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Clitoris
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===Spotted hyenas=== [[File:Spotted hyena in Kenya.jpg|thumb|With a [[Genitourinary system|urogenital system]] in which the female urinates, mates and gives birth via an enlarged, erectile clitoris, female spotted hyenas are the only female mammals devoid of an external vaginal opening.<ref name="Glickman" />]] While female [[spotted hyena]]s were sometimes referred to as pseudohermaphrodites<ref name="Roughgarden 2004 pp37-40" /> and scientists of ancient and later historical times believed that they were hermaphrodites,<ref name="Roughgarden 2004 pp37-40" /><ref name="Glickman" /><ref name="Rosenzweig" /> modern scientists do not refer to them as such.<ref name="Glickman" /><ref name="Blumberg" /> That designation is typically reserved for those who simultaneously exhibit features of both sexes;<ref name="Blumberg" /> the genetic makeup of female spotted hyenas "are clearly distinct" from male spotted hyenas.<ref name="Glickman" /><ref name="Blumberg" /> Female spotted hyenas have a clitoris 90 percent as long and the same diameter as a male penis (171 millimetres long and 22 millimetres in diameter),<ref name="Roughgarden 2004 pp37-40" /> and this [[pseudo-penis]]' formation seems largely [[androgen]]-independent because it appears in the female fetus before differentiation of the fetal ovary and [[adrenal gland]].<ref name="Glickman"/> The spotted hyenas have a highly erectile clitoris, complete with a false [[scrotum]]; author John C. Wingfield stated that "the resemblance to male genitalia is so close that sex can be determined with confidence only by palpation of the scrotum".<ref name="Wingfield 2006 p2023" /> The pseudo-penis can also be distinguished from the males' genitalia by its greater thickness and more rounded glans.<ref name="Glickman" /> The female possesses no external vagina, as the labia are fused to form a pseudo-scrotum. In the females, this scrotum consists of soft adipose tissue.<ref name="Wingfield 2006 p2023" /><ref name="Glickman" /><ref name="Szykman"/> Like male spotted hyenas with regard to their penises, the female spotted hyenas have small [[penile spines|spines]] on the head of their clitorises, which scholar {{ill|Catherine Blackledge|pl}} said makes "the clitoris tip feel like soft sandpaper". She added that the clitoris "extends away from the body in a sleek and slender arc, measuring, on average, over 17 cm from root to tip. Just like a penis, [it] is fully erectile, raising its head in hyena greeting ceremonies, social displays, games of rough and tumble or when sniffing out peers".<ref name="Blackledge 2003 p90" /> [[File:Anatomischer Anzeiger (1922) (18006271698).jpg|thumb|upright=1.8|'''Male and female reproductive systems of the spotted hyena, from Schmotzer & Zimmerman, ''Anatomischer Anzeiger'' (1922)'''. Abb. 1 (Fig. 1.) ''Male reproductive anatomy''. Abb. 2 (Fig. 2.) ''Female reproductive anatomy''.<ref name="Schmotzer" /> Principal abbreviations (from Schmotzer & Zimmerman) are: '''T''', testis; '''VD''', vas deferens; '''BU''', bulbus urethrae; '''Ur''', urethra; '''R''', rectum; '''P''', penis; '''S''', scrotum; '''O''', ovarium; '''FT''', tuba Fallopii; '''RL''', ligamentum uteri; '''Ut''', uterus; '''CC''', corpus clitoridis. Remaining abbreviations, in alphabetical order, are: ''AG'', glandula analis; ''B'', vesica urinaria; ''CG'', glandula Cowperi; ''CP'', corpus penis; ''CS'', corpus spongiosum; ''GC'', glans clitoridis; ''GP'', glans penis; ''LA'', musculus levator ani; ''Pr'', praeputium; ''RC'', musculus retractor clitoridis; ''RP'', musculus retractor penis; ''UCG'', canalis urogenitalis.]] {{anchor|Genital grooming}} Due to their higher levels of androgen exposure during fetal development, the female hyenas are significantly more muscular and aggressive than their male counterparts; social-wise, they are of higher rank than the males, being [[Dominance (ethology)|dominant]] or dominant and [[Alpha (ethology)|alpha]], and the females who have been exposed to higher levels of androgen than average become higher-ranking than their female peers. Subordinate females lick the clitorises of higher-ranked females as a sign of submission and obedience, but females also lick each other's clitorises as a greeting or to strengthen social bonds; in contrast, while all males lick the clitorises of dominant females, the females will not lick the penises of males because males are considered to be of lowest rank.<ref name="Szykman"/><ref name="Carey"/> The female spotted hyenas urinate, copulate and give birth through the clitoris since the urethra and vagina exit through the clitoral glans.<ref name="Wingfield 2006 p2023" /><ref name="Glickman" /><ref name="Blackledge 2003 p90" /><ref name="Baskin"/> This trait makes mating more laborious for the male than in other mammals, and also makes attempts to [[Sexual coercion among animals|sexually coerce]] (physically force sexual activity on) females futile.<ref name="Szykman"/> [[Joan Roughgarden]], an [[ecologist]] and [[evolutionary biologist]], said that because the hyena's clitoris is higher on the belly than the vagina in most mammals, the male hyena "must slide his rear under the female when mating so that his penis lines up with [her clitoris]". In an action similar to pushing up a shirtsleeve, the "female retracts the [pseudo-penis] on itself, and creates an opening into which the male inserts his own penis".<ref name="Roughgarden 2004 pp37-40" /> The male must practice this act, which can take a couple of months to successfully perform.<ref name="Carey"/> Female spotted hyenas exposed to larger doses of androgen have significantly damaged ovaries, making it difficult to conceive.<ref name="Carey"/> After giving birth, the pseudo-penis is stretched and loses much of its original aspects; it becomes a slack-walled and reduced prepuce with an enlarged orifice with split lips.<ref name="Rosevear 1974 pp357β358" /> Approximately 15% of the females die during their first time giving birth, and over 60% of their species' firstborn young die.<ref name="Roughgarden 2004 pp37-40" /> A 2006 Baskin et al. study concluded, "The basic anatomical structures of the corporeal bodies in both sexes of humans and spotted hyenas were similar. As in humans, the dorsal nerve distribution was unique in being devoid of nerves at the 12 o'clock position in the penis and clitoris of the spotted hyena" and that "[d]orsal nerves of the penis/clitoris in humans and male spotted hyenas tracked along both sides of the corporeal body to the corpus spongiosum at the 5 and 7 o'clock positions. The dorsal nerves penetrated the corporeal body and distally the glans in the hyena", and in female hyenas, "the dorsal nerves fanned out laterally on the clitoral body. Glans morphology was different in appearance in both sexes, being wide and blunt in the female and tapered in the male".<ref name="Baskin" />
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