Why do some mammals not have testes in a scrotum?

Question

Coming from an evolutionary approach, Is the only purpose of a scrotum to regulate the temperature of the testes?

Knowing all mammals are warm blooded, shouldn't all mammals have testes in a scrotum?

Answer 1

Having descended testes is a derived characteristic within mammals; monotremes and the Afrotheria (including elephants) all retain the ancestral character state (Kleisner, et al., 2010)². Among those mammals with descended testes, these can be ascrotal or scrotal. Testicular descent is hypothesized to have only occurred once within Mammalia, with the ascrotal Laurasiatheria. Descended ascrotal testes are found in cetaceans, phocid seals, hippos, tapirs, rhinos, and some bats. Descended scrotal testes are found in horses, pigs, camels, and Carnivora.

Since basal mammals would presumably have to regulate testicular temperature just as much as derived mammals, the temperature regulation hypothesis seems to not hold up. So the real question is: why have a scrotum? One hypothesis has to do with evolution of fast locomotion (e.g., galloping).

According to Frey (1991, 40)⁴:

The strong flexions and extensions of the vertebral column during gallop should cause intense fluctuations of intra-abdominal pressure. Fluctuations of intra-abdominal pressure severely impede continuous flow of blood in the abdominal veins. Periodically reduced venous drainage resulting in fluctuations of intra-testicular pressure would impair the process of spermiohistogenesis, which is dependent on an absolutely constant pressure within the testis.

Chance (1996)⁵ suggests that the temperature hypothesis might represent a secondary adaptation:

Because in the human male, scrotal testes function optimally at temperatures below that of the body, much speculation, and a considerable amount of research, has gone into attempting to see what (metabolic) advantage might accrue from this lower temperature, without considering the possibility that this is a secondary adaptation to an enforced external position.

Kleisner, K., Ivell, R., and Flegr, J. 2010. The evolutionary history of testicular externalization and the origin of the scrotum. J Biosci 35:27-37

Frey, R. 1991. Zur Ursache des Hodenabstiegs (Descensus testiculorum) bei Säugetieren. Z Zool Sys Evolut-Forsch 29:40–65

Chance, M.R.A. 1996. Reason for externalization of the testis of mammals. J Zool 239:691–695