|
The Mane
|
I found a couple of articles about the lion's mane shared by @chui_ on Carnivora, and I though they are extremely interesting.
I have discussed before with @peter and also with @Spalea about the simmilarities about lions and men and how they fascinate us as a consequence of that simmilarity, whether we realize it or not. Reading this study, one more time I see confirmation of how much like us are them, or better yet, how much like them we as men are. Enjoy. The first, Of Lion Manes and Human Beards: Some Unusual Effects of the Interaction between Aggression and Sociality: By D. Caroline Blanchard. Abstract The function of manes in lions has been a topic of scientific interest since Darwin (1871) suggested that it provides protection in intraspecific fights. Recent experimental studies on wild lions have emphasized the role of female selection, but analyses of specific attack behaviors and targets, and the social consequences of manelessness for lions living in very hot climates suggest that male manes may indeed mitigate the outcomes of intraspecific male attack and thus serve a permissive function for multi-male + female groups, facilitating protection of prides against take-overs and infanticide by nomadic males. Humans also have unusual structural protections for the head, face and neck, areas that are especially accessible during intraspecies attack, and highly vulnerable to damage. One of these, the beard, consists of coarse hairs that grow indefinitely, but only for males, and only during and following puberty; suggesting that it, like the lion's mane, may serve as protection in intraspecies male fights. Such structural protections may reflect a specific combination of lethal weaponry and social life-style, particularly when these are developed so rapidly that they are not accompanied by the evolution of complex attack-inhibiting social behaviors. Patterson, 2004, p. 141 Quote:Lion manes are obviously a sexually selected trait. Manes are restricted to males, contribute little to the lion's general economy of life, and apparently present a number of real costs that would otherwise make it a candidate for elimination by natural selection…..Still, given the interest that people have focused on lions over the last 30,000 years, it is remarkable that we do not know whether manes are the product of male–male contest, or female choice, or possibly of bothAggression, Social Systems and Evolution In terms of phylogeny, aggression is among the oldest of evolved behavior patterns (Blanchard and Blanchard, in press). Exemplars of aggression have been reported in animals without a central nervous system; in a host of invertebrates; and in each of the seven classes of vertebrates, including the most primitive; agnatha, hagfish and lampreys (Malmqvist, 1983). There is an emerging consensus that one major function, i.e. adaptive consequence, of aggression, across animal species, is resource control (e.g. Wilson, 1971), with the further provision that aggression typically occurs in the context of competition from conspecifics over such resources. Winning may be highly adaptive because it results in immediate or longer—term enhancement of access to resources that are important for that species (Blanchard and Blanchard, 1984; Moynihan, 1998). Resources and their distribution are also major factors in the development of species-typical social systems (Rubenstein, 2009), of which within-species aggressive behaviors are one, important, component. Some species such as mice are particularly opportunistic and may show rapid and dramatic changes in social structure in accord with habitat alterations (Bronson, 1979; Gray and Hurst, 1997), a flexibility that may stem in part from their long-term status as human commensals, with its strong requirement of rapid adjustment to host-initiated changes (Blanchard et al. in press). However, the ecological conditions under which most mammalian species have evolved are less variable than those associated with human habitations and human geographic movements, and the social systems evolving in most habitats appear also to be more conservative and resistant to change than are those of mice. Evaluation of the rate of change in evolved characteristics of animals in response to important alterations in ecological conditions constitutes a difficult field of study, but some estimates may be obtained when a fossil record is available that permits information on relatively specific ecological changes. For example, it has been estimated that reductions in selective behavioral responsivity to rattlesnakes and in resistance to the venom of these snakes by California ground squirrels both decline over periods measured in the tens of thousands of years after rattlesnakes have disappeared from the squirrels’ habitats (Coss et al., 1993). As this example suggests, when ecological changes make some of the existing characteristics of species less adaptive than previously, the replacement of these by more adaptive characteristics can be a lengthy and variable process; moreover, maladaptive as well as adaptive aspects of the emerging behaviors may be relevant to the patterns that ultimately emerge. With complex behavior patterns such as those involved in social relationships an even longer time-frame might be needed than for venom resistance and other purely physiological adaptations. However, following long-term exposure to diverse situations, even closely related species would be expected to show behavioral differences based on what is most adaptive in their particular habitats: Rubenstein (2009, p. 243) gives the example of plains zebras (Equus burchelli), living in an environment in which close proximity of food and water allow female zebras to consistently form groups, in turn permitting males to vie for harems in this home range. In contrast, Grevy's zebra (Equus grevyi) inhabit locales where food and water are widely dispersed and scarcer, such that females forage alone and must travel between feeding and watering areas. In this species males form territories along the traveling routes, gaining breeding access to females that pass through or linger within the territory. In both the harem and territorial social systems, zebra males attempt to control access to females in breeding condition, but the time-frame and the location of these efforts vary with the conditions under which these animals have evolved. A Particular Case in Point: Felids Some of these considerations may be relevant to a question that was raised at the very beginning of the scientific study of the evolution of behavior (Darwin, 1871): What is the basis for the evolution of the lion's mane? Is this related to the social systems of lions, which are unique among felids? There are about 40 species of felids, all stemming from a split from other stem-line carnivores about 10–15 million years ago (mya). Animals that can be identified with existing species emerged over a period from about 10–12 mya (for ocelots) to very recently, possibly within historic times. Felids range in size from the black-footed cat (about 3 lbs) to the Siberian tiger, weighing about 200× more, and are world-wide in distribution, excepting only Antarctica and most remote islands. Felid social systems are relatively similar across species: Adult animals tend to be solitary except for females and their young, and amicable adult encounters tend to be connected with reproduction. A great deal is known of the specifics of aggression in Felids, due in large part to the work of Paul Leyhausen, who worked with domesticated cats and with other felid species in captivity, at the Max Planck Institute for Behavioral Physiology, at Wuppertal. Large felids, of the genus Panthera have also been the subjects of extensive field work, which has generally affirmed Leyhausen's conclusions about conservation of many aggressive behaviors and facial expressions across felid species. Briefly, Leyhausen (1978) described intraspecific aggression in these animals as a behavior that is capable of producing great damage, as all felids have weapon systems that have evolved to facilitate their roles as predators but are used also in within-species fighting. Leyhausen also indicated that aggression in felids is more regulated by effective defenses than responsive to the submission signals that are quite effective in reducing intra-group fighting in many canid species, such as wolves and dogs. By selecting animals and arranging situations, Leyhausen was able to polarize attack and defensive behaviors in domesticated cat subjects, or, alternatively, to maximize attack motivations for both combatants. In a highly polarized attack-defend situation, the attacker stands tall and advances directly toward the victim while emitting low growls. The defending animal crouches down, or, with a mixture of aggressive and fearful motives, may assume the classic “Halloween cat” stance with arched back, and erected hairs. As the attacker approaches contact, the defending animal may assume a contorted posture, half on its back but facing the other cat, enabling both fore- and hindpaws to be drawn up and opposed to the oncoming attacker. This stance conceals the nape, the major target site for intraspecific offensive attack. Both fore- and hindpaws may lash out at the attacker and disembowelment of the attacker is possible, particularly if it attempts to reach over to bite at the partly supine defender's nape. However, when both animals are highly motivated to attack, a frontal approach is typically utilized by both, resulting in animals facing each other and delivering forepaw blows largely toward the head and neck of the other. Cat social systems provide a strong enabling factor in the dangerousness of aggression in these animals. Because of their solitary life-style, there is little reason for a felid under attack from a member of its own species to stay and receive bites and blows from the highly developed weapon systems that are characteristic of all felid species. The only context in which fighting may be strongly adaptive is mating, where flight, even if successful in avoiding injury, results in a substantial reduction in a male's extended reproductive fitness. Given that felids, particularly the larger ones such as tigers and leopards, kill large prey, it might be assumed that fighting over such prey would constitute a frequent occasion for fighting. However, the solitary life-style of these animals also acts to reduce conspecific encounters over prey. Lion Social Systems Lions are a different story. First, with the exception of the domesticated cat, which is not yet a truly separate species as it still freely interbreeds to produce fertile offspring with its wild ancestors (Driscoll et al., 2007), the lion (Panthera leo) is the most recent cat species to emerge. While the earliest lion-like cat may date to the late Pliocene, cave paintings indicate that males had no manes (Barnett et al., 2006). Modern maned lions appear to have descended from a single population that arose about 200 thousand years ago, possibly in Africa, and spread throughout the northern hemisphere, replacing earlier lion-like cats in Europe as recently as 10–15,000 years ago (Yamaguchi et al., 2004). Although Yamaguchi et al. (2004) suggest, based on a comparative analysis of food abundance factors in living species, that the earlier Holartic cave lion might have been group-living as are modern lions, there is no direct evidence that this was the case. Thus this particular mode of sociality, highly unusual for felids, may have been characteristic of Holarctic lions (Panthera leo spelaea) or, it may have evolved, along with the male lion's mane, only in modern lions. The classic hypothesis for a relationship between group-living and the development of manes is that manes reduce the dangers associated with fighting within lion groups (Darwin, 1871; Ewer, 1968; Schaller, 1972; Blanchard and Blanchard, 1984). Insofar as group-living is adaptive, serious injuries resulting from fighting within the group would be maladaptive for both the attacker and the injured animal, encouraging the latter to leave the group or reducing its ability to join in protecting the group from attack by outsiders. This is a particularly important consideration for lions, in that take-over of prides by nomadic lions entails infanticide by the new pride males (Loveridge et al., 2006) providing an extremely direct and strong adaptive consequence for success or failure at take-over, which in turn reflects the strength and numbers of the male cohorts on each side of the battle. A recent proliferation of nature documentaries plus the development of YouTube and other online sources has made videos of fighting in a variety of animals available. There are literally dozens of lion fighting videos online, and some of these provide clear footage of adult males fighting, with continuity, in nonenclosed areas that appear to be the animals’ natural habitats. Although the films were certainly selected for dramatic impact, and it is sometimes difficult to ascertain whether any external provocation might have been involved, it seems highly unlikely that the animals’ actual fighting behaviors were trained or shaped. Moreover, a variety of animals and videotaping sources appear to have been involved, substantially reducing the possibility that the clips present a systematically biased view of the actual behaviors involved in serious fights in lions. The form of both attack and defense in male lions appears to be highly consistent across these clips, particularly demonstrating the mutual frontal attack seen in other felids. The only time that an attacked male voluntarily exposes its back to an attacker is when it is fleeing. Even then, as the pursuer comes close, the pursued animal typically turns toward it, presenting its own weapons. Even a young cub, attacked and ultimately killed by an adult male, turns to face its attacker, flipping on its back in a move reminiscent of defensive domestic cats, as contact becomes imminent. Arguments Against the Mane as Protection This view of male lions’ manes as protective has recently been challenged by findings regarding wound locations on wild males, females, and young; by studies measuring approaches to dummy male lions with varying mane characteristics; and by observations of maneless lions in the Tsavo area of Kenya. With reference to the first of these, West and Packer (2002) examined sites of “lion-inflicted wounds” on a large sample of lions, sorting them into four regions; neck/shoulder, as delineated by the perimeter of the mane; face; trunk; and legs. They found that wounds were fairly evenly distributed over all four regions in females and subadults, and over three, excluding the area covered by the mane, in adult males. Moreover, females and subadults were equally likely to survive wounds to each body part. These figures are not easy to reconcile with those of Schaller (1972) who describes three direct observations of lions killing lions (p. 189): Of these, two involved bites to the nape of the neck, while the third was a bite to the lower back, severing the vertebral column. A fourth example, not witnessed but examined while the victim was still alive, albeit expiring, involved a pride male with multiple deep wounds, including a broken saggital crest, and a penetrating wound to the chest, with tatters of his mane scattered over a 3 × 10 meter area (p. 48). These examples do not indicate that other sites are not involved in attacks of one lion on another, but they do suggest that bites to the head and nape, if they reach these targets, are likely to be especially lethal. Moreover, in 259 observations of aggressive acts between female lions and cubs at kills, about 75% involved slaps at the head and neck, suggesting that these are indeed targets, albeit of blows that appear less likely to leave wounds (Schaller, 1972, p. 134). It might also be noted that wound sites do not necessarily have the same meaning as do targets: If, as in domestic cats, female and subadult lions have behavioral defenses that serve to protect the neck and shoulder by interposing teeth and claws to the attacker, a lack of disproportionate wounds on these sites may simply reflect the success of such defenses. Indeed, if the face (one of the four areas measured) had a relatively even share of wounds (West and Packer, 2002) even though it is much the smallest of the four areas evaluated, this may suggest that faces were wounded disproportionately to their area, perhaps in consequence of a facing defense that served to protect the neck and shoulders. Our own observations of adult male lions suggest that facial wounds are very common (see Figure Figure1).1). At any rate, the argument that a lion's mane is protective applies to animals that have manes, i.e. adult males. If such protection were equally important for females and subadult males, and this was indeed the basis on which manes have evolved, then presumably females and subadult males would have manes as well. They do not. Figure 1 Adult male with one half of his beard removed.  *This image is copyright of its original author A second finding relates to approach and avoidance responses of wild lions to dummies set up to have long or short manes, and, light or dark manes. Females approached the dark-maned model, and the long-maned model, proportionately more than did males (West and Packer, 2002). Female choice of both is consonant with findings that dark-maned males have higher testosterone levels than light-maned males, and that mane length (and other aspects of quality) reflect health as well as genetic differences. Male avoidance of these particular dummies may well reflect the same factors, and is also consonant with a view that mane quality provides protection against attack, providing abundantly maned lions an advantage in agonistic situations. Sociality in Maneless Lions A particularly interesting factor in the relationship between lion manes and lion social systems is that there are two groups of lions in which males show much less abundant manes; Asian lions in the Gir forest of India, and the Tsavo lions of lowland Kenya. Due to human pressures on their habitat, the Gir lions are basically a remnant population, albeit with relatively high genetic diversity (Sachdev et al., 2005) but the Tsavo lions are doing well, and lack of a mane does not appear to have had any effect upon their reproductive efficiency (Gnoske et al., 2006), or on their predatory capacity: Two of these male lions were notorious for killing over 100 workers on the Kenya-Uganda railway, at the end of the 19th century (Patterson, 2004). Gir lions have substantially scantier manes than those in Africa, with the exception of the Tsavo region, while the Tsavo lions are typically characterized as maneless. Abundantly-maned male lions disproportionately overheat when the ambient temperature is high (West and Packer, 2002), strongly suggesting that the relatively poor manes of lions in both areas constitutes a direct evolutionary response to the heat-based maladaptiveness of manes in these local climates. Tsavo, lying between the upland plateau of Kenya and the coast, is within three degrees of the equator and has no cool months of the year. The Gir forest has year-round high levels of both heat and humidity, producing a climate where “discomfort from heat and humidity” is rated as high or extreme for 10 of the 12 months of the year (BBC weather service). These climate factors, taken in conjunction with direct measures of overheating of lions with manes in hot weather, appear to provide a sufficient and direct explanation for the manelessness of Tsavo and Gir males. If lion manes arise because they serve as a permissive factor in the context of multi-male prides, what happens to multi-male prides when males do not have manes? In particular, if mane reduction in these lions is an evolutionary response to long-term high heat and humidity in the environment, alterations in their social systems may be secondary to such male mane changes, i.e. represent an effect of mane changes on sociality. Surveying 13 lion groups of known age and sexual composition in Tsavo, Kays and Patterson (2002) reported that only 2 of the 13 groups had more than one male, and both of these were nomadic, male-only groups. Moreover, in only one of the two were the animals adults. The male-female prides surveyed had an average of more than seven females each, indicating that resource scarcity was not important in group size, and could not account for the presence of only a single male per group. These demographics constitute a very striking difference from lion prides in other areas of Africa. Schaller (1972; Table II) notes that in 14 Serengeti lion prides, there were between two and four adult males each, with no pride having only a single adult male. Schaller did not quantify mane characteristics in these males. However, in the Kays and Patterson (2002) study, mane scores based on length, thickness, and color of hair were determined for males of four prides and three nomad groups. Mean scores were somewhat higher for the pride males than for the nomad groups, with the latter containing a higher proportion of young males. However, these scores overlapped completely, ranging from 0 to 24 for the pride males and 0 to 24 for the nomad group males, suggesting that female choice based on male mane characteristics was not a major factor in the composition of prides. While this finding may seem somewhat at variance with West and Packer's (2002) finding that females tended to approach male dummies with darker/longer manes, it emphasizes that the motivations involved in such approaches are not clear. The appearance of a strange male in a female's pride territory may well elicit approaches based on motivations other than sexual interest (e.g. Schaller, 1972, p. 53), including assessment of the risk posed to herself and her offspring. The Gir lions show an even more deviant pattern of sociality, compared to those in most areas of Africa. In the Gir forest “..male and female lions lead separate lives and rarely associate with each other… Prides are composed of related females, their young, and subadult male offspring…” (Sunquist and Sunquist, 2002, p. 293). Thus in both Tsavo and Gir, the scantiness or lack of manes in male lions is associated with a particular change in sociality, in which adult males do not simultaneously associate with both females and each other. Notably, the difference does not involve a reduction in association with females per se, as the Tsavo lion prides typically include one male and several females, nor does it involve a lack of male associations, as both Tsavo and Gir males often associate in male-only groups. The crucial feature appears to be whether multiple males and females associate in relatively long-term relationships, i.e. prides. In the areas where they occur, these prides are crucially involved in successful lion reproduction. In Kruger, Smuts (1982) noted that prides without males in constant attendance failed to raise any cubs. As females are the primary hunters in most prides (Schaller, 1972), this failure to rear cubs probably is not due to cub starvation in prides without males, but instead is the consequence of infanticide by nomadic males (Bertram, 1978; Packer and Pusey, 1983). This suggests that multi-male, mixed-sex lion prides enhance reproductive success by protecting the pride from take-over and subsequent infanticide by other males. If this analysis is valid, the adaptiveness of male manes may be to allow males within prides to fight, while reducing the chance of lethal contact with the particularly vulnerable sites that are normally protected by the mane. There is some difference of opinion (or observation) about the levels of aggression by male lions in a courtship context, with some authorities reporting vigorous fighting (Guggisberg, 1961) while others (Schaller, 1972) do not. Insofar as success in male–male fighting influences access to females in estrus, the male mane would have an obvious adaptive advantage in this context. However, fighting in contexts other than mating can also provide an arena for realization of the adaptiveness of manes in pride males. In particular, lions show little restraint of aggression when feeding, showing few inhibitions about slashing and biting fellow pride members: Schaller (1972; p. 135) suggests that this is due to their lack of a true dominance hierarchy, related to the generally solitary life-styles of felids. There is no reason to believe that such disputes are less common, or potentially less damaging, in male–male groups. However, if fighting within a nomadic multi-male group is sufficiently damaging as to injure or drive away group members, this has little or no immediate impact on the reproductive success of males in the group, although it might reduce the probability of future success in taking over a pride. In contrast, for a pride male to be injured or driven off makes the pride immediately and substantially more vulnerable, and the infanticide that typically follows a successful take-over may completely eliminate the progeny of the pride males (Packer et al., 2009). So, to go back to the question posed in the introduction to this article, is it male–male fighting, or female choice, or both, that makes the lion's mane adaptive? In favor of a view that male–male contests are made more adaptive by manes are findings that: (1)Fatal injuries from lion fights often involve the head and nape. (a)There is some evidence for targeting of these sites. (b)Lion fighting involves frontal confrontations such that damage in these areas might be expected in the absence of evolved targeting and ©These areas are particularly vulnerable, in terms of immediate mortality in response to bites and blows. (2) When manes are reduced, likely by environmental factors, multi-male, mixed-sex prides are vanishingly rare. The West and Packer (2002) findings do suggest some female choice of more abundantly maned males, in that: (1)Females approached heavily-maned dummies more than did males. (2)Also, field studies indicate that female lions do exercise a substantial degree of mate choice, although it is not known if this reflects male mane characteristics. (3)Mane quality is a sensitive index of health in lions. Putting these findings together, it seems most like that “both” is the best answer, but with the further provision that female choice appears to be based on a characteristic that is independently adaptive in terms of the social characteristics of most lions. That is, the mane “honestly advertises” the quality of a trait that is adaptive for males (see Johnstone, 1995 for a discussion of this view of sexual section). Indeed, if the mane were selected simply on the basis of female choice, albeit with the result that male offspring with high levels of this trait would similarly prove attractive to females, this would raise a legitimate issue of why only female lions, of all the many felid species, show such a choice. The relationship between multi-male mixed-sex social groups and male manes might conceivably have arisen as a coincidence. However, the “coincidence” explanation is substantially challenged by findings that mane reductions in the Gir and Tsavo lions, that appear to be the result of persistent hot or hot and humid environments known to produce overheating in maned lions, are associated with a virtual elimination of multi-male prides. Lions….and Men? Humans are fascinated with lion manes. This may reflect nothing more than a human propensity to be intrigued by unique or unusual features of the natural world. It is possible, however, that something more is involved. The lion's mane is a highly unusual, sexually dimorphic, male feature, arising about the time of puberty, found in a species that is more social than are its phylogenetic relatives, and in possession of weapons that may be lethally applied to others of its kind: The feature covers an area that is directly accessible in frontal confrontations, may be a particular target of attacks, and is highly vulnerable, as indexed by potential mortality when wounded. Morris (1968), in “The Naked Ape” raised the intriguing question of why humans have less, or shorter and thinner, hair on most body sites. Perhaps an even more intriguing question is why humans, absolutely uniquely in the animal kingdom, have two sexually dimorphic hair patches that grow indefinitely. Moreover, one of these patches, the beard, begins to grow during puberty, and is thickest in young adulthood and full maturity, tending to decline in old age. Like the lion's mane, male beards are widely assumed to be somewhat adaptive in the context of providing a visual aid to identification of gender at a distance; in advertising social dominance; or as sexually attractive to women (Barber, 1995). As evidence for the latter is inconsistent (Feinman and Gill, 1977; Hatfield and Sprecher, 1986), and the accuracy and value of other presumed adaptive functions of beards are debatable (Barber, 1995), the near-universal existence of male beards in humans, albeit variable in individual magnitude, remains something of a mystery. What about a protective function, similar to that of the lion's mane? Figure Figure11 provides a very good look at the facial and neck structures covered by beards, as well as some estimation of the degree of coverage they may confer. These areas are front and center in aggressive confrontations and may be especially vulnerable to blows when weapons are not used, or to blows from the “blunt force” weapons that have been available throughout much of hominid evolution. The “glass chin” phenomenon is well known in boxing, and direct blows to the front and sides of the neck as well as the area just under the nose can be particularly lethal. Both are particular targets of attack in unarmed combat techniques (Schillingford, 2001). This protective feature of human beards is also recognized in the Technical and Competition Rules of the international Amateur Boxing Association, which prohibit beards in boxing matches3. In addition, based on extensive observations of encounters in young children, the child psychologist Margaret Manning has suggested (personal communication) that the head/face/neck area is a specific target of attack in young children. This is particularly interesting as combat training is unlikely to be involved in childrens’ preferences, and it may provide something of a functional or adaptive basis for the nonsexually dimorphic, evergrowing hair patch that covers the dorsal, side, and back sections of the head. Notably, if left to grow, head hair also covers the back and sides of the neck, leaving the front of the neck, the most vulnerable aspect in frontal confrontations, exposed except in pubertal and postpubertal males. The many points where manes and beards show parallels also bring up the interesting question of why other mammalian species with potentially lethal weaponry fail to develop such protections. One answer, already mentioned, is that they may be much less necessary in species with solitary life-styles. Another is that a structure such as the horns or antlers of ungulates may simultaneously serve as weapon, target site, and protection against this specialized form of intraspecific attack. Indeed, such “protective” weaponry is much more common in gregarious than in solitary species of African bovids (Estes, 1992). However, many other social species have weapons that are clearly capable of causing lethal conspecific damage, but have no notable structures to protect highly vulnerable sites. One possible reason for this omission may be that animals with both dangerous weapons and a long and consistent evolutionary history of sociality, such as most canids, tend to have a clearer within-group dominance structure than do lions, including behavior mechanisms (“submission”) that reduce intraspecific attack (MacDonald and Sillero-Zubiri, 2004). Humans do certainly have both dominance hierarchies and submissive behaviors, but these appear to vary widely across cultures and their existence and extent in precultural humans is difficult to estimate. While there appears to be a linear dominance hierarchy in male chimpanzees (Goldberg and Wrangham, 1997), this appears not to be so in bonobos (Paoli et al., 2006); these having equal claim to the ancestral line from which humans split. Perhaps lions and humans came to their need for protection against lethal intraspecific attack through slightly different routes; Lions with long-term lethal weapons but lately evolving a social life-style, whereas people are from a line with substantial, though not clearly delineated, sociality, but lately evolving the development and use of lethal weapons. Both species may have encountered the combination of a social life-style and lethal weaponry too rapidly to evolve innately-organized social mechanisms that protect individuals from the damaging consequences of intra-group attack. This view implies that structural protections; here, long, coarse, and abundant hair growth in relevant sites, may be more easily or rapidly evolved than are complex behavioral changes. Such an issue may be resolvable, given current developments in molecular genetics. However, it is only one of many potential questions following from the present analysis. It seems likely that a combination of experimental and ethological approaches may be necessary in order to gain a better understanding of the complex evolutionary interplay between sociality and other behavioral and structural characteristics of particular species.
This one is by Peyton West. It's a long, but good read. There are some images in the pdf, but I can't post those. You can see them anyway in the link.
The Lion’s Mane Neither a token of royalty nor a shield for fighting, the mane is a signal of quality to mates and rivals, but one that comes with consequences The African lion is one of the world’s most admired and best studied species, yet its most striking feature has long been a mystery: Why do lions have manes? Charles Darwin, who knew almost nothing about lions, was one of the first to suggest an answer, writing, “The mane of the lion forms a good defence [sic] against the one danger to which he is liable, namely the attacks of rival lions.” This unsupported hypothesis prevailed until 1972, when George Schaller published his seminal work, The Serengeti Lion. Schaller suggested that males bore sumptuous manes to signal their quality as prospective mate, similar to the displays of several other polygamous species. Although these two hypotheses were not mutually exclusive, scientists tended to favor one or the other. When I began my research in 1995 neither theory had been systematically tested.
Craig Packer introduced the question to me in a casual conversation about potential thesis projects, months before I started graduate school at the University of Minnesota. “There are really two big mysteries left about the big cats,” he said. “Why did saber tooth tigers have saber teeth and why do lions have manes?” I remember thinking that there wasn’t much I could do about saber tooth tigers, but the lion’s mane—I was hooked. The possibility of answering such a basic question was exactly the reason I got into science in the first place. I soon joined Craig’s lab despite his warning: “it’s not an easy project….” Three basic features guide any thinking about the lion’s mane. First, the mane is sexually dimorphic (only males have manes); second, the mane begins development at puberty; and third, the mane is highly variable both within and between populations. Manes vary in color from almost white to deep black and in overall size from the slightest “Mohawk” and side-whiskers to a long, thick coat that covers the shoulders and chest. Furthermore, individual manes are not uniformly sized or colored but are often a patchwork of lengths and hues. These features are consistent with the idea that the mane is a product of sexual selection. Most sex-selective traits are sexually dimorphic, begin development at puberty and are highly variable. According to the theory of sexual selection, such characteristics evolve under the stress of competition for mates. Sexually selected traits can increase reproductive success in two ways. The first, known as male-male competition, increases the ability of males to compete against other males for females. Traits in this category include armor to protect males from opponents, weapons to disable opponents or signals of fighting prowess that males use to assess opponents. Generally, males with more exaggerated features are better competitors. The second role of sexually selected traits, mate choice, increases male attractiveness to females. Traits of this sort, such as bright coloration, long feathers or elaborate calls, usually relate to the male’s condition. Females that prefer more “ornamented” males may obtain benefits directly, in the form of more offspring, or indirectly, through better genes for their offspring. One of our objectives was to determine whether the mane functioned in male-male competition, mate choice or both. Serengeti Story When Craig said studying the mane would be challenging, he knew what he was talking about. He has studied lions in the Serengeti National Park in Tanzania for almost 30 years and endured all sorts of grueling ordeals in the name of scientific exploration. His hard work made my job easier though, because thanks to his efforts and those of other scientists, there is a vast database on the Serengeti lions. Not only has this work answered most questions about lion behavior, but demographic and physiological data let us study the heritability of traits and other questions that are difficult to answer for wild populations. Studying sexual selection in the field, in a long-lived species like the lion, would have been impossible without this prior research. To start with, knowledge of lions’ social structure allowed us to refine our hypotheses about sexual selection. Female lions live in prides consisting of related females and their dependent offspring. As the cubs grow, young females typically join their mothers’ pride, and young males form “coalitions” and disperse to look for their own pride. This creates a system in which a small group of males can monopolize many females, leading to severe reproductive competition. Predictably, males compete intensely for mates, and they compete on two levels. At the group level, male coalitions vary in size, and larger coalitions sire more offspring than small coalitions. Individuals within a coalition also compete: If a male discovers an estrous female, he will jealously guard her and prevent her from mating with his companions. As Craig and his colleagues discovered, this behavior skews the paternity rates for individuals in larger male coalitions. In contrast, female lions are egalitarian. Unlike some social carnivores such as wolves and hyenas, all of the adult females in a pride reproduce, and female lions don’t have a dominance hierarchy, which often dictates reproductive success in other species. Furthermore, a key attribute of lion society is that females breed synchronously, which means that there are often more estrous females available at one time than there are resident males. Males cannot usually defend more than one female at a time, but they willingly mate with additional females if possible. Thus, if estrous females outnumber males, the “excess” females—those that aren’t actively guarded—are free to choose among coalition males. The bottom line is that this social system provides opportunities for sexual selection based on male-male competition and mate choice. This combination is not entirely surprising. Although historical studies of sexual selection focused on one or the other hypothesis, more recent work demonstrates that the two mechanisms often operate together. Twenty years of field observations also helped answer our next question: What kind of trait would be most useful to lions? With lethal claws and teeth, fighting is very costly, even for the victor. For this reason, just as Darwin suggested, males might benefit from a shield to protect them during fights. However, avoiding the fight altogether would be a greater advantage; thus, males would benefit from a signal that conveys a rival’s fighting ability. From the females’ perspective things are slightly more complicated. Unlike many mammals, male lions play an important role in raising offspring, but they are also utterly intent on their own reproductive fitness. When a new coalition of males joins a pride, they immediately kill or evict the offspring of the previous males. This behavior accounts for more than 25 percent of cub deaths and is a major variable in female reproductive success. In the short term, a group of females can fend off infanticidal newcomers, but the pride’s resident males bear most of the responsibility for protecting young lions. The displacement or loss of a male coalition generally leads to 100 percent mortality of any unweaned cubs. Females would thus benefit from a signal that advertised a male’s ability to fight off would-be usurpers. Males also help feed the pride. Although male lions are often depicted as parasites, lying around while females do all the work, males are extremely capable hunters of a key prey species: the Cape buffalo. Buffalo are large and slow, and hunting them depends less on the speed and agility evinced by females and more on the weight and strength characteristic of males. A buffalo will satiate a big pride, and this species is the most important prey throughout much of the lion’s range. For females, such contributions are critical because starvation is another common cause of death among cubs. Any trait that advertises male hunting ability or contains general information about a male’s nutritional status would be valuable. This knowledge allowed us to refine our thinking before beginning our fieldwork. We hypothesized that the mane might function in any of three ways: as a shield against injury, as a signal of the male’s ability to fight and protect his cubs (essentially the same thing), or as a sign of the male’s nutritional status. The physiological attributes of hair support the idea that it could convey such signals. Hair growth depends on a variety of factors including, among other things, hormones, health and nutrition. In sexual selection terms, hair is “condition -dependent,”meaning that its appearance is often related to the underlying condition of the animal. More specifically, hair growth and pigmentation are influenced by testosterone, which in turn is related to aggression and might be an indicator of fighting ability. Additionally, malnourished and sick mammals often develop rough, unhealthy looking hair, and poor nutrition, such as copper and zinc deficiencies, can inhibit hair growth and pigmentation. The Meaning of the Mane Our first goal was to address the maneas-a-shield hypothesis, which makes two simple predictions: that the mane is an effective barrier against the teeth of rival lions, and that males with longer or darker manes are injured less frequently or less severely. Unfortunately, these predictions are almost impossible to verify. Fights between lions are rarely witnessed and individuals are seldom seen regularly enough to assess the frequency with which they are wounded. Instead, we generated two related hypotheses that were testable. First, we predicted that if the mane’s primary function was protection, the “mane area,” or the areas of the body covered by mane hair, would be a special target during fights and that most lion-inflicted wounds would be found there. Second, we predicted that wounds to the mane area would be more serious and more likely to be fatal. We addressed our hypotheses by combing the records for injuries and eliminating those that were not inflicted by other lions. The resulting database consisted of the locations and survival rates for wounds to males, females and subadult lions. These data did not support the mane-as-shield hypothesis. Wounds to the mane area were no more frequent or lethal than those to other parts of the body. The observations were true not only for adult males but also for females and subadults, indicating that the manes of adult males did not introduce bias by obscuring wounds from view. It seems that a lion’s teeth provide more than enough incentive to avoid tangling with the front end. Finding little evidence to support the mane-as-protection hypothesis, we turned to the idea that the mane functions primarily as a signal, asking specifically: What ecological trends predict mane length and darkness? A critical first step was to quantify objectively the length and darkness of a lion’s mane. For this task we turned to our photographic archives, which included pictures of virtually every male lion to appear in our study area since the project began in 1966. While in the field, we continued to photograph males every six months to document new animals and record any changes in their manes. We then recruited undergraduate students, who were informed of the general nature of our work but knew nothing about the individual animals, to “grade” the pictures for length and darkness. At least five students graded each picture; we then eliminated the low and high scores and averaged the remainder. These measurements became the backbone of our research. We first used these data to address several long-standing questions about lion manes. We ascertained, for example, that manes in the Serengeti generally begin developing at just under one year and continue growing until males reach 4.5 years of age. The mane gains pigment rapidly during this time, until the color becomes more stable about a year after growth ends. It continues to darken at a slower rate throughout life. We also demonstrated that the agerelated increases in length and color mirrored the increase in testosterone during adolescence. A welcome surprise was that the manes of individual males were not always constant over time; although the pattern of sharp gains in length and color followed by slow darkening was typical, the manes of some lions became llighter or shorter, or changed back and forth. These results were inspiring because they gave further credence to the idea that the mane is condition-dependent—this kind of variation in sexually selected traits often reflects changes in the underlying condition of the animal. More generally, our analysis revealed that mane length and darkness are correlated with several ecological factors. In males older than five years, mane length was most closely associated with injury. Prior to starting the analysis, we knew anecdotally that the manes of injured males were often reduced and could fall out altogether, and we now discovered that injured males also had more subtle reductions in mane length. This fact is significant because it suggests that mane length might signal a male’s current fighting ability—injured males should be less able or less aggressive fighters. Mane color proved more interesting still: In addition to the age effect, we found that males with darker manes had higher levels of testosterone, suggesting greater aggression, and were on average better fed throughout the year, suggesting either general dominance or superior hunting ability. These results implied that both length and color provided interesting information for other lions, and that both males and females would benefit from using it. Shorthand for Quality Signaling theory predicts that if information is available animals will take advantage of it, but demonstrating the truth of this principle can prove challenging. Our next step was to look for evidence in our long-term records that males and females were actually using the information contained in the manes. For males, this proof was particularly difficult to get. We knew from previous research that dominance relations in male coalitions do occur—in coalitions consisting of three or more males, generally only two males fathered all of the offspring—but we were unable, because of incomplete photographic data, to link these relations to mane characteristics. Instead, we turned to an experimental protocol used by previous lion biologists: playbacks. This technique uses recordings of natural animal sounds to mimic situations that we would otherwise seldom witness. We broadcast the roars of single, unfamiliar females to coalition males in order to evoke male competition for access to an estrous female. Because the first male to reach an estrous female generally ends up guarding and mating with her, we reasoned that whichever male led the group would most likely be dominant. If mane color or length were indications of dominance, we predicted that the male with the darker or longer mane would be the first to the loudspeaker. Experiments with 13 resident male coalitions uncovered a revealing and surprising result. Mane length had little relation to “dominance,” as measured by our tests, but males with darker manes were significantly more likely to win the race to the female, suggesting that mane darkness does predict male dominance. We next turned to females, and here our long-term data proved more useful. Because “excess” estrous females choose their own mate, we looked among the records for situations in which a male mated with more than one female in the course of an hour, assuming that at least one of the females was there by choice and was therefore exercising a preference. We found 14 examples of this situation for which we also had good data on the mane characteristics of all the males in the coalition. Again the results contained a surprise. Like coalition males, females appeared to place little value on the length of the mane—in only seven of these examples did the male in question have the longest mane of his coalition. However, color was again a critical factor. In 13 of the 14 observations, the females mated with the male whose mane was darkest. The consistent results indicated that mane darkness played a role in sexual selection, but they left some nagging questions. Why didn’t lions pay attention to mane length when it could betray recent injury? And how did lions respond to strangers, whose arrival had such potentially disastrous consequences? Observational data were inadequate because such meetings are rare, occur mostly at night and are impossible to predict. Thus, we again turned to active experiments, presenting to the real lions “dummies”—two plush, life-sized toy lions that differed only in their mane—to see if mane characteristics influenced the lions’ behavior. Fool Me Once… We were optimistic about this approach. Ecologists commonly use dummies to study sexual selection in other species, and another graduate student from the Packer lab had successfully used a lion mount (prepared by a taxidermist) in earlier experiments. However, getting the actual dummies was a problem. We couldn’t find a source for large, realistic stuffed lions, and custom-made toys were prohibitively expensive. Then, in a stroke of serendipity, Craig was contacted by a documentary filmmaker, Brian Leith, who wanted to make a film about lions. Brian was captivated by the experiments we were planning, and in no time at all he discovered Anna Club Plush, a Dutch company that was willing to donate stuffed toy lions made to our specifications. Within a few months, four beautifully plush, life-sized male lions arrived in the Serengeti. We christened them Romeo (short dark mane), Lothario (short blonde mane), Julio (long dark mane) and Fabio (long blonde mane). In each experiment we presented a choice between two dummies to single-sex groups of adult lions. Lothario and Fabio helped us test the importance of mane length, and we used Julio and Fabio to test the effect of mane darkness. The manes were attached with Velcro, a feature that allowed us to switch manes and control for any differences between individual dummies. Once we found a group of lions, we waited until dusk (when lions are more active), set up the decoys downwind (to remove any effect of scent) and broadcast recordings of hyenas at a kill. This cue evoked a speedy response from the lions, who gathered to scavenge a meal. As they approached, the lions quickly noticed the two “strangers” and their attention shifted to the dummies. At that point, we turned off the sound and watched. The early experiments were nail-biting affairs. Would the protocol work? Could the dummies fool real lions? Happily, it was clear from the start that we were getting good data. After a quick start toward the loudspeaker, the real lions became much more cautious when they caught sight of the plush ones, stopping for a careful look every few feet before proceeding. Upon reaching the dummy, their first act was often to sniff under its tail. In order to eliminate the effect of scent on their behavior, we noted on which side the lions approached the dummies because they usually made that decision at a distance of 100 meters or more. Thus, a female approach on the side of the dark-maned dummy counted as a preference for the darker mane. Although the initial experiments were promising, it took three years to get enough data to draw meaningful conclusions: To our great surprise, the lions soon became habituated to the dummies. Lions that had seen them before—even years before—were never really fooled again. Their behavior was much less cautious and they often failed to approach at all. Because lions live in fission-fusion groups, the project became much, much more complicated. Even if three out of four lionesses were dummy “virgins,” we couldn’t test the group—all four had to be naive. Furthermore, because the tests of males required resident coalitions (nomadic males would flee rather than approach the strange males), we had to expand beyond our study area to get an adequate sample size. But our reward for all this work was a fascinating set of data. Similar to the long-term mating records, females in this test strongly preferred dark manes, approaching on the side of the black-maned dummy nine out of ten times. They approached the longer-maned dummy in only seven of ten trials (a non-significant result). Similarly, males were sensitive to mane darkness, avoiding the darker-maned dummy in five out of five trials. However, unlike the playback study, the tests with Lothario and Fabio showed that males were extremely sensitive to mane length; they avoided the longhaired dummy in favor of the shorthaired one in nine out of ten tests. The different results arose from the different contexts for the two experiments: Whereas the earlier study tested dominance within a coalition, the plush lions simulated interactions between unfamiliar males. Because mane length can indicate short-term quality in the form of recent fighting success, this signal may be more relevant when deciding to challenge an unfamiliar opponent than when dealing with a well-known, long-term confederate. We had established that the mane acts as a signal to other lions, but what were the actual benefits of having or preferring dark or long manes? A further look at our long-term records filled in these gaps. Although mane length had no detectable link with overall fitness, mane darkness was a significant factor. Males with darker manes spent more of their lives residing with a pride and were more likely to survive when wounded. Furthermore, their offspring were more likely to reach their second birthday (which also benefits females that chose dark-maned males) and less likely to be wounded, suggesting that darker maned males provide better protection from other lions (the most common cause of injury). Degree of Cost The analysis showed several benefits of having a dark mane. So why don’t all males have them? In other words, what prevents “dishonesty” among males who are wooing prospective mates? This is a common question in the study of sexual selection. Scientists generally answer that the production or maintenance of such a phenotype must be so costly that only superior males can afford it. So what is the cost of a black mane? Heat. Since the early 20th century, naturalists such as Frederick Selous noted that lions in different regions had different mane characteristics, and they even linked this variety to temperature. Males in colder, higher-altitude habitats tend to have bigger manes than those in hot, humid climates, and all lions are extremely sensitive to heat. Larger animals have more difficulty with high temperatures because of their higher ratio of volume to surface area, and many lion behaviors seek to minimize heat stress. Sleeping in the day and limiting most activity to the night is one example; others include lying on their backs to expose their thinly skinned bellies, resting on high rocky knobs to catch the breeze, and panting after exertion or large meals. Unlike dogs, lions do not have cool wet noses, and unlike people, they don’t sweat. Their only means of thermoregulation are breathing (panting) and radiating heat from the skin. In this context, the mane is a handicap because it prevents the efficient dissipation of heat. Furthermore, dark hairs are thicker than light hairs, creating a better insulator, and dark surfaces absorb more solar energy than light ones. These facts suggest that heat stress might be the most significant cost associated with the mane, and that lions with long, dark manes would be most affected. Testing these hypotheses was challenging—we couldn’t exactly use a thermometer on wild lions. However, the technology of infrared thermography, which can measure the precise surface temperature of a distant object, held some promise. The latest generation of such cameras was sufficiently portable to bring to the field, and a manufacturer, Flir Systems Incorporated, agreed to let us rent one of the pricey devices at a discount. We spent three months snapping infrared pictures of all the adult males in the area as well as many females and subadults. Our first step was to compare the average surface temperatures of the sexes, reasoning that if the mane had a thermoregulatory cost, males would be hotter than females. This turned out to be true, but there was a problem: Males are 50 percent larger than females and might be hotter just because of a greater ratio of volume to surface area. Unable to solve this conundrum in the Serengeti, we traveled to Tsavo National Park in Kenya, where male lions were reportedly “maneless.” Tailed by a crew from National Geographic, we took a month-long detour in search of males that lacked manes but retained the size advantage over females. If Tsavo males and females showed different temperatures, then we could infer that lion temperature was a function of size. But if the sexes had similar surface temperatures in Tsavo, then the difference in temperature seen in Serengeti lions must be a result of their manes. We discovered that many of Tsavo’s adult males did have extremely small manes, although not all were maneless. In support of our heat-stress hypothesis, Tsavo was noticeably hotter and more humid than the Serengeti, and the lions there appeared to be more challenged by the climate. For example, we observed the novel behavior of scraping away the topsoil before lying down, presumably to find cooler earth underneath. Yet aside from their manes and their heat-related adaptations, Tsavo’s lions were virtually identical to Serengeti lions in appearance and behavior, and the sizes of prides and coalitions were the same. While in Tsavo, we took the opportunity to perform several dummy tests and found that these males behaved in exactly the same way as Serengeti lions, appearing intimidated by longer and darker manes. And despite the challenges of unfamiliar surroundings, thick obscuring brush and the lack of radio collars, we took many thermal images. What we found was that Tsavo males, despite being bigger than Tsavo females, were no hotter. Thus, we concluded that the temperature difference between males and females in the Serengeti derived from the heat cost of the mane rather than that of a larger body. Back in the Serengeti, we next compared males’ body temperatures to their mane scores to find precisely which mane characteristics were influential. Mane length had little effect, but males with darker manes were significantly hotter than those with lighter manes, even after controlling for ambient temperature, wind, humidity and prior activity. This connection supported our prediction that males with darker manes paid a higher price in terms of heat stress. In addition, we confirmed that males with darker manes had higher proportions of abnormal sperm (the link between testicle temperature and sperm production is well known). We also found that unlike light-maned males, dark-maned males reduced their food intake in hotter weather. Lions with big bellies (from eating big meals) tended to pant more and had higher surface temperatures, suggesting that gorging also causes heat stress. Males with dark manes are already more compromised by the heat, and thus must eat smaller meals when temperatures rise. Once we had linked individual mane phenotypes to heat stress, we revisited some of our long-term data to see if similar effects existed at the population level. We wondered whether mane characteristics varied with small-scale changes in habitat or with seasonal differences in ambient temperature. The answer was yes to both questions. Males born into prides in the Serengeti woodlands, the warmest habitat in our study area, had shorter manes throughout life regardless of the climate they lived in as adults. Similarly, males that reached adolescence in warmerthan-average years maintained shorter manes over the course of their lives regardless of their place of birth. Finally, males from the Ngorongoro Crater, the coolest part of our study area, had significantly darker manes. These results emphasize the importance of heat for determining mane traits and for the species in general: Such sensitivity to the negative consequences of hyperthermia argues that lions may be living at the edge of their tolerance for heat. This possibility raises an interesting conservation issue. Global warming is real, and as research continues to uncover the negative effects of climate change on wildlife, an exquisite sensitivity to heat takes on an importance that is more than academic. Broadly speaking, we predict that the continued rise in average temperatures in East Africa will lead to fewer lions with long, dark manes. This shift may negatively affect industries such as tourism or legal sport hunting. More important is the result of such a change on the lions. In the evolutionary past, climate change may have driven to extinction species with sexually selected traits they could no longer afford. Although the mane’s phenotypic plasticity makes this scenario unlikely in lions, certain behavioral or physiological adaptations could become necessary. Such changes are impossible to predict, but any effects on a keystone predator like the lion have the potential to influence whole ecosystems. Our research emphasizes the potential consequences of climate change and argues for the importance of behavioral studies to detect its more subtle effects. 
@Tshokwane How is it that those studies don't support the idea that a lion's mane offers protection, when we have a number of videos showing the opposite, like this one? One of these lions at Amboseli National Park (near Mount Kilimanjaro in Kenya) tried really hard to kill the other lion with a bite to the neck, but it didn't work, and this has got to be one of the strongest attempts by any big cat to try killing a lion by biting its mane that I've seen!
Not to mention that there is at least one expert who does support the notion that the lion's mane does offer protection, like Dereck Joubert?  (03-20-2020, 10:41 PM)BorneanTiger Wrote: @Tshokwane How is it that those studies don't support the idea that a lion's mane offers protection, when we have a number of videos showing the opposite, like this one? One of these lions at Amboseli National Park (near Mount Kilimanjaro in Kenya) tried really hard to kill the other lion with a bite to the neck, but it didn't work, and this has got to be one of the strongest attempts by any big cat to try killing a lion by biting its mane that I've seen! You give me the opportunity because you quote "Tshokwane" that our colleague Ezequiel in this forum cannot answer you....He passed away on june 14 2019 from cancer.....I didnt saw any info or anouncement from the forum.....Forgive me if i didnt saw it RIP "Tshokwane"  (03-20-2020, 11:14 PM)Mohawk4 Wrote:What ? I'm just learning what you say, I didn't know that... I liked to swap with him very much, he was a perfect moderator. Very, very sad to learn that. RIP "Tshokwane" and thoughts to his family.(03-20-2020, 10:41 PM)BorneanTiger Wrote: @Tshokwane How is it that those studies don't support the idea that a lion's mane offers protection, when we have a number of videos showing the opposite, like this one? One of these lions at Amboseli National Park (near Mount Kilimanjaro in Kenya) tried really hard to kill the other lion with a bite to the neck, but it didn't work, and this has got to be one of the strongest attempts by any big cat to try killing a lion by biting its mane that I've seen! (03-20-2020, 11:14 PM)Mohawk4 Wrote:(03-20-2020, 10:41 PM)BorneanTiger Wrote: @Tshokwane How is it that those studies don't support the idea that a lion's mane offers protection, when we have a number of videos showing the opposite, like this one? One of these lions at Amboseli National Park (near Mount Kilimanjaro in Kenya) tried really hard to kill the other lion with a bite to the neck, but it didn't work, and this has got to be one of the strongest attempts by any big cat to try killing a lion by biting its mane that I've seen! That was sad news, I and some others knew about his situation but I think, that no-one here was sure if he was too ill to participate or if that, what you now told, had happened. Truly sad news, he was a nice person and a good poster here. I didn´t know him too good, but that is how I remember him. I´m happy that I had the opportunity to change a few words with him, while he was still among us. If someone close to him reads this, I simply want to express my respect for his memory and my sincere condolences, a good man passed away too soon.
Tshokwane loved and followed Majingilane coalition for years....In a group of this coalition one of his colleagues made the anouncement....
 *This image is copyright of its original author
 
RIP. I had a few interactions with him on the forum. He was very logical in the way he analyzed lion behavior and I agreed with him on a lot of his posts. Sad news. How old was he?
ALL
Tshokwane was a giant lion hunted by many. A fitting name for our lion mod, I thought. He told us about his situation about a year ago. After his last post in the mod thread, we were unable to contact him. The news (thanks Mohawk) is devastating. We'll start an Im Memoriam thread soon. (03-20-2020, 11:14 PM)Mohawk4 Wrote:(03-20-2020, 10:41 PM)BorneanTiger Wrote: @Tshokwane How is it that those studies don't support the idea that a lion's mane offers protection, when we have a number of videos showing the opposite, like this one? One of these lions at Amboseli National Park (near Mount Kilimanjaro in Kenya) tried really hard to kill the other lion with a bite to the neck, but it didn't work, and this has got to be one of the strongest attempts by any big cat to try killing a lion by biting its mane that I've seen! Last time he was on was June 12. I talked to him a lot on Google+ years back about these big cats, and he was a really cool guy also smart. This is him on Twitter. RIP.. https://twitter.com/majingilane  (03-20-2020, 11:14 PM)Mohawk4 Wrote:(03-20-2020, 10:41 PM)BorneanTiger Wrote: @Tshokwane How is it that those studies don't support the idea that a lion's mane offers protection, when we have a number of videos showing the opposite, like this one? One of these lions at Amboseli National Park (near Mount Kilimanjaro in Kenya) tried really hard to kill the other lion with a bite to the neck, but it didn't work, and this has got to be one of the strongest attempts by any big cat to try killing a lion by biting its mane that I've seen! I just realised that from here. This is shocking. |
| Users browsing this thread: |
| 1 Guest(s) |