Mating modifications imposed by the land environment
Greater problems arise on land than in water. Eggs produced by truly terrestrial creatures are either retained in the parental body during their development or must be fully protected from drying up. Protective membranes must be tough indeed. More importantly, however, sperm cells must still be deposited where they can swim toward the eggs, for they cannot survive or function except in a watery solution of dilute salts. In all terrestrial creatures, except those that return to water to breed, sperm can survive only in the body of the male or female organism. All insects, therefore, must mate in order for eggs to be fertilized, and all have appendages at the rear of the body that serve as a copulatory device capable of being used even when in flight. Sperm is injected into the female’s duct or storage sac, either for immediate fertilization or for later use. The queens of bees, ants, and termites, in fact, mate once and for all during a nuptial flight and thereafter use the stored sperm to fertilize all the eggs they subsequently produce.
The land vertebrates have to cope with much the same breeding circumstances as the insects. Man is more aware of these procedures because they happen mostly in much larger creatures and also because he has some fellow feeling for them. Reptiles, birds, and even the most primitive surviving mammals—namely, the platypus and spiny anteater of Australasia—produce yolky eggs encased in a more or less rigid calcareous shell. Moreover, within the shell, a thick layer of albumen surrounds the egg proper. Both the albumen and the shell are added after the ovum leaves the ovary and during its passage down the oviduct. Fertilization must take place, if at all, as the eggs enter the oviduct, for neither the albumen nor the shell can be penetrated by spermatozoa. Sperm must therefore be introduced into the female and must be able to make their way up to the end of the oviduct, which is a very long journey for so small a cell. An enormous number must begin the journey to make sure that some will reach the goal.
Sexual anatomy
In reptiles and birds of both sexes, as in amphibians and fish, a single opening to the exterior serves jointly for both the intestine and reproductive duct. This is the cloaca, or vestibule. Nevertheless, copulation of a sort occurs in all three groups of terrestrial vertebrates: the reptiles, birds, and mammals. With the exception of man, the male always mounts the female from the rear or back, and in both reptiles and birds the cloacal openings are pressed closely together to form a continuous passage from one individual to the other. With one exception, the archaic tuatara (Sphenodon) of New Zealand, all present-day reptiles have an erectile penis, derived from the cloacal wall, that delivers the sperm into the proper duct. One mating may serve for a long time, and there are cases known in which female snakes have laid fertile eggs after months and sometimes years of isolation in captivity. On the other hand, a penis of any sort is lacking in most kinds of birds, and the pressing together of the cloacal apertures seems to serve well enough. The most advanced copulatory procedure is that of mammals. In mammals the cloaca has become replaced by separate openings for the reproductive duct and intestine, respectively. Eggs have become microscopic, devoid of shell, yolk, and virtually all albumen, although they still need to be fertilized as they enter the upper end of the oviduct. A well-developed, erectile penis is always present in the male for the ejaculation of stored sperm well up the reproductive passage of the female. Accordingly, the two sexes have become strikingly differentiated anatomically, with regard to delivery of sperm, compared with the seemingly primitive anatomical equipment of birds.
Courtship
The coming together of two members of the opposite sex is a necessary preliminary to mating. It may be accomplished by two individuals independently of any larger congregation, or it may result from two individuals pairing off within a breeding population that may have assembled even from the ends of the earth. In the one the problem is to find one another; in the other the problem is to find the appropriate place, called the staging area. In both cases timing and some sort of navigation are important. Mass assembly appears to be the more effective, although a local crowd of any kind of animal may be an open invitation to predators, human or otherwise, and may on occasion become disastrous.
The searching out of a solitary individual by another of the opposite sex can be a difficult matter. In the dark depths of the ocean, for instance, where fish and other marine life forms are extremely scarce and scattered, the chance of encounter is rare indeed. The small angler fish (Photocorynus spiniceps) that cruise around at great depths are most unlikely to meet a member of the opposite sex at a time or place when the female happens to be ready to shed her eggs. As a form of insurance to this end, however, any small, young male that happens to meet a large female, apparently at any time, immediately fastens on to her head or sides by his jaws and thereafter lives a totally parasitic existence sustained by the juices of the female body. Sperm thus becomes available at any time the female may produce eggs to be fertilized.
On land this individual procedure of searching out is common among insects and the more predatory mammals. Male crickets and cicadas sound their familiar signals, by night or by day, which attract any females within hearing distance. More remarkable are those insects and other creatures that produce living light, in some cases for no apparent purpose but in others, such as the firefly, for signalling between the sexes in the dark of summer nights. The male individuals, always more dispensable than females, fly freely at considerable risk, flashing their light at regular intervals. The light of the female, perched more safely on some tall grass, winks back as though it were a landing light, and so they come together. Each of the several species of firefly has its own flash code, or rhythm, and any wasteful attempt at interspecies mixing is avoided. On the same principle, female moths send their personal perfume into the night air, and those males that detect the scent fly toward the source, the winner taking all. Mammals also depend mainly on their sense of smell, being generally colour blind, not too attentive to sound, and, apart from the grazing and browsing creatures, mainly active at night. The scented sex appeal of a cat in heat, whether domestic or wild, excites all the males in the neighbourhood and, with or without the sound of voice, male and female come quickly together in the dark. In all of these, courting is mostly uncalled for since only ready-to-mate individuals are involved in this sexual searching in the dark.
Courting is necessary whenever the male is a supplicant. A female may not be ready to mate, and stimulation in the form of dance or song may be required to create the mood; or, as is commonly the case, there is a surplus of available and eager males, and one must be chosen among many. However it may be, courting is most practiced not only when the female is in command of the final outcome but also when the mating procedure presents certain difficulties. A small male spider dances before a larger and ever ravenous female in an effort to induce her sexual interest rather than her hunger. Birds especially, however, depend on courtship as a preliminary to mating. The mating of birds represents copulation in its simplest form, without benefit of significant anatomical devices. Bird wings are a poor substitute for arms in a sexual embrace. Consequently the fullest cooperation between male and female is essential to success. In most birds a long-lasting, often lifetime, bonding becomes established between a male and female, a bonding that is usually reinforced by ritual behaviour at certain intervals, particularly during the onset of each breeding season and on various occasions when the individuals meet after short periods of separation. In some species a new mate may be taken each season or, as in sparrows, a general promiscuity may prevail.
One important aspect of courtship concerns the question of recognition. In gull colonies, for instance, members of the opposite sex look very much alike, and, at least to humans, the various individuals of one sex or the other may appear exactly the same. The advantages, with regard to successful production, incubation, and rearing of eggs and young, of permanent or semipermanent mate selection, however, are as great in gull colonies as elsewhere. The preliminaries to such a mutual selection not only establish a bond, by various posturings, but also establish the many small idiosyncrasies of action that add up to individuality and make one bird distinguishable among many within a colony, at least to its mate.
Many different forms of sex-oriented behaviour have consequently evolved among birds, depending on the character and particular needs of the various species. Penguins apparently not only look alike to human observers but also to themselves. Penguins seemingly have trouble even distinguishing between the sexes. Being unable to dance or sing, though they can make a lot of noise, male penguins can do little more than offer a pebble to a prospective female. If she accepts it as a token contribution to nest making, the match is on. If it is rejected, the suitor may have picked an unready female or even another male. In the case of most birds, however, the male can either sing, particularly the smaller kinds, or can strut and dance, with wings and feathers displayed, and some species, such as the lyre bird, continue to enchant the female by sight and sound together. In general, the need for physical mating has led to courtship and an emotional bonding between mating pairs throughout much of the animal kingdom at the higher level, particularly among birds and mammals. These are primarily utilitarian functions relating to the survival of the species, but in their fullest expression they represent what seem to man to be among the finest attributes of life.