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penis is the primary sexual organ that male and hermaphrodite animals use to inseminate sexually receptive mates (usually females and hermaphrodites respectively) during copulation. Such organs occur in many animals, both vertebrate and invertebrate, but males do not bear a penis in every animal species, and in those species in which the male does bear a so-called penis, the penes in the various species are not necessarily homologous. For example, the penis of a mammal is at most analogous to the penis of a male insect or barnacle.
The term penis applies to many reproductive intromittent organs, but not to all; for example the intromittent organ of most cephalopoda is the hectocotylus, a specialised arm, and male spiders use their pedipalps. Even within the Vertebrata there are morphological variants with specific terminology, such as hemipenes.
In most species of animals in which there is an organ that might reasonably be described as a penis, it has no major function other than intromission, or at least conveying the sperm to the female, but in the placental mammals the penis bears the distal part of the urethra, which discharges both urine during urination and semen during copulation as the occasion requires.
Most male birds (e.g., roosters and turkeys) have a cloaca (also present on the female), but not a penis. Among bird species with a penis are paleognathes (tinamous and ratites), Anatidae (ducks, geese and swans), and a very few other species (including flamingoes and chickens). A bird penis is different in structure from mammal penises, being an erectile expansion of the cloacal wall and being erected by lymph, not blood. It is usually partially feathered and in some species features spines and brush-like filaments, and in flaccid state curls up inside the cloaca. The Argentine Blue-bill has the largest penis in relation to body size of all vertebrates; while usually about half the body size (20 cm), a specimen with a penis 42.5 cm long is documented.
While most male birds have no external genitalia, male waterfowl (Anatidae) have a phallus. Most birds mate with the males balancing on top of the females and touching cloacas in a “cloacal kiss”; this makes forceful insemination very difficult. The phallus that male waterfowl have evolved everts out of their bodies (in a clockwise coil) and aids in inseminating females without their cooperation. The male waterfowl evolution of a phallus to forcefully copulate with females has led to counteradaptations in females in the form of vaginal structures called dead end sacs and clockwise coils. These structures make it harder for males to achieve intromission. The clockwise coils are significant because the male phallus everts out of their body in a counter-clockwise spiral; therefore, a clockwise vaginal structure would impede forceful copulation. Studies have shown that the longer a male’s phallus is, the more elaborate the vaginal structures were.