The complexities of intraspecific competition are illustrated through elegant work on horned beetles (Emlen 2000). Beetle horns are rigid extensions of the exoskeleton and have evolved repeatedly within this order of insects. These horns are generally only expressed in males and are used in combat with other males for access to resources and/or females. These resources are in discrete, readily defensible patches, and the horns allow males to defend these sites and to mate with the females found there. The jousting contests between males can be dramatic and the winner is rewarded with mating privileges.
These horns, however, do not come without cost. They constitute a large investment in energy, and as much as 10% of body mass may be devoted to them. Horn growth prolongs development time and risk of larval mortality, and there is a trade-off between horn development and the ability to fly. Finally, the development of horns leads to lowered visual acuity and to smaller eyes. Nevertheless, it appears that since larger horns are useful in gaining access to females, the reproductive benefits gained from large horns offset the costs of production and maintenance.
The ability of males to grow horns is based on larval nutrition. Both final adult body size and the size of horns in males depend on the amount of food they consume as larvae. Males exposed to poor conditions as larvae are small and do not produce horns, whereas well-fed males become large adults able to produce horns. Horn production shows an insignificant level of heritable genetic variation (Moczek and Emlen 1999).
These complexities and trade-offs are illustrated through studies of dung beetles of the genus Onthophagus. The basic life history of these beetles is as follows. After finding a patch of dung, a female digs a tunnel in the soil beneath the dung. She then buries dung below ground to provide nutrition for the larvae. Females spend days inside a single tunnel, pulling down pieces of dung to various blind ends of tunnel branches, each with its "brood ball" of dung. A single egg is laid at the top of a brood ball, and a single larva develops in isolation within the brood ball.
Females mate repeatedly with males during the time of egg laying. Male reproductive behavior consists of securing their unique access to females in the tunnels. The large, horned males guard the tunnels and fight other males trying to approach the females. Larger males with larger horns win these fights. For two males of the same size, the one with the larger horns usually wins. Thus long horns provide males with significant advantages.
However, all is not lost for a small male. Although they are not adept at guarding entrances to tunnels or winning jousts, small males have other tactics. They attempt to slip undetected past the large males, or they dig side tunnels that intercept a guarded tunnel well below ground. Thus hornless males may manage to sneak undetected into guarded tunnels and mate with the female. If such a "sneaker" male is caught, he is chased out by the resident male, who then returns to the female and mates with her. This dilutes or displaces sperm from the sneaker male. Sneaker males actually do best when they have no horns at all, since horns get in the way of sneaking! Success for a small male depends on rapid and undetected entry into tunnels, and horns cause vibrations as they scrape against tunnel walls, which would alert the resident male. Finally, as mentioned earlier, beetles with very large horns tend to have undeveloped eyes and/or a diminished flying ability due to smaller wing size and greater body weight. Since dung is a temporary resource, another component of fitness is the ability to disperse and locate new dung heaps. As the dung resource diminishes, some of the males, as larvae, will receive a less-than-optimum amount of food. They will be small and lack horns. Thus, although they would be poorly equipped to defend a female in a burrow, they are well adapted to disperse and find a new dung pile.
Both intraspecific competition and sexual selection are played out on the fields of manure. Male-female competition in dunnocks
Another aspect of intraspecific competition is competition between the sexes as expressed through mating systems. Obviously ecological conditions can influence the mating behavior of individuals. As conditions change, a cooperative pair or an animal society may quickly dissolve into a set of competing individuals. The following study (Davies 1992, 1995) illustrates some of the complexities in what appears to be a simple pair-bonded mating system.
The dunnock (Prunella modularis), formerly known as the hedge sparrow, is not a true sparrow but an accentor. In Old English "dun" means brown and "ock" means little. Thus P. modularis is the archetypal little brown bird. It seems unremarkable as it shuffles about under the bushes collecting tiny insects for its young. The impression gained is of harmonious cooperation. The Reverend Morris (1856, in A History of British Birds) admired this species so much that he urged his parishioners to emulate its behavior. The Reverend, as quoted by Davies (1992), found this bird to be "unobtrusive, quiet and retiring, without being shy, humble and homely... sober and unpretending in its dress ... while still neat and graceful..." Due to its extraordinary mating system, however, the Reverend Morris would hardly have been pleased had his congregation followed the example of these birds. Although a dunnock territory can contain one seemingly harmonious pair, a female in the territory next door may be mating with two males, or a male may be mating with two females.
The conventional view of pair formation in songbirds is that males first set up territories and advertise for mates by singing. Females then choose among male territories. However, in dunnocks females defend their own territories against other females, occupying exclusive areas with little overlap among neighbors. Females settle independently of the males, based on the quality of the territory. Males then compete to defend the females from other males.
The fact that males and females set up territories independently results in a wide variety of mating behaviors. In some cases, a single male defends one female territory (producing monogamy); at other times a male defends two adjacent females (producing polygyny). In other cases two males share the defense of one female (polyandry), or several adjacent females (polygynandry).
If two males share a territory with a single female, the dominant or alpha male (usually the older one) tries to evict the beta male. If he is unsuccessful, the two share defense of the joint territory. Some males wander for weeks before finding a permanent home. Sometimes a beta will overlap with two alphas. The larger the territory of the female, the harder it is for a male to monopolize her. The larger the male territory, the more likely he is to have two females.
Once a female has built her nest, she solicits copulations from males. Mating begins 3-7 days before the first egg is laid, and lasts up to completion of the clutch of 3-5 eggs. One egg is laid per day during this period. Monogamous males chase off neighboring males who are interested in mating. When two males share a territory, the alpha male follows the female everywhere to prevent her from copulating with the beta male. A female often maneuvers to throw off the alpha male. She then solicits the beta for mating. Females have many tricks, and seem intent on preventing exclusive mating by the alpha.
The act of copulation was described by Selous (1933) as "bizarre." "The hen elevated her rump and stood still, when the male, hopping up, made little excited and very wanton-looking pecks in this region, that is to say the actual orifice. There was actually no mistaking the nature and significance of the actions, rather lecherous, as it seemed to me. This is a very remarkable thing ... but I do not understand it" (Selous 1933, pp. 107-9). During the pecking described by Selous, the female cloaca makes pumping movements, resulting in the ejection of a small droplet of fluid. As soon as this occurs the male copulates with her.
Davies (1992) analyzed the droplets and discovered that they contain masses of sperm from the previous matings. Like other birds, female dunnocks store sperm. Females with two males may copulate up to six times per hour. As a result, there will be a pool of sperm in the female's cloaca and vagina for much of the mating period. The male's pecking stimulates the female to eject this pool of sperm to make way for his insemination and give his own sperm a better chance of being stored.
After the excitement of the mating period, life on a dunnock territory becomes peaceful during incubation. Females incubate the clutch of eggs alone for 11-12 days. Males help with chick feeding for 11-12 more days in the nest and for two weeks after fledging until the young become independent.
The results of DNA tests on the blood of the chicks and the parents showed that the female is the mother of all chicks in the nest. In monogamous and polygynous territories, the male was the father of all chicks. When two males guarded a territory with one female, however, if the female mated with both males, paternity was mixed. The alpha male fathered 55% of the brood and the beta male 45%.
From the female perspective, when both males mated with her they both helped feed the brood. If she mated exclusively with the alpha male, only he helped feed the brood. Males, however, cannot recognize their own young. If they have mated with the female, they will help raise the young, even in cases where they did not happen to father any of the chicks. If a beta male gained a large share of the copulations he more readily helped with feeding of the chicks. Similarly, if an alpha male was removed experimentally, allowing the beta a larger share of the matings, the beta male worked harder to feed the chicks than did the alpha. Thus, males varied their parental effort in relation to their chance of paternity, not simply according to dominance rank.
For a female, polygyny is the least desirable situation. She must share the help of one male with another female, and some of her chicks often starve to death. A polyandrous female was the most successful because she had the help of two males in raising her young. This explains why the females sneaked around and tried to get the beta males to mate with them. They hoped for future help in rearing of the young.
For a male, the situation is reversed. In polygyny, although each female is less productive, the combined output of two females often exceeds that of one female in monogamy.
Thus the mating system reflects an intraspecific competitive battle between the sexes in dunnocks. Behavioral and genetic studies of other bird species show that the dunnock mating system is not unusual. However, extra-pair copulations and fertilizations vary across species for reasons only poorly understood (Petrie and Kempenaers 1998, Blomquist et al. 2002). Extra-pair paternity is much less frequent in non-passerine than in passerine birds (Birkhead et al. 2001). For example Blomquist et al. (2002) found that in western sandpipers (Calidris mauri) only 5% of all chicks were the product of extra-pair matings. Nevertheless the application of molecular techniques to behavioral studies has allowed us to ask new questions about the mechanisms of intraspecific competition.
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