Non-adaptive adoptions of nestlings in the colonial lesser kestrel: proximate causes and fitness consequences

We examined the causes, costs and benefits of adoption in the altricial lesser kestrel Falco naumanni. Specifically, we tested the intergenerational conflict hypothesis, proposed to explain adoption in some birds. Adoptions involved 76% of the nests and 51% of the nestlings at a mean age of 25 days (12 days before fledging). Nest-switching nestlings were not in poorer body condition, more parasitized or younger than their siblings, and body condition and prey delivery rates of their parents did not differ from those of other parents. In the foster nest, adopted nestlings did not benefit from higher feeding rates or a prolongation of the nestling period. They did not have fewer nest-mates or achieve higher rank within the new brood. Thus, adopted nestlings did not improve their body condition and survival. Adult lesser kestrels seemed unable to finely discriminate beween their own and alien chicks. Foster parents bore the cost of an increase of prey delivery rates, although it did not affect their survival or subsequent reproductive performance. Therefore, our results do not support the intergenerational conflict hypothesis, and suggest that adoption in this species is non-adaptive. Traditionally, the lesser kestrel bred in cliffs where movement among nest-sites was restricted. Nowadays, about half of the colonies are in tiled roofs which facilitate nest-switching by nestlings. The high rate of adoptions may thus be explained as reproductive errors associated with the recent occupation of a new breeding habitat. Received: 3 May 1996 / Accepted after revision: 19 January 1997     

Food access, brood size and filial cannibalism in the fantail darter, Etheostoma flabellare

We compared the occurrence of filial cannibalism in fed and starved male fantail darters (Etheostoma flabellare). All males in the experiment consumed eggs, and 56% ate all of their eggs. A male's initial body condition did not explain the number of eggs that he ate. Neither did non-fed males eat more eggs than fed males. Fed males were able to maintain better body condition during the experiment, but the change in body condition also depended on the number of eggs eaten. Thus, males who ate more eggs were able to maintain better body condition.The most important determinant of whether or not a male ate all of his eggs was his initial egg number. Males with small egg masses ate all of their eggs whereas males with large egg masses were only partial cannibals. There was, however, no difference in the total number of eggs eaten by total and partial cannibals. We conclude that eggs are only partially eaten for energetic reasons. We also suggest that small egg masses are completely consumed because the costs of caring for a small egg mass may exceed the expected reproductive benefits of a small egg mass. Received: 26 January 1996 / Accepted after revision: 2 November 1996     

Male-biased maternal expenditure and associated costs in fallow deer

This study tested whether fallow deer mothers, Dama dama, bias their investment towards sons and, thus, whether sons are more costly to produce than daughters. Young (2 years) and old (≥3 years) hinds were analysed separately. Old hinds who raised sons accumulated less body mass than those who raised daughters, during the period between late gestation and the end of lactation. This difference in body mass persisted to the following spring. Mothers who had raised sons gave birth later and their offspring's pre-winter mass was lower the following year than for mothers who had raised daughters. These results indicate higher expenditure for hinds who raise sons and support theories of male-biased maternal investment. However, young mothers with sons and those with daughters did not differ in reproductive performance the following year. One reason might be that young mothers are close to the maximum level of maternal expenditure, since they are still growing, and cannot invest any extra resources in sons. Received: 28 August 1997 / Accepted after revision: 5 April 1998     

Testis size variation in frogs: testing the alternatives

While sperm competition may be a major factor affecting relative testis size in vertebrates as a whole, additional hypotheses have not been given much attention in frogs. This is important because sperm competition is relatively uncommon in frogs and relative testis size varies in frogs that do not have multiple-male breeding systems. This paper tests two additional hypotheses for differences in relative testis size among frogs: relative clutch size (number of eggs/snout vent length) and androgen level. Testis size was measured in 90 species of frogs belonging to five families. Relative testis size was found to be positively correlated with relative clutch size in species that lack sperm competition. Mean androgen levels of species also positively covaried with relative testis size. However, there was no correlation between relative testis size and level of male agonistic behavior among species, despite other work indicating that testosterone levels are positively correlated with agonistic behavior in at least some species. These findings suggest that a number of factors in addition to sperm competition are important in the evolution of testis size in male frogs. Received: 17 January 1997 / Accepted after revision: 23 June 1997     

Variation among populations of the sailfin molly in the rate of concurrent multiple paternity and its implications for mating-system evolution

We examined patterns of concurrent multiple mating in a live-bearing poeciliid fish, the sailfin molly (Poecilia latipinna). We tested whether the probability of multiple paternity was related to female body size or fertility and whether the rate of multiple paternity varied among four populations that differed in their distributions of female body size and fertility. We analyzed data on mother and offspring genotypes for three polymorphic allozymes by three techniques, including a maximum-likelihood estimator that accounts for sampling error in both parental and offspring allele frequencies. The estimated rate of multiple paternity varied between 0.09 and 0.85, and the rate in one population varied seasonally between 0.33 (spring) and 0.85 (autumn). The variation in these rates was not associated with variation in body-size distributions among populations but was closely associated with variation in size-specific fertility: populations with greater variation in female fertility had higher multiple-paternity rates. Within two populations, logistic regression revealed that individual females of larger body size and greater size-specific fertility were more likely to carry multiply sired broods. This result is consistent with observations made in one of the populations 5 years earlier. In general, the results strongly suggest that the mating system varies markedly among conspecific populations of sailfin mollies and that larger, more fertile females are the objects of intermale competition. Received: 6 May 1996 / Accepted in revised form: 5 December 1996     

Brood sex ratio is dependent on female mating status in polygynous great reed warblers

Females capable of adjusting the sex ratio of their offspring should be more fit than females lacking such an ability. In polygynous birds where breeding success in males is more strongly influenced by body size and/or attractiveness than in females, females might produce more sons when predicting good conditions or when mating with attractive males. Polygynous great reed warbler, Acrocephalusarundinaceus, males direct most of their feeding effort to the primary (first-hatching) nest and in these nests increase their feeding effort in relation to the brood sex ratio (proportion of sons). Therefore, with the expectation of well-nourished sons, we would predict that females which start breeding first within harems might produce more sons than those which start breeding later, and in anticipation of sons with good genes, that females mated to polygynous males might produce more sons than females mated to monogamous males. I took blood samples from hatchlings and determined the sex using DNA markers. The sex ratio of primary (monogamous and polygynous primary) broods is more male-biased (mean 0.58 males, n = 50) than that of secondary (polygynous secondary and tertiary) broods (mean 0.46, n = 25). Moreover, in the secondary broods with the largest clutch (five eggs), in which offspring are most likely to suffer food shortage, the sex ratio was distinctively female biased (mean 0.33, n = 10). In the primary broods, sex ratio was correlated to harem size. The results suggest that great reed warbler females modify the brood sex ratio to produce both well-nourished sons and sons with good genes, but the former effect is probably stronger than the latter factor. Received: 11 March 1998 / Accepted after revision: 23 May 1998     

Disease resistance: a benefit of sociality in the dampwood termite Zootermopsis angusticollis (Isoptera: Termopsidae)

The benefit of sociality in relation to disease susceptibility was studied in the dampwood termite Zootermopsis angusticollis. Although contact with high concentrations of fungal conidia is lethal, the survivorship of nymphs exposed to spore suspensions ranging from 6 × 10⁶ to 2 × 10⁸ spores/ml of the fungus Metarhizium anisopliae increased with group size. The survivorship (measured as LT₅₀) of isolated individuals ranged from 3.0 to 4.8 days, but infected nymphs living in groups of 10 and 25 individuals survived significantly longer (5.6–8.3 and 5.6–9.1 days, respectively). In most cases, there were no significant differences in the survival distributions of the 10- and 25-termite groups. When nymphs were infected with concentrations of 7 × 10¹–7 × 10⁴ spores/ml and allowed to interact with healthy nestmates, fungal infections were not contracted by the unexposed termites. Moreover, infected termites benefitted from social contact with unexposed nestmates: their survival rates were significantly higher than those of infected termites living with similarly infected nestmates. Allogrooming, which increased in frequency during and after exposure to conidia, appeared to remove potentially infectious spores from the cuticle, thus increasing termite survivorship. These results suggest that allogrooming plays a crucial role in the control of disease and its death hazard in termites. The infection-reducing advantage of group living may have been significant in the evolution of social behavior in the Isoptera. Received: 18 March 1998 / Accepted after revision: 31 May 1998     

Parentage and paternal care: consequences of intersexual selection in Savannah sparrows?

Empirical relationships between parentage and male parental care are commonly interpreted in the context of life-history models that consider increased offspring survivorship as the only benefit of paternal effort. However, indirect benefits associated with male care can also influence a male's response to cuckoldry: if females allocate paternity according to their prior experience with male parental care, it may pay for males to provision extra-pair young in early broods. Here, I assess the relationship between first-brood parentage and paternal care in a population of Savannah sparrows (Passerculussandwichensis) where a male's fertilization success in the second brood appears to be influenced by his prior parental performance. Based on the multi-locus DNA fingerprinting of 17 first broods, male feeding effort was influenced by parentage (percent of brood resulting from within-pair fertilizations) but not by brood size, male mating status (monogamous versus polygynous), timing of breeding (hatching date), structural size (wing length) or condition (mass). Males provided more care to broods that contained few within-pair young. This result supports the idea that males provision young to increase their future mating success, but alternative hypotheses involving male quality and timing of breeding cannot be excluded. Received: 13 August 1996 / Accepted after revision: 22 February 1997     

Body mass and parental decisions in the Antarctic petrel Thalassoica antarctica : how long should the parents guard the chick?

In Procellariiformes, the parents guard the chick after it has attained homeothermy. This strategy may reduce the probability that a small chick is taken by predators, but is costly as only one parent can forage at a time. The decision to leave the chick may therefore be a compromise between the chick's vulnerability to predators, the body condition of the parent on the nest and whether the foraging parent returns in time. We studied how the number of days that parents guarded the chick was related to the body mass of the parent at the nest and the time the foraging parent spent at sea in the Antarctic petrel Thalassoica antarctica. We also examined how the body mass of the parent on the nest and the duration of the foraging trips influenced the chicks' body condition at the end of the guarding period. When the foraging parent did not return to the nest in time to relieve its mate, the number of days the parent on the nest kept guarding the chick was positively related to its body mass on arrival in the colony. The number of days the foraging parent spent at sea was positively related to the body mass of its mate, but those that returned in time had a shorter stay at sea relative to their mate's body mass than those that did not return before their mate had left. Apparently, both the body mass of the parent at the nest and the ability of the foraging parent to adjust its stay at sea to the mate's body mass is important for the number of days the parents guard the chick and also the chick's body condition at this point. The inability to return to the nest before the mate has left may be the result of needing a minimum amount of time at sea to find food, or because some parents having low foraging success and therefore prolong their stay at sea. Received: 10 October 1997 / Accepted after revision: 14 March 1998     

Antler asymmetry and immunity in reindeer

Fluctuating asymmetry (FA) measures an individual's ability to undergo identical development in bilaterally symmetrical characters and may indicate sensitivity to environmental stress. FA in ornamental characters is often positively related to parasite intensities, which are important environmental stressors. Parasites affect and are affected by several parts of the immune system, and the ability to resist parasites may be signalled via FA in ornaments. In this study we examined reindeer antlers, which show FA, demonstrated to be caused by parasite infections. We measured antler FA, immune parameters (i.e. densities of different classes of leukocytes, IgG levels and abomasal lymph node numbers) and intensity of abomasal nematodes in free-ranging 1.5-year-old male reindeer slaughtered in the early part of their rutting period. We found a relationship between parasite intensity and immune parameters suggesting that our measures of immune activity reflect density of current parasite infections. More important, these immune parameters were associated with FA in both the main beam length and numbers of antler tines. The immune parameters were, however, only weakly correlated to antler size. This indicates that FA, but not size, of antlers grown during exposure to a multitude of environmental stressors may reveal information about individual immunity that can be important for host-parasite interactions. Antler FA may therefore communicate an individual's quality during the rut in reindeer. Received: 30 March 1998 / Accepted after revision: 15 August 1998     

Different antipredator behaviour in two anuran tadpoles: effects of predator diet

Recent investigations have indicated that animals are able to use chemical cues of predators to assess the magnitude of predation risk. One possible source of such cues is predator diet. Chemical cues may also be important in the development of antipredator behaviour, especially in animals that possess chemical alarm substances. Tadpoles of the common toad (Bufo bufo) are unpalatable to most vertebrate predators and have an alarm substance. Tadpoles of the common frog (Rana temporaria) lack both these characters. We experimentally studied how predator diet, previous experience of predators and body size affect antipredator behaviour in these two tadpole species. Late-instar larvae of the dragonfly Aeshna juncea were used as predators. The dragonfly larvae were fed a diet exclusively of insects, R. temporaria tadpoles or B. bufo tadpoles. R. temporaria tadpoles modified their behaviour according to the perceived predation risk. Depending on predator diet, the tadpoles responded with weak antipredatory behaviour (triggered by insect-fed predators) or strong behaviour (triggered by tadpole-fed predators) with distinct spatial avoidance and lowered activity level. The behaviour of B. bufo in predator diet treatments was indistinguishable from that in the control treatment. This lack of antipredator behaviour is probably related to the effective post-encounter defenses and more intense competitive regime experienced by B. bufo. The behaviour of both tadpole species was dependent on body size, but this was not related to predator treatments. Our results also indicate that antipredator behaviour is largely innate in tadpoles of both species and is not modified by a brief exposure to predators. Received: 22 August 1996 / Accepted after revision: 31 January 1997     

Oligogyny by unrelated queens in the carpenter ant, Camponotus ligniperdus

Multilocus DNA fingerprinting and microsatellite analysis were used to determine the number of queens and their mating frequencies in colonies of the carpenter ant, Camponotus ligniperdus (Hymenoptera: Formicidae). Only 1 of 61 analyzed queens was found to be double-mated and the population-wide effective mating frequency was therefore 1.02. In the studied population, 8 of 21 mature field colonies (38%) contained worker, male, or virgin queen genotypes which were not compatible with presumed monogyny and therefore suggested oligogyny, i.e., the cooccurrence of several mutually intolerant queens within one colony. Estimated queen numbers in oligogynous colonies ranged between two and five. According to the results of the genetic analysis, most of the queens coexisting in oligogynous colonies were not closely related. Pleometrosis is very rare and queenless colonies adopt mated queens both in the laboratory and field. Therefore, the most plausible explanation for the origin of oligogynous colonies in C. ligniperdus is the adoption of unrelated queens by orphaned mature colonies. The coexistence of unrelated, but mutually intolerant queens in C. ligniperdus colonies demonstrates that oligogyny should be considered as a phenomenon distinct from polygyny. Received: 18 December 1997 / Accepted after revision: 20 June 1998     

Changes in male guppy courting distance in response to a fluctuating light environment

In the guppy (Poecilia reticulata), effective courting by a male requires visual contact with the female. Therefore, environmental light intensity may affect male display behavior, particularly initial courtship distance. We found that male guppies courted at exact and predictable distances from the female given a particular light level, both in field and laboratory studies. In lower light levels (<0.1 μmol m⁻² s⁻¹), for example at dawn, dusk, or under heavy canopy, males court females at closer and less variable distances (<3 cm). At higher light levels, which occur during most of the day and with less canopy cover, males often court from twice or three times further out. Light levels over guppy streams change over relatively short time periods and ranges, correlating with variation in courtship distances. Laboratory manipulations of irradiance confirmed that courtship distance depends on illumination. Hence, courtship distances may be set by the effect of lighting on signal efficiency, minimization of energy or time expenditures, or predation risk. Received: 16 December 1997 / Accepted after revision: 8 August 1998     

Estimating paternity with spatial behaviour and DNA fingerprinting in the striped plateau lizard, Sceloporus virgatus (Phrynosomatidae)

The association between spatial proximity and paternity was studied in a population of the striped plateau lizard, Sceloporus virgatus. The relationship between estimated mating success and male phenotypic traits was examined for a sample of 55 males. DNA samples were obtained from 13 female-offspring families. The males with the closest spatial proximity to each female were tested as possible sires within each family. Fingerprinting with two multilocus hypervariable minisatellite probes revealed a strong correspondence between male-female spatial proximity and actual paternity. Paternity could be assigned for 72 of the 100 hatchlings. Most hatchlings with identifiable sires were attributed to a male with the highest category of spatial proximity to the mother. However, there was a low to moderate level of multiple paternity within clutches, and for some clutches probable sires could not be identified even though the most likely behavioural candidates were tested. Thus, nonterritorial males or other males lacking strong social and spatial relationships with females may achieve some degree of reproductive success. Analysis of mating success revealed that male success increased with body size, up to a point beyond which larger size conferred no advantage. Received: 7 January 1997 / Accepted after revision: 16 June 1997     

A same-sex stepparent shortens a prebreeder's duration on the natal territory: tests of two hypotheses in Florida scrub-jays

Prebreeders of the Florida scrub-jay (Aphelocoma coerulescens) are less likely to be found on their natal territories with a same-sex stepparent than with parents or an opposite-sex stepparent. We tested two models that had been proposed to account for this sexual asymmetry. The dominance hypothesis states that stepparents perceive same-sex prebreeders as competitors, primarily for a mate, so behave aggressively toward them. The pair-formation hypothesis states that prebreeders remain home to pair eventually with the opposite-sex stepparent. Predictions from these two models were tested by analyses from a quarter-century of records on a study population at Archbold Biological Station and by new behavioral field observations at the Station. Results clearly rejected the latter and strongly supported the former hypothesis. No prediction from the pair-formation hypothesis was confirmed: no pairing by a prebreeder of either sex has ever occurred with a stepparent; remaining home was equally frequent in age-one males with stepmothers and genetic mothers, and in females with stepfathers and genetic fathers; and the same results were found in age-two prebreeders. By contrast, the data strongly supported the dominance hypothesis. Field data showed higher aggression rates by stepfathers to male prebreeders and stepmothers to female prebreeders than by parents to their same-sex genetic offspring. Fewer age-one males remained home with a stepfather than with the genetic father, and the same was found for age-one females with a stepmother and genetic mother; at age two, the effect occurred only in female prebreeders. The two hypotheses make different predictions about prebreeders with two stepparents versus with both parents, thus providing a critical test. The dominance hypothesis correctly predicted decreased duration at home by age-one males and females; data for age-two females were in the direction of predicted difference but not significant; and, as in other tests, no effect was found for age-two males. We propose that age-two male prebreeders remain home despite elevated aggression from stepfathers because these prebreeders retain the possibility of budding from or inheriting the natal territory. Received: 16 July 1997 / Accepted after revision: 26 June 1998     

Maternal investment in Antarctic fur seals: evidence for equality in the sexes?

Studies of the otariids (fur seals and sea lions), a highly sexually dimorphic group, have provided conflicting evidence of differential maternal expenditure in male and female offspring and, thus, suggestions that they conform to predictions of investment theory are equivocal. Since the mid-1970s, a diversity of research on Antarctic fur seals (Arctocephalus gazella) including studies of their reproductive ecology, lactation energetics, and foraging behaviour have been conducted at Bird Island, South Georgia that have resulted in one of the more complete and diverse data sets for any species of otariid. These long-term data were reviewed to determine whether there was any evidence to support that differential maternal expenditure occurred in Antarctic fur seals. Most of the data examined were collected during five consecutive austral summers from 1988 through 1992 and included years in which local food resources were abundant and scarce. We were unable to detect differences in the sex ratios of pups at birth or sex-biased differences in growth rates estimated from serial data, the number of foraging trips made, the duration of attendance ashore, diving behaviour, suckling behaviour, or milk consumption in any year and in the duration of foraging trips or age at weaning in 2 of 3 years. In addition, we found no evidence of greater reproductive costs between mothers with sons or daughters relative to their reproductive performance the following year. In contrast, sex-biased differences were only found in the duration of foraging trips in 1990, the age at weaning in 1988, and consistently in growth rates estimated from cross-sectional data. We suggest that differential maternal expenditure does not occur in Antarctic fur seals because male pups probably do not gain greater benefit from additional maternal expenditure than female pups. After weaning, males experience a period of rapid juvenile growth over 3–4 years during which time body mass nearly trebles. This growth will almost certainly be dependent upon available food resources then rather than on any maternal expenditure received over the first 4 months of life and, thus, the assumptions of the Trivers and Willard hypothesis are probably invalid for Antarctic fur seals. Received: 10 July 1996 / Accepted after revision: 3 March 1997     

Sex linkage among genes controlling sexually selected traits

Using literature data on reciprocal crosses I estimated the influence of sex-chromosomal genes on morphological and behavioral traits. To determine a special role of the sex chromosomes for sexually selected traits, I compared the estimated influence of X-chromosomal genes on sexually selected traits with their estimated influence on traits not under sexual selection. About one-third of the phenotypic variation in sexually selected traits is caused by X-chromosomal genes. There was, in contrast, no significant influence of X-chromosomal genes on traits that were classified as not sexually selected. Sexually selected traits thus seem to be influenced significantly more by X-chromosomal genes than traits not under sexual selection. Though this differential influence of X-chromosomal genes cannot readily be explained with current theoretical knowledge, it may shed some light on X-linked hybrid sterility and on the discussion between arbitrary and good-gene models for the evolution of female choice. Received: 30 January 1998 / Accepted after revision: 20 June 1998     

Past reproductive success affects future habitat selection

Correlational studies have shown that an individual's past reproductive success often increases its breeding site fidelity (i.e., the tendency to return to a previously occupied location), suggesting that individuals use their reproductive experience to assess habitat quality. However, the causality of the relationship between reproductive success and site fidelity is still uncertain. In a field experiment, the effect of mating success on site fidelity was isolated from potential confounding variables in a territorial dragonfly, the eastern amberwing (Perithemis tenera). The experiment controlled for site quality, intrinsic characteristics of males, previous territorial experience at the site, arrival order, and territorial evictions. Males that were prevented from mating were much more likely to change sites the following day than control males that were allowed to mate. This result was not affected by age, the amount of time a male spent on the site, or mortality. These results imply that individuals use their own reproductive success to assess the quality of the habitat. The benefit to an individual of using its reproductive success to determine habitat quality is discussed relative to other sources of information. Received: 31 May 1996 / Accepted: 31 January 1997     

Parentage and sex allocation in the facultatively polygynous ant Myrmicatahoensis

Most social groups have the potential for reproductive conflict among group members. Within insect societies, reproduction can be divided among multiple fertile individuals, leading to potential conflicts between these individuals over the parentage of sexual offspring. Colonies of the facultatively polygynous ant Myrmicatahoensis contain from one to several mated queens. In this species, female sexuals were produced almost exclusively by one queen. The parentage of male sexuals was more complex. In accordance with predictions based on worker sex-allocation preferences, male-producing colonies tended to have low levels of genetic relatedness (i.e., high queen numbers). Correspondingly, males were often reared from the eggs of two or more queens in the nest. Further, over half of the males produced appeared to be the progeny of fertile workers, not of queens. Overall investment ratios were substantially more male biased than those predicted by genetic relatedness, suggesting hidden costs associated with the production of female sexuals. These costs are likely to include local resource competition among females, most notably when these individuals are adopted by their maternal nest. Received: 3 March 1998 / Accepted after revision: 20 June 1998