Social and mating system of cooperatively breeding laughing kookaburras (Dacelo novaeguineae)

DNA fingerprinting was combined with field observations over four breeding seasons to investigate the social structure and mating system of the laughing kookaburra (Dacelo novaeguineae). Groups comprised a socially dominant pair and up to six helpers of either sex. Helpers were always recruited from young hatched in the group. Territorial inheritance, which is a feature of other cooperative breeders and an oft-cited benefit of philopatry, did not occur. Helpers only attained dominant status in an established group by dispersing into a vacant dominant position in that group. However, helpers could also form new groups by excising a new territory, often through a ”budding” process. The mating system was overwhelmingly monogamous. There were no cases of extra-group parentage in a sample of 140 nestlings; within groups of three or more birds, dominance predicted parentage almost perfectly (99.2% of 129 nestlings), irrespective of whether helpers in the group were related to one or both dominant birds. This is contrary to predictions from models of reproductive skew, possibly because they currently fail to incorporate the willingness of females to share reproduction among males. Received: 15 May 1999 / Received in revised form: 2 November 1999 / Accepted: 6 November 1999     

Ectoparasites of house sparrows (Passer domesticus): an experimental test of the immunocompetence handicap hypothesis and a new model

Conspicuous secondary sexual traits may have evolved as handicap-revealing signals or as badges of status. We present results of an experiment using males of the sexually dimorphic house sparrow (Passer domesticus), that support the idea that the male-specific bib can be both a handicap-revealing signal and a reliable badge indicating the physical condition of the bird. In a test of the immunocompetence handicap hypothesis, wild-caught adult male house sparrows were studied in captivity. Birds implanted with elevated doses of testosterone were more dominant, had higher circulating levels of both testosterone and corticosterone and they also harboured relatively larger ectoparasite loads. Higher parasite loads were also associated with individuals showing lower immunocompetence and larger changes in bib size. A new model for immunocompetence effects in sexual selection is introduced, integrating actions that the hypothalamopituitary axis exerts on gonads, adrenals and the thyroid gland. The ”integrated immunocompetence model” synthesizes both the ”handicap” (i.e. survival-decreasing) and ”badge of status” (i.e. survival- enhancing) models for evolution of secondary sexual traits. Received: 15 May 1999 / Received in revised form: 2 November 1999 / Accepted: 6 November 1999     

Sexual selection on mature male parr of masu salmon (Oncorhynchus masou): does sneaking behavior favor small body size and less-developed sexual characters?

In many salmonid species, males exhibit morphological dimorphism associated with alternative mating behaviors. ”Precocious males” have a small body size with little or no development of sexual characters and adopt sneaking to gain access to females, while ”migratory males” of large body size and well-developed secondary sexual characters fight. We quantified selection on precocious male parr of masu salmon (Oncorhynchus masou) under simulated natural conditions to examine the contribution of morphology to sneaking success. In contrast to the prediction that sneaking behavior favors small body size, we detected selection favoring relatively large body size for sneaking success. This selection pressure was caused by the dominance hierarchy within parr and may have been facilitated by indifference of dominant migratory males to parr. Unlike the secondary sexual characters exhibited by migratory male salmon, such as the hooked snout and humped back, no morphological characters other than body size contributed to the reproductive success of masu salmon parr. This non-contribution may have been responsible for the lack of development of sexual characters in precocious males. Received: 15 November 1999 / Accepted: 20 May 2000     

The cost of host egg damage caused by a brood parasite: experiments on great spotted cuckoos (Clamator glandarius) and magpies (Pica pica)

Adult great spotted cuckoos, Clamator glandarius, frequently damage one or more eggs of their magpie host, Pica pica, without removing or eating them. The presence of damaged host eggs could signal parasitism thereby increasing the probability that the parasitic egg is ejected. This hypothesis was tested by experimentally introducing a model cuckoo egg with or without damaged host eggs. Magpie responses to experimental parasitism did not differ significantly between treatments implying that damaged host eggs are not used by magpies to assess parasitism. We followed the fate of magpie eggs naturally damaged by the great spotted cuckoo or experimentally damaged by us. Host response was very similar for naturally or experimentally damaged host eggs, but varied significantly according to the type of egg damage, eggs being removed more frequently when pecked than crushed, while cracked eggs were never removed. However, the egg damage that most readily causes egg removal is albumen leakage. Received: 30 November 1998 / Received in revised form: 7 June 1999 / Accepted: 12 June 1999     

Duration of paternal care in pine engraver beetles: why do larger males care less?

Male pine engravers, Ips pini (Coleoptera: Scolytidae), assist their mates during reproduction by removing the debris that accumulates while females excavate oviposition tunnels in the phloem tissue of host tree bark. Although duration of paternal care and male reproductive success are positively correlated, large males leave their mates and brood sooner than small males. We address two hypotheses to explain the earlier departure of larger males from their breeding galleries: (1) females oviposit most rapidly when paired with large males, thereby reducing the length of time that paternal care increases male reproductive success, (2) larger males have better prospects for future reproduction, and thus leave their galleries in search of new breeding opportunities sooner than smaller males. Contrary to the first hypothesis, when females were paired either with large or small males, there was no effect of male size on their rate of oviposition. Consistent with the second hypothesis, males that initiated breeding galleries were larger than males from the general population. In addition, large males flew farther and faster on flight mills than small males, which may indicate that large males have an advantage in locating suitable breeding sites. Thus, we suggest that large male pine engravers leave their galleries earlier than small males because large individuals have better prospects for future reproduction. Received: 30 November 1997 / Accepted after revision: 23 May 1998     

Effects of territory quality, food availability and sibling competition on the fledging success of oystercatchers (Haematopus ostralegus)

We investigated the fledging probability of oystercatcher, Haematopus ostralegus, chicks as a function of hatching order, brood size, territory quality and food availability. Sibling dominance was related to the hatching order in both low- (’leapfrogs’) and high-quality (’residents’) territories. Differences in hatchling mass might have aided the establishment of a dominance hierarchy, since breeders produced small late eggs and hatchlings. These mass differences were most pronounced in leapfrogs, and in large broods in years with lower food availability (’poor’ years). Late hatchlings fledged less often and with lower body masses compared to early hatchlings in all situations. Leapfrogs produced smaller broods and hatched their broods more asynchronously in poor years than leapfrogs breeding in years with more available food (’good’ years) and residents breeding in both poor and good years. Large brood sizes resulted in lower survival of hatchlings in poor years. These results favour the ’brood reduction’ hypothesis. However, contrary to the expectations of this hypothesis, hatching order also affected fledging success in residents. Moreover, large brood size resulted in higher survival of hatchlings in good years, particularly in residents. Thus, although large broods experienced losses due to sibling competition in some years, they nevertheless consistently produced more fledglings per brood in all years, both as leapfrogs and residents. We believe this effect is due to parental quality correlating with initial brood size. Most leapfrogs, at best, fledged one chick successfully each year, losing chicks due to starvation. Nevertheless, leapfrog broods were reduced in size after hatching significantly less quickly than resident broods. These results suggest that breeders lay and hatch insurance eggs to compensate for unpredictable losses due to the high predation rates on both nests (ca 50%) and chicks (ca 90%), in accordance with the ’nest failure’ hypothesis. Received: 14 February 2000 / Revised: 27 September 2000 / Accepted: 10 June 2000     

Frequency and timing of extrapair fertilisation in the polyandrous red phalarope (Phalaropus fulicarius)

In sequentially polyandrous bird species, where females mate with different males in succession during a single breeding season, sperm stored by females can occasionally lead to extrapair fertilisations (EPFs) in clutches cared for by the second and subsequent males. Thus, we predicted that in red phalaropes (Phalaropus fulicarius) – a sex-role-reversed, sequentially polyandrous, arctic breeding shorebird – EPFs would occur more frequently in clutches laid later in the breeding season. We used multilocus DNA profiling to examine the frequency and timing of EPFs in a population of red phalaropes breeding in the Canadian high arctic. Using a technique to determine parentage without maternal DNA, we inferred that 6 of 70 chicks in 18 broods resulted from EPFs – one extrapair chick in each of 6 broods. These results were supported by a further analysis using microsatellite DNA. As predicted, broods containing EPFs hatched from clutches laid significantly later in the season than did broods containing no EPFs. The difference in median hatch dates between broods with and without EPFs was 9.0 days, or 38% of the entire egg-laying period in that season. For the whole breeding season, we estimated that 6.5% of chicks were sired by extrapair males, which is similar to extrapair paternity estimates for other sex-role-reversed birds, but relatively low compared to the majority of socially monogamous species studied so far. Received: 24 August 1998 / Received in revised form: 18 January 1999 / Accepted: 24 January 1999     

Low paternity in the hornet Vespa crabro indicates that multiple mating by queens is derived in vespine wasps

Queen mating frequency was studied in the European hornet, Vespa crabro, by analyzing four DNA microsatellite loci in 20 workers from each of 14 nests. Queens were found to be predominantly singly mated (9/14), although double (4/14) and triple mating (1/14) also occurred. For most multiply mated queens, paternity was significantly biased with the majority male fathering on average 80% of the female offspring. The population-wide effective mating frequency was therefore low (1.11), and sister-sister relatedness high (0.701 ± 0.023 SE). Low effective mating frequency in Vespa, in combination with data from other vespines, suggests that high paternity frequency is derived in the group. Some problems with the non-detection of fathers, where the queen was not sampled or shared alleles with males, are analyzed. Received: 16 November 1998 / Received in revised form: 29 March 1999 / Accepted: 12 April 1999     

The effect of queen-worker conflict on caste determination in the bumblebee Bombus terrestris

Endocrine analyses were used to investigate the well-known association between queen production and the onset of worker reproduction (termed the competition phase, CPh) in Bombus terrestris. Larvae that reached the age of 5 days before the CPh had a worker-like profile: low juvenile hormone (JH) biosynthesis rates and low JH hemolymph titers. In contrast, larvae that reached the age of 5 days during the CPh had a queen-like profile: high JH biosynthesis rates and high hemolymph JH levels. Larval fate could be manipulated by transplanting egg cells into host colonies with different social structures. There was a steep rise in JH production in larvae transplanted into colonies near or during the CPh. This indicates that during colony development, larvae switch from the ”worker developmental pathway” to the ”queen developmental pathway,” and that the switch is socially regulated. In small rearing groups, larvae reared with queens before the CPh developed into workers, whereas those reared with queens after the CPh developed into queens. Variation in worker type (naive or experienced) did not affect caste determination. Therefore, we hypothesize that queens produce a pheromone that directly inhibits queen differentiation by larvae. We also present two alternative scenarios that explain the timing of gyne production in B. terrestris, one based on ecological constraints and the other based on queen-worker competition. Received: 20 October 1999 / Received in revised form: 18 December 1999 / Accepted: 23 January 2000     

Breeding synchrony and extra-pair mating in red-winged blackbirds

Using data from a 6-year paternity study of red-winged blackbirds, I tested the hypotheses that increased nesting synchrony should either promote extra-pair mating by increasing the advantage of extra-pair mating to females, or decrease extra-pair mating by constraining males from seeking extra-pair copulations. Contrary to these hypotheses, the occurrence of extra-pair paternity did not vary with nesting synchrony over the breeding season, or vary with the number of synchronous nests within territories or within marshes, or with nesting order on territories. However, for nearly all nests with extra-pair young, there were fewer females synchronous with that nest on the cuckolder's territory than on the territory of the cuckolded male. This “advantage” of a synchrony difference was less pronounced for older males that cuckolded younger males, particularly when the two males were not neighbors. Collectively, these results suggest that breeding synchrony affects extra-pair mating by affecting mate guarding, but that breeding synchrony alone can not be used to predict which females are more likely to engage in extra-pair mating, nor with which extra-pair males they will mate. Understanding why extra-pair mating by older males is less affected by breeding synchrony may explain much about both the proximate and ultimate causes of extra-pair mating in red-winged blackbirds. Received: 7 June 1996 / Accepted after revision: 25 November 1996     

Response thresholds to sucrose predict foraging division of labor in honeybees

Division of labor, where thousands of individuals perform specific behavioral acts repeatedly and non-randomly, is the hallmark of insect societies. Virtually nothing is known about the underlying neurophysiological processes that direct individuals into specific behavioral roles. We demonstrate that sensory-physiological variation in the perception of sucrose in honeybees measured when they are 1 week old correlates with their foraging behavior 2–3 weeks later. Workers with the lowest response thresholds became water foragers, followed with increasing response thresholds by pollen foragers, nectar foragers, bees collecting both pollen and nectar, and finally those returning to the colony empty (water<pollen<nectar<both<empty). Sucrose concentrations of nectar loads were positively correlated with response thresholds measured on 1-week-old bees. These results demonstrated how the variable response thresholds of a sensory-physiological process, the perception of sucrose, is causally linked to the division of labor of foraging. Received. 28 June 1999 / Received in revised form: 2 November 1999 / Accepted: 20 November 1999     

Polygyny in the red-winged blackbird: do females prefer monogamy or polygamy?

Five hypotheses have been proposed to explain polygyny in the red-winged blackbird (Agelaius phoeniceus). We categorized the hypotheses into three groups based on female preference for unmated versus monogamously mated males: (1) the “polygyny threshold” model, “sexy son” hypothesis and the “asynchronous settlement” model, which assume that females prefer unmated males to mated males on breeding situations of homogeneous quality; (2) the “neutral mate choice” hypothesis, which assumes that females have no preference; and (3) the “cooperative female choice” model, which assumes that females prefer monogamously mated males to unmated males. We tested the direction of female preference in two field experiments. In both experiments, newly settling females were given a choice of two adjacent territories, one defended by an unmated male and the other by a monogamously mated male. Male mating status was randomized with respect to the variation in territory quality and male quality. Early in the breeding season, significantly more females settled with the unmated males than with the mated males. Although more females settled with the unmated males than with the mated males late in the breeding season, the difference was no longer significant. Female settlement late in the season appeared to be related to the tenure of the resident females: the new females avoided territories where the resident females were in early stages of their nesting, but settled on territories where the resident females were in late stages. The pattern of female settlement shows that females prefer unmated males to mated males. The preference is consistent with the polygyny threshold model, sexy son hypothesis and the asynchronous settlement model, and inconsistent with the neutral mate choice hypothesis and the cooperative female choice model. For this reason, the latter two hypotheses are unlikely to explain the occurrence of polygyny in our population of red-winged blackbirds. Received: 1 December 1994 / Accepted after revision: 28 October 1995     

Kin conflict over caste determination in social Hymenoptera

We argue that caste determination, the process whereby females in the social Hymenoptera develop into either queens or workers, is subject to kin-selected conflict. Potential conflict arises because developing females are more closely related to their would-be offspring than to those of other females. Therefore, they may favour becoming queens contrary to the interests of other developing females and of existing queens and workers. We suggest two contexts leading to potential caste conflict. The first occurs when queens are reared in a reproductive phase following an ergonomic phase of worker production, while the second occurs when queens and workers are reared simultaneously. The first context assumes that workers' per capita contribution to colony survival and productivity falls with rising colony size. A critical feature influencing whether potential conflict is realized is the extent to which developing females can determine their own caste (“self-determination”). Self-determination is facilitated when female larvae control their own food intake and when queen-worker size dimorphism is low. We know of no strong evidence for actual conflict over caste fate arising in the first context. However, stingless bees and polygynous ants with excess queen-potential larvae that are either forced to develop as workers or are culled as adults demonstrate actual caste conflict in the second context. Caste conflict does not preclude caste regulation for “the good of the colony”, but such regulation is contingent on either the absence of potential conflict or on developing females losing control of their caste fate. Received: 22 March 1999 / Received in revised form: 15 June 1999 / Accepted: 19 June 1999     

Is asynchronous hatching adaptive in herring gulls (Larus argentatus)?

Hatching asynchrony commonly induces a size hierarchy among siblings and the resultant competition for food between siblings can often lead to starvation of the smallest chicks within a brood. We created herring gull (Larus argentatus) broods with varying degrees of hatching synchrony by manipulating the timing of incubation while maintaining the originally laid eggs. The degree of hatching asynchrony affected sibling size hierarchy at the time of hatching of the last-hatched ”c-chick.” In unmanipulated broods, there was no disadvantage of being a c-chick. However, when asynchrony was experimentally increased, we found reduced survival of the c-chick only in the exaggerated asynchronous experimental group. The effects were observable only during the first 10 days of chick life. We recorded no cases of the chicks dying of starvation. Furthermore, behavioral observations indicated that there was no sibling competition, and no selective feeding of larger sibs in the study colony. We propose that the observed lower survival rates of c-chicks in exaggerated asynchronous broods resulted from their lesser motor abilities, affecting their chances of escaping predators. Fledging success for the whole colony was generally high and almost half of all pairs fledged all three chicks, which is indicative of a good feeding environment. We argue that normal hatching asynchrony is a favorable solution in a good feeding environment, but that increased asynchrony reduces breeding success. We do not view asynchrony in the herring gull as an adaptation for brood reduction and propose instead that it may come about because there has been selection for incubation to start before clutch completion. Received: 14 April 1999 / Received in revised form: 20 October 1999 / Accepted: 23 January 2000     

DNA sequence analysis identifies genetically distinguishable populations of harp seals (Pagophilus groenlandicus) in the Northwest and Northeast Atlantic

Harp seals (Pagophilus groenlandicus Erxleben, 1777) comprise three populations based upon whelping areas in the Greenland Sea, White Sea, and Northwest Atlantic. The last comprises two subpopulations, one whelping in the Gulf of St. Lawrence (“Gulf ”) and one on the pack ice of the southern Labrador/northern Newfoundland coastal shelf (“Front”). A total of 40 female seals from the four whelping areas were collected during the 1990 and 1992 whelping seasons. DNA sequence variation was examined in a 307 bp region of the mitochondrial cytochrome b gene. Eleven variable nucleotide positions defined 13 genotypes: a significant fraction of the genotypic variance (F _ST=0.12, or 0.09 as measured by Weir's coancestry coefficient θ) is attributable to differentiation between Northwest and Northeast Atlantic populations. There was no significant differentiation between the two whelping areas in the Northwest Atlantic, or between the Greenland Sea and White Sea. These findings suggest significant reproductive isolation exists between trans-Atlantic breeding populations. Received: 18 January 1999 / Accepted: 22 February 2000     

Helpers and egg investment in the cooperatively breeding acorn woodpecker: testing the concealed helper effects hypothesis

In cooperatively breeding acorn woodpeckers (Melanerpes formicivorus), helper males have a large positive effect on fledging success in good acorn crop years but only a small positive effect in poor acorn crop years, while helper females exhibit the opposite pattern. Based on these findings, we tested the “concealed helper effects” hypothesis, which proposes that laying females reduce investment in eggs (with respect to their size, number, or quality) in a way that confounds helper effects and results in an absence of a relationship between helpers and breeding success. Results generally failed to support the hypothesis. Mean egg size was positively related to temperatures during the 10 days prior to egg-laying and negatively related to the food supply as indexed by the prior fall’s acorn crop, but there were no significant differences vis-à-vis helpers except for interactions with the acorn crop that only partly corresponded to those predicted. With respect to clutch size, females laid larger clutches when assisted by female helpers, opposite the pattern predicted. Although our results suggest that egg size is adjusted to particular ecological circumstances, we conclude that neither egg nor clutch size is adjusted in a way that confounds the apparent effects of helpers, as proposed by the concealed helper effects hypothesis.     

Why do snakes have eyes? The (non-)effect of blindness in island tiger snakes (Notechis scutatus)

Large (to >1 m), diurnally active tiger snakes (Notechis scutatus) are abundant on Carnac Island, near the coast of Western Australia. Our behavioural and mark-recapture studies provide the first ecological data on this population, and reveal a surprising phenomenon. Many adult tiger snakes have had their eyes destroyed, apparently during nest defence, by silver gulls (Larus novaehollandiae). This loss of vision did not reduce the snakes' body condition (mass relative to length), or their rates of growth or survival (measured over a 12-month period). Blind male snakes trail-followed females, and mated successfully. Thus, destruction of a major sensory modality had no detectable effect on these predators. This result is strongly counter-intuitive, but mirrors an earlier report of congenital blindness (without ill-effects) in American viperid snakes. Similarities between the two systems (island populations, highly venomous snakes, reliance on sessile prey) clarify the circumstances under which the loss of vision does not reduce an organism's viability. These natural experiments support Gans' hypothesis of “momentarily excessive construction” in that the snakes possess a complex organ system that they do not actually require for successful feeding, survival or reproduction. Received: 30 November 1998 / Received in revised form: 24 April 1999 / Accepted: 9 May 1999     

Competition and group size in Thomas’s langurs (Presbytis thomasi): the folivore paradox revisited

Among primates, group size is highly variable. The standard ecological model assumes that better predation avoidance as group size increases favours living in larger groups, whereas increased travel costs and reduced net food intake due to within-group competition for resources set the upper limit. Folivorous primates, however, tend to defy this generalisation in that some live in small groups despite low costs of feeding competition. To resolve this ’folivore paradox’, it has been suggested that folivore group size is limited by social factors such as male harassment or infanticide, or that females can disperse more easily and thus maintain group size near optimum levels. In this paper, we examine the effects of group size on home range size, day-journey length, activity budget and diet in wild Thomas’s langurs (Presbytis thomasi), which live in one-male multi-female groups with a limited life cycle. We examined only data from the stable middle tenure phase when factors such as the strength of the breeding male or the way in which groups were formed did not influence ranging and activities. During this phase, group size affected day-journey length and home range size, and had a minor effect on diet, but did not influence time spent feeding or resting, allogrooming or birth rates. Hence the upper limit to group size during the middle tenure phase in Thomas’s langurs is not set by feeding competition. The folivore paradox is not due to frequent female dispersal in Thomas’s langurs. The timing of female dispersal is not as expected if it serves to keep group sizes near the ecological optimum, and groups seem to be below this optimum. Instead, female reproductive success is presumably maximised in small to mid-sized groups because larger groups show a clear trend to experience higher risk of take-over, often accompanied by infanticide. Because females can redistribute themselves among nearby groups when groups reorganise each time a new male starts up a new group, females can keep the group small. Thus, a social factor, risk of infanticide, seems to provide the selective advantage to small group size in Thomas’s langurs. Received: 29 July 1999 / Revised: 17 November 1999 / Accepted: 15 October 2000     

Carbon content and nutritional condition of sardine larvae (Sardina pilchardus) off the Atlantic coast of Spain

As part of a “European Sardine/Anchovy Recruitment Program” (SARP), sardine larvae (Sardina pilchardus) were sampled off the Atlantic coast of Spain through the spawning season from March to June. The larvae were analysed for carbon and nitrogen content as a measure of nutritional condition and survival potential. There was no significant diel variation in larval carbon content, but there was a small significant diel variation in nitrogen; the absence of a strong diel signal in elemental composition was ascribed to the overnight retention of the gut contents. There was an increase in carbon content with increase in body length which reached an asymptote at ∼40% carbon content at a larval length of 20 mm. It is argued that larvae with a carbon content of <25% of body weight were nutritionally stressed, with the smaller larvae (<10 mm in length) appearing to be more vulnerable to food limitation. Although larvae with the lowest age-specific carbon content (poorest condition) occurred on the cruise with the lowest food availability, there was no consistent relationship between carbon content and food availability. While the successive monthly estimates of carbon content revealed differences in potential recruitment between months, these were not related to the birth-date distribution of the surviving juveniles. Received: 23 November 1998 / Accepted: 3 March 1999     

Fine structure of the “button setae” in the deep-sea pelagic copepods of the genus Euaugaptilus (Calanoida: Augaptilidae)

 Many species of the carnivorous copepod family Augaptilidae, a primarily meso- and bathypelagic group, possess specialized “buttons” on the setae of their maxilla and maxilliped, which have been thought to be a simple cuticular outgrowth. The fine structure of the button setae in three species of Euaugaptilus was examined by light- and electron microscopy from samples collected in the eastern Indian Ocean and the subtropical western North Pacific. The buttons are arranged in two rows along the inner surface of the setae. There are differences in the shape, size, and arrangement of buttons among the three species, Euaugaptiluslaticeps, E.longimanus, and E.magnus. The button setae have an elaborate internal structure consisting of “setal shaft”, “stalk”, “disc”, and “outer membrane”, with microtubule bundles in the setal lumen, but have no muscular tissues. The disc has only pin-point contact to the core of the stalk, but has connection over a wider area with the fibrous tissue surrounding the core, and the contact area between the stalk and the setal shaft is also small. These structures suggest that the disc will move freely on the apex of the core of the stalk and the stalk may be movable at its connection to the setal shaft. All these components of the button setae may function in concert to fit the surface of the discs firmly to the surface of a prey, and to absorb the energy generated by the struggling prey. Received: 30 November 1999 / Accepted: 19 May 2000