Body size effects on vertical climbing among chimpanzees

I analyzed observations from a yearlong study of the positional behavior of Pan troglodytesat the Mahale Mountains National Park to determine whether there are detectable differences in behavior between large and small individuals. Analysis was complicated by a weak correlation between body size and social rank. To factor out rank effects, I performed two types of analyses, depending on the type of data: (1) multiple regressions or (2) comparisons of similarly ranked animals of different body size. With social rank effects accounted for, larger males fed lower in the canopy, fed on the ground more often, fed preferentially among food tree species with smaller adult heights, and climbed significantly less often than smaller males did. Contrary to expectation, large males utilized smaller weight-bearing structures than small males did. These results suggest that large males minimized climbing versus optimizing support diameters, perhaps because vertical climbing is disproportionally expensive for larger animals. The large body weight of chimpanzees compared with other primates suggests that minimizing altitude changes, and therefore vertical climbing, is an important consideration in budgeting daily energy expenditures.     

Social rank and body size as determinants of positional behavior inPan troglodytes

A yearlong study of the positional behavior ofPan troglodytes at the Mahale Mountains National Park yielded 571 hr of observation.Cant (1987) articulated four predictions concerning the relationship between body weight/branch diameter and positional behavior based on the classic suspensory-ape paradigm. He noted that only two were supported by orangutan data. Three of these four hypothesis were not supported by chimpanzee data, as follows: there was no significant difference between the three largest males and the four smallest males in (1) branch diameters (5.8 cm vs 5.2 cm) nor (2) in the percentage of arm-hanging (13.6% vs 12.1%); and (3) large males did not arm-hang significantly more often than small males in any of three support diameter categories. The fourth hypothesis, that arm-hanging should be more common among smaller branches, was supported: arm-hanging as a percentage of all posture rose from 2.5% to 8.3% to 24% as stratum size decreased from >10 cm to <3 cm. The possibility that the first three hypotheses failed because of confounding effects of a correlation between body size and social rank was examined. Multiple regressions were done on 6600 2-min instantaneous observations on focal individuals. With social rank effects factored out, larger individuals preferentially utilized smaller, rather than larger supports. When positional mode frequencies were compared between large and small males matched for social rank, large males exhibited a lower frequency of arm-hanging than small males. An unexpected result was that social rank more consistently predicted branch diameter choice than body size. The most profound trend was for high ranking males to use larger supports, even though they spent more time in the terminal branches. These results suggest that (1) suspensory behavior is functionally related to small branch diameters; (2) chimpanzees do not prefer smaller branches, rather they are forced into them by food choice limitations; and (3) social rank more profoundly affects chimpanzee behavior than body weight.     

Mating system of the lefteye flounder,Engyprosopon grandisquama

Underwater observations were conducted on the reproductive behavior and mating system of the lefteye flounder,Engyprosopon grandisquama, off Nagashima, southwest of Kyushu Island, Japan. Two types of males were found: large males, which defended territories against other large males, and small males, which did likewise but only against smaller males. Large males established territories which encompassed or ovelapped the home ranges of 1 or 2 cohabitant females. Territories of the small males, in which a smaller female often maintained a home range, overlapped those of large males. Pair spawning occurred around sunset. Mating of large males with cohabitant females was observed 36 times and that of a small male with a smaller female once. Pair formation was assorted by body size, paired males being larger than females in most cases. Thus, inE. grandisquama, gigamous large males were common, small males occurring within the former's erritorial boundaries mating with smaller females.     

Effects of body size and social context on the arboreal activities of lowland gorillas in the Central African Republic

The objectives of this 27 month study were to document the positional behaviors used by lowland gorillas (Gorilla gorilla gorilla) in the Central African Republic and to compare the effects of body size dimorphism on the use of arboreal substrates. During this study, despite their great size, all gorillas used trees regularly. Predictions concerning the relationship of body size to arboreal behavior were generally upheld. Small branch and suspensory activities were rare for silverbacks. Females used smaller and multiple substrates and suspensory postures more frequently than males. Although females foraged in the periphery of trees, males stayed close to the cores and rarely used terminal branches. In addition to body size, this study found that party size, social rank, and tree structure all influence an animal's substrate choice and subsequent positional activities. Lone males typically remained in the cores of trees where substrates are large. Group males may have been forced to use all parts of trees because others were present. Lone males used small crown trees which provided easy access to terminal branch foods. Males and females foraging together used larger trees (containing more feeding sites) than single sex groups. Female positional behavior may have been affected by the presence of males. When apart from males, females used the cores of trees and larger substrates more than when foraging with males. As habitat and social context both influence substrate use, they should be considered essential components of body-sized based interpretations of the behavior of fossil or extant species. © 1995 Wiley-Liss, Inc.     

Is bigger better? Male body size affects wing‐borne courtship signals and mating success in the olive fruit fly,Bactrocera oleae (Diptera: Tephritidae)

Variations in male body size are known to affect inter‐ and intrasexual selection outcomes in a wide range of animals. In mating systems involving sexual signaling before mating, body size often acts as a key factor affecting signal strength and mate choice. We evaluated the effect of male size on courtship displays and mating success of the olive fruit fly, Bactrocera oleae (Diptera: Tephritidae). Wing vibrations performed during successful and unsuccessful courtships by large and small males were recorded by high‐speed videos and analyzed through frame‐by‐frame analysis. Mating success of large and small males was investigated. The effect of male–male competition on mating success was evaluated. Male body size affected both male courtship signals and mating outcomes. Successful males showed wing‐borne signals with high frequencies and short interpulse intervals. Wing vibrations displayed by successful large males during copulation attempt had higher frequencies over smaller males and unsuccessful large males. In no‐competition conditions, large males achieved higher mating success with respect to smaller ones. Allowing large and small males to compete for a female, large males achieve more mating success over smaller ones. Mate choice by females may be based on selection of the larger males, able to produce high‐frequency wing vibrations. Such traits may be indicative of “good genes,” which under sexual selection could means good social‐interaction genes, or a good competitive manipulator of conspecifics.     

Optimal climbing speed explains the evolution of extreme sexual size dimorphism in spiders

Several hypotheses have been put forward to explain the evolution of extreme sexual size dimorphism (SSD). Among them, the gravity hypothesis (GH) explains that extreme SSD has evolved in spiders because smaller males have a mating or survival advantage by climbing faster. However, few studies have supported this hypothesis thus far. Using a wide span of spider body sizes, we show that there is an optimal body size (7.4 mm) for climbing and that extreme SSD evolves only in spiders that: (1) live in high‐habitat patches and (2) in which females are larger than the optimal size. We report that the evidence for the GH across studies depends on whether the body size of individuals expands beyond the optimal climbing size. We also present an ad hoc biomechanical model that shows how the higher stride frequency of small animals predicts an optimal body size for climbing.     

Ecological context and the importance of body and gnathopod size for pairing success in two amphipod ecomorphs

Ecological context generates interspecific variation in mating behavior by imposing differential constraints on the action of sexual selection, but operation of these constraints in nature is not well understood. We used field and laboratory studies to examine the importance of body size and size of sexually dimorphic appendages, the gnathopods, for pairing success in two freshwater amphipod species within the Hyalella azteca species complex. We focused on a large-bodied species found in habitats where ecological factors tend to favor large body size, and a small-bodied species in habitats where small body size is favored by size-selective predation. A field study indicated that although male pairing success was greater for larger males in both species, pairing success increased throughout the range of variation in male size in the large species, whereas, in the small species, pairing success was low for smaller individuals, but roughly constant across intermediate to larger sizes. A laboratory mate choice experiment was consistent with the field study, finding that females of the large species exhibited a preference for larger males that was independent of absolute male size, but females of the small species were indifferent when choosing between males of intermediate to larger size. Species also differed in the direction of sexual size dimorphism in the field, with males being the larger sex in the large species but the smaller sex in the small species, a pattern consistent with the species differences in mate preference. Large gnathopod size relative to body size was associated with enhanced pairing success across all body sizes for the large species, but, in the small species, large gnathopod size enhanced pairing success only in smaller males. Species differences in mating behavior appear consistent with change driven by differing forms of the interaction between sexual and natural selection.     

Tree structure and sex differences in arboreality among western lowland gorillas (Gorilla gorilla gorilla) at Bai Hokou, Central African Republic

The aim of six months of research in 1995 on the gorillas (Gorilla gorilla gorilla) at Bai Hokou, Central African Republic, was to study the effects of tree structure, relative numbers of arboreal feeding sites, and sex differences in body size on arboreal foraging. The analysis presented here also documents inter-annual variation in fruit availability and climbing by silverback gorillas by comparing the 1995 results to those from earlier research, 1990–1992. This analysis suggests that female gorillas maintain similar levels of arboreality in fruit-rich and fruit-poor seasons and years, but silverbacks may be more terrestrial when fruit is scarce or difficult to access. Trees of different shapes present different numbers of feeding sites to bigger males and smaller females. Male climbing is affected by the availability of fruit, and small trees with narrow crowns that lend easy access to fruit from the core. This study suggests that the energetics of vertical climbing and biomechanical constraints imposed by small branch feeding sites in the periphery of trees may constrain the arboreal behavior of male gorillas. Fine-tuned comparisons of food availability, tree structure, and variation in social context of behavior across habitats, will assist efforts to understand differences in ecology among populations and species of African apes.     

Body size and social dominance influence breeding dispersal in male Pachydiplax longipennis (Odonata)

1. Dispersal behaviour can be affected by an individual's phenotype, by the environmental or social context they experience, and by interactions between these factors. Differential dispersal propensities between individuals may also be an important modifier of functional connectivity between populations. To assess how a key trait, body size, affected both social interactions and dispersal behaviour, this study examined the relationship between body size, antagonistic interactions, and breeding dispersal in male dragonflies (Pachydiplax longipennis) across a seasonal decline in adult body size. 2. During a seasonal peak in male body size in this study, dispersers were smaller than non‐dispersers. Later in the season, the body size of dispersers and non‐dispersers did not differ. 3. Focal observations found that body size was related to competitive dominance, large males engaged in aggressive chases more often and smaller males were more frequently pursued. 4. These results indicate that when large males were present, small males were more likely to disperse suggesting that dispersal is a tactic adopted by social subordinates in this context. If breeding dispersal is typically undertaken by subordinate males, functional connectivity between populations may be less than estimated from absolute dispersal rates.     

Mate size and breeding success in a monogamous cichlid fish

Synopsis The patterns of mate size and parental care of a monogamous cichlid fish,Cichlasoma maculicauda, were studied in Gatun Lake, Panama. Males defend territories which serve as courtship and nest sites. Within a population most mates in pairs are of equal size rank. In each pair the male is larger than the female, probably because most mature males are larger than most mature females. Clutch size increases with female body size. Male size affects breeding success in two ways. First, larger males provide nest sites less susceptible to destructive wave action. Second, young of larger males grow faster than young of smaller males. Large males defeat small males in contests for position in feeding areas, and this may provide their young with better feeding conditions. In the laboratory young growth rates increase with food abundance, and at high levels of food surpass those observed in nature. Fast growth of young reduces their vulnerability to predators and should allow parents to breed more often. Young survival rates improve with the size of the parents, so that larger fish raise more offspring at each breeding attempt. These observations suggest why preference for large mates should occur.     

Sex-specific life history patterns in bluegill (Lepomis macrochirus): interacting mechanisms influence individual body size

The ultimate body size that an individual fish achieves can be a function both of direct effects of growth or indirect effects associated with the timing of sexual maturation (and associated energetic tradeoffs). These alternatives are often invoked to explain variation in body size within and among fish populations, but have rarely been considered simultaneously. We assessed how resource availability and timing of maturation interact to influence individual body size of bluegill (Lepomis macrochirus). Resource availability (high and low food) and the social structure of the population (presence or absence of large, mature males) were varied in experimental ponds. Food ration affected growth (larger fish in the high food treatments) and the social structure of the population affected timing of maturation (early maturation of males in the absence of large males). Treatment effects, however, were sex-specific; males responded to the social structure of the population and females were more responsive to resource availability. We also found individuals that became sexually mature were smaller than those that remained immature, although results were sex-specific and resource dependent. For males, individuals that matured were smaller when resources were limited; mature and immature females showed no difference in body size regardless of food ration. We show that both resource availability and the processes that control timing of maturation interact in sex-specific ways to influence body size of bluegill. These results suggest that a more robust explanation for variable body size requires consideration of sex-specific interactions between ecological (food and growth) and evolutionary (timing of maturation) mechanisms.     

Paternal body size affects reproductive success in laboratory-held zebrafish (<Emphasis Type="Italic">Danio rerio</Emphasis>)

Across many fish species, large females tend to exhibit higher individual reproductive success due to elevated fecundity and the provisioning of better conditioned eggs and offspring compared to small females. By contrast, effects of paternal body size on reproductive success are less well understood. We disentangled the maternal- and paternal-size dependent effects on reproductive output and early life history in zebrafish (Danio rerio). In the laboratory, females and males from four size categories (small, medium-sized, large and very large) were allowed to spawn freely in a full factorial design with 10 replicates per size combination. As expected, larger females produced more eggs and better conditioned offspring compared to smaller females. Male body size further contributed to zebrafish reproductive success: offspring sired by large males exhibited higher hatching probability and these offspring also hatched earlier and larger than offspring fertilized by small males. However, the largest males experienced lower mating success and received fewer eggs than males of the smaller size classes. While male body size substantially affected reproductive success in zebrafish, it remained unclear whether and to what degree direct paternal effects (e.g., related to sperm quality) or indirect paternal effects stemming from differential allocation patterns by females were the mechanism behind our findings. Answering this question constitutes an important future research topic.     

Male size does not affect the strength of male mate choice for high-quality females in Drosophila melanogaster

Theory predicts that the strength of male mate choice should vary depending on male quality when higher-quality males receive greater fitness benefits from being choosy. This pattern extends to differences in male body size, with larger males often having stronger pre- and post-copulatory preferences than smaller males. We sought to determine whether large males and small males differ in the strength (or direction) of their preference for large, high-fecundity females using the fruit fly, Drosophila melanogaster. We measured male courtship preferences and mating duration to show that male body size had no impact on the strength of male mate choice; all males, regardless of their size, had equally strong preferences for large females. To understand the selective pressures shaping male mate choice in males of different sizes, we also measured the fitness benefits associated with preferring large females for both large and small males. Male body size did not affect the benefits that males received: large and small males were equally successful at mating with large females, received the same direct fitness benefits from mating with large females, and showed similar competitive fertilization success with large females. These findings provide insight into why the strength of male mate choice was not affected by male body size in this system. Our study highlights the importance of evaluating the benefits and costs of male mate choice across multiple males to predict when differences in male mate choice should occur.     

Male Mating Success in the Mountain Dusky Salamander,Desmognathus ochrophaeus: Are Small,Young, Inexperienced Males at a Disadvantage?

Young, small, sexually-inexperienced males suffer lower mating success than older, larger, more experienced males in many species of animals. I compared the mating success of male Desmognathus ochrophaeus (Amphibia: Caudata: Plethodontidae) reared to maturity in the laboratory with that of males collected as adults in the field. Courtship trials were staged in the laboratory between single males and females. Laboratory-reared males were significantly smaller in body size than adult males collected in the field, were certainly inexperienced sexually and were probably younger. No difference was found between laboratory-reared males and field-collected males in any measure of mating success, although the former apparently produced smaller sperm masses. I conclude that newly-mature male D. ochrophaeus are not discriminated against by females in the context of mate choice, and do not need to “rehearse” courtship in order to obtain successful inseminations. However, newly-mature males are probably at a disadvantage in terms of aggressive and sperm competition with larger males.     

Mate choice in female convict cichlids (<Emphasis Type="Italic">Amatitlania nigrofasciata</Emphasis>) and the relationship between male size and dominance

We examined how male size and fighting ability influence a female’s mate assessment process and her eventual mate choice in the monogamous convict cichlid, Amatitlania nigrofasciata. Females always chose the larger of two males when they were allowed to see a larger male next to a smaller one and when a larger male defeated a smaller one in a fight. They did not differentiate between large and small males when they did not see the two males together nor did they choose a dominant over a subordinate male when both were the same size. We suggest that females select on the basis of male size because it is a better predictor of both direct and indirect benefits (i.e., future competitive interactions and foraging ability) than dominance behavior only. Despite selecting one male over the other early in the courtship period, females continued to visit both males until spawning. Our evidence suggests that this assessment behavior more closely resembles a bet-hedging tactic rather than the female’s indecision as to which male will be her mate.     

Resource availability modulates the effect of body size on reproductive development

Within-species variation in animal body size predicts major differences in life history, for example, in reproductive development, fecundity, and even longevity. Purely from an energetic perspective, large size could entail larger energy reserves, fuelling different life functions, such as reproduction and survival (the “energy reserve” hypothesis). Conversely, larger body size could demand more energy for maintenance, and larger individuals might do worse in reproduction and survival under resource shortage (the “energy demand” hypothesis). Disentangling these alternative hypotheses is difficult because large size often correlates with better resource availability during growth, which could mask direct effects of body size on fitness traits. Here, we used experimental body size manipulation in the freshwater cnidarian Hydra oligactis, coupled with manipulation of resource (food) availability to separate direct effects of body size from resource availability on fitness traits (sexual development time, fecundity, and survival). We found significant interaction between body size and food availability in sexual development time in both males and females, such that large individuals responded less strongly to variation in resource availability. These results are consistent with an energy reserve effect of large size in Hydra. Surprisingly, the response was different in males and females: small and starved females delayed their reproduction, while small and starved males developed reproductive organs faster. In case of fecundity and survival, both size and food availability had significant effects, but we detected no interaction between them. Our observations suggest that in Hydra, small individuals are sensitive to fluctuations in resource availability, but these small individuals are able to adjust their reproductive development to maintain fitness.     

Male body size and condition affects sperm number and production rates in mosquitofish,Gambusia holbrooki

Sperm number is an important predictor of paternity when there is sperm competition. Sperm number is often measured as maximum sperm reserves, but in species where mating is frequent, males will often be replenishing their reserves. Thus, variation in how quickly males can produce sperm is likely to be important in determining male success in sperm competition. Despite this, little is known about how male size, body condition or diet affects sperm production rates. We counted sperm number in large and small Gambusia holbrooki (eastern mosquitofish) after 3 weeks on either a high or low food diet. Sperm number was significantly higher in both larger males and in well‐fed males. We then stripped ejaculates again either 1, 2, 3, 4 or 5 days later to investigate subsequent sperm production. The rate of sperm replenishment was influenced by an interaction between size and diet. Large, well‐fed males had consistently high levels of sperm available over the 5 days (i.e. rapid replenishment), whereas small poorly fed males showed consistently low levels of sperm availability over the 5 days (i.e. slow replenishment). In contrast, large, poorly fed and small, well‐fed males increased their sperm numbers over the first 3 days (i.e. intermediate replenishment). Our study highlights that when mating is frequent and sperm competition is high, size and condition dependence of maximal sperm number and of sperm production rate might both contribute to variation in male reproductive success.     

Aggression and RHP in the Northern Swordtail Fish, Xiphophorus cortezi: The Relationship Between Size and Contest Dynamics in Male–Male Competition

Current theory predicts that contest outcome, as well as decisions on whether to initiate a contest, escalate during a contest or retreat are decided by asymmetries in resource holding potential (RHP) and/or expected payoffs between contestants. In this investigation, dyadic contests were staged between male swordtail fish (Xiphophorus cortezi) where individuals were paired based on cumulative fight records and were ranked at the end of the trials in order to approximate RHP. Size was the only asymmetry that I did not attempt to control for and as a result, I was able to determine the relationships between size, contest initiation, escalation and outcome. Individuals changed their contest initiation strategy based on their size relative to that of their opponents, and contrary to predictions, the smaller of the two males in each contest was more likely to initiate the conflict than was the larger male. However, the larger of the two males was more likely to win and standard length proved to be a moderate predictor of an individual's final rank. Regardless of size, initiators fared poorly, winning only 31% of the contests. In instances where the smaller males won the contests, they were no more likely to initiate the encounter than was the larger male. However, when small males did win, fights lasted longer, suggesting that in some cases smaller males may be able to outlast their opponents.     

Few, small, and male: multiple effects of reduced nest space on the offspring of the solitary wasp, Euodynerus (Pareuodynerus) posticus (Hymenoptera: Vespidae)

The size of the preexisting wood cavities used as nests by aculeate Hymenoptera is expected to have consequences on fitness parameters such as offspring number and size. We evaluated the consequences of using small and large (three-times more voluminous) trap-nests by the solitary wasp, Euodynerus (Pareuodynerus) posticus (Herrich-Schaeffer). Following life-history and sex allocation theories, a number of non-mutually exclusive hypotheses were formulated: i.e. small nests either produce smaller or fewer offspring and/or more males, the cheaper sex. Wasps built about 28% more, but shorter brood cells in large nests, although their volume was still much higher in large nests. Adult males had smaller body size in small nests, but female size did not differ between large and small nests, possibly as an adaptive response against the future higher foraging costs of size-reduced females. Sex-ratio was often biased towards males in small nests. Mortality did not differ between large and small nests. We conclude that E.?(P.) posticus females would benefit from using larger nests, but that the sex-ratio would be probably overall unbalanced if females would not use also smaller, male-oriented tunnels.     

Social control of terminal phase transition in primary males of the diandric wrasse, <Emphasis Type="Italic">Halichoeres poecilopterus</Emphasis> (Pisces: Labridae)

In many diandric fishes, large territorial males with bright body coloration (terminal phase (TP) males) are derived either from initial phase (IP) females that change sex to male or from IP primary males that change color and behavior, but do not change sex. The mechanism controlling the transition of IP primary males into TP males is not well understood. We conducted cohabitation experiments to examine social conditions favoring TP transition by primary males in the diandric wrasse, Halichoeres poecilopterus. IP primary males always started TP-specific sexual behavior in the presence of a smaller subordinate, and subsequently acquired TP body coloration. In contrast, primary males under subordinate conditions often performed female-like sexual behavior. In pairs with similar body sizes, both individuals initiated TP male behavior. The results suggest that TP transition in primary males may be closely related to a dominance relationship (or size order) within social groups, as it is in the case of sex change by females.