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2012 COURSE DATES: AUGUST 4 – 17, 2012 - Sirenian International

2012 COURSE DATES: AUGUST 4 – 17, 2012 - Sirenian International

2012 COURSE DATES: AUGUST 4 – 17, 2012 - Sirenian International

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polyandrous - often mating with several males over a<br />

period of several hours. When Hartman (1979) compares<br />

the mating behavior of manatees to that of elephants, he is<br />

writing from an ultimate evolution perspective. In other<br />

words, he is hypothesizing that this aspect of the mating<br />

system evolved millions of years ago in an ancestor shared<br />

by both the manatee and the elephant. Since sirenians and<br />

proboscideans are two of only four extant orders that share<br />

a common ancestor among them, manatees are often<br />

compared to elephants using the ultimate evolution<br />

perspective [NOTE: The other two orders contain the<br />

hyraxes and the aardvarks, which are rarely compared to<br />

manatees in the literature]. Another promiscuous aspect of<br />

the manatee mating system is scramble-polygyny, where<br />

multiple males attempt to mate with the estrous female, but<br />

- without overt competition. Although males aggregate on<br />

the estrous female and jockey for the best position - they<br />

exhibit little agonistic behavior. Interestingly, male<br />

dugongs appear to be more agonistic during mating events.<br />

They set up territories and exhibit lek mating behaviors<br />

(Anderson 1997). What perspective would we use to<br />

compare mating strategies between manatees and dugongs?<br />

Timing: Let's assume that our observation was of a<br />

mating herd. That is, the group of manatees in the center<br />

of the cove consisted of 1 estrous female and 3 males.<br />

Why was the first manatee, the one originally sighted in the<br />

resting hole, not involved in the mating herd? Looking at<br />

the situation from a proximate perspective, there are<br />

several possibilities, and all involve timing. Suppose the<br />

resting manatee was a female. If she was sexually mature,<br />

but not in estrous, the mating herd would have no interest<br />

as she would not be producing an estrous signal. An<br />

estrous signal is the proximate cause of the mating herd<br />

behavior. It's "how" the males know the female is ready to<br />

conceive. Similarly, if the female were sexually immature,<br />

she could not be in estrous and therefore would not be<br />

sending a signal. Scientists have only recently answered<br />

the question of when a female manatee becomes sexually<br />

mature, thanks to the development of a new aging technique<br />

by Miriam Marmontel, et al. (1990). Since manatees<br />

continuously regenerate new teeth throughout their lives<br />

(Domning and Hayek 1984), they cannot be aged by their<br />

dentition like many other marine mammals. But, by<br />

looking at growth layers in manatee ear bones, we are now<br />

reasonably confident that female manatees in Florida reach<br />

sexual maturity between the age of 3 and 4 years - most<br />

giving birth to their first calf at age 4 (Marmontel 1995).<br />

Questions of "how" the behavior of signaling develops in<br />

females as they mature fall under the proximate<br />

development perspective.<br />

On the other hand, if the resting manatee was a<br />

male, why wasn't he attracted to the estrous female in the<br />

middle of the cove? He could have been either sexually<br />

immature or sexually inactive. Using the presence or<br />

absence of sperm in the testes as an indicator, Hernandez et<br />

al. (1995) found that sexual maturity (proximate<br />

development) varied among Florida male manatees with<br />

5<br />

both size and age with some males becoming<br />

physiologically mature as young as 2 years and as small as<br />

237 cm. But, from a proximate cause perspective, they<br />

also found that the reproductive system varied in<br />

functionality among mature male manatees depending on<br />

season in Florida, with little evidence of spermatogenesis<br />

present during winter months (December <strong>–</strong> February). In<br />

many mammals, reproductive activity varies seasonally<br />

with photoperiod, or the number of light hours per day.<br />

The pineal gland is usually the organ associated with<br />

behavioral changes affected by photoperiod. However, no<br />

pineal gland has ever been found in manatees or dugongs<br />

(Ralph et al. 1985, W. Welker personal communication<br />

2000). From an ultimate evolution perspective, it is<br />

interesting that the literature is unclear regarding the<br />

existence of a pineal gland in elephants (Ralph et al. 1985).<br />

Whether the resting manatee was inactive or<br />

immature, his timing would have been out of sync with the<br />

female and the estrous signal would have no effect on his<br />

behavior. From an ultimate function perspective, we say<br />

that those manatees whose sexual behavior is triggered at<br />

the appropriate time (i.e. when both the male and female are<br />

sexually mature, active, and receptive) are more<br />

reproductively successful than manatees that waste energy<br />

on futile sexual encounters. From an ultimate evolution<br />

perspective, there have been some behaviors observed in<br />

Antillean manatees that might be associated with seasonal<br />

spermatogenesis (G. Smith unpublished data). More<br />

studies are necessary before we can determine if seasonality<br />

affects the reproductive behavior of manatees in Belize.<br />

Parental Care: The final aspect of reproductive<br />

problem solving discussed here is parental care. Like many<br />

mammals, female manatees invest considerable time and<br />

energy into a relatively small number of offspring as their<br />

reproductive strategy. From an ultimate function<br />

perspective, data collected by scientists in Florida suggest<br />

that those females that invest 2 years of parental care in<br />

each offspring prior to becoming pregnant again are more<br />

successful than other females (Marmontel 1995). Although<br />

calves begin eating on their own within 3 months of birth,<br />

they continue to nurse periodically (Hartman 1979) as they<br />

grow and learn migration routes from their mothers (R.<br />

Bonde, personal communication 1999). When we ask<br />

"why" this behavior exists, we are asking ultimate function<br />

questions. Perhaps calves need the extra protein and fat<br />

provided by mother's milk during developmental years. Or,<br />

perhaps it takes calves almost two years to learn the routes<br />

to warm water effluents and good foraging grounds<br />

necessary for survival through the temperate winters in<br />

northern and central Florida.<br />

In an ongoing study of Antillean manatees in the<br />

Southern Lagoon of Belize, Buddy Powell is also seeing<br />

mother-calf pairs remain together for long periods of time<br />

(www.wesave.org/manatee/). When we compare this<br />

behavior between the Florida and Antillean subspecies, we<br />

are using the ultimate evolutionary perspective. On the<br />

other hand, "how" the mother-calf pair remains together

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