Nestling American robins compete with siblings by begging

Summary The evolution of intense begging by dependent nestling birds has recently been the subject of several theoretical papers. The interesting problem here is that nestlings should be able to communicate their nutritional status to parents in ways that are less costly energetically and less likely to attract predators. Thus, conspicuous begging behaviour is thought to have evolved as a result of either competition among nestmates or the manipulation of their parents to provide more food than would otherwise be favoured by selection. We studied sibling competition for parental feedings in the American robin (Turdus migratorius). We demonstrate that the probability that an individual nestling received food was related to several indices of begging. When we experimentally prevented parents from feeding part of their brood, both the intensity of begging and the number of feedings subsequently received by food-deprived nestlings increased. Furthermore, the begging intensity of those nestlings that were not food-deprived also increased in response to the begging of their hungrier siblings.Offprint requests to: R. Montgomerie     

Parental care of nestlings by male red-winged blackbirds

Summary Nestling feeding by males is less common among birds with polygynous mating systems than in monogamous species, because of the pronounced trade-off between parental behavior and the attraction of additional mates. In this study, however, we found that male red-winged blackbirds (Agelaius phoeniceus) commonly assisted females in feeding nestlings in several Ontario marshes. Male parental care was additional to that provided by females, and it significantly enhanced the fledging success of nests (Table 2). Male redwings did not help to feed all nests on their territories: primary and secondary nests were much more likely to receive male parental care than tertiary and later nests. Contrary to expectation, male parental care was not restricted to the nests of primary females: a greater proportion of secondary than primary nests were assisted (Tables 4a and b). The presence of new females settling on the territory at the same time that a resident female had nestlings, resulted in males deferring parental care until a later brood. This suggests that males trade off the recruitment of females against parental care to an existing brood. Although the number of nestlings fledging from a male's territory was strongly influenced by the number of females in the harem, males could additionally increase their reproductive success by feeding nestlings in one or more nests on their territories (Fig. 2). The reproductive success of females was significantly enhanced by male assistance in feeding nestlings (Table 3). However, those females not receiving male assistance on territories of feeding males did not suffer a significantly reduced reproductive success in comparison to females on territories of non-feeding males (Table 2). Males varied considerably in the quality of brood care given. We therefore suggest that the quality of male parental care may be a factor considered by females in choosing a breeding situation.     

Begging behavior and food acquisition by brown-headed cowbird nestlings

Understanding the selective forces that limit the exaggeration of begging signals is a critical issue in understanding the evolution of begging behavior. I studied the begging behavior of nestlings of the brown-headed cowbird (Molothrus ater), a brood parasite. In the nests of indigo buntings (Passerina cyanea), brown-headed cowbird nestlings received approximately twice as much food per hour than their host nestmates. I tested three hypotheses for the mechanism by which cowbirds acquired more food than their bunting nestmates: the size advantage hypothesis, the signal exaggeration hypothesis, and the novel begging behavior hypothesis. I found support for the hypotheses that cowbirds acquire more food as a result of their larger body size, and due to the exaggeration of begging signals that are not dependent on body size. I did not find support for the role of novel begging behaviors in cowbird food acquisition. These results suggest that food acquisition by host chicks in unparasitized nests could be increased by the exaggeration of begging signals. Recent work suggests that such exaggeration may be limited by the risk of nest predation, but further studies are needed. Received: 12 December 1997 / Accepted after revision: 29 December 1997     

Evidence of kin-selected tolerance by nestlings in a siblicidal bird

Behaviorally dominant members of blue-footed booby (Sula nebouxii) broods can effect siblicide by restricting access of subordinate siblings to parents providing food. In spite of their capacity for siblicide, dominant chicks permit subordinates to feed during short-term food shortage; in fact, the proportion of the food that the dominant takes is independent of the total amount delivered in older chicks. A model of optimal food distribution suggests that dominant chicks maximize their inclusive fitness with this pattern, rather than by satisfying their own food requirements and leaving what remains for the subordinate sibling. The indirect reproductive potential represented by a chick's sibling appears to have influenced the evolution of siblicidal brood reduction in this species.     

How the mountain chickadee procures more food in less time for its nestlings

Summary Parent mountain chickadees (Parus gambeli) visit the same foraging location on consecutive flights more frequently than expected by chance. This pattern of repetitive return to a foraging location does not increase the frequency of capturing the same prey on consecutive flights. The quantity of prey captured on a visit also does not change significantly with repeated visits to a foraging location. Foraging time, however, is significantly shorter during repeated visits to a foraging location. Foraging time is therefore a major determinant of whether a run of consecutive flights to a location occurs. Decreased foraging time is also a benefit of selecting the same prey type on consecutive flights. Over the course of a day, the prey types returned on the greatest percentage of flights are returned multiply per flight. These prey are usually small and are probably present at high densities in the environment. Large prey size is the second best predictor of frequency of a prey type in the diet. Short foraging time to capture a prey is the third best predictor. Thus prey which can be readily obtained in large numbers, even if they are small prey, are preferentially returned to older nestlings. The mountain chickadee uses information on foraging time to decide when to move to a new foraging location or when to pursue a different type of prey. These decisions should increase the amount of time available for prey gathering.     

Great spotted cuckoo fledglings are disadvantaged by magpie host parents when reared together with magpie nestlings

The post-fledging period is a critical phase for juvenile survival, and parental care provided during this period is a key component of avian reproductive performance. Very little is known about the relationships between foster parents and fledglings of brood parasites. Here, we present the results of a 5-year study about the relationships between fledglings of the non-evictor brood parasitic great spotted cuckoo (Clamator glandarius) and its magpie (Pica pica) foster parents. Sometimes, great spotted cuckoo and magpie nestlings from the same nest can fledge successfully, but most often parasitic nestlings outcompete host nestlings and only cuckoos leave the nest. We have studied several aspects of cuckoo post-fledging performance (i.e. feeding behaviour, parental defence and fledgling survival) in experimental nests in which only cuckoos or both magpie and cuckoo nestlings survived until leaving the nest. The results indicate that great spotted cuckoo fledglings reared in mixed broods together with magpie nestlings were disadvantaged by magpie adults with respect to feeding patterns. Fledgling cuckoos reared in mixed broods were fed less frequently than those reared in only cuckoo broods, and magpie adults approached less frequently to feed cuckoos from mixed broods than cuckoos from only cuckoo broods. These results imply that the presence of host's own nestlings for comparison may be a crucial clue favouring the evolution of fledgling discrimination; and furthermore, that the risk of discrimination at the fledgling stage probably is an important selection pressure driving the evolution of the arms race between brood parasites and their hosts.     

Provisioning of nestlings by dunnocks,Prunella modularis,in pairs and trios compensation reactions by males and females

Summary The mating systems in a population of dunnocks included monogamy, polygyny, polyandry and polygynandry. Broods were provisioned by lone females, by females with the part-time help of one male, by females with the full-time help of one male (pairs) or by females with the part-time or full-time help of two males (trios). We compared provisioning of nestlings in these different systems and used the results of natural and experimental removals to examine how individuals reacted to the provisioning by others. In pairs, males and females provisioned at similar rates (Fig. 2a). In trios, females and alpha males fed chicks at the same rate, and at a higher rate than beta males (Fig. 2b). Females attained maximum male help when two males shared provisioning equally (Fig. 6). Females were more likely to achieve this when they were able to escape alpha male guarding during the mating period and exercise some control over mating access by the two males (Fig. 7). Female provisioning did not increase when their help was reduced from two males to one (Fig. 4 a), but it did increase when male help was further reduced to either one male's part-time help (Fig. 5 a) or no male help (Fig. 3 a). Beta males increased their provisioning rate when alpha males were removed from a trio (Fig. 4b). Alpha males may have increased their rate when beta males were removed but any reaction was small. When one male was removed from a trio, the remaining male's provisioning rate was as expected for a male in a monogamous pair. When reactions occurred, they were not sufficient to compensate for the loss of the removed adult and nestling weight was reduced. We discuss the relevance of these results to the maintenance of parental care by pairs and trios.     

An experimental study of competition for food between male and female nestlings of the red-winged blackbird

Summary Ten experimental broods of red-winged black-bird (Agelaius phoeniceus) nestlings, experimentally manipulated to contain two males and two females of similar age, were observed with the aid of video cameras to determine (a) which nestling characteristics were most important in influencing its chances of being fed and (b) if males and females differed with respect to these characteristics. Nestlings that could reach highest while begging were most successful at obtaining food from parents during individual feeding bouts. However, while there was a tendency for nestlings that begged earlier to be offered food first, a nestling's position in the nest during these bouts did not influence its feeding success. Males begged more than often females and were offered more food by parents. Males were also more likely to be fed when they begged. This was most likely because males, being larger than females, were able to reach higher while begging. Neither sex was more likely to beg sooner or occupy certain positions in the nest. However, while males that had been transferred from other nests received less food than natal males, there were no differences in food acquisition between transferred and natal females. Differences in the relative success of males and females under different conditions may help explain differences in fledgling sex ratios observed in many dimorphic species.     

Local and meso-scale patterns of recruitment and abundance of two intertidal crab species that compete for refuges

Interference competition for limited habitat or refuges is known to produce density-dependent mortality and generate patterns of micro-habitat distribution. While in mobile species the outcome of interference at a local scale can usually be determined from differences in body size and behavior, the population-level consequences of such interactions vary depending on rates of settlement and recruitment at a site, which are not directly correlated to local reproductive success. Previous experimental studies in central Chile demonstrated that interference competition for refuges is the primary factor driving microhabitat segregation between the predatory crabs Acanthocyclus gayi and Acanthocyclus hassleri, with the latter species monopolizing galleries inside mussel beds and excluding A. gayi to rock crevices. Between April 2001 and March 2006 we quantified monthly recruitment rates in artificial collectors at 17 sites over 900 km of the central coast of Chile. Results show that recruitment rates of A. hassleri are almost two orders of magnitude lower than those of A. gayi, and that they are tightly and positively correlated among sites across the region, suggesting that at scales of kilometers larval stages of these species are affected by similar oceanographic processes. Total crab densities per site were also positively correlated between species and strongly associated to mussel cover, with overall low crab densities at all sites where mussel cover was lower than about 60%. At all sites with mussel cover >60%, the ratio of A. gayi to A. hassleri density progressively decreased from recruits (2.6) to juveniles (0.5) to adults (0.04), overcoming initial differences in recruitment rates. The relative success of the inferior competitor at sites with low mussel cover does not appear to provide a potential mechanism favoring regional coexistence through dispersal to other sites (“mass effects”), because their densities were lower than at sites of high mussel cover. Yet, at many sites of low mussel cover the dominant competitor is virtually absent, allowing A. gayi to attain larger population sizes at the scale of the region. Thus, the factors limiting the dominant competitor from successfully utilizing other microhabitats seem to be the most critical factor in promoting both local and regional coexistence between these species.     

Habitat-mediated variation in predation risk by the American marten

The probability of prey encounter, attack, capture, and kill are often hypothesized to depend on habitat structure, but field evidence in terrestrial systems is rare. We tested whether predation efficiency by the American marten (Martes americana) and fear of predation by their primary prey, the red-backed vole (Clethrionomys gapperi), differed between 20- to 50-year-old regenerating forest stands and older uncut stands. Our results showed that the frequency of prey encounter, prey attack, and prey kill were higher in old uncut forests, despite the fact that small-mammal density was similar to that in younger logged forests. These differences in predation efficiency were linked to higher abundance of coarse woody debris, which seems to offer sensory cues to martens, thereby increasing the odds of hunting success. Red-backed voles in regenerating forest stands exhibited increased wariness compared to voles living in old uncut forest, suggestive of a behavioral response to habitat-mediated variation in predation risk.     

Identifying climate change refugia for South American biodiversity

Refugia-based conservation offers long-term effectiveness and minimize uncertainty on strategies for climate change adaptation. We used distribution modelling to identify climate change refugia for 617 terrestrial mammals and to quantify the role of protected areas (PAs) in providing refugia across South America. To do so, we compared species potential distribution across different scenarios of climate change, highlighting those regions likely to retain suitable climatic conditions by year 2090, and explored the proportion of refugia inside PAs. Moist tropical forests in high-elevation areas with complex topography concentrated the highest local diversity of species refugia, although regionally important refugia centers occurred elsewhere. Andean–Amazon forests contained climate change refugia for more than half of the continental species’ pool and for up to 87 species locally (17 × 17 km² grid cell). The highlands of the southern Atlantic Forest also included megadiverse refugia for up to 76 species per cell. Almost half of the species that may find refugia in the Atlantic Forest will do so in a single region—the Serra do Mar and Serra do Espinhaço. Most of the refugia we identified, however, were not in PAs, which may contain <6% of the total area of climate change refugia, leaving 129–237 species with no refugia inside the territorial limits of PAs of any kind. Our results reveal a dismal scenario for the level of refugia protection in some of the most biodiverse regions of the world. Nonetheless, because refugia tend to be in high-elevation, topographically complex, and remote areas, with lower anthropogenic pressure, formally protecting them may require a comparatively modest investment.     

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     

Feeding by larvae of intertidal invertebrates: assessing their position in pelagic food webs

One of the leading determinants of the structure and dynamics of marine populations is the rate of arrival of new individuals to local sites. While physical transport processes play major roles in delivering larvae to the shore, these processes become most important after larvae have survived the perils of life in the plankton, where they usually suffer great mortality. The lack of information regarding larval feeding makes it difficult to assess the effects of food supply on larval survival, or the role larvae may play in nearshore food webs. Here, we examine the spectrum of food sizes and food types consumed by the larvae of two intertidal barnacle species and of the predatory gastropod Concholepas concholepas. We conducted replicated experiments in which larvae were exposed to the food size spectrum (phytoplankton, microprotozoan and autotrophic picoplankton) found in nearshore waters in central Chile. Results show that barnacle nauplii and gastropod veligers are omnivorous grazers, incorporating significant fractions of heterotrophs in their diets. In accordance with their feeding mechanisms and body size, barnacle nauplii were able to feed on autotrophic picoplankton (<5 microm) and did not consume the largest phytoplankton cells, which made the bulk of phytoplankton biomass in spring-summer blooms. Balanoid nauplii exhibited higher ingestion rates than the smaller-bodied chthamaloid larvae. Newly hatched C. concholepas larvae also consumed picoplankton cells, while competent larvae of this species ingested mostly the largest phytoplankton cells and heterotrophic protozoans. Results suggest that persistent changes in the structure of pelagic food webs can have important effects on the species-specific food availability for invertebrate larvae, which can result in large-scale differences in recruitment rates of a given species, and in the relative recruitment success of the different species that make up benthic communities.     

The systematic position of some boring sponges (Demospongiae,Hadromerida) studied by molecular analysis

A phylogenetic analysis of some bioeroding sponges of the family Clionaidae (Order Hadromerida) was performed to resolve some taxonomic problems both at the specific and the supraspecific level using the D2 and D3 regions of 28S rDNA. Species belonging to the genera Cliona, Cliothosa, Spheciospongia (fam. Clionaidae) and Diplastrella (fam. Spirastrellidae) from the Mediterranean Sea and Celebes Sea (Indonesia) were analysed. In the phylogenetic tree, the species clustered on two main branches, one comprising Cliona celata, C. rhodensis, C. utricularis, and Cliothosa hancocki, and the other made up of C. viridis, C. nigricans, C. schmidti, C. jullieni, Spheciospongia solida and S. vagabunda. Above the species level, data do not support the separation of the genus Cliothosa from Cliona, while they do support the inclusion of some massive boring species, previously assigned to the genus Spirastrella, in the family Clionaidae. At the species level, data demonstrated the genetic identity of taxa C. viridis and C. nigricans, in spite of their considerable morphological differences. In contrast, the yellow species commonly attributed to C. celata are probably to be considered as a complex of sibling species with a number of distinct taxa present in the Mediterranean. Data also showed the identity of the Mediterranean and Pacific populations of C. schmidti, suggesting the status of a Tethyan relict for this species.     

Can pigeons be fooled about the actual release site position by presenting them information from another site?

Summary The aim of the experiment was to test the hypothesis that pigeons depend on route- and/or site-specific airborne parameters to establish their position relative to the loft. Pigeons were transported to the release site with free access to the environmental air. They were then enclosed in large airtight containers filled with air from the release site and either transported via the loft to a release site in the opposite direction with respect to the loft (experimental birds) or via the loft back to the same site (control birds). Before release the pigeons (still inside the containers) were made anosmic by a local anesthetic applied to the olfactory membranes through the nostrils. Vanishing bearings of experimental birds were on the average in a direction opposite to home while the vanishing bearings of control birds were homeward oriented. For this initial orientation the pigeons seem to rely on airborne cues obtained at a site where they last had access to ambient air or cues obtained from the air inside the sealed containers. Irrespective of the treatment the bearings pooled with respect to north still show a residual orientation to the NE. There was no difference either in the homing speeds or in the homing times of anosmic control and experimental pigeons. We therefore have to assume further mechanisms guiding the pigeons home in addition to a possibly olfactory one.     

A Case for Using Plethodontid Salamanders for Monitoring Biodiversity and Ecosystem Integrity of North American Forests

Abstract: Terrestrial salamanders of the family Plethodontidae have unique attributes that make them excellent indicators of biodiversity and ecosystem integrity in forested habitats. Their longevity, small territory size, site fidelity, sensitivity to natural and anthropogenic perturbations, tendency to occur in high densities, and low sampling costs mean that counts of plethodontid salamanders provide numerous advantages over counts of other North American forest organisms for indicating environmental change. Furthermore, they are tightly linked physiologically to microclimatic and successional processes that influence the distribution and abundance of numerous other hydrophilic but difficult-to-study forest-dwelling plants and animals. Ecosystem processes such as moisture cycling, food-web dynamics, and succession, with their related structural and microclimatic variability, all affect forest biodiversity and have been shown to affect salamander populations as well. We determined the variability associated with sampling for plethodontid salamanders by estimating the coefficient of variation (CV ) from available time-series data. The median coefficient of variation indicated that variation in counts of individuals among studies was much lower in plethodontids (27%) than in lepidoptera (93%), passerine birds (57%), small mammals (69%), or other amphibians (37–46%), which means plethodontid salamanders provide an important statistical advantage over other species for monitoring long-term forest health.