted T-cell-mediated immune response better than comb size, indicating that the carotenoid-based colouration of this ornament might reliably signal this aspect of male quality.

Nematode parasites reduce carotenoid-based signalling in male red grouse

Carotenoids determine the yellow-red colours of many ornaments, which often function as signals of quality. Carotenoid-based signalling may reliably advertise health and should be particularly sensitive to parasite infections. Nematodes are among the commonest parasites of vertebrates, with well-documented negative effects on their hosts. However, to date, little is known about the effects that these parasites may have on carotenoid-based signalling. Tetraonid birds (grouse) exhibit supra-orbital combs, which are bright integumentary ornaments pigmented by carotenoids. We tested the effect of the nematode parasite Trichostrongylus tenuis on signalling in free-living male red grouse Lagopus lagopus scoticus. We show that experimentally reduced nematode infection increases plasma carotenoid concentration and comb redness, demonstrating for the first time that nematodes can influence carotenoid-based signals.

Carotenoid-based colouration and ultraviolet reflectance of the sexual ornaments of grouse

Among the most familiar sexual signals are red, yellow, and orange sexual traits pigmented by carotenoids. Many birds can detect near-ultraviolet (UV) light, and UV signals can play key roles in mate choice. Grouse (Tetraonidae) exhibit bright carotenoid-dependent sexual ornaments, their supra-orbital combs, which to humans appear orange-red. Combs also reflect in the UV, which is not visible to humans but is likely to be visible to grouse. In male red grouse Lagopus lagopus scoticus, we show that comb UV reflectance decreases with increasing comb size and redness. By removing the epidermis of combs, where carotenoid pigments are, we show that the UV reflectance is a property of the dermis, underneath the red pigmented epidermis. Carotenoid pigmentation of combs acted as a mask to reduce reflectance by the dermis in the range 400–550 nm and in the UV, 300–400 nm. Patagium skin (non-ornamental skin under the wing) also reflects in the UV, but epidermis removal on this bare part tended to reduce UV reflectance, whereas removal of the red epidermis of combs increased UV reflectance. Males in better condition (greater body mass relative to size) had bigger and redder combs, but with less UV. Thus, carotenoid pigments of grouse combs are deposited on a white background with significant UV reflectance, which can influence how the signal is perceived by conspecifics. Carotenoid-based traits exhibit UV reflectance in a number of species, but how UV reflectance and carotenoid pigmentation influence colour remains little known for integumentary ornaments compared to plumage traits. UV vision is not uncommon in birds and other animals, so future studies should investigate how UV reflectance influences the perception of carotenoid-based signals of quality.

Elevated spring testosterone increases parasite intensity in male red grouse

The expression of testosterone-dependent sexual traits might signal the ability of their bearers to cope with parasite infections. According to the immunocompetence handicap hypothesis (IHH), such signals would be honest because physiological costs of testosterone, such as a reduced ability to control parasite infections, would prevent cheating. We tested whether testosterone would affect the outcome of a standardized parasite challenge in red grouse, using a main parasite of the species, the nematode Trichostrongylus tenuis. We caught males in spring, removed their nematode parasites, and implanted them with testosterone or empty implants, as controls. After 1 month, they were reinfected with a standard dose of infective T. tenuis parasites. When challenged, testosterone males had relatively less globulin relative to albumin plasma proteins than control males, an indication that they had experienced increased physiological stress. Testosterone-treated males had significantly more T. tenuis parasites than controls in the next autumn and also had more coccidia and lost more weight than controls. Testosterone-treated males nevertheless benefited from their elevated spring testosterone: they had bigger sexual ornaments than controls both in spring and autumn, and they tended to have a higher pairing and breeding success than controls. Our results supported the IHH in showing that elevated testosterone impaired the ability of males to cope with a standardized challenge by a dominant parasite. Testosterone thus plays a key role in mediating trade-offs between reproductive activities and parasite defense, and testosterone-dependent comb size might honestly signal the ability of red grouse to control T. tenuis infection.

Interactions between population processes in a cyclic species: parasites reduce autumn territorial behaviour of male red grouse

The causes of population cycles fascinate and perplex ecologist. Most work have focused on single processes, whether extrinsic or intrinsic, more rarely on how different processes might interact to cause or mould the unstable population dynamics. In red grouse (Lagopus lagopus scoticus), two causal mechanisms have been supported: territorial behaviour (changes in autumn aggressiveness) and parasites (parasite induced reduction in fecundity). Here, we report on how these two regulatory processes might interact, by testing whether the parasite suspected to cause the grouse cycles, the nematode Trichostrongylus tenuis, reduces male autumn territorial behaviour. We either treated males with an anthelmintic, to remove parasites (dosed or D-males), or challenged them with infective T. tenuis larvae, to increase parasite intensity (challenged or C-males). We first show that dosing was effective in removing T. tenuis parasites, while parasite intensities increased in challenged birds during the autumn. Because old males initially had more parasites than young males, the treatments generated greater differences in parasite intensity in old than in young males. We also show that various aspects of territorial behaviour (increase in testosterone-dependent comb size in autumn, territorial call rate, likelihood of winning territorial interactions and over-winter survival) were significantly higher in dosed than in challenged males, but in old birds only. Our data thus supported the hypothesis that parasites reduce male aggressiveness during the autumn territorial contests, and could thereby influence recruitment. Our results also highlight that the territorial behaviour of young males, which have fewer parasites, is not as limited by parasites as that of old, previously territorial males. We discuss the implications of these findings for our understanding of the processes regulating red grouse populations and causing their complex, unstable population dynamics.

Testosterone and autumn territorial behavior in male red grouse Lagopus lagopus scoticus

In many bird species, males exhibit territorial aggression outside the breeding season, when testosterone concentrations are low and may not regulate territorial behaviors. The hormonal regulation of aggression at this time of year has only been studied in passerine birds. Here, we investigated the role of testosterone in the regulation of aggression in a non-passerine bird, the red grouse Lagopus lagopus scoticus. Male red grouse are aggressive in early spring when breeding starts, in autumn when they establish territories, and sporadically through much of the winter. We first describe seasonal variations in plasma testosterone concentrations and in the size of males' sexual ornaments, their red combs, which relates to aggressiveness. Testosterone concentrations and comb size were correlated. Both increased in autumn to a peak in October, and then increased again in spring, to a greater peak in early April. Secondly, we experimentally investigated the effects of testosterone, and of an anti-androgen (flutamide) used in combination with an aromatase inhibitor (ATD), on autumn territorial behavior. Males were treated with either empty implants, as controls (C-males), testosterone implants (T-males), or with flutamide and ATD implants (FA-males). One month after implanting, both T- and FA-males had higher concentrations of testosterone than C-males. Comb size, aggressive call rate, and response to playbacks of territorial call all significantly increased in T-males. However, the increase in testosterone in FA-males did not increase comb size or aggressive behavior. In the following spring, after the content of implants was used, FA-males had significantly lower testosterone than C-males, and had a reduced seasonal increase in comb size. The results suggest that testosterone plays a significant role in regulating red grouse aggressive behavior in autumn. However, the observation that flutamide and ATD treatment did not reduce territorial behavior, suggests that estradiol may also be involved in the regulation of non-breeding aggression.

Testosterone, immunocompetence, and honest sexual signaling in male red grouse

The expression of sexual ornaments has been suggested to reliably indicate individual quality, such as the ability to cope with parasites and diseases. The Immunocompetence Handicap Hypothesis (IHH) states that testosterone-dependent ornaments honestly signal such quality because of physiological costs associated with testosterone, such as impaired immune function. We tested predictions of the IHH both correlatively and experimentally in red grouse Lagopus lagopus scoticus. Male grouse exhibit supra-orbital red combs whose size is testosterone-dependent. We found that comb size was not correlated to infection intensity by two parasites (coccidia and the nematode Trichostrongylus tenuis), but it was significantly positively correlated with condition and T-cell-mediated immunity (the ability to mount a primary inflammatory response). We manipulated testosterone by means of implants and re-caught males after a month to investigate the effects on comb size, condition, immunity, and parasite load. Males implanted with testosterone had increased comb size, lost more condition, and had lower T-cell-mediated immunity than control males. Increased testosterone also resulted in a significant increase in coccidia infection intensity but had no effect on T. tenuis burden. The results are consistent with predictions of the IHH and suggest that comb size honestly indicates immunocompetence and males' ability to cope with certain parasites. Females could thus benefit from choosing mates based on the expression of this sexual trait.

Sexual ornamentation relates to immune function in male red grouse Lagopus lagopus scoticus

Sexual ornaments may enable females to discriminate amongst potential mates and choose those having either fewer parasites, or those which are more immunocompetent, or better able to cope with parasites. Tetraonid birds exhibit supra‐orbital combs that function in both intra‐sexual competition and mate choice. Through a correlative and experimental approach we investigated whether comb size of male red grouse Lagopus lagopus scoticus was related to infection intensity by their main parasite, the gastro intestinal nematode Trichostrongylus tenuis, and to immune function. We first looked at the relationships between immune function, parasite infection and condition. We found that spleen mass, an indirect measure of immune response, correlated positively with T. tenuis load, and negatively with condition. Cell‐mediated immunity, a measure of immune defence, correlated positively with condition, and although not significantly related to T. tenuis load, increased when nematode parasites were experimentally reduced. Secondly, we investigated whether comb size was related to condition, T. tenuis load or immune function. Comb size was not significantly related to T. tenuis infection and did not change significantly after nematode parasite removal. However, males with bigger combs were in better condition, had a lighter spleen, a lighter spleen than expected from their T. tenuis load and had greater cell‐mediated immunity. The findings suggest that comb size relates to immune function rather than T. tenuis parasite infection intensity. Males with bigger combs are likely to be of higher phenotypic quality because they are more immunocompetent and might be better able to cope with the detrimental effects of parasites.

A condition dependent link between testosterone and disease resistance in the house finch.

Testosterone has recently been proposed as a link between male quality and health and the expression of sexual traits. We investigated the relationship between testosterone and measures of the individual condition and health of males in a natural population of house finches (Carpodacus mexicanus). We also conducted a captive experiment in order to test for the effects of testosterone on resistance to coccidia, which is a common parasite of house finches. Free-living males in better condition had higher testosterone levels and lower corticosterone levels than free-living males in poor condition. In our captive experiment, increased testosterone accelerated the rate of coccidial infection as compared with sham-implanted or gonadectomized males. Although the differences were not significant, free-living males infected with coccidia had lower levels of testosterone and higher levels of corticosterone than males that were not infected. Thus, experimentally elevating testosterone levels in captive males resulted in a higher percentage of infected males, while free-living males with coccidial infection had low testosterone levels. This apparent discrepancy between captive and free-living males in the association of testosterone and disease may be explained by the condition dependence of testosterone. These results suggest that the testosterone-dependent sexual traits reliably indicate male overall condition and health and, thus, females could benefit from assessing potential mates based on these traits.

Spatial arrangement of kin affects recruitment success in young male red grouse

Models have shown that population cycles might be driven by time lags resulting from positive feedback between kin structure and population change, coupled with negative feedback between density and population change. One such model operates through kin favouritism facilitating the recruitment of young cock red grouse. We investigated whether recruitment by young cocks depended on the presence and spatial arrangement of elder relatives in the territorial population. We used molecular genetic estimates of relatedness, and checked for effects of covariates including natal territory size, hatching date, body size, parasite burdens and local density. Philopatric recruitment by cock red grouse led to the formation of clusters of contiguous territories owned by kin. The probability that an individual young cock would establish a territory increased with the number of kin in his father's cluster. This pattern might have been due to genetic quality determining both recruitment success and the size of the paternal cluster. If so, there should have been a positive correlation between a young cock's probability of recruitment and the number of his relatives in the population, irrespective of their spatial distribution. This did not occur and so the effect of cluster size is unlikely to have been confounded by genetic quality. The only morphological measure correlated with recruitment success was supraorbital comb size. The results are consistent with the prediction that kin tolerance affects recruitment but were at the level of the individual within years, rather than the population among years. Hence an experimental test of the kin favouritism hypothesis for population cycles, by manipulation of relatedness in populations among years, is now required.

Spatial distribution of genetic relatedness in a moorland population of red grouse (Lagopus lagopus scoticus)

Abstract Several aspects of the ecology and biology of red grouse ( Lagopus lagopus scoticus ) could prevent the complete admixture of genes within and between populations. Male red grouse display a high degree of natal philopatry, are territorial, and show less aggression to kin than to non-kin. Such factors acting in combination predict limited male-mediated gene flow, which will promote social structure within a population by the formation of stable kin clusters, and facilitate a rapid rise in allelic coancestry and/or inbreeding. In this study we utilize hypervariable microsatellite polymorphisms to examine the extent of social affiliation between relatives in a moorland population of grouse from NE Scotland. Levels of genetic relatedness between individual male red grouse occupying territories at Glas Choille in the spring and autumn of 1995 were examined, and kin clusters delimited. Nine kin groups (mean size =2.4 individuals) were identified prior to breeding in the spring, which increased to 11 kin groups (mean size =4.0 individuals) when territories were reformed in the autumn. The majority of those individuals that were recruited into the adult population during the autumn already had a first-order male relative established, supporting the hypothesis that recruitment is facilitated by behavioural interactions among relatives. The demographic and population genetic consequences of philopatric recruitment and kin clustering are examined and discussed.

Spatial distribution of genetic relatedness in a moorland population of red grouse (Lagopus lagopus scoticus)

Several aspects of the ecology and biology of red grouse (Lagopus lagopus scoticus) could prevent the complete admixture of genes within and between populations. Male red grouse display a high degree of natal philopatry, are territorial, and show less aggression to kin than to non-kin. Such factors acting in combination predict limited male-mediated gene flow, which will promote social structure within a population by the formation of stable kin clusters, and facilitate a rapid rise in allelic coancestry and/or inbreeding. In this study we utilize hypervariable microsatellite polymorphisms to examine the extent of social affiliation between relatives in a moorland population of grouse from NE Scotland. Levels of genetic relatedness between individual male red grouse occupying territories at Glas Choille in the spring and autumn of 1995 were examined, and kin clusters delimited. Nine kin groups (mean size =2.4 individuals) were identified prior to breeding in the spring, which increased to 11 kin groups (mean size =4.0 individuals) when territories were reformed in the autumn. The majority of those individuals that were recruited into the adult population during the autumn already had a first-order male relative established, supporting the hypothesis that recruitment is facilitated by behavioural interactions among relatives. The demographic and population genetic consequences of philopatric recruitment and kin clustering are examined and discussed.

A two-dimensional individual-based model of territorial behaviour: possible population consequences of kinship in red grouse

A two-dimensional simulation model of territory establishment and maintenance by individual male red grouse investigated whether differences in aggressive behaviour between kin and non-kin might cause population cycles or other demographic instability. Two behavioural phenotypes were envisaged. Intolerant cocks were equally aggressive to all neighbours, while kin-tolerant cocks were less aggressive to their close kin. The model's dynamics showed that: (i) pure populations of intolerants reached low but quite stable densities; whereas (ii) pure populations of kin-tolerant cocks attained higher densities with larger fluctuations; while (iii) populations of kin-tolerant cocks able to disperse showed even wider fluctuations. Mixed populations of intolerants and kin-tolerants co-existed at intermediate frequencies, with marked fluctuations. Thus kin-tolerant behaviour destabilised the model, giving it the potential to bifurcate to a cycle, but cycles were not observed. Two variants of the model altered grouse behaviour in response to changes in density, by changing either their kin tolerance or their number of dispersal attempts. Sensitivity tests showed these variants often produced density cycles, with periods around 6–15 years. We concluded that: differential aggressive behaviour between kin and non-kin during territorial disputes can destabilise a two-dimensional model population that has a life history similar to red grouse; dispersal may have a big effect on the stability of such model populations; some realistic variant models produced density cycles with periods and amplitudes comparable to those observed in some studies of wild red grouse.

Annual cycle of plasma concentrations of growth hormone in red grouse (Lagopus lagopus scoticus).

Abstract The circulating concentration of growth hormone (GH) was determined at monthly intervals in male and female red grouse maintained under ambient temperature and photoperiod. In both sexes the variations in plasma GH level showed distinct unimodal patterns. The highest concentrations of plasma GH were observed in spring and early summer (April, May, and June), while the lowest levels were found in autumn and winter (during November and January in females and December in males).

A nonparametric measure of intraclass correlation

A nonparametric measure of intraclass correlation based on the probability of certain types of concordances among the observations and which can be estimated from ranked data is proposed. One estimator whose variance can be estimated in an unbiased way is shown to be asymptotically normally distributed. Properties of the measure and its estimator are studied for a normal model; in particular the method is shown to provide a relatively powerful test of the null hypothesis of zero intraclass correlation in a normal population. The intraclass correlation coefficient is often used as a measure of the degree to which individuals from the same family resemble one another in some variable such as height or weight. From a random sample of families and a random selection of individuals from each family, both the between and within family components of variance can be estimated and an estimate of the intraclass correlation obtained. When the observations are ranks this approach is not feasible since direct estimates of the components of variance are not available. An example occurs in a study of aggressiveness in male red grouse (Moss, Watson & Parr, 1974). One measure of aggressiveness of an individual bird is a dominance rank, the position of that bird in the dominance hierarchy of a given group of birds. Comparison of the degree of familial resemblance in different populations is complicated by the fact that the data are ranks and that dominance hierarchies can be variable in size with different family sizes. The development of standard test statistics, such as the Kruskal-Wallis statistic (Kruskal, 1952), as measures of intraclass correlation is unsatisfactory because of the complicated dependence on the structure of the sample; see ? 6. In this paper a measure of intraclass correlation is proposed which overcomes this problem. It is independent of the structure of the sample and it can be estimated in an unbiased way from ranked data. The interclass rank correlation coefficients, Kendall's Xr and Spearman's p (Kendall, 1955), can be interpreted in terms of probabilities of concordances among the observations from a bivariate distribution. In response to Hill (1974), Kerridge (1975) has used this probabilistic interpretation of r in an analysis of Football League results. The measure presented in the present paper is based on concordances among observations from a population in which individuals occur in families. Its properties for a general class of populations are discussed and some particular results for a one-way classification model with normally distributed effects are presented. The power of the associated test is assessed for a normal model for a range of sample structures.