Aggression in female Red-Winged Blackbirds: A strategy to ensure male parental investment

Summary Female Red-winged Blackbirds (Agelaius phoeniceus) are often aggressive towards conspecific females during the breeding season. We hypothesize that the function of female-female aggression in this species is to guard the nonshareable portion of the male's parental investment.The investment-guarding hypothesis predicts that a female should be more aggressive toward another female evincing interest in mating with the territory-owning male than toward a female simply perching within the male's territory. Results of mount presentations to females with active nests confirmed this prediction. Nesting females attacked a stuffed conspecific female mounted in a precopulatory, soliciting posture significantly more often than a mount in a normal, perched posture.The male's nonshereable parental care consists of provisioning his young, and most of this care is invested in the brood of his primary (first-to-nest) female. It is therefore predicted that primary females should be more aggressive than secondary (later-nesting) females. Female mount presentations also confirmed this prediction. Primary females attacked the soliciting mount significantly more often than secondary females.     

Sexual selection based on egg colour: physiological models and egg discrimination experiments in a cavity-nesting bird

It has been proposed that the blue-green bird egg colourations of many avian species may constitute a sexually selected female signal that males can use to modulate their parental investment. A fundamental untested assumption for the validation of this hypothesis is that males can accurately assess differences in the colour of eggs. A recent review suggests that this could be particularly problematic when egg clutches were located within a dimly lit nest cavity, due to limitations of the visual system in low light conditions. Here, we first used a photoreceptor noise-limited model of colour discrimination ability that accounts for visual performance under low light conditions to study whether a typical cavity-nesting passerine, the spotless starling Sturnus unicolor, can discriminate their eggs under the ambient illumination in their nest-holes. Secondly, we tested the validity of model predictions with behavioural data collected in two egg discrimination experiments performed in this species. Estimated egg detectability depended entirely on model assumptions about visual limitations linked to light intensity. Starlings would not be able to discriminate egg differences in their nests if the model was based on the assumption that light intensity limited detectability, whereas they could potentially perceive as different many possible pairwise clutch comparisons if the model assumption was that light intensity did not limit detectability. Results of behavioural experiments fitted the prediction of the visual model where light intensity did not limit detectability. Our results suggest that photoreceptor noise-limited colour models based on stimulation of single photoreceptors cannot, at present, be used to predict egg discrimination ability in spotless starlings under low light conditions. Future studies aiming to test egg discrimination constraints in the frame of the sexual selection hypothesis should therefore combine both modelling and behavioural experiments to determine which are the components of the models that produce the mismatch with the behavioural conditions.     

Neighbor–stranger discrimination in Audubon's shearwater (Puffinus l. lherminieri) explained by a “real enemy” effect

Neighbor–stranger discrimination (NSD) occurs when animals respond with more aggression to strangers than to territorial neighbors. NSD has been reported in many species that defend multi-purpose territories for breeding and foraging, but it is rare among species that defend other types of territories. For birds that defend only their nest sites, there is no experimental evidence for NSD, and observational studies have provided mixed results. In a colony of Audubon's shearwaters (Puffinus l. lherminieri), I played back the calls of a neighbor and a stranger to males defending nest sites. Subjects responded with longer calls to playbacks of strangers than to those of neighbors. In shearwater colonies, strangers are often birds looking for future breeding sites. In contrast, there is no evidence that established breeders compete with their neighbors for any resources. Shearwaters should benefit from NSD because strangers represent a “real enemy” and established neighbors do not.     

Acoustically mediated neighbor recognition in the North American bullfrog,Rana catesbeiana

Summary Playback experiments were conducted to determine whether territorial male bullfrogs (Rana catesbeiana) were capable of discriminating between advertisement calls of neighbors and strangers. Territorial males adjacent to subject males were removed and replaced with a speaker which broadcast calls of a stranger or the removed male. Ten of 11 males responded more strongly to calls of strangers than to calls of neighbors. When advertisement calls of neighbors were broadcast to subject males from positions within the removed neighbor's territory and from positions opposite the subject male, all 9 males tested responded more strongly to calls of a neighbor broadcast from a new position than to playbacks of the same call broadcast within the removed neighbor's territory. Because male bullfrogs recognize familiar calls and associate these calls with a particular direction or location, these data provide the first evidence for acoustically mediated neighbor recognition in frogs.     

Large male advantage for access to females: evidence of male-male competition and female discrimination in a territorial salamander

Summary In the natural habitat visited in this study, adult male red-backed salamanders (Plethodon cinereus) found near females were significantly larger than adult males found alone; there was no evidence for size assortative associations. Previous research has indicated that females associate preferentially with males that occupy high quality territories and that larger individuals are more successful at obtaining higher quality areas. In laboratory experiments, when resource quality was held constant and males were restrained so that male-male interactions were prohibited, females discriminated behaviorally between large and small males by spending more time visually and chemically assessing larger males and more time apparently attempting to leave the chamber when near smaller males. Females were found near the larger males at the end of the trials significantly more often than predicted by chance. In a separate experiment in which only females were restrained, larger males spent significantly more time in aggressive postures when paired with smaller males than when alone with the female. Smaller males spent significantly more time in submissive postures when paired with larger males and more time near the female when alone. Therefore, large body size of males may positively affect both intra- and intersexual interactions and, ultimately, mating success of male P. cinereus.     

Independent effects of male and female density on sexual harassment, female fitness, and male competition for mates in the western mosquitofish Gambusia affinis

Operational sex ratio (the ratio of sexually active males to fertilizable females) has a major influence on male competition for mates and male–female interactions. The contributions of male and female density per se to mating system dynamics, however, are rarely examined, and the fitness consequences are often inferred rather than quantified. Male mosquitofish (Gambusia affinis) compete aggressively and frequently harass females for copulations, a behavior thought to reduce female fitness. Female fitness can also be reduced by increases in female density, which may affect food availability, cannibalism rates, and chemical interactions between females. I manipulated male and female densities of G. affinis to measure their effects on male–male aggression, male harassment toward females, and female fitness. I found that males chased rivals more often and attempted fewer copulations when female density decreased, but surprisingly male density had no significant effect on the frequency of these male behaviors. In contrast, males’ agonistic displays toward other males increased with male density, but display behavior was unaffected by female density. These results suggest that male and female density do not always contribute equally or at all to the patterns of behavior we observe. Female fitness declined as female density increased, the opposite pattern expected if male harassment is costly to females. This suggests that a strong, negative effect of female density overwhelmed any potential costs of male harassment. Sources of female density dependence and the consequences of changes in male and female density to patterns of male behavior are discussed.     

Sparring,relative antler size,and assessment in male caribou

Summary There are two kinds of antler combats in male deer. Fighting is rare, violent, occurs between matched males, and can cause injury and death. Sparring is common, usually gentle, often occurs between unmatched males, and involves no risk of injury. We recorded 1308 sparring matches and only 6 fights between wild male woodland caribou (Rangifer tarandus caribou) in 2 years of study. In the 713 sparring matches between males with unequal antlers, the animal with smaller antlers initiated close to half of the encounters, but terminated nearly 90% of them. That was true whether a sparring match involved two adults, two yearlings, or an adult against a yearling. We argue that the best interpretation of that shift in the decisions of the smaller-antlered male is that sparring serves to assess a partner's weight and strength relative to one's own. Thus, through sparring, antlers could be used to allow a form of tactile (proprioceptive) assessment of fighting ability, which could be the basis for subsequent visual assessment at a distance. Offprint requests to: C. Barrette     

Host selection,location, and association behaviors of anemonefishes in field settlement experiments

Nine anemonefish species were reared in the laboratory, and individuals were released in the field (Lizard Island, Australia and Madang, Papua New Guinea) at different distances and orientations away from natural (anemone species the fishes are found with in nature) and unnatural species of host anemones. Experiments were conducted to examine factors that could affect the settlement behaviors of the fishes: current velocity, distance and orientation of the fishes to the anemones, chemical vs visual cues, and presence of conspecific or heterospecific fishes. The fishes were usually attracted toward natural host species of anemones but not towards unnatural host anemone species nor to pieces of dead coral. Host selection during settlement provided the best explanation for the host specificity patterns displayed by anemonefishes in nature. The fishes used chemical cues released from the anemones to identify and locate the appropriate host species and could effectively locate the anemones from a maximum distance of 8 m downstream. Fishes released upstream or to the side of anemones (where anemone chemicals were assumed to be reduced or absent) were much less successful in locating anemones. The ability of the fishes to locate natural host anemones was strongly reduced when there was no water current. The presence of resident anemonefishes on host species of anemones did not influence the attraction behavior of anemonefishes released downstream from the anemones. Once the released fishes got close to or entered the anemones, the resident fishes would generally bite and chase them until the recruits left the anemones. Most fishes were not stung upon initial contact with the anemones.     

Long-term experiments on lifespan, reproductive activity and timing of reproduction in the Arctic copepod Calanus hyperboreus

Several experiments were conducted with starved and fed females of the Arctic copepod Calanus hyperboreus (1) to investigate their lifespan, reproductive period, egg production and egg viability; (2) to study the effect of origin, i.e., Atlantic and Arctic Waters in the Greenland Sea, on the timing of reproduction; and (3) to study the effect of time of collection on the onset of reproductive activity as a first approach to study control mechanisms of the reproductive cycle. Females collected in October produced up to 1,000 eggs and had a maximum lifespan of 164 days without feeding, whereas fed females produced up to ca. 6,000 eggs and survived up to 806 days. These observations support earlier assumptions that females were multiannual-iteroparous, i.e., capable to spawn in successive years, which would be unique for calanoid copepods. In starved females, clutch size decreased significantly with each spawning event. Viable eggs were produced during most of the life time. There was no difference in the timing of reproductive activity between females from the West Spitsbergen Current and the Greenland Sea Gyre. Fed and starved females collected in May and June began to spawn circa 2 and 4 months after collection, respectively, whereas females collected in August and October started spawning at the same time, in the middle of October. This indicates initiation of reproductive activity in the field in August, coincident with the descent into deep waters. Potential cues for the untimely spawning of females collected in spring and ‘unnatural’ feeding in fall experiments are discussed. Their large size, robustness and combination of different types of diapause in their life cycle make C. hyperboreus a good model organism to study diapause control mechanisms.     

华南土壤对五氯苯酚吸附的行为及其控制因素研究

应用批量吸附实验法研究了华南地区7个红壤剖面对PCP的吸附最ΓPCP与剖面深度的关系，以及与土壤的理化性质、化学组成、矿物组成和形态的关系。双变量积矩相关分析发现吸附量ΓPCP与土壤pHw值、TOC呈非常显著相关，与AAOFe、AAOAI和DCBAI呈显著相关，但偏相关分析和剔除法逐步线性同归分析清楚显示，控制吸附最ΔPCP的4个最为重要因素是土壤pHw值、TOC、非晶态氧化铝（AAOAI）和砂粒矿物相（主要是石英）含量，其次是粘土矿物（主要是高岭石和水云母）、CaCO3和晶态氧化铁矿物（DCBFe）的含量。随着深度增加，土壤对PCP的吸附量ΓPCP明显下降。不同的剖面，吸附量有较明显差别，其与土壤pHw值和TOC等含量变化明显相关。     

The effects of mating on infanticide and pup discrimination in white-footed mice

Summary A laboratory study was conducted with white-footed mice (Peromyscus leucopus) to determine (1) if cohabitation and/or successful mating inhibits infanticide, and if so for how long, and (2) if males and females can discriminate between their own (familiar) and strange (unfamiliar) pups. Infanticidal males and females that were paired for 15 days and did not reproduce were not inhibited from killing pups, whereas infanticidal males and females that were paired until they reproduced were inhibited from committing infanticide at 4 days postpartum. Neither males nor females discriminated between familiar and unfamiliar pups with regard to engaging in infanticide. Approximately 50% of wild-caught males and females killed pups in the laboratory. Infanticidal pairs bred significantly sooner than did noninfanticidal pairs. The results are consistent with the promiscuous nature of P. leucopus males and territoriality of P. leucopus females. Offprint requests to: Donna M. Cicirello, Kellogg Biological Station, 3700 E. Gull Lake Drive, Hickory Corners, MI 49060, USA     

Nutritional and related experiments on laboratory maintenance of three species of symbiont-bearing,large foraminifera

Factors were examined that affect survival and growth of two common species of large foraminifera from the Red Sea,Amphisorus hemprichii Ehrenberg andAmphistegina lobifera Larsen, 1976. The former is host for dinoflagellate and the latter for diatom zooxanthellae. Experimental conditions were modeled on conditions at 25 m during spring at Wadi Taba, Gulf of Elat, Israel, the season and site where the experimental organisms were collected between 1983 and 1988. The two species responded quite differently in nutritional experiments.A. hemprichii grew, on average, 0.270 mm in diameter in 3 mo on a diet ofNitzschia subcommunis Hustedt,Chlorella sp. (clone AT) orCylindrotheca closterium Rabenhorst isolated from their native habitat. Unfed controls did not grow. In contrast, unfed populations ofA. lobifera grew as well or better than those that were fed unialgal diets. Growth of both species was enhanced on particular mixed algal diets. Both species required photosynthetically active symbionts. Even when fed weekly and supplied with nutrients, neither species survived in the dark. All individuals ofA. hemprichii died after 8 wk incubation in the dark;A. lobifera survived longer, but all were dead by 13 wk. The highest growth rate ofA. hemprichii (0.037 mm wk^–1) was obtained when they were fed, the medium was enriched, and the medium was changed weekly. All other conditions being the same, growth rate dropped to 0.009 mm wk^–1 when the medium was changed every 3 wk. In contrast,A. lobifera grew fastest when the medium was changed every 3 wk. Food or enrichment with nitrate or phosphate did not stimulate growth (0.03 mm wk^–1) over that of the controls. Specimens ofMarginopora kudakajimensis Gudmundsson from Japan, another dinoflagellate-bearing species, were also tested. They grew best (0.02 mm wk^–1) when cultured in light, in media enriched with nitrate and phosphate changed weekly, and fed. All three species withdrew nitrate and phosphate from the medium in chemostat experiments.     

Effects of temperature on success of (self and non-self) fertilization and embryogenesis in Diploria strigosa (Cnidaria, Scleractinia)

The effects of seawater temperature on the degree of success of self-fertilization versus cross-fertilization and on embryogenesis were investigated in the scleractinian coral Diploria strigosa (Dana 1846). Gametes from nine colonies were collected from the Flower Garden Banks reefs, a set of coral reefs in the northern Gulf of Mexico, ~110 nm offshore from Texas, United States. Gametes from the coral colonies were combined in all possible combinations and placed in 30°, 31°, and 32°C water baths on successive nights, respectively. Each cross, whether selfed or out-crossed, and each temperature treatment, was replicated twice. A high frequency of successful self- and cross-fertilization was observed at all temperature treatments and in both blocks. Higher temperatures, however, commonly produced numerous developmental aberrations during embryogenesis of the larvae. Thus, although fertilization rates can remain high under high temperature conditions, if temperatures remain high for several days, embryonic development and larval viability may be expected to decrease dramatically. We propose that the success of coral larval development may be diminished in areas where abnormally high sea surface temperatures occur during the spawning season. We also propose that highly successful selfing may enhance the abundance of locally adapted genotypes, which in turn may be advantageous where reefs, such as the Flower Garden Banks, are relatively geographically isolated.     

Reproductive trade-off in male Antarctic krill,Euphausia superba

Freshly caught male and female Euphausia superba from the same swarm exhibited different rates of mortality subsequent to capture. Mortality was significantly higher for reproductive males (100%, n=68) than for females (3%, n=186) within the first 3 d of capture. Total lipid and triacylglycerol levels in male, female and juvenile Euphausia superba were analysed and compared. All reproductive male krill analysed from this swarm had low lipid levels (1 to 3% dry weight) with negligible triacylglycerol stores (0 to 2% of total lipid). Somatic lipid stores in female and juvenile krill ranged from 8 to 30% of which up to 40% was triacylglycerol. The levels of algal sterols in the digestive gland of males, females and juveniles indicate that all krill had been feeding recently. An analysis of the sex ratio of krill catches derived from data collected over seven summers from the Prydz Bay region showed a decrease in the proportion of males with increasing size. There was a sharp decline in numbers of male krill once they attained a length of 51 to 55 mm. Low lipid levels in redroductive male krill may be due to reproductive costs. The resulting low storage-lipid levels are accompanied by high mortality in male krill.     

Age- and state-dependent reproductive effort in male mountain goats, Oreamnos americanus

In mammalian polygynous mating systems, male reproductive effort consists mainly of male–male competition and courting of females, which entail substantial somatic costs. Males are thus expected to adjust their reproductive effort according to their age and condition. In this study, we examined how activity budgets of male mountain goats (Oreamnos americanus), a polygynous ungulate, varied with age in a marked population over two periods: (1) summers 1995–2006 and (2) ruts 2004–2006. We then assessed if the proportions of time spent in male–male competition and courtship behaviors were influenced by age-specific body mass and social rank during the rut. Males spent most of their time foraging and resting during summer, and rested more and foraged less with increasing age. During the rut, pronounced shifts in activity budgets occurred as juveniles (1–2 years) increased time spent foraging, whereas adults (≥3 years) increased standing and time spent in social interactions at the expense of foraging. At old age, reproductive effort either stabilized or decreased slightly, providing weak support for the ‘mating strategy–effort’ hypothesis, predicting that courtship behaviors should peak in prime-aged males. Age-specific body mass did not affect time spent in male–male competition, but was positively related with time spent in courtship behaviors, providing support for the ‘individual quality’ hypothesis, predicting that males with more resources at the start of the rut should spend more time in mating-related activities. Age-specific social rank did not affect reproductive effort. Surviving to prime age while increasing mass each year should thus allow male ungulates to gain greater ability to court estrus females.     

Preliminary rearing experiments on the larvae of Sergestes lucens (Penaedia,Natantia, Decapoda)

Sergestes lucens Hansen, a mesopelagic shrimp fished commercially in Suruga Bay, Japan, was successfully reared from egg to post-larval stage V under laboratory conditions. Chaetoceros ceratosporum and Artemia nauplii were found to be satisfactory food in the laboratory during rearing. Growth, mortality, food preference, and feeding and swimming activities during the various developmental stages were investigated. Temperature changes greatly affected the speed of development and the mortality of the larvae. The optimum temperature range for larval development was 18° to 25°C. The growth rate (length) of larval stages was as rapid as 0.16mm/ day at 20 °C and 0.21 mm/day at 23 °C. The larvae first started feeding on phytoplankton at elaphocaris stage I, and then gradually became predators in the post-larval stages. It is suggested that the critical period for the species occurs in the elaphocaris stages. Environmental data, vertical distribution of the species, and data obtained from laboratory experiments suggest that the fluctuation in the abundance of S. lucens is greatly influenced by the water temperature at around 50 m from June to August. Feeding mechanisms observed in the post-larval stages are described.     

Parasite faunas,testosterone and secondary sexual traits in male red-winged blackbirds

Summary We examined associations among parasite infections, secondary sexual traits and testosterone in male red-winged blackbirds sampled at the start of the breeding season. Parasites quantified included ectoparasitic lice and mites and endoparasitic blood protozoans, nematodes, trematodes and cestodes. Secondary sexual traits that we quantified included body size, epaulet size and color, song repertoire size and song switching rate, and behavioral responses to male and female models. Overall we found few significant associations between parasites and secondary sexual traits, between secondary sexual traits and testosterone, or between parasites and testosterone. In addition, most parasite taxa appeared to infect birds independently, although the low prevalence (<50%) of many of the parasites meant that our sample sizes were too small to detect weak associations. Our most promising results were obtained for ectoparasitic mites, which tended to occur on birds uninfected with other parasites, on birds with longer epaulets, and on birds with higher levels of testosterone. Epaulet length and testosterone are both probable correlates of dominance in this species. Further research will be required to determine whether there is a causal link between the immunosuppressive effects of testosterone and the mite infections, and between testosterone, epaulet length and male mating success. Correspondence to: P. Weatherhead     

Dominance rank,female reproductive synchrony,and male reproductive skew in wild Assamese macaques

In groups with multiple males, direct mate competition may select for the evolution of dominance hierarchies that sort males into a queue for access to fertile females. The priority-of-access (PoA) model proposed by Altmann in 1962 makes explicit predictions about the resulting paternity distribution based on an interaction between male dominance rank and the overlap of female receptive phases. Here, we investigated whether the logic of the PoA model predicted the distribution of paternity across ranks in a seasonal breeder with high reproductive synchrony over six consecutive mating seasons. We studied 18 males that resided in a group of wild Assamese macaques (Macaca assamensis) in their natural habitat at Phu Khieo Wildlife Sanctuary, Thailand, between 2006 and 2011 with 5 to 13 conceptions per season. We assessed whether mate guarding increased paternity success, described “short-term” deviations from predicted paternity distribution, and examined how these are related to the number of competitors and fertile females. We determined genetic paternity of 43 (93 %) offspring born into the study group and found reproductive skew to be relatively low with 29 % alpha male paternity in accordance with the high degree of female reproductive synchrony observed. Short-term deviations from expected paternity distribution over ranks were not explained by the number of resident males or the number of conceiving females or their interaction. Within the limits of this study, these results suggest that even if males cannot discern female fertile phases, if reproduction is seasonal, and if reproductive synchrony is high, males may also compete directly over access to females.