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Encyclopedia of Evolution.pdf - Online Reading Center

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Other reproductive and related activities. <strong>Evolution</strong>ary<br />

explanations are also available for reproductive systems that<br />

seem very strange to human observers. In sea horses and some<br />

other fishes and amphibians, males may carry the fertilized<br />

eggs in their mouths or in special pouches until they hatch.<br />

These examples <strong>of</strong> “fathers giving birth” are not simply weird<br />

stories <strong>of</strong> nature. In aquatic animal species with external fertilization<br />

(where females deposit eggs and the males deposit<br />

semen on the eggs), there is at that moment no question <strong>of</strong><br />

paternity. However, if both <strong>of</strong> them leave, some other male<br />

could deposit semen, which might affect the outcome <strong>of</strong> paternity<br />

in at least some <strong>of</strong> the hatchlings. The male, by gathering<br />

the eggs and carrying them, can prevent other males from<br />

fertilizing them. Male brooding, far from being an example<br />

<strong>of</strong> males showing care to their <strong>of</strong>fspring, may actually be an<br />

example <strong>of</strong> evolution in response to male–male competition.<br />

<strong>Evolution</strong>ary hypotheses might also help to explain<br />

homosexual behavior. Though not a reproductive system,<br />

it is derived from reproductive systems that the animals<br />

already have. Homosexual behavior, both between males<br />

and between females, is widespread in the animal kingdom.<br />

Examples include bonobos, penguins, dolphins, macaques,<br />

baboons, and rhesus monkeys. In some cases, it is connected<br />

with male aggressive behavior, as in razorbill birds. Animals<br />

with homosexual proclivities can pass on their genes, if they<br />

are homosexual only part <strong>of</strong> the time; or, their relatives can<br />

pass on the genes if the genes are not expressed. Its widespread<br />

occurrence among animals suggests that homosexual<br />

behavior may have a genetic basis.<br />

Concealed ovulation. Humans are one <strong>of</strong> 32 primate<br />

species with concealed ovulation. In many animal species,<br />

the female advertises ovulation, sometimes with conspicuous<br />

swellings, scents, and behaviors. The males, whether<br />

in a harem or a promiscuous breeding system, know which<br />

females to mate with and when, to maximize their chances <strong>of</strong><br />

producing <strong>of</strong>fspring. However, in humans, men do not know<br />

when women ovulate—nor do the women. Therefore men<br />

cannot choose to mate with women only during ovulation.<br />

Two major explanations for the evolution <strong>of</strong> concealed<br />

ovulation have been <strong>of</strong>fered. One, proposed by evolutionary<br />

biologists Richard Hamilton and Katharine Noonan, has been<br />

called the “daddy at home” theory. Concealed ovulation keeps<br />

the male home, because he cannot know when his mate is fertile<br />

and has to keep copulating with her over long periods <strong>of</strong><br />

time in order to produce <strong>of</strong>fspring. If he stays home he is more<br />

likely to help provide for, and raise, the <strong>of</strong>fspring. Concealed<br />

ovulation might also discourage the man from finding other<br />

mates for reproductive purposes, because he cannot know<br />

which <strong>of</strong> them, at any given time, are fertile. This explanation<br />

posits an association <strong>of</strong> concealed ovulation with monogamy.<br />

The other explanation, <strong>of</strong>fered by evolutionary biologist<br />

Sarah Blaffer Hrdy, has been called the “many fathers”<br />

theory. She suggests that concealed ovulation makes it impossible<br />

for a male to know which <strong>of</strong>fspring are really his, or,<br />

more to the point, which ones are not, therefore which ones<br />

to kill. The females never benefit from having their <strong>of</strong>fspring<br />

killed, therefore concealed ovulation might be the female’s<br />

way <strong>of</strong> keeping males from killing her <strong>of</strong>fspring. Male goril-<br />

reproductive systems<br />

las, in a harem system, know which <strong>of</strong>fspring are theirs, and<br />

they kill others; male vervet monkeys, in a promiscuous system,<br />

do not know which <strong>of</strong>fspring are theirs and seldom kill<br />

them. This explanation suggests an association <strong>of</strong> concealed<br />

ovulation with promiscuity.<br />

Nothing could be easier, it would seem, than to simply<br />

look at the data to see which explanation is correct. It turns<br />

out not to be quite so simple. Swedish evolutionary biologists<br />

Birgitta Sillén-Tullberg and Anders Møller gathered data on<br />

primate breeding systems and the occurrence <strong>of</strong> concealed<br />

ovulation. They found that nearly all monogamous primates<br />

(10 out <strong>of</strong> 11) had concealed ovulation. This would seem to<br />

confirm the daddy-at-home theory. However, the reverse is<br />

not true. Of the 32 primate species with concealed ovulation,<br />

22 are not monogamous. According to Sillén-Tullberg and<br />

Møller, concealed ovulation may have evolved eight different<br />

times in different primate lineages, and it may not be possible<br />

to formulate a single explanation for it.<br />

Sex Ratios<br />

In most populations <strong>of</strong> animals, and <strong>of</strong> plants that have separate<br />

sexes, there is an equal number <strong>of</strong> males and females.<br />

The sex ratio (number <strong>of</strong> males per female) is about 1:1. This<br />

occurs because if one <strong>of</strong> the sexes is less abundant, each individual<br />

<strong>of</strong> that sex can have more <strong>of</strong>fspring than an individual<br />

<strong>of</strong> the other sex, which creates a selective advantage for the<br />

less abundant sex. This is an example <strong>of</strong> frequency-dependent<br />

selection. This occurs only when <strong>of</strong>fspring <strong>of</strong> the two sexes<br />

are about equally expensive for the parents to produce. In<br />

some special cases, the sex ratio departs from 1:1.<br />

• Among honeybees, the queen is related to each <strong>of</strong> her sons<br />

and daughters by a relatedness <strong>of</strong> 0.5. The queen would<br />

therefore benefit from producing an equal number <strong>of</strong> sons<br />

and daughters. The workers, however, may be related to<br />

one another by a factor <strong>of</strong> 0.75, but to their brothers by<br />

only 0.5. The queen may lay equal numbers <strong>of</strong> male (haploid)<br />

and female (diploid) eggs, but the workers kill many <strong>of</strong><br />

the males, resulting in a sex ratio considerably less than 1:1.<br />

• In some insects, the larvae parasitize hosts by living inside<br />

<strong>of</strong> them. When they become mature, the only other members<br />

<strong>of</strong> their species that they may encounter may be those<br />

that emerged from the same host, which are likely to be<br />

their siblings. In such situations, it is advantageous for the<br />

parent who lays the eggs to produce only enough males to<br />

fertilize the females, resulting in a sex ratio considerably<br />

less than 1:1.<br />

Reproductive systems, in general, encourage the production<br />

<strong>of</strong> genetically varied <strong>of</strong>fspring. These systems can take<br />

on many and varied forms over evolutionary time and can<br />

be influenced by factors that are unrelated to reproduction.<br />

Reproductive systems therefore defy complete explanation or<br />

even adequate classification.<br />

Further <strong>Reading</strong><br />

Asa, Cheryl S., and Carolina Valdespino. “Canid reproductive biology:<br />

An integration <strong>of</strong> proximate mechanisms and ultimate<br />

causes.” American Zoologist 38 (1998): 251–259.

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