I was recently asked a reasonable and intelligent question on elephants. One thing led to another, and after a bit of research I discovered the fascinating world of elephant masturbation. If you want to collect sperm from a (captive) elephant, how do you do it? Luckily youtube provides the answer. It seems that manual stimulation of the penis (1) just doesn't do it for elephants, and (2) is physically dangerous (read on), so you have to stick your arm into the animal's rectum and vigorously stimulate its prostate gland. How vigorously? Watch the video. The eventual result - and I'm not entirely sure how long this takes - is ejaculation...
Of course male elephants don't (ordinarily, so far as we know) have any trouble ejaculating when interacting sexually with other elephants, and may ejaculate prematurely during their efforts to achieve successful penetration (Bagemihl 1999). It can take (presumably inexperienced) males a bit of rooting around with the flexible penis before finding the vulva. As is well known, the elephant vulva is not located just beneath the tail as is the case in most mammals, but down on the ventral surface and well between the legs, meaning that the vagina is very long (70-90 cm). This unusual position means that pregnancies are sometimes unnoticed and even that females have on occasion been mistakenly identified as male: Shoshani (1991) wrote of a case where a female was thought to be a male throughout her 25 years of life and was only found to be female on her death. Why elephants are built like this has been a long standing question.
In order for mating to be successful then, the penis has to be both long and flexible. Naturally S-curved (with the tip pointing upwards), it can be up to 1 m long (some sources say up to 1.5 m) and have a girth of 16 cm. It has a Y-shaped external urethal opening and is controlled by a large levator penis muscle. The clitoris is also large, at a whopping 40 cm, and is manipulated by a levator clitoris muscle. Perhaps the most interesting bit of information (provided in the video by elephant reproduction expert Robert Hermes) is that, not only does manual stimulation fail to work on elephants, it's also physically dangerous. Apparently there are parts of the elephant penis that, when touched, cause the organ to flick around with enough force to knock over an unwary veterinarian or even inflict a black eye. Don't believe me? Watch the second video. Or... check the technical literature (Schmitt & Hildebrandt 1998, Portas et al. 2007).
And, ha ha ha, while this is all very amusing, the reason that sperm needs to be collected from elephants is of course for use in insemination and in building breeding stocks. Bull elephants are relatively few in captivity, and those that exist seem to suffer from an unusually high number of fertility problems. This has initiated recent ultrasonographic work that seeks to find and identify good sperm donors (Hildebrandt et al. 2000). So... now you know. By the way, penises and vaginas are all very interesting, but does anyone actually know how an elephant's trunk works?
If you're particularly interested in elephants be sure to read the recent piece on Jeheskel Shoshani.
Refs - -
Bagemihl, B. 1999. Biological Exuberance. Animal Homosexuality and Natural Diversity. St. Martin's Press, New York.
Hildebrandt, T. B., Hermes, R., Pratt, N. C., Fritsch, G., Blottner, S., Schmitt, D. L., Ratanakorn, P., Brown, J. L., Rietschel, W. & Göritz, F. 2000. Ultrasonography of the urogenital tract in elephants (Loxodonta africana and Elephas maximus): an important tool for assessing male reproductive function. Zoo Biology 19, 333-345.
Portas, T. J., Bryant, B. R., Göritz, F., Hermes, R., Keeley, T., Evans, G., Maxwell, W, M. C. & Hildebrant, T. B. 2007. Semen collection in an Asian elephant (Elephas maximus) under combined physical and chemical restraint. Australian Veterinary Journal 85, 425-427.
Schmitt, D. L. & Hildebrandt, T. B. 1998. Manual collection and characterization of semen from Asian elephants (Elephas maximus). Animal Reproduction Science 53, 309-314.
Shoshani, J. 2000. Anatomy and physiology. In Shoshani, J. (ed) Elephants. Checkmark Books (New York), pp. 66-81.
With six years of phd work on theropod dinosaurs behind him, Darren Naish mostly spends long hours in the library, hunched over his laptop. But he gets out sometimes, and picks up litter and pursues exotic lizards across the British countryside, aiming all the while to publish his technical work on obscure Cretaceous dinosaurs. He also messes around with pterosaurs, swimming giraffes, British big cats and stuff like that. He has given up on the stupid idea of being a dedicated academic and ekes out a living as a technical consultant, editor and author. He can be contacted intermittently at eotyrannus (at) gmail dot com. For more biographical info go

Comments
Oh.My.God. That was unexpected.
I'd like to point out that my reasonable and intelligent question was IN NO WAY related to elephant masturbation techniques.
Posted by: Carpworld | June 25, 2008 8:38 AM
Yeah yeah SURE it wasn't :)
Posted by: Bad Bunny Boingo | June 25, 2008 8:50 AM
How much time do you spend each day reading primary literature? 25 hours?
Clitoridis. Latin declension is occasionally unpredictable.
Posted by: David Marjanović | June 25, 2008 8:54 AM
26.
Yes, the correct name is erectores clitoridis, but the reference I was looking at at the time (Shoshani 2000) used the vernacular 'levator clitoris'. I can live with this.
Posted by: Darren Naish | June 25, 2008 9:05 AM
Oh, I employ a female elephant to do that...
Posted by: Tengu | June 25, 2008 11:24 AM
... I will never, ever, ever, ever, ever understand what drives people to go into Zoologic Medicine.
Posted by: Nentuaby | June 25, 2008 1:50 PM
WHY, Darren? WHY?
Also, I think the phrase should be changed to "hung like a pachyderm."
Posted by: Zach Miller | June 25, 2008 2:43 PM
When I was in Egypt, someone remarked he thought tending to camels was the lowest job possible; lower than cleaning toilets. I would have to say this job is worse than tending to camels.
Posted by: Michele | June 25, 2008 3:50 PM
FOAF once had a job at the Ministry of Agricultures artificial insemination place.
His job, would you believe? was Milking the Bull.
And he had to wear a rubber suit for it too. (But no prostrate prodding, mercifully, just...)
Posted by: Tengu | June 25, 2008 4:48 PM
Thanks a lot, I just recommended your site to a colleague the other day. I can only imagine what they will think :)
Posted by: Dr. Tachyon | June 25, 2008 5:21 PM
The things I learn on this site LOL
Yours truly is one of the most brilliant, informative, and DIVERSE blogs out there.
This was definately not what I expected to learn about this morning, and I feel I should now have a sock on my door or something while surfing hehehehe
Posted by: Traumador the Tyrannosaur | June 25, 2008 8:02 PM
How did they figure this technique out and who was the brave soul who first ventured to try it? If there is ever a medal awarded to heroes of conservation biology . . .
Posted by: cfrost | June 26, 2008 3:28 AM
>Why elephants are built like this has been a long standing question.
Would it be so that their babies don't have a fatally long drop on birth? I've seen footage of elephant birth (quite rare apparently) and the mother didn't appear to crouch, so that even from its lower starting point the baby hits the ground with quite a thud.
cheers
Adam
Posted by: Adam Yates | June 26, 2008 3:36 AM
Shouldn't there at least be some Barry White on the video as a backing track?
Posted by: Dave Hone | June 26, 2008 4:35 AM
Adam, "Hi"! And secondly, I'd be surprised if that was it - female giraffe happily drop their spawn from about 6-7 ft (2 m) to the ground and those babies are far more spindly with breaky bits like legs and necks compared to a relatively robust elephant calf. It might well have an effect, but I'd be very surprised if that was the whole reason.
Posted by: David Hone | June 26, 2008 9:52 AM
.. I was thinking giraffes too. Furthermore, if (a speculative 'if', but one that I can kid myself into accepting) the genital morphology of modern elephants arose at the base of Elephantimorpha (the mammutidan + elephantidan clade), the most basal forms we know of were, while fundamentally elephanty in shape and proportions, not too big. The holotype adult (c. 26 yr old) Eritreum, for example, had a shoulder height of 1.2-1.3 m (Shoshani et al. 2006), which is much less than that of, say, most cattle. More basal proboscideans are of course even smaller, but then they are not elephantine in proportions and it seems easier to assume that they had a different urogenital system.
I don't know about you, but I find it difficult to think that mammutids and so on were fundamentally different from extant elephants in this regard... though I am speculating. The way to have a go at testing this is to look long and hard at proboscidean pelves. I've done this and don't see much that gives the game away as goes the morphology of the urogenital system but, hey, I'm no expert (incidentally, there is a pretty good literature on fossil elephant pelves, some of it devoted to sexual dimorphism). We do know that Mammut exhibited the same sort of sexual dimorphism in the pelvis as do living elephants (Green 2006), perhaps implying the same sort of soft-tissue morphology.
Ref - -
Green, J. L. 2006. Chronoclinal variation and sexual dimorphism in Mammut americanum (American mastodon) from the Pleistocene of Florida. Bulletin of the Florida Museum of Natural History 46, 29-59.
Shoshani, J., Walter, R. C., Abraha, M., Berhe, S., Tassy, P., Sanders, W. J., Marchant, G. H., Libsekal, Y., Ghirmai, T. & Zinner, D. 2006. A proboscidean from the late Oligocene of Eritrea, a "missing link" between early Elephantiformes and Elephantimorpha, and biogeographic implications. Proceedings of the National Academy of Sciences 103, 17296-17301.
Posted by: Darren Naish | June 26, 2008 11:20 AM
Do elephants have any particular structures or set-ups to keep their testes cool? They are way in there...
Posted by: Darby | June 26, 2008 12:31 PM
It's unclear from the diagram: does the elephantine prostate surround (and in later life expand to constrict) the urethra as in homo sap?
Just how widespread (so to speak) is that particular feature of unintelligent design, anyhow?
Posted by: Pierce R. Butler | June 26, 2008 1:05 PM
It's a pity no mastodonts appear to've had the good taste to freeze themselves so we could look up their naughty bits.
Posted by: Andreas Johansson | June 26, 2008 2:31 PM
Darby: how do elephants keep their testes cool? Good question - I don't know the answer, but you'll be very interested to hear that some workers have argued that the testicular position present in the group is a result of the aquatic ancestry most recently supported by Gaeth et al. (1999) and Liu et al. (2008)... go here.
Pierce: the elephant prostate (in both Elephas and Loxodonta) differs from that of humans in being a paired structure situated entirely dorsal to the urethra, rather than being a ring-like structure that surrounds the urethra (and each half of the organ is tri-lobed). Prostate infections in elephants are recorded but (so far as I can tell from a quick check of the literature) prostate problems do not cause constriction of the urethra. For unknown reasons, the prostate of Loxodonta is about twice as big as that of Elephas and has irregularly shaped internal cavities of unknown function. Source: the Hildebrandt et al. paper cited above.
Refs - -
Gaeth, A. P., Short, R. V. & Renfree, M. B. 1999. The developing renal, reproductive, and respiratory systems of the African elephant suggest an aquatic ancestry. Proceedings of the National Academy of Sciences 96, 5555-5558.
Liu, A. G., Seiffert, E. R. & Simons, E. L. 2008. Stable isotope evidence for an amphibious phase in early proboscidean evolution. Proceedings of the National Academy of Sciences 105, 5786-5791.
Posted by: Darren Naish | June 26, 2008 4:41 PM
The headline would be funnier if "you" were italicized, instead of "do".
The secret to cost-effective elephant semen harvesting is to find someone who really likes doing it -- but only let them watch, and charge them for it.
Posted by: Nathan Myers | June 27, 2008 2:13 AM
Isn't it simply the ancestral condition? I remember looking at a paper (perhaps in Evolutionary Theory, in any case very long ago) that said elephants and tenrecs have higher rates of evolution because they, well, don't keep their testes all that cool.
Posted by: David Marjanović | June 27, 2008 6:45 AM
Re: internal testes..
Nope, based on what's present in marsupials and other taxa, a scrotum is basal for therians at least. Werdelin & Nilsonne (1999) examined this issue and argued for multiple independent losses of the scrotum in eutherians. However, the phylogeny they mapped their characters on to would now be considered very out of date. Fully internal testicles (the so-called testicondy condition) seem to charaterise all afrotherians and I've seen some discussion of the possibility that testicondy is a synapomorphy of the group. The paper proposing rapid evolution resulting from hot testicles is by the same authors (Werdelin et al. 1999).
Ref - -
Werdelin, L. & Nilsonne, A. 1999. The evolution of the scrotum and testicular descent in mammals: a phylogenetic view. Journal of Theoretical Biology 196, 61-72.
- ., Nilsonne, A. & Fortelius, M. 1999. Testicondy and ecological opportunism predict the rapid evolution of elephants. Evolutionary Theory 12, 39-45.
Posted by: Darren Naish | June 27, 2008 7:47 AM
Oh hold on - you might have meant the ancestral condition inherited from other afrotherians, not from other non-afrotherian placentals. Sorry, I misunderstood, and had forgotten the earlier statement about internal testes resulting from an aquatic phase unique to proboscideans or tethytheres.
Posted by: Darren Naish | June 27, 2008 7:50 AM
No, I did mean the ancestral condition for therians. Aren't marsupial scrota cranial of the penis? Do monotremes have one? And the name Scrotifera suggests that (eu)lipotyphlans have testicondy, too...
Concering Werdelin et al., it's nice to see how reliable my memory is :-)
Posted by: David Marjanović | June 27, 2008 5:10 PM
I may be entirely wrong about this, but testicont animals should have extenstive vascularization of the inguinal cavity and therefore the vascular structures associated with the urogenital system. It's likely simply that the system found in female animals is developed in male animals. If I recall correctly, testicont males actually "attempt" to exteriorize their genitals during development, but then retract them. Prior to this, the vascular system common to females and prenatal males (first trimester in humans as a frame of reference) does not diverge, and genes should kick in. I am not sure. This may simply argue that testicont males are femininized.
Posted by: Jaime A. Headden | June 29, 2008 1:13 PM
Jaime: The expression "kick in" in the context of testicles makes me cringe involuntarily.
Posted by: Nathan Myers | June 29, 2008 11:19 PM
Great post, Darren... Are you planning to take over Olivia's place as Dr Tatiana?
Posted by: luca | June 30, 2008 7:33 AM
Link doesn't work.
Posted by: David Marjanović | July 5, 2008 5:27 AM