Sex allocation in a monomorphic seabird with a single-egg clutch: test of the environment, mate quality, and female condition hypotheses

Sex allocation theory posits that mothers should preferentially invest in sons when environmental conditions are favorable for breeding, their mates are of high quality, or they are in good body condition. We tested these three hypotheses in rhinoceros auklets (Cerorhinca monocerata), monomorphic seabirds that lay a single-egg clutch, in 2 years that differed in environmental conditions for breeding. Results supported the environment and mate quality hypotheses, but these effects were interactive: offspring sex was independent of paternal traits in the poor year for breeding, while females mated to larger and more ornamented males reared more sons in the better year. Conversely, offspring sex was unrelated to female condition, as indexed by hatching date. We propose that good rearing conditions enable females to rear sons possessing the desirable phenotypic attributes of their mates. Results also supported two critical assumptions of sex allocation theory: (1) dimorphism in offspring condition at independence: daughters fledged with higher baseline levels of corticosterone than sons and (2) differential costs of rearing sons versus daughters: mothers rearing sons when environmental conditions were poor completed parental care in poorer condition than mothers rearing daughters in the same year and mothers rearing either sex when conditions were better. These novel results may help to explain the disparate results of previous studies of avian sex allocation.     

Genetic evidence for mixed parentage in nests of the emu (Dromaius novaehollandiae)

Parentage in emus (Dromaius novaehollandiae) was examined by microsatellite analysis using four independent loci. Of 106 chicks sampled in one breeding season from 18 nests, 54 (51%) were not fathered by the nesting male, 12 (11%) were not from the observed mate of the sitting male, and 9 (8%) represented intra-specific brood parasitism, having no alleles in common with either nest parent. Some males (11%) fathered all chicks in their nests, but the majority showed high levels of cuckoldry. Those males commencing incubation earliest in the season tended to have the highest levels of paternity in their own nests. These results reveal a high frequency of extra-pair fertilisations and resultant cuckoldry in a predominantly socially monogamous bird and support recent reports which have described the emu mating system as a complexity of polyandrous, promiscuous and monogamous behaviour. Parentage assignment of chicks resulting from extra-pair fertilisations revealed an evenly scattered pattern of paternity that did not show any particular male dominance in reproductive success. These results lead to a reassessment of behavioural observations of emus, the consequences of parentage distribution, and theories about mating systems and sexual selection. The frequency of extra-pair copulations and intra-specific brood parasitism suggests patterns of descent that differ greatly from those implied by social monogamy. Received: 27 November 1999 / Received in revised form: 8 January 2000 / Accepted: 8 February 2000     

Nitrogen versus phosphorus limitation in a subtropical coastal embayment (Moreton Bay; Australia): Implications for management

An approach combining nutrient budgets, dynamic modelling, and field observations of phytoplankton and nitrogen (N₂)-fixing Lyngbya majuscula following changes in wastewater N loads, was used to demonstrate that Moreton Bay is potentially phosphorus (P) limited. Modelling and nutrient budgeting shows that benthic N-fixation loads are high, allowing the system to overcome any potential N-limitation. Phytoplankton biomass has shown little change from 1991 to 2006 in the sections of Moreton Bay most impacted by wastewater effluents, despite a large reduction in wastewater N loads from 2000 to 2002. This is consistent with modelling that also showed no reduction in primary productivity associated with reduced N loads. Most importantly, there have been rapid increases in the occurrence of N-fixing L. majuscula in Moreton Bay as wastewater P loads have increased relative to wastewater N loads. This is also consistent with modelling. This work supports the premise that there may be fundamental differences in nutrient limitation of primary production between subtropical and temperate coastal systems due to differences in the importance of internal nitrogen sources and sinks (N-fixation and denitrification). These differences need to be recognised for optimum management of coastal systems.     

Distributional ecology and food habits of the banded blennyParaclinus fasciatus (Clinidae): a resident in a mobile habitat

An ecological study of the banded blennyParaclinus fasciatus (Steindachner, 1876) showed that the fish is tolerant of physical/chemical conditions in the shallow subtidal area of Apalachee Bay, Florida (USA). Population density ofP. fasciatus was unrelated to physical/chemical characteristics and seagrass biomass at field stations. A strong correlation was found between numbers ofP. fasciatus captured and abundance of unattached red algae (drift algae) at the field sites, although seasonality in population density was related to summer spawning. Spawning began at approximately 28°C. The life span of the clinid is probably little longer than 1 yr. Long-term variation in population size was also demonstrated.P. fasciatus was found to be a highly specialized carnivore which consumes only a few species of amphipods and shrimps; however, major seasonal variation in diet occurred. Variations in food habits were related to known patterns of prey abundance in Apalachee Bay. The most important prey species taken byP. fasciatus, includingCymadusa compta andHippolyte zostericola, are associates of red algae masses, further substantiating the close functional relationship of the clinid with its algal habitat which moves about the bay. The driftalgae habitat is probably an important source of shelter, food, and dispersal for the banded blenny. The eurytolerance ofP. fasciatus makes it particularly well adapted for life in an unusual, mobile habitat.Gontribution No. 173 of Harbor Branch Foundation, Inc.     

Respiratory responses of the estuarine mysid Neomysis integer (Peracarida: Mysidacea) in relation to a variable environment

The euryhaline mysid Neomysis integer (Peracarida: Mysidacea) is a common member of the hyperbenthos of the upper reaches of European estuaries. In the East Looe River Estuary (Cornwall, England), this species experiences extensive tidal and seasonal changes in temperature (3 to 15 °C) and salinity (1 to 34‰). In this investigation, the effects of temperature (5, 10 and 15 °C) and salinity (1, 10, 20 and 30‰) on the oxygen consumption of male and female N. integer are reported, and are related to field measurements to identify the adaptive responses of the respiratory physiology to such a variable environment. The general responses were similar for each sex; however, at any given temperature/salinity combination, male N. integer consumed more oxygen than females. The general trends were increased oxygen consumption with increasing temperature (Q₁₀ values ranged from ∼1.7 to 2.5) and decreased oxygen consumption with increasing salinity. Temperature and salinity interacted at high water antagonistically to minimise changes in mysid oxygen-consumption. When related to tidal fluctuations in temperature and salinity experienced by N. integer inhabiting the East Looe River Estuary, the results reveal how the respiratory physiology of this species is adapted to its variable environment. Received: 16 June 1998 / Accepted: 15 December 1998     

Fluoride distribution in the environment along the gradient of a phosphate-fertilizer production emission (southern Brazil)

Airborne fluoride was determined in the rainwater, surface soil and groundwater along a gradient of emission of a phosphate fertilizer factory in Rio Grande, southern Brazil. Concentrations of fluoride in rainwater and groundwater achieved 3 mg l⁻¹ and 5 mg l⁻¹, respectively, and were dependent on pH. The fluoride deposited from emissions accumulated in a superficial horizon of soil in quantities comparable to those in the manufactured end-products—up to 23,000 mg kg⁻¹. Fluoride distribution in the environment is controlled by physical–chemical parameters of emission, rain intensity and soil properties. The highest fluoride concentrations were registered at a close distance of up to 2 km from the factory. The distribution of fluoride in groundwater resembled the same distribution in rainwater due to the high permeability of the local soils. Fluoride penetration to the groundwater also depended on the type of vegetation cover. The groundwater in woodland areas was less affected by contamination of fluoride than in the grassland areas, most probably because of the influence of eucalyptus throughfall, which increases the pH of wet precipitates.     

Biomagnification of trace elements in a benthic food web: the case study of Deer Island (Northern Yellow Sea)

In this study, we investigated the concentrations of ten trace elements (Cu, Zn, Mn, Se, Ni, Cd, Cr, Pb, Hg, and As) and their trophodynamics in a benthic food chain of Deer Island, Northern Yellow Sea. The concentrations of Cu, Zn, Mn, Se, Ni, Cd, Cr, Pb, Hg, and As in the food chain ranged from 3.2 to 23.2, from 71 to 227, from 7.4 to 45.6, from 0.44 to 5.80, from 0.73 to 7.60, from 0.14 to 1.65, from 0.68 to 6.70, from 0.08 to 1.86, from 0.08 to 1.18, and from 0.24 to 3.92 mg kg^?1 dry weight, respectively. Among these trace elements, the linear regression between the log-transformed concentrations of Hg and Cd and δ¹⁵N values showed statistically significant increase (p<0.05) with the slopes of 0.134 and 0.144, indicating biomagnification of Hg and Cd occurred in the benthic food chain of Deer Island. While the linear regression for other eight trace elements (Cu, Zn, Mn, Se, Ni, Cr, Pb and As) were characterised by extensive scatter with non-significant correlation coefficients (R ²=0.002–0.235) and slopes (p=0.079–0.875), indicating there were not biomagnified or biodiluted of these trace elements.     

Fertilisation success in the fly Dryomyza anilis (Dryomyzidae): effects of male size and the mating situation

Male fertilisation success in relation to male size and the mating situation (ordinary pair formation with a single, nonvirgin female vs. take overs) was examined in the fly Dryomyza anilis. In ordinary matings, large males achieved higher fertilisation success than small ones when they were the second to mate with the female. Take overs differ from ordinary pair formation in that the second male experiences intensified sperm competition. This is because in take overs the female is not able to discharge any of the sperm inseminated by the first male as she usually does before a new mating. Compared with ordinary matings, take overs reduced the fertilisation success of the second male by 8–10%, whereas that of the first male was 7–14% higher in take overs. Even though the intruder was always larger than the paired male his superior fertilisation success did not compensate for the effect of the sperm already present in the female. In D. anilis, males can increase their fertilisation success by tapping the female's external genitalia with their claspers or having several copulation bouts per mating. Thus, in a take over, the intruder could respond to the intensified sperm competition by performing more tapping sequences per copulation bout or more copulation bouts per mating. In matings observed in the wild, males performed more tapping sequences after a take over than after pair formation with a single female, although the difference was not significant. The results show that there are differences in fertilisation success between males of different size. In addition, different mating situations can result in considerable variation in the fertilisation success of an individual male. Higher fertilisation success for the first male after a take over may be significant, in particular, for the reproductive success of small males, which frequently lose their females to large males.     

Trade-off between fitness components in males of the polygynous butterfly Callophrys xami (Lycaenidae): the effect of multiple mating on longevity

The differential costs of mating paid by males and females influence the nature and strength of sexual selection. In butterflies, males invest a relatively large amount of time and resources in each mating, but male survival costs of mating have not been demonstrated. I present the results of experiments designed to measure the effect of different aspects of mating on male longevity in the polygynous butterfly Callophrys xami. In experiment 1, I compared the longevity of pairs of males that produced similar amounts of spermatophore, but that mated at different rates, a different numbers of times, and that produced spermatophores at different rates, and found that the longevity of ”low-mating-rate” males was not different from that of ”high-mating-rate” males. In experiment 2, the longevity of virgin males was not significantly different from that of multiply mated males. In experiment 3, I used resource-limited males resulting from experimental food limitation of last-instar larvae; resource-limited virgin males lived significantly more days than resource-limited multiply mated males. Since ecological costs of mating (e.g., disease transmission, predation risk) were excluded in the experiment, diminished male longevity was a product of physiological costs of sexual interactions. These results suggest that the cost of ejaculate production is an important cause of longevity reduction when there are resource limitations; however, the role of other possible physiological costs of mating in longevity reduction is still unknown. Received: 21 March 2000 / Accepted: 26 August 2000     

Mitochondrial DNA Variability in the Endangered Razorback Sucker (Xyrauchen texanus): Analysis of Hatchery Stocks and Implications for Captive Propagation

The razorback sucker ( Xyrauchen texanus ) is a large, long-lived catostomid fish endemic to the Colorado River drainage of western North America, endangered because of recruitment failure. Efforts to preserve the species have emphasized artificial propagation and reintroduction. Given the importance of maintaining genetic diversity in such a program, we examined mitochondrial DNA diversity in a source population (Lake Mohave, Arizona-Nevada) and three hatchery-produced year classes (1987, 1989, 1990). The source contained considerable variation, indicated by high haplotype diversity ( ĥ = 0.97) and a large number of unique haplotypes (17 in 25 individuals). Diversity also was high in the 1987 ( ĥ = 0.89, 6 haplotypes in 10 individuals) and 1989 hatchery-produced year classes ( ĥ = 0.91, 7 in 11), but significantly lower in the 1990 year class ( ĥ = 0.71, 4 in 10). Low diversity in the last class was likely because of differences among females in fecundity, viability of progeny, or both. Because natural populations have collapsed throughout the species' range, we must identify methods that preserve the most diversity. We examined three potential alternatives: standard hatchery propagation, natural spawning in predator-free environments, and protective custody of larvae collected from the lake with reintroduction after growth to a size likely to survive. The last is the preferred alternative and should be pursued as the most cost-effective option for preserving genetic diversity in the razorback sucker.     

Between-group contest competition for food in a highly folivorous population of black and white colobus monkeys (Colobus guereza)

In theory, between-group contest (BGC) competition for food can greatly influence female social relationships and reproductive success in primates, but few studies have investigated whether such effects occur and, if so, under what ecological conditions. There is evidence that adult male black and white colobus monkeys (Colobus guereza, “guerezas”) defend the food their mates need against other groups, suggesting that BGC competition is important in this species. Using data on feeding, ranging, vegetation patterns, and intergroup encounters between six neighboring guereza groups, I provide evidence that the highly folivorous guerezas at Kanyawara, Kibale National Park, Uganda, engaged in BGC competition over unevenly dispersed, relatively high-quality feeding areas or “core areas”. Intergroup aggression was common, and groups’ home ranges overlapped. Groups were more likely to initiate high-level aggression if they encountered another group within or near their core area, and groups that initiated and won encounters often fed in the same areas in which losing groups had fed. Guerezas fed selectively on species with contagious (clumped) distributions and concentrated their feeding efforts in areas of the forest that contained the most food (core areas). Groups could be ranked in a linear dominance hierarchy, and group rank number was inversely related to the quantity and quality of food in groups’ core areas. This study not only provides good evidence that BGC competition occurs in primates but it also reinforces the idea that folivore food resources may be worth defending.     

The effect of pup vocalisations on food allocation in a cooperative mammal, the meerkat (Suricata suricatta)

In the meerkat (Suricata suricatta), a cooperative mongoose, pups follow potential feeders while the group is foraging and emit incessant calls when soliciting food from them. In contrast to a ’stationary’ brood of chicks, in which nestlings are fed at a fixed location, meerkat pups are ’mobile’ and become spread out. The question arises whether meerkat pups that experience different constraints to those facing chicks have evolved similar begging strategies. This paper describes the vocalisations that meerkat pups emit in the context of begging and investigates the influence of these calls on food allocation by older group members and on the behaviour of littermates. Meerkat pups use two types of calls when soliciting food from a potential feeder. The most common is a ’repeat’ call, which pups emit continuously when following an older forager over several hours a day. In addition, when a potential feeder finds a prey item, the pups next to it emit a bout of calls with increased calling rate, amplitude and fundamental frequency, termed ’high-pitched’ calls. Observations, together with playback experiments, showed that more prey was allocated to pups that called longer and more intensely. The pup closest to a feeder was almost always fed. The probability of emitting high-pitched calls did not depend on the time since a pup had received food, and the change from repeat to high-pitched calls occurred suddenly. The main function of the high-pitched call, therefore, does not appear to be to signal a pup’s hunger state. More likely, the two calls, in the context of begging, may be an adaptation to energetic constraints in a mobile feeding system. Pups, which are dispersed during foraging, may emit repeat calls over long periods to prevent potential feeders from eating all the prey themselves. At the moment a potential feeder finds prey, pups may give the more intense high-pitched calls to direct feeders to bring the food item to them and not to a littermate. Therefore, unlike the stationary feeding system where chicks emit one type of begging call when the feeder approaches the nest, meerkats, with a mobile feeding system, have evolved two discrete types of vocalisations in the context of begging. Received: 22 November 1999 / Revised: 1 July 2000 / Accepted: 17 July 2000     

Division of labour in a crisis: task allocation during colony emigration in the ant Leptothorax unifasciatus (Latr.)

Division of labour during colony emigration is widespread in ants. An important problem is how tasks are allocated during colony movement from one nest site to another. The generally favoured view is that emigrations are organised by a minority group of individuals, which either work unusually hard at tasks (elites) or have the exclusive task of carrying out the emigration (moving specialists). Five consecutive emigrations of a Leptothorax unifasciatus (Latr.) colony showed that the number of transporters, i.e. the individuals that took an active part in the emigration by transporting brood and ants, was smaller than it would have been if allocation of this task was random during each emigration. However, single emigrations of another three colonies, for which the spatial distribution and behaviour of the workers had been observed for a week prior to the emigration, demonstrated that the transporters did not form a homogeneous group. They differed in their spatial positions and tasks before the emigration. There was also no evidence that transporters worked harder or less hard than their nestmates before the emigration. Therefore, the individuals which carry out emigrations in L. unifasciatus colonies appear to be neither moving specialists nor elites. We propose that task allocation during emigrations of L. unifasciatus colonies is based on a feedback mechanism that involves learning.     

Division of labor in a dynamic environment: response by honeybees (Apis mellifera) to graded changes in colony pollen stores

 A fundamental requirement of task regulation in social groups is that it must allow colony flexibility. We tested assumptions of three task regulation models for how honeybee colonies respond to graded changes in need for a specific task, pollen foraging. We gradually changed colony pollen stores and measured behavioral and genotypic changes in the foraging population. Colonies did not respond in a graded manner, but in six of seven cases showed a stepwise change in foraging activity as pollen storage levels moved beyond a set point. Changes in colony performance resulted from changes in recruitment of new foragers to pollen collection, rather than from changes in individual foraging effort. Where we were able to track genotypic variation, increases in pollen foraging were accompanied by a corresponding increase in the genotypic diversity of pollen foragers. Our data support previous findings that genotypic variation plays an important role in task regulation. However, the stepwise change in colony behavior suggests that colony foraging flexibility is best explained by an integrated model incorporating genotypic variation in task choice, but in which colony response is amplified by social interactions. Received: 17 October 1998 / Received in revised form: 11 March 1999 / Accepted: 12 March 1999     

Morphological differences between females of different <Emphasis Type="Italic">Jaera</Emphasis> species (Crustacea: Asellota: Isopoda) in the White Sea: a possible solution to an old problem

It is well known that females of “Jaera albifrons” species complex cannot be determined on the basis of morphological analysis. In this work, an attempt to separate females of three Jaera species on the basis of morphometric characters, the length of so-called “coupling zones” (CZ), was made. Coupling zones are those parts of the partners’ bodies that contact during copulation. Female CZ is a region between the posterior edge of the pleotelson and the border between the fourth and the fifth thoracic segments, where openings of dorsal vaginas are situated. Male CZ extends from the anterior edge of the cephalon to the first abdominal segment carrying the copulative organ. Absolute and relative lengths of CZ were used. The latter was calculated as ratio of absolute length of CZ to body length. Two settlements of Jaera in the White Sea were studied for 3 years. Species composition of these populations was quite different (the first one was dominated by J. ischiosetosa and the second by J. albifrons and, 1 year by J. praehirsuta). Males of different species were shown to be distinct in terms of absolute and relative length of CZ. Frequency distributions of absolute and relative CZ length of females differed significantly in the two settlements. The experiments based on male choice in mixed samples were organized. They revealed that females chosen by males of different species differ significantly by CZ parameters.     

Spatial Components of Bird Diversity in the Andes of Colombia: Implications for Designing a Regional Reserve System

Abstract:  Beta diversity, or the turnover in species composition among sampling sites in a region, is an important criterion for obtaining adequate representation of regional biodiversity in systems of protected areas. Recently, the additive model for partitioning regional (gamma) diversity (in opposition to the multiplicative model) has been proposed because it allows a direct measure of the contribution of beta diversity to gamma diversity. We determined avian beta diversity along latitudinal (among neighboring river drainages) and elevational axes in a 1347-km² region on the western slope of the Central Cordillera of the Colombian Andes, where a regional system of protected areas is being designed. We then compared avian beta diversity between sites based on rapid versus long-term (>1 year) inventories and between fragmented sites versus continuous forest. Overall, beta diversity represented 63.1% of gamma diversity among 16 sites. Elevational differences in species composition accounted for 43.3% of regional diversity, whereas differences among drainages accounted for 19.8%. A complementary cluster analysis showed that sites grouped by elevational zones. Rapid inventories overestimated beta diversity because of sampling effects, but the effect was biologically small. Estimators of species richness derived from species accumulation curves provided a useful alternative to compensate for undersampling in short-term surveys. Forest fragmentation increased beta diversity because of differential local extinction of populations. Nevertheless, in our region, forest fragments contributed to gamma diversity because they contained complementary sets of species. More importantly, they contained populations of special-interest species. Although the region is relatively small, our analyses indicate that spatial differentiation of the biota is an important factor for deciding number and location of protected areas in the Andean region.     

Consequences of communal gall occupation and a test for kin discrimination in the aphid Tamalia coweni (Cockerell) (Homoptera: Aphididae)

This study investigated the consequences of communal gall induction on individual and group fitness in the aphid Tamaliacoweni. The possibilities that kin discrimination and foundress density are factors favoring communal gall occupation were examined. Clonally produced aphid foundresses were collected to create two treatments: clonal groups and groups of less closely related individuals. These were confined on suitable host plant tissue to compare their respective propensities towards communal behavior. There were no significant differences in the frequencies of communal gall occupation; therefore, active kin discrimination by T.coweni foundresses apparently does not play a role in their communal behavior, within the context of this experiment. In a second experiment, aphid foundress density on the host plant was manipulated in three treatments and was correlated with the frequency of communal gall occupation. Individual fitness was inversely related to the mean number of foundresses per gall. These results suggest that communal gall occupation does not necessarily represent mutual cooperation but may instead be the outcome of competition for limited gall sites on the host plant. However, natural selection at the clonal level may favor communal gall occupation under the conditions of resource limitation and high within-group relatedness. Received: 26 September 1997 / Accepted after revision: 28 March 1998     

Importance of heterotrophic planktonic communities in a mussel culture environment: the Grande Entrée lagoon,Magdalen Islands (Québec,Canada)

Mussel culture in coastal environments relies on the availability of food of sufficient quality and quantity. Both to determine this availability and to examine impacts that this aquaculture practice may have on the environment, it is important to have good knowledge of the type of plankton communities present in aquaculture sites. It is usually thought that phytoplankton make up the bulk of mussel diet in many of these sites. Here we show that the Grande-Entrée lagoon [Magdalen Islands, Gulf of St Lawrence (GSL), Canada], where commercial mussel culture has been on-going since 1980, differs from this pattern. Heterotrophic protists dominate for most of the summer-early fall season (apart from short diatom bursts), with a high average biomass of 160 mg C m⁻³. The dominance of small-sized phytoplankton cells (notably green algae), low nutrient concentrations (e.g. 0.3 μM NO₃⁻ on average) and high biomass of heterotrophic protists (mostly naked ciliates and tintinnids) all point to the importance of the microbial food web in this shallow marine environment. Sustained cultivation of suspended mussels in the lagoon suggests that these heterotrophic protists could be an important source of food for the mussels, supplementing the small amount of phytoplankton present.     

Maternal energy expenditure does not change with flight costs or food availability in the pied flycatcher (Ficedula hypoleuca): costs and benefits for nestlings

We manipulated parental work load without changing brood size in a population of pied flycatchers Ficedula hypoleuca by removing two primaries (7 and 9) from each wing of females, thus reducing wing area and increasing flight costs. At other nests, we offered supplementary food in the form of live mealworms (10–20 g daily from hatching) to reduce brood demand and thus parental foraging costs. Other nests were left as controls. The daily energy expenditure of females feeding 12-day-old nestlings was measured with doubly labelled water D₂ ¹⁸O. Females in both treatments expended the same amount of energy, fed at the same rate and had similar body masses to birds in the control group. No effect of treatment on male mass and feeding effort was detected. More nestlings, however, died in nests of handicapped females. Nestlings of handicapped females had significantly lower body mass and haematocrit values than nestlings in food-supplemented nests, with nestlings in control nests occupying an intermediate position. The effects of both treatments on nestling mass, haematocrit values and mortality rates were only noticeable in nests infested with mites. Maternal energy expenditure is apparently constrained and offspring pay the costs imposed by reduced provisioning rate or increased demand caused by ectoparasites, while receiving benefits when food supply improves. The presumption that avian reproductive costs derive from changes in a flexible energy output may not be met in many cases. Received: 24 October 1998 / Received in revised form: 15 March 1999 / Accepted: 26 April 1999