Fieldfare (Turdus pilaris) breeding success in relation to colony size,nest position and association with Merlins (Falco columbarius)

Summary Clutch size, nestling production and breeding success were studied in colonial Fieldfares (Turdus pilaris) in a subalpine birch forest during ten breeding seasons. Reproductive success was highest for central pairs in large colonies; such pairs benefited most from communal defence against nest predators. Fieldfares and Merlins (Falco columbarius) usually bred in association. Fieldfares breeding away from Merlins had lower breeding success than pairs associated with Merlins, which also benefited by reduced nest predation. Fieldfares apparently chose to nest near Merlins, which had already laid eggs when the thrushes started nest-building.     

Communal nesting, kinship, and maternal success in a social primate

Communal nesting, where several mothers regularly pool and cooperatively rear offspring, is unusual in mammals. This type of crèching behavior is especially rare among primates, with the notable exceptions of humans, some nocturnal strepsirrhines, and—as we show in this study—black-and-white ruffed lemurs (Varecia variegata). Here, we combine data on nesting behavior, genetic relatedness, and infant survivorship to describe variation in ruffed lemur infant care and to examine the potential benefits of ruffed lemur communal breeding. Reproductive events were rare, and females produced litters (synchronously) only once in 6 years of observation. We show that not all mothers participate in communal crèches, but those that did had greater maternal success; communal breeders spent more time feeding and their offspring were more likely to survive. Although cooperating mothers were often related, females also cooperated with non-kin, and those who shared infant care responsibilities had greater maternal success than mothers who did not participate. If there is indeed a causal link between maternal cooperation and reproductive success, this unusual behavior, like that of human communal rearing, may have evolved via some combination of kin selection and mutualism.     

Reproductive success in a low skew, communal breeding mammal: the banded mongoose, Mungos mungo

In most cooperatively breeding species, reproduction is monopolised by a subset of group members. However, in some species most or all individuals breed. The factors that affect reproductive success in such species are vital to understanding why multiple females breed. A key issue is whether or not the presence of other breeders is costly to an individual’s reproductive success. This study examines the factors that affect the post-parturition component of reproductive success in groups of communal-breeding banded mongoose (Mungos mungo), where up to ten females breed together. Per-litter reproductive success was low (only 18% of pups survived from birth to independence). Whilst singular breeding was wholly unsuccessful, there were costs associated with breeding in the presence of increasing numbers of other females and in large groups. Synchronisation of parturition increased litter success, probably because it minimises the opportunity for infanticide or decreases competitive asymmetry between pups born to different females. There was no evidence of inbreeding depression, and reproductive success was generally higher in litters where females only had access to related males within their group. I conclude that communal breeding in female banded mongooses represents a compromise between the benefits of group-living and communal pup care on the one hand, and competition between females to maximise their personal reproductive success on the other. Such conflicts are likely to occur in most communal breeding species. Whilst communal breeding systems are generally considered egalitarian, negative effects of co-breeders on individual reproductive success is still an issue.     

Sex-specific patterns of lifetime reproductive success in single and repeat breeding steelhead trout (Oncorhynchus mykiss)

All other things being equal, the lifetime reproductive success (LRS) of iteroparous and semelparous individuals should scale with the number of breeding seasons. Deviations from this relationship may occur for many reasons, including age- or size-related fecundity or life history trade-offs, which may differ between sexes. We used 19 brood years of DNA parentage analysis in a small (N = 4–143 year⁻¹) wild, unexploited population of steelhead trout (Oncorhynchus mykiss) to compare the LRS of individuals that spawned only once [“one time spawners” (OTS), N = 355 male, 371 female] to those spawning twice [“repeat spawners” (RPS), N = 13 male, 49 female]. Female RPS had nearly twice the LRS of female OTS (1.17 offspring per female vs 0.91 offspring per female), whereas male RPS had nearly three times the LRS of male OTS (1.54 offspring per male vs 0.57 offspring per male). Female RPS produced slightly more adult offspring during their second breeding season than their first (0.78 vs 0.82 offspring per female); however, male RPS produced all of their adult offspring in their second breeding season (0 vs 1.54 offspring per male). The additional growth in body size of males between breeding seasons may give them an advantage in their second breeding season, but the lack of offspring produced in their first season suggests a trade-off between survival and future reproduction that was not expressed in females.     

Alternative male mating tactics in a cichlid, Pelvicachromis pulcher: a comparison of reproductive effort and success

Pelvicachromis pulcher is a small African cichlid which breeds in holes. Males may either reproduce monogamously (pair males), polygynously (harem males), or be tolerated as helpers in a harem territory (satellite males). These helpers share in defence of the territory against conspecifics, heterospecific competitors and predators. There are two male colour morphs that are fixed for life and are apparently genetically determined. These differ in their potential mating strategy. Red morph males may become harem owners, while yellow morph males may become satellite males, and males of both morphs may alternatively pair up monogamously. We compared the reproductive effort and success of these three male reproductive strategies. Effort was measured as attack rates, time expenditure and the risk of being injured or killed when attacking competitors or predators of three sympatric fish species. Reproductive success was measured by observing how many eggs were fertilized by each male when this was possible, and by using genetic markers. The number of fry surviving to independence of parental care was used as a criterion of success. The reproductive success of harem males was 3.3 times higher than that of pair males and 7 times higher than that of the average satellite male. Dominant satellite males, however, were as successful as monogamous pair males, using the measure of fertilized eggs. To our knowledge, this has not been found previously in any fish species. Both harem and pair males had lower parental defence costs per sired offspring, however, than males using the alternative satellite tactic. Defence effort was significantly related to the risk of injury. Received: 17 January 1996 / Accepted after revision: 9 June 1997     

The production of sexual and asexual larvae within single broods of the scleractinian coral, Pocillopora damicornis

The genotypic compositions of two populations of the brooding coral, Pocillopora damicornis, were studied by exhaustive sampling of adult colonies on a fine scale in southern Taiwan. In addition, 100 larvae were randomly selected among more than 1,000 brooded larvae collected from a single broodparent within each population. Using 7 polymorphic microsatellite markers, both populations were found to be highly clonal, and 7 clonal lineages were characterized. One clonal lineage (C1) dominated both study areas and comprises 54.9% of all colonies sampled, while any of the other 6 clonal lineages represented no more than 5%. Among the 100 larvae randomly selected for genotyping from each broodparent, the extent of clonal reproduction was high, and only 29 and 6 larvae, respectively, were found to be genotypically different from their broodparent. Among the 35 genotypically distinct larvae, 33 were thought to be derived from sexual reproduction, and 2 were assumed to be clonal propagules that had undergone somatic mutations. Two genotypically identical larvae were also found in one of the 2 sexually derived larva arrays, indicating the possible existence of polyembryony. The high proportions of clonality in both adult colonies and brooded larvae suggest that asexually produced larvae might significantly contribute to the recruitment of local populations. The dense clonal population of P. damicornis in the study area favors the quick recolonization view of clonal propagules after disturbances.     

Past reproductive effort affects parental behaviour in a cichlid fish,Cichlasoma nigrofasciatum: a comparison of inexperienced and experienced breeders with normal and experimentally reduced broods

Defensive and parental care behaviour of convict cichlids that differed in past effort was compared. Before testing, some fish were bred three times while others were not bred. Age was held constant; all individuals in this study were approximately 20 months old (±2 months) at test time. Furthermore, half of the pairs in this study had their broods experimentally reduced by 50%. Results indicated that past effort across breeding attempts affects investment in the current brood. Experienced pairs were more aggressive toward a model predator than inexperienced parents. However, no major differences were observed in depreciable care (i.e. fanning). Contrary to previous studies, brood size had minor effects on parental care. This discrepancy could be due to the age of the parents; individuals in this study were significantly older than fish tested in previous studies. The results support parental investment theory and suggest that past effort is not only important within breeding episodes but also within an animal's lifetime.     

Sex ratio,monogamy and breeding success in the Midas cichlid (Cichlasoma citrinellum)

Summary The Midas cichlid (Cichlasoma citrinellum) is a large, aggressive monogamous fish from the Great Lakes of Nicaragua. It lives in an ecosystem where breeding sites are at a premium and where breeding success can be low. I tested the strength of the pair bond by manipulating the sex ratios in captive groups, using ratios of 1:1, 2:1, 4:1, 1:2 and 1:4 (:). No polygamy was seen at any of the ratios. Departure from a ratio of 1:1 decreased breeding success for those pairs that formed. Both sexes appeared to enforce the pair bond but did so in different ways: females attacked males and females equally but males attacked other males significantly more than they did females. These results, coupled with field observations, indicate the Midas cichlid is obligated to monogamy by the demands of competition for breeding sites, the need for two parents to protect the fry and by energetic constraints.     

Patterns of larval and early juvenile growth in a semiterrestrial crab, Armases angustipes (Decapoda: Sesarmidae): comparison with a congener with abbreviated development

In a semiterrestrial and estuarine tropical crab, Armases angustipes Dana (Grapsoidea: Sesarmidae), changes in biomass (measured as dry mass, W; carbon, C; nitrogen, N; and hydrogen, H; per individual) and relative elemental composition (C, N, H, in percent of W; C:N mass ratio) were studied during development from an early egg stage through hatching, the complete larval phase, metamorphosis and the first juvenile crab stage (CI). In the megalopa and CI, growth was measured also within the moulting cycle, and biomass and elemental composition were determined in cast exuviae. From an early egg stage to the freshly hatched larva, A. angustipes lost about 20% of W, 29% of C, 5% of N and 32% of H. Proportionally higher losses in C than in N were reflected also in a significantly decreasing C:N mass ratio (from 5.02 to 3.74). These results indicate that lipids mobilised from yolk reserves represented the principal metabolic substrate for embryonic energy production, while proteins were catabolised at a much lower rate. The present data of growth and exuviation are compared with previously published data from a congener, A. miersii Rathbun, which has an abbreviated and facultatively lecithotrophic mode of larval development (with three instead of four zoeal stages; stages I and II in principle independent of food). When growth is measured as an increase in the final (premoult) biomass of successive developmental stages, both species show an exponential pattern. Within the moulting cycles of the megalopa and the first juvenile, both species show parabola-shaped growth curves, with a rapid biomass increase in postmoult and intermoult stages, and losses in the premoult phase. Thus, the two Armases species show, in general, similar patterns of larval and early juvenile growth. However, the initial size of eggs and larvae is about four times larger in A. miersii, and its biomass remains higher throughout the period of larval and early juvenile development. A. angustipes is able to partially make up for this difference, as it has an additional zoeal stage, and its megalopa and CI stages show higher relative biomass increments (in percent of initial values). Due to this compensatory growth pattern, A. angustipes reaches in its CI stage about half the biomass of a juvenile A. miersii. When exuvial losses of megalopae and juveniles are compared between these two species, A. miersii shows higher biomass losses per individual (corresponding with its larger size), but lower relative losses (C, N, H, in percent of late premoult body mass or in percent of previously achieved growth increments). Differences in larval and early juvenile growth and in the exuvial losses of megalopae and juveniles of these two congeners are discussed in relation to their differential ecology, life history and reproductive strategy.     

Cooperation in breeding by nonreproductive wrens: kinship,reciprocity, and demography

Summary Stripe-backed wrens (Campylorhynchus nuchalis) often live as adults in large groups on permanent, communally defended territories. Nonbreeding adults cooperate in rearing the young of a single breeding pair; this aid substantially increases the reproductive success of the breeders. In a 6-year study in Venezuela of a completely colorbanded population of 25–30 groups, most adults participated in breeding only as helpers and priority to breeding status was strictly age-determined. Detailed behavioral observations at breeding nests with nestlings showed that, in a sample of 100, helpers nearly always contributed as much to the care of young as breeders. Further, aid-giving does not vary systematically with relatedness of ycung to helpers or with probability of future reciprocation by young. Young being raised are most often at least half siblings of helpers, but seldom return aid to adults that helped raise them. Even adopted helpers collaborate fully. Patterns of demography and dispersal show slow turnover of breeders, delayed reproduction, and a viscous population structure.Application of Hamilton's condition for selection for aid-giving reveals that most individuals in this population can maximize inclusive fitness in the first 2 years by helping instead of breeding. Variation in helping effort and in age of first breeding is related to variation in natal group size and competition resulting from variable demographic neighborhoods in different years or in different parts of the population. Because reciprocation in the form of specific alliance formation among nonreproductives is uncommon, nonspecific reciprocity between cohorts and kin selection account well for the observed pattern of age-dependence in first breeding. Nondiscriminating helping in this population is associated with stable monogamous pair boncs, stable territory boundaries and group membership, strict seniortiy for breeding position, high viscosity and consistent effectiveness of aid. Under these circumstances, very simple behavioral rules amounting to nearly automatic helping seem sufficient to confer critical inclusive fitness gain on helpers.     

Male care,mate switching,and future reproductive success in a double-brooded passerine

Summary In species with male parental care, polygyny may reduce the parental effort provided by a male, and hence impose a cost on the fitness of his mates, because of decreased growth, survival or health of offspring. I examined a cost of polygyny in the green dart-poison frog, Dendrobates auratus, a species with male parental care in which both male polygyny and mate guarding by females occurs (Summers 1989). All D. auratus males seen carrying tadpoles in a marked area were followed and the pools where they deposited their tadpoles were recorded. Males frequently deposited more than one tadpole in the same pool (in 25% of the observed depositions a male deposited a tadpole in a pool where he had previously deposited at least one other tadpole). Experiments involving manipulation of tadpole densities in pools typically utilized by D. auratus (calabash husks and treeholes) showed that increasing tadpole number had a strong negative effect on both growth rate and survivorship, indicating that polygyny can impose a cost on the fitness of a male's offspring. Hence, females do face a potential cost, in terms of reduced offspring growth and survivorship, when their mates mate polygynously and care for the offspring of other females.     

Black Rats, Island Characteristics, and Colonial Nesting Birds in the Mediterranean: Consequences of an Ancient Introduction

Abstract: The devastation of island faunas by alien species has been instrumental in raising concerns about the global threat to biological diversity. Colonial nesting species, often restricted to islands, have been affected severely. Eradication of introduced species as a means to alleviate the problem is usually done with little or no understanding of the mechanisms governing interactions between introduced and native species. Such an understanding could help target management action. We analyzed how island area, rock substrate, bird species biology, and presence of an introduced species, the black rat (   Rattus rattus ), interact to explain the distribution and abundance of colonial nesting birds on a set of 72 islands from six archipelagos in the western Mediterranean. Rats were introduced to this region over 2000 years ago, and these communities have had time to reach an equilibrium. Using general linear models, we show that rats have affected species distributions more on the smaller islands and on islands with an acidic or neutral rock substrate; larger bird species are more resistant. On the smaller islands, where rat densities are highest, larger colonial birds are negatively affected. On larger islands, where rat densities are lower and fluctuate form year to year, larger colonial birds can maintain healthy populations despite the presence of rats. Although rats caused local extinction or reduction in bird abundance, the presence of islands varying in size and/or substrate allowed most archipelagos to retain their suite of colonial nesting bird species, despite a presumably reduced abundance for most species.     

Biochemical composition and energy reserves in the larvae and adults of the four british periwinkles Littorina littorea,L. littoralis,L. saxatilis and L. neritoides

The biochemical composition of the adult body tissue is similar in Littorina littorea, L. littoralis, L. saxatilis and L. neritoides. In the newly crawling metamorphosed young of L. littoralis and L. saxatilis, the biochemical composition is similar to that of the mature adults. The newly released planktonic veliger larvae of L. littorea and L. neritoides, which represent and earlier stage of development than the newly crawling young, have a neutral lipid level (mean, 16.5% of ash-free dry flesh weight) approximately three times the level (5.4%) in the newly crawling young of L. littoralis and L. saxatilis. Otherwise the biochemical composition of free-swimming larvae is similar to that of newly crawling young and adults. Neutral lipid is apparently utilised by L. littoralis and L. saxatilis larvae during larval development and metamorphosis. It is suggested that neutral lipid is the major energy reserve of Littorina veliger larvae whereas in the adults, as exemplified by L. littorea, both lipid and carbohydrate are important as energy reserves.     

Cooperative breeding and immunity: a comparative study of PHA response in African birds

Cooperatively breeding birds might be expected to suffer from higher costs of parasitism than pair-breeding species because of two aspects of their ecology which should facilitate horizontal transmission and possibly select for higher parasite virulence: first, they interact regularly with more individuals than pair-breeding species, and second, these individuals are commonly close relatives that could share similar resistance alleles. This hypothesis predicts that cooperative breeders should invest relatively more in immune defence than closely related species which breed in pairs. I tested this prediction comparatively in African birds by examining the response of the immune system to the mitogenic lectin, phytoahemagglutinin (PHA response) in relation to cooperative breeding. Among 66 species, of which 18 breed cooperatively, PHA response was significantly higher in cooperatively breeding species. This association appeared not to be confounded by body size, clutch size, nest position, coloniality or similarity owing to common phylogenetic descent. These results suggest that cooperatively breeding birds may have been selected to invest more than pair-breeders in defences against parasites. If so, then additional costs of philopatry and helping behaviour might be imposed on breeders, helpers and offspring.     

Larval paternity and male reproductive success of a broadcast-spawning gorgonian, Plexaura kuna

Male reproductive success of the broadcast-spawning gorgonian, Plexaura kuna Lasker, Kim and Coffroth, 1996, was measured in June 1994 and June and July 1995 at two sites in the San Blas Islands, Panamá in order to determine the environmental and biotic factors affecting individual reproductive success. Developing embryos were collected in the field during natural spawning events, and paternity determined using randomly amplified polymorphic DNA markers. Analyses of F₁ progeny from defined laboratory matings established that the markers were inherited in Mendelian fashion, and allowed the determination of the zygosities of the markers. P. kuna is clonal, but male reproductive success was not strictly proportional to clone size. Proximity to females appeared to have a greater effect on male reproductive success than clone size, and on both reefs the most successful male clone was the one closest to the spawning female clone. Current direction and transport of gametes by eddies explained variation in paternity assignments between nights. Clonal propagation allows clones to grow and spread toward each other, and may enhance male reproductive success. Received: 1 April 1997 / Accepted: 11 February 1998     

Fighting success and attractiveness as predictors of male mating success in the black field cricket, Teleogryllus commodus: the effectiveness of no-choice tests

Females are generally assumed to prefer larger, more dominant males. However, a growing number of studies that control for male-male competition have shown no correlation between dominance and attractiveness. Aggressive males can interfere with female mate preference either by physically coercing females into mating or by driving submissive males away and restricting mate choice. The most common method of assessing female mate choice is by using simultaneous two-choice tests. These control for male-male interactions, but usually interfere with physical and chemical cues involved in mate selection or alter male behaviour. They are therefore unsuitable for many study species, especially insects. Another method is the no-choice test that measures a females latency to mating when placed with a single male as an indication of male attractiveness. No-choice tests control for male-male aggression while allowing full contact between pairs (they allow actual mating to be directly observed rather than to occur based on a correlated behaviour). So far, however, no study has confirmed that males that entice females to mate sooner actually enjoy increased longer-term mating success. As such, the accuracy of no-choice tests as a method of examining mate choice remains untested. Here, we used no-choice tests on the black field cricket, Teleogryllus commodus, to show that (1) females did not prefer males that won fights (dominant males), and (2) latency to mating predicts actual mating success. We have clearly demonstrated the usefulness of no-choice tests and, considering the advantages of this method, they should be more often considered for a wider variety of taxa.Communicated by D. Gwynne     

Racketed tail of the male and female turquoise-browed motmot: male but not female tail length correlates with pairing success,performance, and reproductive success

Both males and females of many avian species maintain elaborate plumage traits, and elaborate monomorphic plumage may convey adaptive benefits to one or both sexes as inter- or intraspecific signals. Both sexes of the turquoise-browed motmot (Eumomota superciliosa) are elaborately plumed with long racket-tipped tail. I investigated whether the racketed tail functions as a sexually selected signal in one or both sexes by testing the predictions that males and/or females with the largest tails have: (1) greater pairing success, (2) greater reproductive performance (clutch-initiation date, clutch size, and hatching success), and (3) greater reproductive success. Yearling males with longer denuded rachises (wires) on the central tail feathers had greater pairing success. In addition, adult males with longer wires paired with females who laid larger clutches, had greater hatching success independent of clutch size, and fledged more young. There was no relationship between female tail plumage and pairing success, reproductive performance, or fledgling success. These results are consistent with the hypothesis that male tail plumage functions as a mate choice or status signal, but that the tail of the female does not function in a sexually selected context. I discuss alternative hypotheses for the evolutionary maintenance of the elaborate female tail plumage.     

Distribution,habitat preferences and breeding periods in the crustaceans Sesarma cinereum and S. reticulatum (Brachyura: Decapoda: Grapsidae)

The distribution and habitat preferences of Sesarma cinereum and S. reticulatum were studied in the region of Beaufort, North Carolina, USA. S. cinereum inhabits the supralittoral zones of marshes characterized by high salinities (mean=27.9%) and sandy substrates, while s. reticulatum were found to prefer eulittoral marshes with brackish salinities (mean=16.2%) and silty substrates. Major differences have also been found in the cycling of egg production between these two species. Populations of S. cinereum produce approximately 4 to 6 egg batches in close synchrony with the lunar phase, whereas populations of S. reticulatum produce only from 2 to 3 batches of young per breeding season. It appears that female S. cinereum invest less time in carrying their eggs (approximately 1 lunar month), while female S. reticulartum invest a greater amount of time (eggs carried for approximately 45 days). These and other specific differences in reproductive strategies suggest that populations of S. cinereum are responding to density-independent selective pressures, while populations of S. reticulatum exhibit characteristics of density-dependent selective effects.