Parental investment decision rules and the Concorde fallacy

Summary Investment theory states that animals should base their parental investment decisions on expected benefits, and not on whether or not past investment will be wasted. Otherwise, they would comnit the Concorde fallacy. If reproduction has a cost, however, then past investment and expected benefits are necessarily confounded. Assuming a cost of reproduction, animals will be selected to maximize their remaining lifetime reproductive success, subject to a tradeoff between present and future reproduction (Williams' principle). We extend Williams' principle and develop an experimental design that would allow past investment and expected benefits to be varied independently. This design illustrates the importance of the value of the brood relative to the value of future reproduction.     

Paternity and condition affect cannibalistic behavior in nest-tending bluegill sunfish

Theory of parental care evolution predicts that a parent should invest more in a brood when its fitness value is greater than alternative investments such as the parent's own survivorship or future broods. In fish, filial cannibalism (eating one's own offspring) is widespread and represents a challenge to parental care evolution. In this study, I investigated filial cannibalism in bluegill sunfish (Lepomis macrochirus). Bluegill are characterized by alternative mating tactics referred to as "parentals" and "cuckolders". Parentals delay maturation, construct nests, court females and provide sole parental care for the developing offspring. Cuckolders mature precociously and parasitize parentals using two tactics called "sneakers" and "satellites". I found that parentals that obtained fewer eggs during spawning appeared more likely to completely cannibalize their brood (total filial cannibalism: P=0.07), regardless of their condition. Among parentals that provided care, partial cannibalism was greater during the egg phase as compared to the fry phase of care, but it was unrelated to brood size. Throughout the care period, parentals in better condition were less likely to partially cannibalize their brood, indicating that parentals use cannibalism to replenish energy reserves. Independent of condition, parentals that were cuckolded more were more likely to eat part of their brood. This relationship was evident only after the eggs had hatched, which is consistent with data showing that parentals can use olfactory cues produced by fry but not eggs to assess their paternity. This latter result proposes that parentals may be selectively culling cuckolder offspring from their nest. These data provide empirical support for parental care theory, and the first evidence for the importance of paternity on cannibalistic behavior.Communicated by M. Abrahams     

Parental investment without kin recognition: Simple conditional rules for parent-offspring behavior

Species differ widely with regard to parental investment strategies and mechanisms underlying those strategies. The passing of benefits to likely offspring can be instantiated with a number of different computational and behavioral systems. We report results from an agent-based model in which offspring maintain proximity with parents and parents transmit benefits to offspring without the capacity of either parent or offspring to 'recognize' one another. Instead, parents follow a simple rule to emit benefits after reproducing and offspring follow a simple rule of moving in the direction of positive benefit gradients. This model differs from previous models of spatial kin-based altruism in that individuals are modeled as having different behavioral rules at different life stages and benefits are transmitted unidirectionally from parents to offspring. High rates of correctly directed parental investment occur when mobility and sociality are low and parental investment occurs over a short period of time. We suggest that strategies based on recognition and bonding/attachment might serve to increase rates of correctly directed parental investment under parameters that are shown here to otherwise lead to high rates of misdirected and wasted parental investment.     

Inadvertent errors and error-constrained optimization: fallible foraging by bluegill sunfish

Summary When the expected reward rate is continuously reduced by foraging in a patch, foragers may adjust their patch persistence times to maximize the average long-term reward rate. The marginal-value model predicts the optimal persistence time for this situation. But real foragers may be unable consistently to achieve a precise persistence time. If the costs of under- and over-persistence differ, or if the resulting distribution of persistence times is skewed, a sufficiently broad persistence-time distribution can substantially shift the actual optimum. Moreover, this error-constrained optimum depends on the variable used by the forager to decide when to leave the patch (e.g., on persistence time per se, cumulative number of prey eaten, or instantaneous feeding rate). Here, we analyze laboratory data from bluegill sunfish (Lepomis macrochirus) foraging on larval-midge prey (Chironomus riparius) in patches of artificial vegetation, and we explore some wider implications of a model that seems to fit the data. The bluegills stayed 4%–157% longer in patches than predicted by the marginal value theorem. This behavior closely matched numerical solutions based on the observed variability of persistence times and the assumption that departures were cued by instantaneous feeding rate. On the other hand, the other two mechanisms that we investigated (i.e., persistence time per se and cumulative number of prey eaten) predict weak to moderate underpersistence relative to the marginal-value predictions, patterns quite unlike those observed. Surprisingly, the instantaneous-rate mechanism yields roughly a 10% lower over-all maximal reward rate than would either of the other two departure-cuing mechanisms. The modeling analysis documents the considerable sensitivity of our results to (1) the departure-cuing mechanism, (2) the shape of the frequency distribution of the departure-cuing variable, (3) the way that the shape of this distribution shifts as its mean changes, and (4) the magnitudes of the foraging parameters. Offprint requests to: P.H. Crowley     

Male size and mating tactic influence proximity to females during sperm competition in bluegill sunfish

Many fishes are characterized by intense sperm competition between males that use alternative mating tactics. In externally fertilizing fishes, males’ proximity to females during spawning can be an important determinant of fertilization success. Here, we assess how mating tactic, body length, speed during streak spawns, and periphery cover affect males’ proximity to females during sperm competition in the externally fertilizing bluegill (Lepomis macrochirus). Bluegill are characterized by three mating tactics referred to as parental, sneaker, and satellite. Parentals are territorial and construct nests, while sneakers use a streaking behavior, and satellites use female mimicry to steal fertilizations from parentals. We show that a small body length is important for sneakers but not for satellites to obtain a close position to the female during spawning. Specifically, smaller sneakers obtain a closer position to females than larger sneakers in part by positioning themselves closer on the periphery of a parental’s nest before streaking but show no difference in the speed at which they streak. The amount of peripheral vegetation around a parental’s nest did not appear to affect proximity of sneakers to females, and there was no relationship between the amount of peripheral vegetation and the frequency of intrusions by either sneakers or satellites. Finally, parentals were farther from the female when a sneaker or satellite intruded than when they spawned alone with the female.     

Solitary nesting as an alternative breeding tactic in colonial nesting bluegill sunfish (Lepomis macrochirus)

Colonial breeding can evolve in response to benefits afforded by clumped individuals, such as reduced predation and increased ease of assessing potential mates. However, colonial breeding can also impose costs such as increased disease transmission or increased cuckoldry. Here, we investigate solitary nesting as a potential alternative breeding tactic in colonial breeding bluegill sunfish (Lepomis macrochirus). Most male bluegill, termed parentals, compete for nesting sites in colonies and then court and spawn with females and provide sole care of the eggs. Although nesting in a colony results in reduced predation and fungal infection of broods, it comes at a cost of increased parasitism by specialized cuckolder males that do not nest. We found that 4.5% of parentals forgo spawning in a colony and instead construct nests solitarily. Solitary males were of similar size and age to colonial males, but were in significantly better condition. Solitary males obtained as many eggs as males nesting in the center of colonies, and significantly more than males nesting on the periphery of colonies. Thus, females do not appear to discriminate against solitary males. Solitary males had smaller ear tabs, a presumed sexually selected character used by parental males in intrasexual competition, than colonial males. Tracking data revealed consistency in nesting tactic (but not nest position within the colony) between spawning attempts. We suggest that solitary nesting represents either a facultative decision made by parental males in top condition at the onset of breeding, or a life history decision to forgo spawning in colonies.Communicated by K. Lindström     

Application of various mathematical models to data from the uptake of methyl mercury in bluegill sunfish (Lepomis macrochirus)

Evaluated and compared are the results of various mathematical models to data from the uptake of methyl mercury in bluegill sunfish (Lepomis macrochirus). Data were obtained from the direct uptake of methyl mercury from water in bluegill at 9°C and at a CH₃HgCl concentration of 0.2 ppb. The models considered are: (1) linear uptake relationship; (2) first-order kinetics, one-compartment or single-exponential model; (3) combination of first-order kinetics and the linear uptake relationship, designated as slope-exponential model; (4) two-compartment or double-exponential model; (5) three-compartment model, including a storage compartment for the build up of contaminant, designated as storage model; (6) three-compartment model, allowing two routes of elimination, designated the “Y” model; (7) an empirical uptake relationship having the form of a rectangular hyperbola.A comparison of the nonlinear models showed that the empirical model had fewer convergence problems in obtaining parameter estimates and its parameter values had less variation. The best fit was made by the “Y” model, the empirical model gave one of the smaller values of the residual mean squares, and other nonlinear models had computational convergence difficulties. The lack of computational problems and the comparitively small residual mean squares resulted in the selection of and the preference for the empirical model for use in describing the individual and interactive effects of temperature and methyl mercury concentration on the direct uptake of methyl mercury from water by bluegill.     

Parental investment in southern elephant seals,Mirounga leonina

Summary The southern elephant seal is among the most sexually dimorphic and polygynous of all mammals: males may be more than 10 times the weight of reproducing females and only the largest 2–3% of males are likely to breed. Current optimization theories of sexual selection predict that evolution would favor greater parental investment in individual males than in females. Because southern elephant seals represent an extreme of polygyny and sexual dimorphism, they might be expected to show a dramatic difference in parental investment in male and female pups. However, in a study of parental investment in elephant seals at South Georgia, using several different methods, we found no such difference after parturition. Mother-pup pairs were immobilized and weighed early in lactation, recaptured near the end of lactation and reweighed. A further 30 pups were weighed an average of five times during lactation to establish the shape of the growth curve and to serve as partial controls for the previous set of animals. Initial post-partum weight in females ranged from 346 to 803 kg (=506, SD=111, n=26). Pup birth weight was related to mothers' post-partum weight in female pups but small females often gave birth to large male pups. Male pups were significantly heavier at birth than females. However, this size difference did not persist. Male and female pups were suckled for the same period, grew at the same rate and were not significantly different in weight at weaning. Mothers lost weight at the same rate regardless of their pup's sex.     

Derived decision rules for pollution control in a general equilibrium space economy

This paper attempts to narrow the gap between economic theory and policy inthe field of environmental pollution control by expanding the traditional general equilibrium model to include the kind of spatial detail that is important for describing pollution. This model is then used to derive theorems which provide the basis for the development of spatially differentiated, tax-based decision rules. In spite of the fact that these rules require no information on either damage costs or control costs, they maintain many of the desirable properties of the more conventional informationally intensive tax policies.     

Parental investment, adult sex ratios, and sexual selection in a socially monogamous seabird

Although most birds are monogamous, theory predicts that greater female parental investment and female-biased adult sex ratios will lower the polygyny threshold. This should result in polygynous mating, unless obligate biparental care or the spatial and temporal distribution of fertilizable females constrains a male’s ability to take advantage of a lowered polygyny threshold. Here we present data on the extent of male sexually dimorphic plumage, adult sex ratios and breeding season synchrony in three populations of a socially monogamous seabird, the brown booby Sula leucogaster. For one of these populations, San Pedro Mártir Island, we also present data on differences in male and female parental investment, mortality and probability of pairing. The extent of plumage dimorphism varied among populations. Sex ratios were female biased in all populations. On San Pedro Mártir Island, parental investment was female biased, females failed more often than males to find a mate, but there was no polygyny. We suggest that on San Pedro Mártir: (1) a period of obligate biparental care coupled with a relatively synchronous breeding season constrained the ability of males to take advantage of a high environmental polygamy potential and (2) the resulting socially monogamous mating system, in combination with the female-biased adult sex ratio, caused females to be limited by the availability of males despite their greater parental investment. Received: 18 November 1999 / Accepted: 24 January 2000     

Parental investment and the secondary sex ratio in northern elephant seals

Summary Data on northern elephant seals, Mirounga angustirostris, bearing on sex ratio theory were collected at Año Nuevo, California, and other Californian and Mexican Islands, during the period 1967 to 1988. The mass of males exceeded that of females by 7–8% at birth and at weaning. The sex ratio was biased to males at birth (51.2%) and was near unity at weaning (49.6% males). The sex ratio did not vary as a function of maternal age or maternal mass except in 6-year-old females, who produced significantly more males. Although sons cost more to rear in energetic terms than daughters, and mothers were more successful weaning the latter, the sex of the pup reared exerted no significant effect on the mother's reproductive performance the following year or on her subsequent survival. These data suggest that parents invest equally in sons and daughters when investment is measured in terms of future reproduction (Fisher 1930) and provide no support for the theory of adaptive shifts in sex ratio (Trivers and Willard 1973). The small sex difference in mass due to maternal effort reflects the fact that females fast during lactation and all energy transferred is from limited body stores. Because of these circumstances, selection for superior condition at the end of the period of parental investment may act more strongly on pups, who have the opportunity to steal milk, than on their mothers.     

Intra-group relatedness affects parental and helper investment rules in offspring care

In any system where multiple individuals jointly contribute to rearing offspring, conflict is expected to arise over the relative contributions of each carer. Existing theoretical work on the conflict over care has: (a) rarely considered the influence of tactical investment during offspring production on later contributions to offspring rearing; (b) concentrated mainly on biparental care, rather than cooperatively caring groups comprising both parents and helpers; and (c) typically ignored relatedness between carers as a potential influence on investment behavior. We use a game-theoretical approach to explore the effects of female production tactics and differing group relatedness structures on the expected rearing investment contributed by breeding females, breeding males, and helpers in cooperative groups. Our results suggest that the breeding female should pay higher costs overall when helpful helpers are present, as she produces additional offspring to take advantage of the available care. We find that helpers related to offspring through the breeding female rather than the breeding male should contribute less to care, and decrease their contribution as group size increases, because the female refrains from producing additional offspring to exploit them. Finally, within-group variation in helper relatedness also affects individual helper investment rules by inflating the differences between the contributions to care of dissimilar helpers. Our findings underline the importance of considering maternal investment decisions during offspring production to understand investment across the entire breeding attempt, and provide empirically testable predictions concerning the interplay between maternal, paternal and helper investment and how these are modified by different relatedness structures.     

Parental investment in two solitary wasps Ancistrocerus adiabatus and Euodynerus foraminatus (Eumenidae: Hymenoptera)

Summary Females of the solitary eumenid wasps Ancistrocerus adiabatus and Euodynerus foraminatus control the adult size of their offspring by the amount of food provded to the larvae. For both species, larger females provision more offspring and collect more food than smaller females. Males of E. foraminatus, upon emergence as adults, fight for control of the nest area. The winning brother remains at the nest and mates with his sisters as they emerge several days later. Males of A. adiabatus are not as agonistic as males of E. foraminatus so that a number of males may remain near the natal nest. When females nest near each other, male-male interactions increase, and the likelihood of sibling mating decreases. Under conditions of clumped nesting, females make larger offspring and invest more in males than when nesting in isolation.     

Parental and alloparental investment in campo flickers (Colaptes campestris campestris): when relatedness comes first

Behavioral Ecology and Sociobiology - Studies with individual rodents have revealed that they display protean (unpredictable) behavior when attacked by an owl. Other studies have revealed that...     

Effect of decision rules in choice experiments on hunting and bushmeat trade

Providing insight on decisions to hunt and trade bushmeat can facilitate improved management interventions that typically include enforcement, alternative employment, and donation of livestock. Conservation interventions to regulate bushmeat hunting and trade have hitherto been based on assumptions of utility- (i.e., personal benefits) maximizing behavior, which influences the types of incentives designed. However, if individuals instead strive to minimize regret, interventions may be misguided. We tested support for 3 hypotheses regarding decision rules through a choice experiment in Tanzania. We estimated models based on the assumptions of random utility maximization (RUM) and pure random regret maximization (P-RRM) and combinations thereof. One of these models had an attribute-specific decision rule and another had a class-specific decision rule. The RUM model outperformed the P-RRM model, but the attribute-specific model performed better. Allowing respondents with different decision rules and preference heterogeneity within each decision rule in a class-specific model performed best, revealing that 55% of the sample used a P-RRM decision rule. Individuals using a P-RRM decision rule responded less to enforcement, salary, and livestock donation than did individuals using the RUM decision rule. Hence, 3 common strategies, enforcement, alternative income-generating activities, and providing livestock as a substitute protein, are likely less effective in changing the behavior of more than half of respondents. Only salary elicited a large (i.e. elastic) response, and only for one RUM class. Policies to regulate the bushmeat trade based solely on the assumption of individuals maximizing utility, may fail for a significant proportion of the sample. Despite the superior performance of models that allow both RUM and P-RRM decision rules there are drawbacks that must be considered before use in the Global South, where very little is known about the social–psychology of decision making.     

Two-goal environmental policy: An integration of micro and micro ad hoc decision rules

A method of predicting and comparing the immediate impact of different environmental policies and of selecting the optimal policy is suggested in this paper. The degradation of water quality by a polluting industry is used to demonstrate this approach. Aggregation over this industry consists of firms with “putty clay” properties in production, and credit constraints in the capital market generate the macrorelations. The macrorelations which depend on the level of the policy variables are expressed in terms of probabilities and moments of the distribution of the microrelations and are used for different planning purposes.     

Reduced disease in offspring: A benefit of coloniality in sunfish

Summary Increased disease and parasitism are a well-documented cost of group living for colonial birds and mammals, but we now show that disease in offspring of fish may be reduced by nesting in colonies. The aquatic fungusSaprolegnia sp., which is a common cause of egg mortality among freshwater fishes, is more prevalent in the nests of solitary than colonial male bluegill sunfish (Lepomis macrochirus). Moreover, fungal infection decreases with nest density in colonies. This may be due in part to a behavioural advantage since colonial males can devote less time to defending eggs and more time to fanning them, which reduces fungal infection. In addition, we demonstrate experimentally that solitary nests become infected at higher rates than colonial nests, even in the absence of parental males. This suggests that colonies are encountered by spores at a lower rate and/or that the large number of nests in colonies dilutes the number of fungal spores per nest. Through one or all of these mechanisms, egg mortality in colonial nests is lowered significantly. Therefore, in some cases, disease may select for group living.     

Effects of within- and among-patch experiences on the patch-leaving decision rules in an insect parasitoid

The present study aimed to address how an insect parasitoid makes patch-departure decisions from various types of host patches and how previous patch experiences in the environment modify this decision-making process. Experiments were done with the parasitic wasp Aphidius rhopalosiphi attacking the grain aphid Sitobion avenae. In the experiments, wasps were observed in a laboratory environment containing several patches of various host densities, and behavioural records were analysed using a Coxs proportional hazards model. Consideration of the effect of the within-patch experience gave a classic pattern of patch-leaving decision rules in parasitoids: A. rhopalosiphi used local information on host quality (i.e. numbers of ovipositions or rejections) and availability (i.e. patch density) to determine departure decision. However, consideration of previous patch experiences provided evidence that these departure rules are fundamentally dynamic, responding to the physiological state of the animal and the information it has about its environment. Results showed that A. rhopalosiphi decreased its tendency to leave the visited patch after an oviposition. However, when a female has already laid several other eggs in the environment, such an incremental mechanism gradually switched to a decremental one. Hence, A. rhopalosiphi responded to egg-load depletion by leaving the visited patches sooner and by depositing a smaller number of eggs in those patches, which probably led to a decreased level of superparasitism. Results also indicated that previous experiences enabled wasps to estimate spatial host distribution and then to adjust their behavioural decisions accordingly. Thus, A. rhopalosiphi was shown to adjust its patch residence time according to the quality and the number of the patches previously visited. These proximate mechanistic rules adopted by A. rhopalosiphi females are discussed in the context of general predictions from optimality models.Communicated by D. Gwynne     

Parental investment, potential reproductive rates, and mating system in the strawberry dart-poison frog, Dendrobates pumilio

We studied the effect of relative parental investment on potential reproductive rates (PRRs) to explain sex differences in selectivity and competition in the dart-poison frog Dendrobates pumilio. We recorded the reproductive behavior of this species in a Costa Rican lowland rainforest for almost 6 months. Females spent more time on parental care than males, and `time out' estimates suggest that PRRs of males are much higher than than those of females, rendering females the limiting sex in the mating process. Males defended territories that provide suitable calling sites, space for courtship and oviposition, and prevent interference by competitors. Male mating success was highly variable, from 0 to 12 matings, and was significantly correlated with calling activity and average perch height, but was independent of body size and weight. Estimates of opportunity for sexual selection and variation in male mating success are given. The mating system is polygamous: males and females mated several times with different mates. Females were more selective than males and may sample males between matings. The discrepancy in PRRs between the sexes due to differences in parental investment and the prolonged breeding season is sufficient to explain the observed mating pattern i.e., selective females, high variance in male mating success, and the considerable opportunity for sexual selection. Received: 9 June 1998 / Received in revised form: 27 March 1999 / Accepted: 3 April 1999