The mane of the African Lion
Throughout history, manes of male lions have inspired admiration and fear yet no agreement has been reached concerning its function.
The modern lion is instantly distinguished from the other living animals of the Panthera (cat) family but the mane on males and by group-living. All members of the Panthera family are thought to share a common ancestor, and yet these features are unique to lions. We can assume that they evolved after the lion separated from other members of the Panthera family and that both are genetic responses to selection pressures.
However, not all lions sport manes; the lions of Tsavo National Park in Kenya are largely maneless!
Three elements must be considered when looking at manes; only males have them; manes start to grow at puberty; and manes vary greatly in both colour and extent, both between and within populations. Such elements lead to the assumption that the mane is, therefore, a product of sexual selection, either for competition between males or in making a male more attractive to females.
Female lions live in prides resulting in severe reproductive competition. This has lead to the formation of coalitions of males working together to take over a pride. Larger coalitions are more likely to be able to oust a sitting dominant male, are more likely to hold the pride tenure for a longer period and are therefore able to produce more cubs.
Females reproduce in synchrony meaning that more than one female may be in oestrus at any one time, allowing several members of the coalition opportunities to mate. This social system, therefore, allows for sexual selection between males as well as in mate choice.
Darwin suggested that manes were used to protect the male during fights, however, there is a greater advantage of avoiding a fight altogether. This requires a signal that can convey the fighting ability.
From the females’ point of view, incoming males kill all the cubs in the pride so that they can start to reproduce and pass on their genes. A female can benefit from a signal that a male is able to protect her cubs. Further, males are an important asset in providing food for the pride. Buffalo are slow prey and require the strength of a male, rather than the speed of a female to catch. This makes a male’s hunting ability and physiological condition very valuable to females
In sexual selection, hair is condition-dependent and its extent and colour are linked to testosterone levels as well as nutritional deficiencies. A study by West (2005) therefore hypothesised that manes may function in any of three ways; as a shield against injury, as a signal of the males fighting ability and as a sign of a males physiological status. The study set out to look at each of these three functions:
If true, the first function, that of shield against injury, would suggest that males with larger manes are injured less frequently or less severely. As fights between lions are rarely witnessed, the study looked at two other ways of testing the hypothesis. Firstly, if the mane is a protection then that would suggest that the mane area is attacked more often and that these areas would, therefore, show more wounds, and secondly, that injuries to these areas would be more severe. The study, however, found no evidence that the mane area was more likely to be attacked, nor with greater severity, and wounds on males showed the same distribution and severity as were found in females and sub-adults that do not have manes.
The study started to look at the second and third possible functions, that of the mane as a signal of fighting ability and physiological condition. It was found that the manes of males of the Serengeti study area changed over time both in length and pigmentation, and also that manes grew longer and darkened with the increase in testosterone during adolescence.
The study also found a correlation between mane length and colour and several ecological factors; in males over five years of age, mane length was closely associated with injury, leading to a possible signal that a short mane was an indication of lower fighting ability.
Testosterone levels were found to be higher in males with darker manes, and therefore a possible signal of increased fighting ability. Further, darker maned males were generally better fed suggesting a higher dominance level or better hunting ability.
Given the possible signalling properties of the mane, the study set out to test whether lions actually used these signals in their social behaviour. Loudspeakers were used to broadcast the sounds of a female in oestrus. Although mane length was not a significant indicator, males with darker manes were far more likely to win the race to the “female”.
Where there is more than one female in oestrus at any given time, the female is able to make a choice as to which male to mate with. The study, therefore, looked at mating situations where a male mated with more than one female within an hour, on the assumption that at least one of the females was there by choice. In only 50% of examples did the females choose a male with the longest mane length, however in all but one case, the female chose the male with the darkest mane.
To try and answer the question why females did not respond to the signal of mane length when a short mane could be an indicator of injury, the study used dummy lions of varying mane length and colour. The tests showed that females would approach a darker maned dummy, but did not take account of mane length. Males avoided the dark maned males, and were extremely sensitive to mane length, avoiding the longer maned dummy. Data from these studies, when added to results from previous studies concluded that darker maned males are more likely to reside in a pride, are more likely to survive the injury, have more surviving offspring, and are less likely to be wounded. Short manes appear to be a signal of short term injury. The study went on to ask the question of why all males do not have dark manes, hypothesising that there must be some sort of cost to growing and maintaining one.
Males in cooler climates tend to have darker manes than those that live in warmer climates, and a male taken from a warm climate to a cool one is likely to show a darkening of their mane as they acclimatise to the new location. Lions are particularly sensitive to heat as they do not sweat and can only dispel heat through their skin, a process made less efficient by having a mane, especially a dark one. It is therefore highly likely that only the strongest, fittest males can afford to maintain a dark mane.
The study goes on to suggest that global warming and rising temperatures will have an effect on lion populations as the heat stress of darker manes becomes too much and the trait will become less common, with negative effects on photographic tourism and legal trophy hunting industries. For the lions themselves, it may become necessary to adapt behaviorally and/or physiologically to overcome the loss of this important sexual selection signal.
So why are the lions of Tsavo maneless? A study of the number of females in prides has refuted the notion that smaller group sizes lead to reduced sexual competition. Further, hunting has been banned within the Park for many years dispelling the idea that the off-take of large maned males has bred smaller manes into the population. As such, maybe there is a higher environmental cost to holding a mane in this hot, arid area? The thornbush habitat may have an effect. Males tend to take part in hunting more often in closed bushy habitats than in open grassy plains where their conspicuous mane makes hunting more difficult. In such habitats, a mane can easily get snagged as they stalk through the Bush impeding movement and making noise. Thirdly, the burrs of the plant Pupalia lappacea are abundant in Tsavo and even the dextrous fingers of the local baboons struggle to entangle them from long hair.
What is not clear is whether manelessness has evolved as an adaptation to some aspect of the Tsavo environment (i.e., genetic cause) or if the thorns and burrs merely pull out the mane as it grows in adolescent males (i.e., environmental cause). If mane hair is pulled out in each generation of male lions, young males would be expected to have larger manes, and individuals in more open habitats would be expected to retain their manes longer. If manelessness is a newly evolved adaptation, the mechanism may involve testosterone (or its derivatives), which is linked to hair growth and hair loss in mammals.
The mane in males develops during their second year and is complete in the fifth; mane colour ranges from blond to black.
