How does the social structure of lion prides prevent incest?

How does the social structure of lion prides prevent incest?

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I assume that lions don't have a concept of incest and they will just mate with any adult lioness in heat, daughter or not.

If so, is incest inevitable as the female cubs will grow up and remain in the pride? Their brothers will be kicked out as they reach mating age. But the females stay, so what is the mechanism that will prevent their fathers from mating with them?

I suppose that their fathers will be driven out of the pride by incoming males sooner or later. But if it doesn't happen soon enough, then what prevents them from mating with their daughters?

It is true that African lions have recently become more prone to inbreeding, largely due to human actions (reasons include limited dispersal and at least temporarily reduced pride sizes). see here

Generally though, lions likely avoid mating with close relatives innately as most animal species do. see e.g. here

A few things about the social structures of lions (ref.):

  • First, many prides are not controlled by a single male. Although the males might often be related to one another.

  • A single male might often not be in control for long enough to mate with his grown daughters.

  • Females do not only mate with the leader of their pride, but mate sometimes with males from outside of the pride.

How does the social structure of lion prides prevent incest? - Biology

Understanding social network structure and position can aid wildlife conservation.

Threatened wildlife populations offer a vital experimental platform for animal SNA.

Linking animal SNA to practice stimulates design of new practical tools and theory.

Many animals preferentially associate with certain other individuals. This social structuring can influence how populations respond to changes to their environment, thus making network analysis a promising technique for understanding, predicting, and potentially manipulating population dynamics. Various network statistics can correlate with individual fitness components and key population-level processes, yet the logical role and formal application of animal social network theory for conservation and management have not been well articulated. We outline how understanding of direct and indirect relationships between animals can be profitably applied by wildlife managers and conservationists. By doing so, we aim to stimulate the development and implementation of practical tools for wildlife conservation and management and to inspire novel behavioral research in this field.

What silly thing has someone tricked you into doing or believing?

My older brother, when I was little, convinced me that this small plant was a tiny tree, and that the tiny things on it were miniature red apples. It was a hot pepper plant I quickly discovered.


my dad told me that if you drop bullets on the floor they go off so one time we were at acadmey and he drop bullets and i started running and mt family asked me why i was running and i told them and they started laughing

Growing up I had an older cousin who was like a sister to me. I believed everything she told me and she abused that knowledge. The most outrageous thing I believed was the following. She convinced be that a large saw used for yard work hanging in the garage actually was used for decapitating people who trespassed. The reason the teeth on it were so large was because the neck bones were hard to saw and the rust was actually dried blood. I was gullible and terrified.

When I was a LOT younger, my family would take trips to se Grandma, about 60 miles north of where we lived.

Grandma lived on an in-town acreage, just down from Reservoir Hill. My cousins and I always used to make the climb up there whenever we visited. We were always told, "look out for the bull!" (My grandma's neighbor did have cows, and sometimes they were out, which made the bull a real concern.) So, with one keeping lookout, up the hill we went.

After a couple of excursions, we got really good at avoiding the cowpies on our way up the hill.

It was good to sit up on the reservoir on a Summer's day, and feel the breeze. It was a calming effect.

Down the hill we went after a while of being up there. Once again, one of us kept lookout. Years passed, and we all got older. Before my grandma passed away at 97, I found out that the only bull that ever existed was in the minds of the people telling the tale.

I get a little chuckle time to time, when I think of it. For years, my grandparents, aunts and uncles really put one over on us kids.


Having developed the necessary model components, performance of the control rule in maintaining a viable lion population under trophy hunting was evaluated through simulation. The MSE process was iterated forward in time, assuming a heavily depleted initial population, and including uncertainties in demography, observation, and compliance to the minimum age rule. At each iteration of the MSE, waiting time data were sampled from a negative binomial distribution, with a mean obtained from the density of male lions predicted by the operating model and a fixed dispersion. Performance was measured by tracking the realized harvest rate and number of males over a 30-y period. Because some noncompliance to the minimum age rule was incorporated in stochastic projections, performance was measured by recording the harvest rate and population size for the total adult male population (i.e., ≥4 y), for which the sustainable harvest rate is ∼0.25. In addition we recorded the quota and success rate (proportion of quota filled) alongside other hunting and sustainability metrics (Table S3).

During the projection period the adult male population size increased at a rate of two per year from around 38 to 100 individuals while simultaneously increasing the quota from around 15 to 22 (Figs. 2 and 3). The hunting success rate increased from close to zero to 76%, with a simultaneous decrease in the waiting time from 30 to 23 d. Thus, despite stochasticity, the control rule is able to converge on a sustainable quota, which remains relatively stable throughout the projection period.

Dynamic system response over time. Median changes in the number of adult males (≥4 y of age) and the quota are shown following initialization of the system. Shaded regions represent 95% CIs across stochastic iterations, taking into account uncertainty in the dynamics, observation, and compliance to the minimum age criterion.

The evolution of matrilineal social systems in fissiped carnivores

We review matrilineal relationships in the societies of fissiped mammalian carnivores, focusing on how the most complex of these may have evolved from simpler systems. Although competition for food is very intense at the trophic level occupied by most carnivores, and although most species of extant fissiped carnivores therefore lead solitary lives, some species show at least rudimentary clustering of maternal kin and matrilineal resource-sharing or transmission of critical resources between generations. The resources shared or transmitted range from individual food items and territories to entire networks of potential allies. The greatest elaboration of matrilineal relationships has occurred in two large carnivores, lions and spotted hyenas, which occur sympatrically throughout much of Africa. The societies of both these species apparently evolved in response to a shared suite of ecological conditions. The highly matrilineal societies of spotted hyenas are unique among carnivores and closely resemble the societies of many cercopithecine primates. The conditions favouring the evolution of matrilineal societies in carnivores include male-biased dispersal, female philopatry, the need for assistance in protecting or provisioning offspring, reliance on large or abundant prey, particularly in open habitat, high population density and kin-structured cooperative interactions that have strong positive effects on fitness.

This article is part of the theme issue ‘The evolution of female-biased kinship in humans and other mammals’.

1. Introduction

Only 10–15% of all species in the mammalian order Carnivora aggregate during some period outside of the breeding season [1]. Because competition for food is very intense at the high trophic levels occupied by most carnivores, the vast majority of extant mammalian carnivores lead solitary lives. Mammalian carnivores represent an extreme case in which the negative effects of resource competition might be expected to outweigh the positive effects of living and cooperating with close kin. In light of the negative effects of intensive resource competition within this order of mammals, it may seem surprising that any matrilineal societies at all occur within the Carnivora, so it is worth exploring how such societies might have evolved. Our review reveals that some tolerance of proximity of close female kin and temporary group formation by close female kin occur even among various solitary carnivores. In these solitary species, we also find some of the rudimentary building blocks of more complex matrilineal societies, such as resource-sharing among close female kin or transmission of critical resources from one generation of females to the next. Furthermore, some species in multiple carnivore families are somewhat to highly gregarious, and several of these species show clustering of maternal kin in space, as well as matrilineal resource-sharing. Elaborate matrilineal societies characterized by a high frequency of nepotistic interactions have evolved in two highly gregarious carnivore species, the lion (Panthera leo) and the spotted hyena (Crocuta crocuta). In both these species, the positive effects on fitness of nepotistic and cooperative behaviour among female kin have been well-documented. Therefore, even in the extreme case of mammalian carnivores, which compete so intensively for food and spatial resources, the benefits of living and cooperating with female kin can favour the evolution of matrilineal societies.

Here we review the literature on matrilineal social systems in fissiped carnivores. ‘Fissiped’ literally means carnivores having toes separated to the base these include all the extant mammalian carnivores except seals, sea lions and the walruses, all of which have toes joined by webbing to transform their feet into paddles (the ‘pinnipeds’). Although most fissiped carnivores are terrestrial, this group also includes otters, which are members of the same family (Mustelidae) that contains badgers and weasels. Various fissiped carnivores show at least rudimentary clustering of maternal kin and matrilineal resource-sharing or transmission of critical resources between generations. We first summarize long-standing ideas about how complex matrilineal social systems most likely evolved from simpler ones in carnivores. We then review the fissiped carnivore species in which rudimentary or complex forms of matrilineal social systems occur, as well as the specific traits and socio-ecological conditions that tend to favour the evolution of matrilineal social systems. We emphasize that many gaps currently exist in our knowledge about the social interactions of many carnivores, especially solitary, forest-dwelling species in the carnivore families Viverridae, Nandiniidae, Mephiditae and Ailuridae. Finally, we conclude by reviewing the ways in which nepotistic and cooperative behaviour among female kin can enhance fitness. In this final section, we focus largely on long-term studies of lions and spotted hyenas in Africa.

2. The evolution of matrilineal societies in carnivores

John Eisenberg [2,3] reviewed evidence suggesting that early carnivores existed in the Eocene at low densities and that they competed for both food and space. Each individual male and female most likely occupied its own home range and defended that space against incursions by all conspecifics. Each female reared her young alone, and the family split up during or shortly after weaning, as the maturing littermates began to compete for food with the mother and with one another. Moreover, within-litter aggression probably appeared early in the evolutionary history of carnivores and promoted the dispersal of family members [2]. An elaboration of this strictly solitary and territorial condition may have occurred when the home ranges of one or more females began to be encompassed by the home range of a single male. Under these conditions, an adult male could impregnate more than one female by excluding other males from his home range. In such solitary species, affiliative behaviour would only be seen during the mating season and during the offspring rearing phase affiliative interactions would otherwise occur infrequently, if at all [4].

Eisenberg [2] argued that various forms of complex sociality in carnivores and other mammals can evolve from one of three types of temporary social units that form during specific phases of the reproductive cycle in solitary mammals. First, male and female conspecifics unite to mate when the female is in oestrus. When selection favours this male–female relationship to endure past fertilization, bonds evolve between mates, and this in turn may lead to long-term monogamous partnerships and shared responsibilities during the rearing of young. The parental family structure is characterized in its extreme form, particularly when dispersal opportunities are limited for subadult offspring, by a bonded pair of adults and their immature descendants. Such protracted monogamous relationships are common among extant canids and social mongooses, many of which engage in cooperative breeding [5–7].

Second, after the birth of a litter to a solitary female carnivore, the littermate offspring form a temporary social unit that persists until they disperse away from their mother. However, selection may act on this social unit to favour littermates who associate closely with one another even after they leave their mother to accomplish goals any solitary individual could not achieve on its own. Examples of this include sibling coalitions that may last for many years among male cheetahs (Acinonyx jubatus), male banded mongooses and male lions, which would be unable to acquire or defend important resources on their own such key resources might include a territory or a group of females [8–11].

Finally, in solitary mammals, the relationship between a mother and the members of her litter endures only until that litter is weaned, after which offspring disperse and the mother resumes her solitary existence. However, selection may act to favour mothers and their offspring who remain together after weaning. This is particularly true for mothers and their daughters, because mammalian dispersal is often strongly male-biased. Mothers, daughters and sisters may then come to tolerate one another and maintain their close associations long after weaning, and in fact throughout their lives, to cooperatively acquire or defend key resources. Eisenberg and colleagues [2,4,5] referred to the temporary social unit composed of a mother and her dependent offspring as a ‘mother–family’, and it is from this unit that the most elaborate matrilineal social systems have evolved in mammalian carnivores, as they have in other orders of mammals, including primates [4]. The fact that carnivores and primates last shared a common ancestor roughly 90 Ma [12] suggests that selection on the mother–family to persist beyond weaning has a long history in mammals when ecological conditions favour its persistence.

3. The occurrence of matrilineal social systems in fissiped carnivores

The families of fissiped carnivores in which some form of matrilineal social systems occurs include Ursidae, Mustelidae, Procyonidae, Canidae, Felidae, Herpestidae, Eupleridae and Hyaenidae (table 1).

Table 1. Summary of the literature on matrilinear characteristics in carnivores.

4. Ursidae

As with other carnivore families, the common ancestor of modern ursids was almost certainly solitary [70,71]. Extant ursids have remained solitary except during the mating season, and when females rear dependent cubs [71]. Despite the largely solitary habits of bears on their individual home ranges, kin-based spatial structures have been found among females in American black bears (Ursus americanus), Asiatic black bears (Ursus thibetanus), grizzly bears (Ursus arctos) and polar bears (Ursus maritimus). Most of these species show a matriline-based spatial distribution in which daughters establish ranges near or overlapping those of their mothers, resulting in spatial clustering of female kin [13–15,72]. Dispersal is male-biased. For example, on average, mother–son grizzly bear pairs are found three times farther apart in space than are mother–daughter pairs [72]. Similarly, genetic relatedness within adult female dyads of polar bears declines significantly with increasing geographical distance [16]). Female polar bears show den area fidelity and a stronger kin structure than males [16,73]. Black bear mothers allow their daughters to occupy portions of their territory, or share a territory with them [13].

5. Mustelidae

Most mustelids are solitary [1] except during mating and when females rear dependent young. Diet and resource distribution play important roles in shaping this solitary behaviour, and the biochemical mechanisms associated with delayed implantation may also contribute [1,74]. However, river otters (Lutra canadensis) show some rudimentary forms of matrilineal sociality. For instance, multiparous females sometimes allow their elder daughter(s) to return to live with them when they start to wean their current pups [24]. Yearling females either return to live with their mother, or they are aggressively expelled permanently from the mother's home range by an older sister. After being accepted back by her mother, an elder daughter becomes a full-time, cohabiting member of the family and a regular companion to the young pups [24].

Giant otters (Pteronura brasiliensis) live in highly cooperative groups led by an alpha pair [21]. Protracted allo-parental care is critical for pup survival in this species [22]. As in ursids, genetic relatedness declines significantly with increasing geographical distance within dyads of adult female, but not male, giant otters [21]. When the dominant female dies, a related female subordinate typically acquires her position and inherits the group and the territory [21,22].

European badgers (Meles meles) are highly flexible and facultatively social. In prime habitat and at high densities, they form groups characterized by both male and female philopatry [17], in which conspecifics that den together in communal burrow systems, called ‘setts’, share a common range [75] within-group relatedness in setts is high [18]. Other forms of cooperation are rare in this species, but allo-parenting may occur among females [19]. In some habitats, females may disperse in coalitionary groups to form new setts (e.g. [20]).

6. Procyonidae

Coatis are the most gregarious Procyonid, living in bands composed of multiple adult females and their offspring. Male coatis disperse after puberty and live either solitarily or temporarily in multi-male groups [25–27]. Males are often excluded from bands except during the mating season, apparently because they may opportunistically prey upon juveniles [28]. The coati diet consists mainly of fruits and invertebrates bands fission into smaller subgroups to forage on fruit. Pairs of females within bands preferentially groom one another and also support one another during infrequent aggression that occurs [29]. Bands split when groups exceed three to five females, with fission occurring along previously established lines of affiliation. In some populations, closely related adult females frequently direct aggression together toward unrelated group-mates in defence of concentrated resources used by their offspring and other kin [30]. Thus, kinship drives cooperative behaviour in these populations, although non-kin may also cooperate elsewhere [25,29].

Throughout their range, raccoons (Procyon lotor) show a high degree of flexibility in their socio-spatial organization. As adults, females may live partially on or close to their natal range, and greater overlap of home ranges is positively correlated with higher relatedness [31]. As in ursids and mustelids, genetic relatedness declines significantly with increasing geographical distance within dyads of female, but not male, raccoons [32]. Similarly, kinkajous (Potos flavus) are generally solitary but are sometimes found in groups composed of a mother–daughter pair plus one or two unrelated males [76].

7. Canidae

Most canids exhibit an enduring bond between mates, in which the male helps the female to rear young. Care of young usually includes provisioning of the female and her litter by the male and other group members [1]. In fact, provisioning the female and young by the male occurs even in those species that do not form packs, such as the arctic fox (Vulpes lagopus) and red fox (Vulpes vulpes). The social organization found in pack-hunting canids (wolves, Canis lupus, Cape hunting dogs Lycaon pictus, dohles, Cuon alpinus and bush dogs, Speothos venaticus) probably evolved through continued interaction between the bonded parents and their offspring after weaning [2,5,77,78]. Both provisioning and pair-bonding are probably phylogenetically old traits within the Canidae both contribute to the successful rearing of the large litters generally borne by female canids. Selection for the formation of larger social groupings appears to have acted to enhance the association between the male–female pair and their offspring.

Because of the importance of food provisioning and territorial defence by males, the social structure in most canids is shared or even dominated by males. However, small, insectivorous canids show little paternal care, and they also show a bias toward female philopatry. For example, in contrast to most other canids, the social organization of the swift fox (Vulpes velox) is based on female territories, which are maintained even in the absence of males [33]. Neighbouring swift foxes are more closely related than non-neighbours, and female clusters are more extensive than male clusters [34]. Relatedness between female holders of adjacent territories is correlated with the degree to which they tolerate home range overlap [34], and neighbours sometimes even share dens. Relatedness also influences the likelihood that an individual will inherit a newly vacated home range [34]. The tolerance shown to neighbouring related swift foxes indicates that these animals can identify their kin and that they maintain long-term relationships with them. This unusual canid social system may have evolved because the importance of food provisioning and territorial defence by males is reduced in swift foxes owing to their reliance on insect food resources [33].

8. Herpestidae

At least 19 species of herpestids are known to be gregarious [35], and at least some indication of matrilineal social structure has been documented in five species: dwarf mongooses (Helogale parvula), banded mongooses (Mungus mungo), meerkats (Suricata suricatta), yellow mongooses (Cynictis penicillate) and white-tailed mongooses (Ichneumia albicauda). As in canids, group living in mongooses appears to be an elaboration of the relationship between a male–female pair and their offspring [79]. Dwarf mongoose and meerkat groups are characterized by male-biased dispersal, female dominance, an alpha pair that controls most reproduction [35] and high levels of intragroup relatedness among females and their descendants [38,41]. Among adult female banded mongooses, intragroup relatedness is often high, particularly in newly founded groups, such as those occurring when individuals disperse together [11,36,80]. In the cooperatively breeding mongooses, allo-care of pups is essential for their survival and may include nursing, guarding, carrying and feeding of young [35]. Helpers are subordinate individuals of both sexes that are often, but not always, related to the latest litter [35,39,81,82]. Both male and female banded mongooses can inherit breeding positions in their natal group, and reproductive skew in both sexes tends to be low in this species [37]. However, in dwarf mongooses [40] and meerkats [42], only female offspring can inherit the alpha breeding position, territory and subordinate helpers from their maternal kin. Reproductive skew in these species tends to be considerably greater than in banded mongooses.

At high densities, the facultatively social yellow mongoose may form groups containing multiple breeding females, which tend to be closely related, though allo-parental care is provided by both kin and non-kin [45]. Similarly, although white-tailed mongooses are typically solitary, at high densities they show range overlap among related adult females and their offspring [35]. Females of this species forage and den alone, but recognize and tolerate one another and act aggressively towards non-group members [44].

9. Eupleridae

This family contains five species of carnivores native only to Madagascar [12], all of which are severely understudied. As in herpestids, group living in euplerids has antipredator and resource defence benefits [46]. Female narrow-striped mongooses (Mungotictis decemlineata) live in small, philopatric groups of related females and their offspring [47]. Group members den together but do not provide allo-parental care [47]. Their social organization is matrilineal, with neighbouring groups sharing the same haplotypes [47].

10. Felidae

The efficiency of hunting and feeding characteristic of felids has facilitated the evolution of more-or-less exclusive home range use and a solitary lifestyle in most species in this family. The primary social grouping in most felids consists of the mother and her offspring. After weaning, the young usually leave voluntarily or are driven away by the mother [83]. However, specific evolutionary precursors to matrilineal societies appear even in solitary felids. For example, as population density increases among wild leopards (Panthera pardus), female home range size decreases, their home ranges show increasing overlap and females show a greater tendency to form matrilineal kin clusters [48].

The lion is the only felid living in mixed-sex groups that habitually hunt, feed and rest together [84,85]. A lion pride consists of several related females and their young adult female pride-mates nurse one another's cubs. Most females remain in the natal pride for life, retaining close associations with 1–20 female relatives, such that female pride members are seldom more distantly related than cousins, sharing on average at least 12.5% of their genes [10]. By contrast, the males that associate with pride females are generally unrelated to them. Females often have close relatives in neighbouring prides, but very few relatives in prides with territories not abutting their own [86]. Relatedness among female lions declines sharply with geographical distance, and mean relatedness is greater in small than large prides [86]. Females occasionally leave their pride to avoid having their cubs killed by new males during pride takeovers. The formation of new prides involves emigration of a group of related females into an area within, or adjacent to, the territory of the natal pride [10]. Females cooperate in group hunts [87] and support one another in aggressive interactions with new cohorts of immigrant males in an effort to prevent infanticide by the males females may be wounded or even killed in these fights. Cooperative defence of cubs appears to be an important aspect of communal rearing among female lions [88].

Given the rare appearance of enduring pair bonds between male and female parents in the family Felidae, complex social organizations could not have derived from a pair and their offspring, as occurred in canids. Instead, the basic social unit of the lion pride emerged from enhanced tolerance among females, based on the continued association of a lioness with her maturing daughters [85,89]. In contrast to canids, male felids do not provision their young, so this falls entirely to the mother and her female kin. The complex social structure in extant lions appears to have been shaped by selection acting on the mother–family as the original temporary social unit [5].

11. Hyaenidae

Two extant hyaenid species show simple forms of matrilineal social organization. Brown hyenas (Parahyaena brunnea) live in small groups, called clans, ranging in size from a single female and her cubs to approximately 14 animals [52]. Clan members share a common territory and den [53,54]. Larger clans consist of extended families that include a female, her adult offspring of both sexes and an immigrant male. Both males and females may provision den-dwelling cubs [52]. Brown hyenas forage solitarily to feed on carrion, fruit and small prey, resources that tend to be rare and widely dispersed, such that they provide food for only a single individual [52–54]. Females often spend their entire lives in their natal clans, and some males stay with their natal clans into adulthood, but dispersal is nevertheless strongly male-biased [52]. Females within clans are more closely related than females between clans and are more closely related than are males with female clan-mates [55].

Like brown hyenas, striped hyenas (Hyaena hyaena) feed mainly on carrion. These animals were long believed to be strictly solitary (e.g. [90]), but recent work has revealed that they sometimes form small aggregations (e.g. [66–69]). Although striped hyenas forage alone, closely related females sometimes share dens in habitats where resource abundance permits [68,69]. Mothers carry food back to the den for their cubs [90–92], and other kin of both sexes have also been observed provisioning den-dwelling cubs [68]. Thus, the social system of striped hyenas appears to be flexible in resource-rich habitats, their groups closely resemble those of brown hyenas, although groups of brown hyenas tend to be larger than do striped hyena groups.

The spotted hyena exhibits by far the most complex matrilineal society found within the order Carnivora. Spotted hyenas live in large clans that typically contain multiple groups of matrilineal kin, as well as one to several immigrant males. Clans, which may contain more than 120 individuals [93], are structured by linear rank relationships [94–96]. Female kin generally occupy adjacent rank positions in the clan's hierarchy, so entire matrilines can be ranked as well as individual hyenas [57]. In contrast to brown and striped hyenas, spotted hyenas are very good hunters, killing up to 95% of their own prey they prey mainly on medium- and large-bodied herbivores [97]. A group of hungry hyenas can reduce a 150 kg antelope to a skull and vertebrae in as little as 13 min, so feeding competition is extremely intense among clan-mates, and an individual's social rank determines its priority of access to carcasses [94,98]. Importantly, food intake limits the rate of reproduction among females [99].

All clan-mates know one another individually, rear their cubs together at a communal den and defend a common territory [58]. Litters, which usually contain only one or two cubs, are smaller in spotted hyenas than in other hyaenids. Maternal kinship is an extremely important determinant of interaction patterns in the societies of spotted hyenas. Females are philopatric whereas males disperse, so females spend their lives associating closely with their female kin [94,100], and they show extensive affiliative and cooperative behaviour toward them (e.g. [56,59,60]). Aggression among wild hyenas is more frequent between than within matrilines [94], although relatedness does not protect females from aggression this and other costs associated with competition between relatives can outweigh the kin-selected benefits of altruism toward relatives in carnivores and other animals [61,101]. Female allies are critical in maintaining access to resources in spotted hyenas [102]. During early ontogeny, young spotted hyenas learn to acquire social ranks immediately below those of their mothers [96,103]. Emergence of a matrilineal social system occurs in both wild and captive spotted hyenas [104]. Matrilineal rank acquisition in both wild and captive environments appears to result from relationships generated between mother–offspring units and other group members (e.g. [105]).

As with lions, it has been hypothesized that spotted hyenas live in groups because cooperative hunting allows these animals to capture larger prey animals or enhance their hunting success, as occurs in pack-hunting canids (e.g. [77]). However, although cooperative hunting does increase the probability of success and allows spotted hyenas to capture larger prey animals, 75% of successful hunts are conducted by lone hyenas [62], and only the largest and most challenging of prey species require more than one hyena for a successful hunt [62,95]. This suggests that cooperative hunting was likely not paramount in the evolution of matrilineal societies in spotted hyenas, although assistance during hunts clearly represents a beneficial side-effect of group living. Instead, the need to defend spatial and food resources from competitors most likely favoured gregariousness among female hyenas. Moreover, the herbivores killed by spotted hyenas represent an abundant and readily renewable food resource that allows these hyenas to reach high population densities. Spotted hyenas often live and hunt in open habitat where competitors, such as lions and members of neighbouring hyena clans, can easily detect kills, which are typically too large to be monopolized by a single individual. These conditions appear to have favoured grouping by females to protect carcasses and also to protect group territories in areas of high population density.

12. Socio-ecological conditions facilitating the evolution of matrilineal societies in carnivores

Although many gaps still exist in our knowledge, particularly about social interactions among small-bodied solitary carnivores, field studies have taught us a great deal about the socio-ecological conditions favouring the evolution of matrilineal social systems in carnivores. These include male-biased dispersal, female philopatry, reliance on large or abundant prey, particularly in open habitat, the need for assistance either in protecting or provisioning offspring or in acquiring and protecting key resources. As in other orders of mammals, natal dispersal in most mammalian carnivores tends to be sexually dimorphic [106,107], with males typically dispersing farther or more frequently than females. In solitary carnivores, both sexes disperse away from the mother's territory, although female offspring tend to settle closer to their mothers than do their brothers. The spatial proximity between mothers and daughters sets the stage for the emergence of nepotistic behaviour among females. For example, as they get older, female leopards and bears relinquish portions of their home ranges to philopatric daughters [13,49].

Patterns of hunting and feeding in terrestrial carnivores, and the resultant tendencies towards clustering of female kin, are strongly influenced by both prey density and density of conspecifics. Both abundant food resources and high conspecific densities encourage reduced dispersal and increased aggregation of female kin in many carnivore families. For example, although dispersal distances are longer among female than male striped hyenas in areas with scarce resources [66], where resource abundance is high striped hyenas exhibit male-biased dispersal, home range overlap and den-sharing among female kin (e.g. [68,69]). Similarly, in both European badgers and giant otters, aggregations of females and inheritance of territorial resources by younger females occur in response to high population density, abundant food resources and the lack of available territories for dispersing offspring [17,21,22,108,109]. The spatial distributions among extant bears appear to be largely due to the patchy distribution of vital resources. However, bears and other solitary carnivores are known to aggregate during periods of, or at places characterized by, abundant food resources (e.g. polar bears [70], raccoons [110], leopards [48]). This may also occur in the other families of solitary carnivores, including Viverridae, Nandiniidae, Mephiditae and Ailuridae, but to our knowledge, it has not yet been described in the literature.

In most carnivores, other than canids and social mongooses, the reproductive roles of adult females, but not males, are dominated by infant care. Females obliged to care for infants on their own tend to have smaller litters than do females in carnivore species in which mates or kin help with offspring care [111]. During the early phases of development in the former species, the mother is solely responsible for protecting and feeding infants when an infant enters the juvenile stages, the mother also plays an important role in its socialization. Protecting infants from infanticidal male conspecifics represents a challenge for female carnivores, who are often considerably smaller than males. In both bears and lions, this need appears to have favoured spatial clustering and resource-sharing by maternal kin. To reduce the risk of infanticide, female bears with young may avoid areas inhabited by adult males [103]. Infanticide has been observed in many carnivores, including American black bears, Asiatic black bears [14], brown bears, polar bears [103] and lions [88].

Variation in social structure among canid species appears to be strongly influenced by the importance to pup survival of food provisioning and territorial defence by males [33]. In pack-hunting canids (wolf, dhole, Cape hunting dog), both group living and cooperative hunting have evolved as adaptive responses to the presence of large prey that cannot be brought down by a single individual (e.g. [77]). Similar adaptations to feeding upon medium and large-bodied ungulates occurred within the Felidae and Hyaenidae, culminating in the evolution of large stable social groups in lions and spotted hyenas. However, whereas in canids, this occurred via the elaboration of the bond between a mated pair and their offspring, in felids and hyaenids, it occurred via the elaboration of the mother–family. More rudimentary forms of the extended mother–family appear in the mustelids, procyonids, euplerids and ursids.

The most elaborate matrilineal societies found in mammalian carnivores are those of lions and spotted hyenas. The widespread African grasslands and abundant herbivores present during the late Pliocene and Pleistocene probably created an environment favouring matrilineal kin aggregations in both these species. The lion evolved between 1 million and 800 000 Ma in Africa [112] similarly, modern spotted hyenas first appear in the fossil record in Africa between 900 000 and 250 000 Ma [113]. Spotted hyenas and lions occur sympatrically throughout much of sub-Saharan Africa, so these two matrilineal societies presumably evolved in response to the same ecological conditions, the most important of which were abundant large-bodied prey, open grassland habitat and considerably higher population densities than is typical of other felids or hyaenids [50]. In open habitats, kills can be detected by competitors over long distances, so the defence of both kills and territorial boundaries demands cooperative action [63]. The spotted hyena is the only extant hyaenid adapted for the hunting of prey, as brown and striped hyenas feed mainly on carrion. Felids other than lions either prefer smaller-bodied prey or live in habitats characterized by denser vegetation than African savannahs [50]. Although it was long believed that the selection pressure favouring the evolution of matrilineal groups among lions was the need for help in capturing large prey, instead the evidence supports the hypothesis that selection favoured female lion kin who helped one another defend young, carcasses and territories [50,88].

Lions compete for carcasses directly with spotted hyenas and, because lions are much larger than hyenas, lions also represent a leading mortality source for hyenas [52,95,114]. Individual hyenas cannot effectively compete with lions for possession of a carcass, or defend themselves from predation by lions, without aid from conspecifics. However, when spotted hyenas join forces with group-mates, they can not only defend their own kills from lions, but they can also sometimes usurp kills made by lions [115]. Under conditions of high population density, effective maintenance of group territories also requires individual females from multiple hyena matrilines to cooperate to defend clan resources against neighbouring hyena clans [116]. Loss of a clan war can result in a substantial reduction of a clan's territory [95] and repeated losses can result in an overall loss of the territory to a neighbouring clan. Cooperative defence of territories appears to offer a similarly important advantage during intraspecific between-group conflicts in many other carnivores, including dwarf mongooses, meerkats, banded mongooses and Ethiopian wolves (Canis simensis, [117]), with the larger of two groups typically winning these disputes (e.g. [118,119]).

13. Fitness consequences of enduring association with maternal kin

Patterns of spatial proximity influence the frequency and outcomes of competitive and cooperative interactions among mammalian carnivores, and this can have profound effects on survival and reproduction, as can enhanced social tolerance among female kin. Inclusive fitness modulates intraspecific competition for resources in carnivores when female relatives are spatially aggregated and likely to interact cooperatively [120]. For instance, the matriarchal spatial distributions characteristic of many bears confer inclusive fitness benefits, because daughters can more easily establish and hold territories without risking dispersal [70]. Dispersal is particularly risky for small carnivores like herpestids [69]. Females need knowledge of local, ephemeral resources including den sites and food sources this knowledge can protect them against both starvation and infanticide [70,88]. Thus, inheriting the dominant position in a natal group can be particularly beneficial for daughters. In meerkats, females gain a higher fitness benefit than do males from acquiring dominant breeding status, and this may explain why traits enhancing competitive ability are more highly developed in females than males [42]. Similarly, inheriting a territory and a breeding position obviates risky dispersal for female giant otters and European badgers [21–23].

Long-term studies of carnivore groups have revealed that the fitness benefits of allo-parental care by females other than the mother range from nil to substantial. Whereas increasing the number of allo-parental females per group correlates negatively with offspring fitness in European badgers [19], they correlate highly positively in meerkats [43] and lions [10,51,121]. Long-term studies have also shown that kin-structured cooperative interactions have strong positive effects on fitness in both lions and spotted hyenas.

The grouping patterns of female lions result from multiple factors, in particular, the demands of protecting young and maintaining a territory [10]. Successful breeding appears to be impossible without a territory in this species. Females compete aggressively against neighbouring prides and the larger pride usually wins between-group encounters [121]. Competition for territories directly affects the fitness of female lions, and larger prides gain and maintain access to the highest-quality habitat. Neighbours have a significant negative effect on female reproductive success and survival, and larger prides are also more likely to maintain control of disputed areas and to improve the quality of their territories [51]. Adult female lions are significantly less likely to be alone when a pride has more neighbours, suggesting sensitivity to the risk of encounter. Interestingly, prides that have recently split, and therefore contain related lions, seldom engage in intergroup territorial competition [51].

Whereas female lions seldom fight with related pride-mates over access to carcasses, female spotted hyenas frequently compete for food resources with their close kin. Kinship fails to protect spotted hyenas from attack when resources are at stake [61]. Some of the most intense competition observed in spotted hyenas occurs between full- and half-sibling littermates as they compete as infants for access to the mother's two teats [122]. Upon the death of an alpha female hyena, her youngest adult daughters may fight viciously with one another to fill the vacant alpha position. Nevertheless, closely related spotted hyenas exhibit strong nepotism and cooperation, both of which yield substantial fitness benefits. Not only do young females inherit their mother's social rank, but they also inherit her entire social network of associates [123]. Maternal kin associate more closely than do non-kin, they are more affiliative toward one another than are non-kin, they hunt together more than do non-kin, and they tolerate one another more readily than non-kin while feeding at kills [52,56,59,60,64]. Close maternal kin are usually the most common partners in coalitionary attacks on unrelated hyenas [61,124]. Furthermore, these animals recognize and respond to distress vocalizations emitted by close maternal kin far more vigorously than they respond to the distress vocalizations of non-kin [125]. In addition to generating these short-term benefits, close associations and coalition-formation among female kin also appear to play a central role in maintaining long periods of stability in the adult female dominance hierarchy [102] they also cause the occasional coup d’état in the hierarchy that overthrows a higher-ranking lineage and thereby improves resource access for an entire matriline [52,65].

14. Conclusion

Evidence of matrilineal social systems is found in many mammalian carnivores, albeit only in rudimentary form in more solitary species. Regardless, it is straightforward to envision how the elaborate matrilineal societies of lions and spotted hyenas evolved from social systems characterized only by such rudimentary building blocks as the temporary mother–family described by Eisenberg [2]. We still know very little about matrilineal kin relationships in many rare or solitary carnivores, particularly small-bodied species occurring in habitats with dense vegetative cover, so this represents an important area for future research. The limits of nepotism in mammalian carnivores should also be explored in future work. The evolution of matrilineal social systems in carnivores has been favoured by high population density, reliance on large or abundant prey, small litter size, male-biased dispersal and the need for assistance in protecting or provisioning offspring or key resources. Once selection has favoured enhanced gregariousness, then the stage is set for the evolution of nepotistic behaviour and kin-structured cooperative interactions among female carnivores, and these in turn can have strong positive effects on fitness.

Data accessibility

This article has no additional data.

Authors' contributions

Each author reviewed the literature on four families and drafted the appropriate sections.


Asiatic lions can often be morphologically differentiated from African lions based on (a) skull characteristics, wherein the Asiatic lions have an extra infraorbital foramen, (b) a typical loose fold of skin on the abdomen known as the belly-fold which is absent in African lions (Oɻrien, 2003), (c) facial characteristics of Asiatic lions, with a more elongated snout and a more sloping forehead giving them a longer profile in lateral view in comparison to the African lions and, (d) males having sparser manes, never covering their ears The mane in the adult lion has the typical “mohawk” style look. (Figure 1, Supplementary Material S1). As part of our long-term research project, we combined our field observations of known lions with techniques developed for African lions to develop criteria for estimating the age of individuals (Supplementary Material S2), which helped us construct their population structure and demographic details.

Figure 1. Face and body profiles of Asiatic lions. (A) Adult male, note the sparser mane (than African lions) that does not cover the ear and the top of the head with a Mohawk look, and the prominent belly-fold and (B) adult female, with a longer sloping snout and side face profile than African lionesses. Also, note the size difference between the male and the female (a consorting pair). Photographs taken by Stotra Chakrabarti.

Between 2001 and 2018, we captured 35 free-ranging lions (including sub-adults that were targeted for understanding dispersal) from different parts of the Gir landscape in order to deploy radio-transmitters or for treatments, and recorded their morphometric details. We found average weights of adult males (n = 7) and females (n = 12) to be 160 (SE 4.7) kg and 116.5 (SE 3.7) kg, respectively (Table 1).

Table 1. Morphometric details of adult Asiatic lions (n = 19 12F, 7 M) captured between 2001 and 2018 for deploying radio-collars. Body length is measured from nose-tip to tail-tip along the curves.

Like tigers and leopards, several local variations in lions based on their mane size and coloration, and coat texture have been recorded from different parts of India and from within Gir (Divyabhanusinh, 2005). Adult male lions are often grouped by local communities into various categories based on the color of their manes that can range from golden yellow (Pinglo), speckled gray (Bhurio) to black (Kamho) (Divyabhanusinh, 2005).

Distribution and Status

The erstwhile range of the modern Asiatic lion, reconstructed mainly from paleontological evidence, literature, art, culture, and shikar (hunting) documents suggest an extensive area from Anatolia, Syria across the Middle East to Eastern India (Kinnear, 1920 Caldwell, 1938 Joslin, 1973). Till the mid-1800s, lions in India inhabited the entire northern Indo-Gangetic Basin in North and Central India and were abundant in the modern states of northern and western India, Bihar and Odisha in the east with the river Narmada being the southernmost boundary (Fenton, 1908 Pocock, 1930 Dalvi, 1969). Subsequently by late 1800's they were exterminated from most of their range because of hunting and habitat loss (Divyabhanusinh, 2005). By 1880s lions were restricted as a single free-ranging population in and around the Barda and Alech hills, Mitiyala, Girnar, and Gir forests in the Saurashtra peninsula of Gujarat (Dalvi, 1969). Although some lions continued to survive in isolated habitat pockets of Iran and Iraq, but these were not viable populations and soon became extinct. By 1888�, hunting and loss of forests due to agricultural expansion and livestock grazing in Saurashtra restricted the lions to a single population in the Gir forests, a patch of about 2,000 km 2 composed of dry deciduous and thorn forest (Divyabhanusinh, 2005).

Driscoll et al. (2002) suggests that about 2,680 (range 1,081𠄴,279) BP, the Kathiawar Peninsula that contained the Gir forests was separated from mainland India by rising sea levels in the Gulf of Khambhat (Gupta, 1972), causing the first genetic bottleneck that isolated the founders of the present Asiatic lion population, compelling them to inbreed for several generations (Oɻrien, 2003). By the time the Gulf water receded and the peninsula became continuous with the mainland, most of the lions from mainland India had become locally extinct providing little chance to the inbred population to enhance their genetic diversity. A second, less-severe but more popularly known bottleneck occurred at the onset of the 19 th century when owing to rampant hunting, Gir lions dwindled to around 㱐 individuals (Edwardes and Fraser, 1907 Kinnear, 1920 Pocock, 1930).

Owing to the timely protection measures taken by the Nawabs of Junagadh who ruled most of the Gir region, lions survived (Divyabhanusinh, 2005) and increased to about 287 by 1936 (Dalvi, 1969). Subsequently, the Government of Independent India enforced a complete ban on lion hunting in 1955 and declared the Gir forests as a Wildlife Sanctuary in 1965. Ensuing protection and habitat management by the Gujarat Forest Department resulted in the lion population increasing steadily (Singh and Kamboj, 1996) to over 500 in the last 2015 total count (Gujarat Forest Department, 2015). The sub-species was also down-listed from the 𠇌ritically Endangered” category of the IUCN Red list in 1990s (Nowell and Jackson, 1996) to 𠇎ndangered” in 2008 (Breitenmoser et al., 2008). Within the past two decades, lions have dispersed into about 13,000 km 2 of agro-pastoral landscape comprising of the Gir Protected Area (Gir PA 1700 km 2 ), Girnar Wildlife Sanctuary (180 km 2 ) and over 11,000 km 2 of human-dominated landscape and coastal scrublands of the surrounding districts of Junagadh, Amreli, Gir Somnath, and Bhavnagar (Ranjitsinh, 2016 Singh, 2017a). Currently, the Saurashtra landscape has a single source population of lions comprised of 񾌀 adult individuals that live within the Gir National Park and Wildlife Sanctuary, and several patchily distributed small sink populations (Pulliam, 1988) of 㱐 individuals each in the human dominated agro-pastoral system (Figure 2). Though these small populations do breed and recruit lions, immigrants from the Gir PA are an essential element for their long-term viability (Banerjee et al., 2010). Radio telemetry (Jhala et al., 2014a) has shown extensive movement between these small populations and with the lion population of Gir PA. Lions thus exist in a classical metapopulation framework in the Saurashtra landscape (Hanski and Gilpin, 1997 Cronin, 2003). Consequently, habitat connectivity that facilitates lion movement between populations is vital for long-term lion persistence in the Saurashtra landscape (Banerjee et al., 2010 Banerjee, 2012).

Figure 2. Lion habitat suitability and current lion distribution across Saurashtra landscape overlaid with Protected Areas, breeding refuges outside the protected areas, important lion movement pathways, and linear infrastructure (major roads and railways). Note the location of Barda WLS that is being considered for lion reintroduction.

While the recovery of Gir lions elucidates a conservation success story it also poses serious challenges for wildlife managers and conservationists in terms of maintaining the future persistence of this subspecies. A population gains security with increasing size and the species becomes secure with increasing number of viable populations (Soulé and Simberloff, 1986). The importance of human free space for large carnivore conservation is undebatable, as conflict with human interests has been the major cause of large carnivore declines worldwide (Woodroffe, 2000). Indeed, lions were often poisoned on livestock carcasses in Gir until recently, when law enforcement became very strict. Currently only 259 km 2 of inviolate space (devoid of human habitation and use) is allocated as Gir National Park for lion conservation in Gujarat. The rest of the protected areas are in the form of wildlife sanctuaries (WLS), reserve forests and protected forests with varying levels of human habitation and legally permitted human use of forest resources (Wildlife Protection Act, 1972), including livestock grazing rights of local semi-nomadic pastoral communities, the Maldharis. With land ownership being primarily private in the landscape outside the PAs, creation of new PAs in Saurashtra is difficult. Since the PAs in the landscape have reached carrying capacity for lions with about 300 individuals (Singh, 1997), maintaining the current population of 500 lions or increasing it can only be achieved by ensuring the continued source value of the Gir PA and by providing dispersal corridors to the several small sink-populations in the agro-pastoral landscape. Coexistence with humans thus becomes an inevitable strategy for maintaining a viable lion population in this landscape. However, the Saurashtra landscape is rapidly transforming due to development of linear infrastructure, expanding urban sprawl, agricultural intensification and changing community values. With increasing lion density in this progressively hostile landscape, a multidisciplinary understanding of lion ecology, conflict resolution, and socio-economic underpinnings is required for maintaining lion-human coexistence.


The earliest attempt to estimate lion population based on shikar records were made by William Rice of the Indian Army in 1850s when he concluded that not more than 300 individuals were left in India (Divyabhanusinh, 2005). Subsequent estimates made by forest and army officials under the rule of Junagadh State figured about 20� lions in between 1905 and 1913. The first lion census based on pugmark counts at waterholes was conducted in 1936 and reported a total of 287 lions (Wynter-Blyth, 1949). Since 1963, the Gujarat Forest Department has estimated lion numbers about every 5 years by a labor intensive, 3-day total count using livestock bait (Singh, 2017b). In this method, a daily record was kept of all lions that visited the baits. Lions feeding on baits remained localized in the vicinity for 3𠄴 days. If, however, lions moved away to another bait site a record of the movement was kept and accounted for to minimize double counts while computing total number of lions. The maximum number of lions recorded on any single day was considered to be the total population.

Both pugmark census and total counts depend on unrealistic assumptions, are error prone as they do not address detection issues, require careful identification of duplications, trained field staff and are resource intensive (Williams et al., 2002). To circumvent these issues, we designed and demonstrated lion abundance estimation in a mark-recapture framework (both conventional and spatially explicit) based on individual identification of lions from their vibrissae patterns, ear notches and permanent body marks (Jhala et al., 1999, 2004 Jhala, 2004 Banerjee and Jhala, 2012). Lions ϡ.5 years were approached within 10� m on foot or from vehicles and photographed. Individual lion details (age, gender, identifying features, associated lions, geographic coordinates, photographs, etc.) were then entered in program LION (Jhala et al., 2005) (Supplementary Material S3) for storing, archiving, identifying and comparing with the lion database so as to generate information useful for abundance estimation and long-term monitoring of demographic parameters and movement patterns.

Lion density was found to be the highest in the Gir PA at 15 (SE 0.1) lions/100 km 2 followed by Girnar WLS [6 (SE 0.7) lions/100 km 2 ] and the human dominated landscape of Saurashtra [2 (SE 0.1) lions/100 km 2 ] (Jhala et al., 2004 Banerjee, 2012 Banerjee et al., 2013). Spatially explicit density of lions in the western part of the Gir PA was positively correlated with tourism hotspots due to artificial food provisioning at these sites (Gogoi, 2015). Due to vegetarian lifestyles of local communities, dead livestock are dumped outside settlements. These carcasses attract large carnivores including lions and leopards (Panthera pardus). To minimize encounters between large carnivores and humans as well as to enhance sighting of lions by tourists, wildlife managers often retrieve such livestock carcasses from forest settlements and dump them at tourist viewing spots. This assured food source increased pride sizes and reduced their home ranges (Gogoi, 2015 Jhala et al., 2016). This distribution pattern caused by subsidized food resources overrides the influence of natural prey and other ecological factors, resulting in local lion densities that are higher than natural densities. We believe that though this practice would enhance tourist viewing but will have serious implications on the social organization of lions, spread of infectious diseases and might cause enhanced predation pressure on wild prey in small pockets harboring artificially enhanced lion density.

Allozyme and microsatellite studies indicate that the Asiatic lions have low genetic diversity due to an isolated, inbred population with a small founder base (Wildt et al., 1987 Oɻrien, 2003). However, random amplified polymorphic DNA analysis showed some levels of polymorphism in Asiatic lions (Shankaranarayanan et al., 1997). Oɻrien et al. (1987) and Wildt et al. (1987) found that Asiatic lions and cheetahs showed a high incidence of morphologically abnormal spermatozoa (79 and 71%, respectively) when compared to free-ranging African lions (25�%) and other species such as bulls Bos spp. and dogs Canis lupus familiaris (20�%). The serum testosterone (a critical hormone for spermatogenesis) was low and Asiatic lions had lower variability in the major histocompatibility complex gene responsible for immunity (Wildt et al., 1987). Todd (1965) attributed dentition abnormalities in Asiatic lions to inbreeding. Decreased heterozygosity likely diminishes reproductive vigor and long-term survival of a population (Oɻrien et al., 1986 Packer et al., 1991).

In order to understand the demographic parameters of Gir lions, 68 adult lions, and 91 cubs from 38 litters were intensively monitored using telemetry and individual lion ID profiles (Banerjee and Jhala, 2012). Records of opportunistic mortality events (n = 228) were used to understand mortality causes. Gir lions apparently increased from about 177 in 1968 to about 523 by 2015 with an r = 0.022 (SE 0.001) translated into an annual population growth of 2.2%. Male: female ratio was 0.63 (SE 0.04) while cub: adult lioness ratio was 0.37 (SE 0.02). Though breeding is observed year round, mating peaked in winter while birth peaked in late summer. Average litter size was 2.39 (SE 0.12). Inter-birth interval was 1.37 (SE 0.25) years (n = 7 lionesses) and was higher [2.25 (SE 0.41) years] when cubs of the previous litter survived to independence. Cub (ρ year) survival was 0.57 (SE 0.04) while survival from cub to recruitment age (3 years) was 51% (SE 4%) with infanticide attributing to 30% (SE 7 %) of mortalities. Average annual survival rate of adult lions (ϣ years) was 0.9 (SE 0.12). Based on records of 228 lion mortalities recorded between 2007 and 2019, we estimated that 30% of the deaths were caused by diseases (Figure 3). Adult lions died primarily due to natural causes (60%), however, human caused mortality was also substantial (32%). Deaths due to falling in open irrigation wells, electrocution by live wires deployed illegally to prevent crop damage from nilgai (Boselaphus tragocamelus) and wild pigs (Sus scrofa) were a cause of concern in the agro-pastoral landscape. These are being addressed by wildlife authorities by subsidizing the construction of parapets around open wells and pulsating solar-powered wildlife fences to agricultural fields.

Figure 3. Causes of mortality among free-ranging Asiatic lions (n = 288). Lions primarily died because of natural causes (60%), while anthropogenic reasons for mortalities (32%) were substantial. Among the natural deaths, 33% were due to canine distemper virus and other diseases.

Banerjee and Jhala (2012) had concluded that demographic parameters of Asiatic lions did not differ from those of African lions, and went on to suggest that there was no evidence of inbreeding depression on vital rates. Subsequently, there have been recorded instances where free-ranging lion cubs were detected with missing and malformed limbs, or were born blind (Supplementary Material S4). These are potential indicators of inbreeding (Oɻrien, 1990), and in nature such handicapped individuals rarely survive to propagate these traits, thereby purging out deleterious alleles from the population over time (Keller and Waller, 2002). Intensive health care of wild lions as practiced in recent times by wildlife managers (between 2001 and 2010, 501 lions were captured and treated by Gujarat Forest Department, Pathak et al., 2002 Meena and Kumar, 2012) ensures survival of many such unfit individuals. Such tampering with natural selection processes can have serious implications on the future survival of wild lions (Banerjee and Jhala, 2012).

Social Organization and Behavior

Though biologists have been observing lions in Gir since 1960's (Joslin, 1973), quantitative data on lion social behavior has only just begun to accumulate (Chakrabarti and Jhala, 2017, 2019). In free-ranging Asiatic lions, prides comprise only of females and their dependent cubs, while adult males (singletons or coalitions) form separate units covering the ranges of multiple female prides (Joslin, 1973 Chellam, 1993). However, adjacent female prides were found to have exclusive territories and such territories remained almost constant over the years (Chakrabarti and Jhala, 2019). Females of a pride rear cubs together in a crຌhe, but estrus synchrony is not as prominent as reported in their Serengeti counterparts (Chakrabarti and Jhala, 2017). Cubs are weaned at 5𠄶 months of age but remain dependent on their natal pride for food till 2𠄳 years of age (Joslin, 1973 Banerjee and Jhala, 2012).

Unlike as reported for egalitarian African lion societies (Packer et al., 1988), Asiatic male lions form hierarchical coalitions wherein every coalition has one dominant male who appropriates 㹰% of all matings and 45% more food from his subordinates from shared kills (Chakrabarti and Jhala, 2017). Owing to such strict linearism in resource appropriation between male partners in the Asiatic lion coalitions, males belonging to coalitions of two acquired higher benefits compared to single and low-ranking males in large coalitions (of Ϣ males). This has resulted in an optimum coalition size of two males in the Asiatic system (Chakrabarti and Jhala, 2017).

Interactions between the two sexes are limited primarily to mating and occasionally on large kills (Meena, 2008 Chakrabarti and Jhala, 2017). Male lions frequently fend for themselves: hunting on their own, scavenging livestock carcasses and kleptoprasitizing kills made by leopards and lionesses (Chellam, 1993 Meena, 2008 Banerjee et al., 2013). Asiatic lions thus form same-sex groups, where each group behaves more like a solitary carnivore and act as independent entities (Chakrabarti and Jhala, 2019). Group sizes are smaller in the Asiatic system with male and female groups averaging at 1.7 (SE 0.2) and 2.5 (SE 0.4) adults, respectively (Gogoi, 2015). Such operational and functional separation between females and males seem to be in contrast with lion societies reported from the Serengeti and Ngorongoro (Schaller, 1972 Bertram, 1978 Packer et al., 1988). However, degrees of male-female interactions akin to that found in Gir have also been reported from lions in the Luangwa valley in Zambia, where hunting of males have severely reduced their numbers and hence, ability to maintain exclusive and all-round access to female groups (Yamazaki, 1996).

Male coalitions (with 𢙒 male partners, n = 7) had an average home range (95% Minimum Convex Polygon) of 120 (SE 19) km 2 , much larger than single males (n = 4) averaging at 31 (SE 3) km 2 . Single males had shorter tenures as territorial breeders [14 (SE 3) months) than coalition males [30 (SE 4) months] (Chakrabarti and Jhala, 2019). For reproducing successfully, a male needs to hold tenure for over 24 months so as ensure that cubs sired by him reach recruitment age and are not killed by infanticidal new territorial males (Schaller, 1972). In cases where resident male(s) were ousted by new male(s), cubs and juveniles 㰘 months of age were mostly killed by the new males or rarely survived when forced to disperse (Chakrabarti and Jhala, 2019).

Chakrabarti and Jhala (2019) hypothesizes that this disparity in group size and male-female association from the lions in the Serengeti can be attributed to plasticity of social behavior in response to the differences in resource availability between the two systems. Asiatic lions subsist on smaller prey (modal prey- chital Axis axis, averaging at around 45 kg) (Meena et al., 2011 Banerjee et al., 2013 Chakrabarti et al., 2016), resulting in heightened intra-group competition for food and ensuing smaller group sizes (Chakrabarti and Jhala, 2017). Furthermore, in the Asiatic system prey species are non-migratory and evenly distributed at reasonably high densities resulting in smaller and seasonally uniform female pride territories (Jhala et al., 2009) and higher lion density. This possibly allows males to maximize their reproductive potential by encompassing many female prides within their home ranges simultaneously. These arguments pertaining to prey- size and availability are in consonance with circumstances prevailing in West and Central African lion populations, where the lack of large prey has been reported to have resulted in small group sizes in lions (Bauer et al., 2003). Furthermore, it has been reported from the woodlands in Kruger that male lions were often found to be loosely associated with a particular pride of females and spent more time patrolling territories, hunting on their own and mingling with other female groups (Funston et al., 1998). Such a system somewhat mirrors the degree of male-female association in the Asiatic lion population, and as postulated by Funston et al. (1998), availability of ample cover seems to be one of the driving mechanisms for such a societal regime. With dense cover that aids in concealment, female lions likely require less assistance from their pride males in safeguarding cubs from marauding, infanticidal males (Funston et al., 1998). Following this argument, Chakrabarti and Jhala (2019) opined that dense cover in the deciduous Gir forests may have also prevented male Asiatic lions from controlling the females and retaining exclusive access to a female group.

In the Asiatic system, although male coalitions encompass multiple female groups, none of the female prides remain exclusive to any particular coalition (Meena, 2008 Banerjee, 2012). Such non-exclusivity of female groups to particular males/coalitions have compelled and allowed females to be promiscuous, where lionesses were found to mate with multiple neighboring (rival) coalitions (Chakrabarti and Jhala, 2019). In systems, where male coalitions have mostly exclusive mating rights over pride females (like in the Serengeti), extra coalition paternity are rare (Gilbert et al., 1991). But in land tenure systems where lionesses encounter multiple male coalitions who can potentially kill unfamiliar cubs, promiscuity likely aids females to familiarize with several males and buffer infanticide (Chakrabarti and Jhala, 2019). Furthering this thought, extra-coalition paternity has been reported from lions in Etosha where a genetic assessment has revealed that 41% of the cubs in the population were borne out of multi-male promiscuous matings (Lyke et al., 2013). The social organization and sexual strategies of lions differ across their entire global range of habitats, highlighting resource-mediated and anthropogenically (hunting pressure) driven behavioral plasticity in lions inhabiting diverse eco-regions (Chakrabarti and Jhala, 2019).

Habitat Needs and Activity

We used VHF, GPS, satellite telemetry and long-term monitoring of known individuals to understand ranging patterns, land tenure, habitat use, and activity patterns of lions (n = 97) across the Saurashtra landscape. Besides obtaining regular fixes (locations ranging from one per hour to one in 3 days), we followed each radio-collared lion on foot and/or a four-wheel drive vehicle continuously for 192� h sessions and carried out an all-behavior sampling [n = ߦ,412 hours of continuous monitoring data from 27 telemetered lions] (details available in Banerjee, 2012 Banerjee et al., 2013 Jhala et al., 2016).

Within the Gir PA, home ranges (95% MCP) of territorial males averaged at 91 (SE 17) km 2 which were more than three times the ranges of breeding females [27 (SE 8) km 2 ]. Lion home ranges in the human-dominated landscape outside the Gir PA were much larger than those inside the PA [territorial male = 832 (SE 42) km 2 breeding female = 169 (SE 57) km 2 ]. Core area (50% Fixed Kernel) of breeding lionesses inside Gir PA [7 (SE 3) km 2 ] were four times smaller than that of breeding lionesses outside the Gir PA [30 (SE 15) km 2 ] (Banerjee, 2012 Chakrabarti, 2018). Larger home ranges of lions in the outer landscape is in accordance to the Resource Dispersion Hypothesis (Macdonald, 1983) attributable to patchy distribution of resources (prey and suitable habitats) in the landscape, while within the Gir PA these are uniformly available.

Banerjee (2012) found average territorial tenure of males (n = 7) to be 36 (range 18�) months while average age at dispersal from natal prides for sub-adult males (n = 6) to be 3.9 (SE 0.13) years. We observed an old displaced male to successfully re-establish another territory and even father cubs after spending some time as a nomad. Average shift between successive territories for adult males was 21 (SE 5) km, while dispersal distance of sub-adult males from their natal territories was 16 (SE 4) km (Banerjee, 2012). Contrary to our expectation, activity patterns of lions within and outside PAs differed very little (Figure 4), attributable to the omnipresent human activities in the landscape and within the PAs (tourism, pilgrimage, grazing of livestock, and commercial activities of Maldharis).

Figure 4. Activity patterns of lions (n = 27 radio-collared lions) inside and outside Gir PA based on continuous (day-night) all behavior sampling data (ߦ,400 h). Major behavioral states have been depicted.

Gir vegetation primarily comprised of thorn and deciduous forests along with evergreen riverine patches (Qureshi and Shah, 2004). These riverine patches were critical lion habitats that provided respite from the summer heat (Jhala et al., 2009). Creation of the 259 km 2 National Park in 1975 after removal of 592 Maldhari families from central part of the Gir PA and recovery of the forest after the cyclone of 1982 has resulted in an increase in shrub (Helicteres isora, Holarrhena antidysenterica etc.) and tree density within the Gir PA (Khan, 1993 Sharma, 1995 Basu, 2013). Wildlife managers believe that this increasing vegetation density makes the habitat unsuitable for lions and their prey (Sinha et al., 2004), and have recommended selective thinning (Singh and Kamboj, 1996). However, wild ungulates of Gir are primarily browsers while domestic livestock are grazers (Dave and Jhala, 2011). Therefore, management interventions of opening habitats (besides removal of exotic invasive weeds like Senna uniflora and Lantana camara that abound in the livestock grazed areas of the PA) should be done only after careful site-specific evaluation.

Within the agro-pastoral landscape outside the PA, core areas of lion home ranges were composed of agriculture and thorn forests (Banerjee, 2012). Home range cores were observed to be farther from villages and townships but were closer to drainage and PAs (Banerjee, 2012 Jhala et al., 2016). Lions were active at night in this human dominated landscape, often venturing into villages and townships to hunt livestock. However, with advent of human activities during the day, lions sought concealment in vegetation cover. Average day time refuge patch size of lions in the human dominated landscape outside the PA was 7.5 km 2 (SE 0.74) but even small patches of vegetation (5𠄷 ha) were used. However, successful breeding by lionesses in this landscape required habitat patches of Ϥ km 2 (Banerjee, 2012). These findings through radio-telemetry highlight the importance of small interspersed vegetation patches characterized by thickets of Prosopis juliflora and Acacia senegal for lion persistence in the larger agro-pastoral landscape of Saurashtra (Figure 2). Remotely sensed time-series data on land cover changes suggests that this agro-pastoral landscape is rapidly being converted into urban setup with increasing development of linear infrastructure (Basu, 2013). Such infrastructure are detrimental for continued lion occupancy of the landscape as they will remove breeding and day-time refuges, as well as hinder dispersal routes between lion populations in the landscape and the PA. If lions are to continue to persist in this landscape, urgent changes in land policy and infrastructure development are required to safeguard these critical habitat patches and their connectivity.

A successful strategy for conserving large carnivores is to maintain a metapopulation structure (Hanski, 1994) within the landscape wherein one to many populations, that are demographically and genetically connected, act as source populations (Chapron et al., 2008 Walston et al., 2010). Preferably the source population habitat(s) for a large carnivore should be inviolate, wherein carnivores can subsist on natural prey and perform their ecological role. For Asiatic lions, such an area is a small National Park (259 km 2 ), that can at best accommodate 25 lions which are demographically not viable by themselves (Banerjee et al., 2010). For tiger reserves in India, a minimum population of 20 breeding females is considered to be viable (Gopal et al., 2007 Chapron et al., 2008). To achieve this, an area of 800𠄱,000 km 2 is required, and has been legally mandated to be made inviolate by incentivized voluntary relocation of human settlements from Tiger Reserves to delineate core areas (Gopal et al., 2007). A similar approach is required for Asiatic lions and an additional area of the Gir WLS needs to be legally demarcated and augmented to the existing National Park so as to cover a total of about 800𠄱,000 km 2 . Land ownership outside the PA is predominantly private and the Gujarat Forest Department has little control over changes in land-use patterns. Therefore, after securing a viable lion population within an inviolate space, protected areas under less stringent categories like conservation and community reserves that permit uses by local communities and safeguard their livelihoods [Wildlife (Protection) Act 1972 (2006 amendment)] should be used to conserve habitat patches within the larger human dominated landscape. Currently, radio-telemetry has shown that lions move across the landscape freely using certain land use categories and topographical features like drainage systems (Banerjee, 2012). However, the expansion of existing roads into heavy traffic highways, railways, and other linear infrastructure is likely to severely curtail such movement. Using lion locations from telemetry (ϩ,000), a habitat suitability map using an ecological niche factor analysis (Hirzel et al., 2002) was prepared and optimal connectivity between lion habitat patches modeled using PATHMATRIX (Basu, 2013 Jhala et al., 2016 Figure 2). These habitat corridors are the minimal requirements for lions to move between habitat patches and maintain the landscape scale metapopulation structure. Infrastructure that cuts through lion habitat patches and corridors needs to be made lion friendly and permeable using wildlife under- and over-passes (Jhala et al., 2016). The only legal provision available to regulate land use conversions in such lion habitats is by the provision of declaring Ecosensitive Zones under the Environment Protection Act (1986). Identified habitat patches and corridors (Figure 2) should be made part of the ecosensitive zone of the Gir PA. Such a declaration by the Government of Gujarat would enable authorities to reduce further losses of these areas to industry, mining and infrastructure while permitting uses that are conducive to lion conservation and local livelihoods. Currently, the Gujarat Forest Department is primarily responsible for lion conservation across the landscape, a responsibility that needs to be shared with various stakeholder agencies including roadways, railways, electricity, and civil administration. Such a multi-collaborative approach would ensure that development and conservation go hand-in-hand and are not always at loggerheads.

Food Habits and Foraging

Until early 1970s, Gir PA was dotted with about 300 Maldhari settlements (nesses) having over 40,000 livestock that formed the staple prey of lions (75% of their diet, Joslin, 1973), while wild ungulate numbers in the PA were few (5,600, Berwick, 1974). In 1975, when Gujarat was under the federal Government rule, about 190 Maldhari families along with their livestock were resettled outside Gir PA. In 1982 Gir experienced a major cyclone that uprooted ߢ.5 million large trees, resulting in the opening of the canopy and increased browse availability for ungulates (Dave and Jhala, 2011). Reduction in competition from livestock (Khan, 1993 Sharma, 1995) coupled with increased food availability by the cyclone and better law enforcement that checked poaching are believed to have resulted in the recovery of wild prey (Dave and Jhala, 2011). Regular monitoring of prey using line transect based distance sampling compared with data on prey estimates from Joslin (1973) and Berwick (1974) show that wild ungulates increased in their numbers till early 2000, and since then have reached stable densities (Jhala et al., 2016). Consequently, proportion of domestic livestock in lions' diet within the PA declined to 52% by the 1980's (Sinha, 1987) and further to 25% (Chellam, 1993 Meena et al., 2011 Banerjee et al., 2013) during the next three decades.

We investigated lion foraging ecology through direct continuous observations on radio-collared lions to record feeding events (Ϧ,000 h observation on 27 lions), and through scat analysis (n = 495). The Saurashtra landscape supports a large livestock population (ߦ.4 million, Junagadh Agricultural University, 2016). With majority of the people being vegetarian combined with the religious sentiment of Hinduism and Jainism, cattle are not consumed for meat. Several charitable cattle camps (locally known as Gaushalas and Panjrapoles) that house old and unproductive cattle are distributed across the landscape. Livestock carcasses are usually dumped at specific locations called haddakhodis outside villages and such Panjrapoles. Carnivores including wolves (Canis lupus pallipes) and striped hyenas (Hyena hyena hyena) within agro-pastoral landscapes rely predominantly on this assured food source (Jhala, 2002) across India. This factor has played a major role in promoting and sustaining the dispersal of lions outside of the PA. Lions are opportunistic feeders and rely both on predation and scavenging. Occurrences of food remains in scats are unable to distinguish between predation and scavenging, and if used alone can overestimate livestock-lion conflict. By using both direct observation on feeding events and scat analysis, Chakrabarti et al. (2016) was able to quantify contribution of dead livestock to lion diet. Chakrabarti et al. (2016) further developed models for estimating biomass consumption from prey occurrences in scats by conducting feeding experiments on lions, correcting previous diet estimates from lion scats that were fraught with considerable biases owing to the use of an incompatible model developed by Ackerman et al. (1984) for pumas (Puma concolor). Optimal foraging models developed by Chakrabarti et al. (2016) suggest that due to constraints of gut fill, passage time and carcass decomposition medium-sized prey like chital comprise of the principal prey for large carnivores, including lions, in tropical systems. Lion diet outside the PA was composed of 25% wild prey and 75% livestock (Banerjee, 2012). However, telemetry data demonstrated that among the total consumption of livestock, 35% was from actual predation while 65% from scavenging (Banerjee, 2012). Rarely were prized productive livestock killed by lions due to the husbandry practice of stall feeding and corralling such livestock during the night (Banerjee, 2012). Farmers were tolerant toward lions in their vicinity and property due to lions acting as effective predators for nilgai and wild pigs that caused substantial crop damage in this landscape.

Lion-Human Conflict and Coexistence

The Gir forests have been inhabited by the Maldharis for the past 200 years (Casimir, 2001). Maldharis have strong ethics and sentiments toward nature and natural resources. They are primarily vegetarian and their major livelihood is livestock husbandry for sale of dairy products. This religious and social background makes them tolerant toward lions, a powerful figure in their folklore and culture. Yet, Maldharis persecute lions to deter them from attacking their stock with sling shots, axes, and staffs. In the past, lions have also been poisoned on livestock kills. The Nawab of Junagadh recognized this threat early on and commenced a livestock depredation compensation scheme to the owners of livestock killed by lions. This scheme has been continued by the Gujarat Forest department and is revised regularly to keep pace with livestock market prices (Supplementary Material S5). Lions loath Maldharis and keep their distance when detected by them and their livestock. The water buffalo (Bubalus bubalis), that constitutes the majority of the livestock (78%) kept by Maldharis, herd together and defend themselves against lions (Banerjee et al., 2013). The husbandry practices of Maldharis are honed over years of experience to minimize losses to predation. Livestock are grazed in forests during the day and corralled in thorn bomas during the night. The herd leaves the boma much after sunrise with one to three herdsmen (depending on the size of the herd) and returns back around sundown. The grazing herd structure is composed of cattle and juvenile male buffalos at the front with prized buffalos in the middle, and herdsmen at leading and trailing ends. During lion attacks, the cattle and juvenile livestock (least expensive) scamper and run, becoming most vulnerable. Adult buffalos form a protective ring, often attacking lions in this formation under directions of the herdsmen, and rarely get killed (Banerjee et al., 2013). Dead livestock of Maldharis are dumped at specific sites and lions use this resource extensively. Radio-collared lions within the PA were observed to make regular excursions to these dump-sites near the nesses in search of free food (Jhala et al., 2016). Therefore, lions do benefit from Maldhari livestock through scavenging opportunities and occasional predation, only when strict law enforcement along with a fair livestock depredation compensation scheme control for lethal retaliation against them. The Maldharis that live in lion habitats benefit from getting free access to forest resources for themselves and their livestock. We found that Maldharis living within the Gir forests made 76 (SE 0.05) % more profits than livestock herders living outside the Gir forests (Banerjee et al., 2013). Thus, the relation between Maldharis and lions is far from harmonious coexistence, it is more of co-occurrence with benefits to both parties that are maintained by a delicate balance through cultural attitudes, strict law enforcement, fair compensation scheme for livestock kills, livelihood benefits to Maldharis and rare attacks on humans by lions. A total 190 lion attacks on humans have been recorded between 2007 and 2016 in the Gir landscape, of which a small proportion (n = 12, 4% 1.3 attacks/year) resulted in human fatalities. While attacks by leopards on humans in the same landscape were 383 between 2011 and 2016, out of which 41 were lethal (ߧ/year). Elephants (Elephas maximus) and tigers cause higher losses to human lives (408 and 34 human deaths/year respectively between 2013 and 2015) across India (answer to un-starred question no. 2581, The Lok Sabha, Government of India, 2017 accessible at Attacks on humans by lions were observed to increase during years of extreme droughts that caused large livestock populations to enter and graze within PAs (Saberwal et al., 1994). Data from telemetered lions show that lions were mostly non-hostile to humans (one in ten thousand encounters translated into an attack, Jhala et al., 2016). Attacks were mostly accidental: lions rarely stalked or targeted humans as prey, but usually attacked in self-defense or when spooked (Banerjee, 2012).

Livestock densities within a PA beyond a threshold were detrimental to native vegetation communities and wild ungulates (Dave and Jhala, 2011). Profuse growth of weeds and unpalatable vegetation were found to grow in the vicinity of ness sites (Dave, 2008). Lions, on the other hand can do well without livestock in their diet and will adjust their densities to natural levels based on the availability of wild ungulates (Schaller, 1972 Van Orsdol et al., 1985) which are reasonably high in Gir PA (63/km 2 Jhala et al., 2016). Therefore, creating additional inviolate space within the Gir PA by relocating the remaining nesses to increase the area under the National Park would not only benefit lions but the entire native biota of the region. At the least, Maldhari ness sites should be rotated every 4𠄵 years to allow native vegetation to recover from the heavy grazing and trampling effects of livestock (Dave and Jhala, 2011).

Banerjee (2012) interviewed 680 local residents in the landscape using structured interviews to gain an understanding on their attitudes toward lions. Besides the common factor of culture and religion that helped foster lion presence in the human dominated landscape, factors related to livelihood benefits differed from those that operate inside the PA. Pastoralists, on the contrary, were not tolerant toward lions because of the losses they incur from lion predation on their livestock and occasional attacks on them when they attempted to deter lion predation on their livestock (Figure 5). Analysis of last 5-year data on livestock kills by lions across the entire Saurashtra landscape (914 villages) suggests an increasing trend in the intensity of depredation (Figure 6). Livestock kills were compensated by the Government, and these helped ameliorate retribution. However, pastoral communities outside the Gir PA were not satisfied with the Government compensation scheme (Banerjee, 2012) since there were no free resources (like for Maldharis inside the PA) and there was a significant deficit between the market rate for livestock and compensation paid for lion predation (Jhala et al., 2018). Rarely do compensation schemes take into account the “lost opportunity cost” and therefore even when compensated at market rates, predation does take a toll on livelihoods (Banerjee et al., 2013).

Figure 5. Attitudes of local people (n = 680 respondents) from 254 villages in the landscape outside the Gir PA regarding the continued presence of lions in their neighborhood. The respondents were categorized based on their livelihoods. The category “others” primarily represent individuals associated with the tourism industry like hoteliers, safari-vehicle providers, etc.

Figure 6. Trend in the intensity of livestock predation (number of predation events/number of villages with predation) by lions in between 2012� within the Saurashtra landscape. The data on livestock kill by lions encompasses a total of 914 villages across the entire range of Asiatic lions.

Economic reasons were found to be the most significant factor shaping people's tolerance toward lions in the landscape. Communities making direct or indirect profit from lions were more tolerant toward them (Figure 5). The two important economic benefits from lions were: (a) their propensity to predate nilgai and wild pigs, both considered as agricultural pests. With no hunting allowed in India, these ungulates can achieve high densities and cause severe local economic losses to livelihoods, (b) presence of lions offered an opportunity for tourism and employment. The Gir PA has a tourism zone where wildlife enthusiasts can visit for a safari, which has encouraged tourist resorts and correlates to flourish in Western Gir. This has economically benefited the local communities residing in this region of Gir. However, not all tourists get to see lions, and the PA management has imposed several restrictions on limited number of vehicles, on-foot access, baiting of lions, etc. Such restrictions are difficult to enforce on private lands across the 13,000 km 2 of lion occupied Saurashtra landscape. Local communities avail this opportunity and conduct “lion shows” outside the PA (Singh, 2017b). Such shows primarily comprise of lions being attracted on private lands through subsidized food (baits/carcasses), while tourists pay the owners of these farmlands to watch lions in action. The tourists often pay exorbitant amounts for these shows as they are guaranteed sightings of lions and granted liberties with them (night photography, watching lions on foot and/or from close proximity) that can be dangerous for tourists as well as lions. However, the profits from such shows are not shared equitably and monopolized by few powerful members of the community. Though considered “illegal,” such lion shows are difficult to control and are a major source of lucrative and easy income for locals across the agro-pastoral landscape. Thus, in our assessment, lion-human coexistence in the human dominated landscape has been possible due to: (a) low lion density (about 2𠄳 lions per 100 km 2 ) (b) low levels of conflict, lions subsist by scavenging dead livestock, predate unproductive cattle (that are reasonably compensated), and rarely attack humans. Problem lions are immediately removed by management (c) economic benefits to local communities through removal of crop pests and revenue generation via lion tourism (d) high level of tolerance of local communities due to religious and cultural attitudes and (e) strict laws and their enforcement against killing of lions. Changes in any of these factors can disrupt the current coexistence. However, since lions continue to increase in density and occupancy across Saurashtra, it is a matter of time before they exceed social tolerance limits. There is a perceived shift in the values of local communities from those of tolerance and reverence toward direct economic gains (Banerjee, 2012). Although attacks on humans are rare, the psychological (L཮ and Röskaft, 2004) and socio-economic consequences of these attacks can be dire for future lion-human coexistence in Saurashtra.

Also, recently, the wildlife authorities have seriously implemented measures to curb “lion shows” by local communities. This may have serious consequences on continued lion persistence in the human dominated landscape, if indeed this action manages to stop such shows. Communities that cannot have direct profits from having lions in their backyards may not be willing to have them there anymore.

Majority of the people in the agro-pastoral landscape of Saurashtra have a positive attitude toward lions (Banerjee, 2012 Meena et al., 2014). This is vital, but a positive attitude by the majority does not necessarily translate into tolerant coexistence, since it is the behavior of the few but resentful people that ultimately determines the dynamics of human-lion interface (Kansky and Knight, 2014). Such behavior is largely determined by a combination of factors relating to their personal situation and experiences, psychological factors and value judgement (Barr, 2003). Understanding biological and social carrying capacities (threshold for human tolerance) for lions thus becomes important in managing coexistence in this multiple-use landscape of Saurashtra. For example, ranches adjacent to Kenya's Tsavo East National Park, lose 3% of their herd's total economic value to lions nonetheless, the ranchers are prepared to tolerate a population of 縦 adult lions whose diet consists of 6% livestock, costing the ranches US$290/lion/year (Patterson et al., 2004). We suggest that lion density outside the Gir PA should be maintained below social carrying capacity and problem lions should be removed immediately from the vicinity of the people. Guidelines for such removals can be adopted from the Standard Operating Procedures developed for tigers and leopards in India (National Tiger Conservation Authority, 2013), keeping in mind the social dynamics of lions (Whitman et al., 2004). Thus, a futuristic and multifaceted policy is required to permit this delicate balance of human-lion coexistence to continue.

A Second Home for Lions and the Mist of Conservation Politics

A single population of an endangered species is susceptible to extinction events caused by environmental and demographic stochasticity (Soulé, 1987). The 1994 outbreak of canine distemper virus (CDV) in the Serengeti killed an estimated 33% of the lion population (Roelke-Parker et al., 1996). An epidemic of such magnitude in Gir could potentially put the Asiatic lion at a high risk of extinction. Gir lions have tested positive for CDV, feline parvovirus, feline herpesvirus, feline immunodeficiency virus and peste des petits ruminants virus (Sabapara, 2002 Ramanathan et al., 2007 Balamurugan et al., 2012). Lions move regularly between habitat patches in the landscape and share space with feral dogs, cats, and other carnivores, creating a condition for the spread of epidemics. A recent infection of canine distemper virus killed a minimum of 28 lions in 2018 as per official records in the eastern part of the PA. However, the actual death toll could be of epidemic proportions, but remains unknown, since many carcasses remain undetected in the wild and investigations were limited only to park authorities.

The threat of extinction due to disease and natural calamities to this single population of lions was recognized early on by the Executive Committee of the Indian Board of Wildlife during a meeting held in Gir in 1956. The first attempt to establish a second population in Chandraprabha in the state of Uttar Pradesh was undertaken in 1957 (Negi, 1969). Though these reintroduced lions initially bred and increased to 11 individuals from the founding population of five, they were subsequently poached out by 1965 (Negi, 1969). After initiation of modern scientific studies on Asiatic lions, Joslin (1985) and Sale (1986) emphasized the need for establishing a second population away from Gir. This was followed by a population-habitat viability analysis workshop in 1993, wherein all stakeholders, including the Government of Gujarat agreed to the need of establishing a second lion population as an insurance against extinction (Ashraf et al., 1995). The Wildlife Institute of India was mandated with the task of identifying a site for establishing this insurance population. From the three potential sites surveyed (Sitamata, Darrah-Jawaharsagar, and Kuno) within the recent historical range of the lion, the area of Kuno Wildlife Sanctuary (345 km 2 ) in the central Indian state of Madhya Pradesh was found most suitable since it was located within an intact forested landscape of about 3,300 km 2 (Chellam et al., 1995).

Substantial efforts were made by both the Government of India and Madhya Pradesh Forest Department in preparing Kuno for lion reintroduction (Johnsingh et al., 2007 Khudsar et al., 2008). Currently Kuno has been declared as an inviolate National Park (700 km 2 ) after the resettlement of 24 villages (1,547 families). A financial investment of about Rs 15 crores (US$ ca. 3.2 million) was done by the Government of India until 2005 for resettlement and management of Kuno (Johnsingh et al., 2006) and an equal amount invested by the Government of Madhya Pradesh. Subsequently, a buffer area of 1,280 km 2 has been added to the Kuno National Park as Kuno Wildlife Division. Better protection, habitat management, and relocation of human settlement along with majority of their livestock, resulted in a substantial recovery of the wild ungulate population. The chital population have exponentially increased from a density of 5 to 68/km 2 within the past 10 years (Banerjee, 2005 Bipin et al., 2013).

Gujarat monopolized Gir lions after they were stripped off their status as India's National Animal in 1973. Lions were promoted as a Gujarat State icon which soon became engrained as a symbol of the pride of the people of Gujarat (Rangarajan, 2001). Indeed, it was due to the efforts of the people of Gujarat that lions have shown an extraordinary recovery for any large carnivore. The local media exemplified and promoted this monopoly (Rahmani, 2013) which was subsequently used as an instrument of political and bureaucratic gain (Dutta, 2019). This new found exclusive ownership of the lions by Gujarat State and its bearing on the public psyche resulted in the Gujarat Government's reluctance to provide a founder stock of wild lions to the State of Madhya Pradesh (Kuno). The Gujarat Forest Department, which is the technical arm of the State Government in matters of wildlife, posed trivial arguments against reintroduction of lions in Kuno (Singh, 2007). However, a landmark judgement was passed by the Supreme Court of India in 2013 [IA No.100 in W.P (C) No.337/1995, accessible at] which directed the Governments of India, Gujarat and Madhya Pradesh to reintroduce lions in Kuno despite contrary arguments of Gujarat. Although this landmark verdict by the apex court was primarily directed toward lion reintroduction, it recognized conservation as an integral part of civilized development and beckoned for applying the “species' best interest standard” for conservation of lions and other endangered species. This judgement strongly places the responsibility on the national and state governments, together with the citizens, to view development through an eco-centric approach and not just with an anthropocentric perspective.

As per the assessment of the committee for lion reintroduction appointed by the Supreme Court through their court order, Kuno National Park can currently hold about 40 lions. The larger forested landscape of about 3,000 km 2 around Kuno, has the potential to support a viable lion population for the long-term. The Kuno lion reintroduction action plan (Ministry of Environment Forests and Climate Change [MoEFCC], 2016) is in consonance with the IUCN/SSC reintroduction group guidelines (IUCN/SS, 2013) and provides operational guidelines to the wildlife managers of Gujarat and Madhya Pradesh States to implement the reintroduction and subsequent management of the lion population. Despite the direction of the Supreme Court in 2013 and an action plan (Ministry of Environment Forests and Climate Change [MoEFCC], 2016) with a clear vision, the reintroduction program is facing a socio-political deadlock for the past 6 years. The program implementation is still being debated between the Ministry of Environment, Forests and Climate Change, Government of India Gujarat Forest Department and the Madhya Pradesh Forest Department.

While lion reintroduction in Kuno was being debated, the Gujarat Forest Department mandated the Wildlife Institute of India to evaluate the potential of Barda Wildlife Sanctuary (Figure 2) as another reintroduction site for lions within Saurashtra, Gujarat. Barda and its adjacent Alech hills (Barda landscape) had lions until the late nineteenth century after which they were locally extirpated (Divyabhanusinh, 2005). Subsequent conversion of forest and grazing lands to agriculture separated Barda from Gir (纀 km). This less permeable habitat matrix along with the initial policy of the Gujarat Forest Department to capture dispersing lions and relocate them back to Gir, prevented recolonization (Ranjitsinh, 2016). The assessment of Barda (Jhala et al., 2014b) suggested that the landscape (410 km 2 comprising of 190 km 2 of Barda WLS, Alech hills and coastal forest patches) could sustain about 25 lions after creating an inviolate area of about 100 km 2 within the Barda WLS, restocking prey, enhancing protection, and restoring habitats. Currently the sanctuary is inhabited by about 1,325 families of Maldharis in 62 nesses and 98% of them are willing to resettle outside Barda (Jhala et al., 2014b). The costs of incentivised, voluntary relocation (Narain et al., 2005) would be close to Rs 200 crore (US$ ca. 28 million). Current wild prey density in Barda is very low and inadequate for sustaining even a single lion pride, but livestock and scavenging opportunities abound. Resettlement of human habitation and building up wild prey is likely to take several years. Establishing a lion population in Barda landscape would be beneficial for lion conservation as well as help conserve the native flora and fauna of this region which is threatened by intense human exploitation. However, a lion population in Barda cannot be considered as a security from catastrophic events like disease epidemics in the Gir landscape due to the geographic proximity of both areas and continuous presence of feral dogs, cats, and livestock in the intervening habitat. Barda, therefore cannot be an alternative solution to lion reintroduction in Kuno (Jhala et al., 2014b). Efforts of the Gujarat Forest Department in conserving a representative lion gene pool through conservation breeding programs (Meena and Kumar, 2012) are important initiatives to pre-empt a catastrophic extinction event within the Gir landscape. However, carnivores bred in captivity over several generation are usually unfit for reintroductions into the wild (Jule et al., 2008). Therefore, we submit that the “species best interest” strategy for securing Asiatic lions in the long-term would be to establish as many free ranging populations as possible within the historic range of the Asiatic lions. Founders for such populations should be sourced from wild Gir lions to capture their existing genetic diversity and subsequently managed as a metapopulation with the Gir lion population (Ministry of Environment Forests and Climate Change [MoEFCC], 2016).

Management Interventions

The contribution of wildlife managers has been the most vital ingredient for the conservation of Asiatic lions. Wildlife management in India is done by the respective State Forest Departments. Their primary role is to manage the PAs in terms of administration, law-enforcement, wildlife conflict mitigation, habitat management, and management of wildlife tourism. Other aspects include community participation through incentives by sharing park revenues in the form �o-development projects,” sensitization of local communities through awareness and education camps, treatment and rescue of wildlife. In this section we succinctly portray the management arena for Gir PA and discuss their strengths and weaknesses under the larger gambit of lion ecology and conservation, tethering with our previous sections.

The total strength of the wildlife department of Gir PA in 2012 was about 688 (Meena and Kumar, 2012). Modern amenities in the form wireless service, good road network, 4- and 2-wheel drive vehicles and arms are available and used by the wildlife authorities. Regular patrolling on-foot and by vehicles has controlled poaching within PAs. However, snaring and electrocution continue to be a major concern for wildlife authorities in the larger landscape. Eight lions were poached in 2007 for meeting the illegal demand of lion bone trade (Singh, 2017b). The wildlife authorities successfully nabbed the poachers and got them convicted in the court of law under the Wildlife (Protection) Act 1972 setting an example that has deterred poaching of lions to a great extent.

The wildlife department has developed competence in veterinary facilities for treating animals in distress at eight facilities across the Gir landscape. Also, lions in conflict or individuals straying into human-habitation too often are captured and rehabilitated. Perception of the public and media to an ailing/injured lion forces wildlife authorities to capture and treat such animals. Within the past decade, medical interventions for treating even minor injuries and ailments in lions have become the norm. As discussed earlier, such actions can tamper with the process of natural selection, and should be undertaken judiciously. The reluctance of wildlife authorities in seeking expert advice on dealing with dangerous situations like CDV outbreaks can have disastrous impacts on the long-term survival of this single population of Asiatic lions.

Gir being a dry deciduous forest tract, water is a major factor that limits the abundance and distribution of animals. Wildlife authorities manage the availability of water in the landscape through provisioning by regular maintenance and filling of artificial waterholes. Weed and invasive species removal is done across the PA, and an area of over 270 km 2 was prescribed for treatment (Meena and Kumar, 2012). Gir PA is prone to fires and the regular management of ߡ,900 km of fire-lines is done annually to contain accidental fires (Meena and Kumar, 2012).

Lion centric tourism within Gir PA is an important source of revenue for the Gujarat Forest Department and about 0.12 million tourists visited Gir PA annually in between 1995 and 2010 (Meena and Kumar, 2012). The number of tourists has substantially gone up in the recent years (0.533 million in 2015 and 0.522 million in 2016). The Forest Department permits tourism in a part of the western Gir WLS by allowing tourist vehicles (accompanied by trained nature guides) to ply over forest roads in eight pre-fixed routes (ranging from 22 to 45 km) after obtaining online permits. In order to reduce tourism pressure inside the Gir PA and to provide tourists with guaranteed opportunities of viewing lions and other wildlife, two safari parks (each of about 4 km 2 , enclosed by chain-link fences) at Devalia (western Gir) and Ambardi (eastern Gir) have been created that house semi-free ranging wildlife including lions. All these activities generate a substantial amount of revenue. For example, in 2016 revenue generated from gate fees was Rs. 102.5 million (~ 1.5 million US$, However, as mentioned earlier, practice of luring lions with artificial subsidies such as carcasses to maximize lion viewing by tourists (Gogoi, 2015) should be discontinued.

With the increase in the extent and magnitude of lion-tourism, hospitality industry has flourished along the periphery of the Gir PA. Within a six-km radius of the tourism circuit of the PA, there are nearly 100 resorts, hotels and guesthouses catering to the needs of tourists. Many such facilities have been totally or partially shut down following a suo motu Gujarat High Court order against illegal and haphazard construction around Gir PA. Meanwhile, the Government of Gujarat has submitted an eco-tourism policy to the High Court proposing to: (i) decline new licenses for hotels and resorts within 1 km of the Gir PA and (ii) levy a new tax known as �o development fee” for conversion of agricultural land to commercial tourism purposes.

With the objective to sensitize the younger generation toward wildlife conservation, nature camps are conducted by the Gujarat Forest Department since 1976. Students from local schools and colleges camp at five designated sites in the PA and are taken on field excursions with trained nature interpreters, interact with wildlife managers through illustrated talks and field demonstrations, and are shown wildlife documentaries (Meena and Kumar, 2012). Additionally, eco-clubs have been established in about 300 schools within the Gir landscape with the aim of spreading awareness related to nature conservation in their localities. The Government tourism facility at Gir has a good interactive interpretation center that is popular amongst the visitors of the park.

In order to garner public support for lion conservation, an �o-development” scheme was initiated in Gir along with six other PAs in India under the India Eco-development Project funded by World Bank's Global Environment Facility. A total of 193 villages have so far been covered in the landscape under this scheme (Singh, 2017a). Under this scheme, repair of village roads, support for self-employment, construction of structures for harvesting water and preventing soil erosion, facilities for education, drinking water, sanitation and improvement of houses are provided and linked to wildlife conservation. Parapets were constructed for about 25,000 open wells that otherwise act as death traps to lions and other wildlife. Members of the local community that have demonstrated wildlife skills are designated as vanya prani mitra (friends of wildlife) and paid a nominal remuneration for assisting with wildlife management activities of the park managers (including information on poaching, fire management, and wildlife conflict resolution). Besides the monetary remuneration, the enhanced social status of the vanya prani mitra is an incentive for community members to strive to become “wildlife friends.” An average amount of Rs. 8.36 million (񾄢,139 US$) was spent by the Forest Department for accomplishing various activities under this scheme between 2006 and 2010.

Another major activity undertaken by wildlife managers is the 5 yearly periodic population enumeration of lions. This exercise is commendable in its extent, effort, and regularity. However, as previously mentioned, the archaic approach of population census through total counts needs to be replaced with modern scientific approaches of animal abundance estimation that explicitly address the issue of detectability and double counts. Such a scientific assessment by an independent agency would also preclude the potential of distorting numbers to create political populations (Darimont et al., 2018) as was done earlier with tiger populations in India (Karanth et al., 2003).

LEAP Prides

Students will be taught about flourishing through the lens of science, philosophy, and their own experience. Through exposure to diverse scientific theories and research on the skills associated with flourishing, to philosophical and religious writings on human happiness and wellbeing, and to various reflective exercises and contemplative practices (e.g., mindfulness and compassion meditation), students will have the opportunity to develop conceptual knowledge, experiential insight, and practical skills related to flourishing.

The Art and Science of Human Flourishing

Human Development and Aging

This pride provides an overview of American government and politics at the national level. In this course, we will first learn about the political institutions that comprise our political system. We will discuss their origins and the ways in which they structure our society. We then will transition into learning about the behavior of both the American mass public and political elites. We will discuss how everyday Americans make political decisions and how their elected representatives respond to the public’s political sentiments.

American Politics: Principles, Processes, and Powers

Rhetoric and Composition

This pride introduces you to current concepts in the maintenance of animal health and the application of scientific information to health-related decision making. You will have the opportunity to apply biomedical sciences to disease prevention programs in companion animals, food animals, and horses. The course will emphasize the biology of infectious diseases, immune response, and toxicology. You will also have opportunities to investigate the wide variety of careers in animal health and the biomedical sciences.

Animal Health and Biomedical Sciences

Effective Speech

Arguing About the World: this pride is reserved for Paterno Fellows aspirants who wish to become better, more capable participants in arguments about some of the most challenging and intractable social problems facing humanity. SOC 5 will examine current social problems such as economic, racial, and gender inequalities social deviance and crime and population, environmental, energy, and health problems. CAS 215 provides an in-depth examination of argumentation in both public and private contexts. The course requires students to investigate the process of researching sound evidence, constructing legitimate argumentative claims, and participating in live debates. Fundamental to this endeavor is a strong attention to research, ethics, and strategy.

This pride is only open to students aspiring to the Paterno Fellows Program. To enroll, contact Barb Edwards at [email protected]

Social Problems


Democracy depends on an informed electorate who engage in critical thinking so as to hold in check potential abuses of power. This pride provides students with the ability to critically assess and create effective discourse.


Effective Speech

In this pride, students will learn the myths of ancient Greece and Rome. These stories include the Olympian gods and goddesses (familiar to many from the Percy Jackson series), and the heroes of epics like Homer’s Iliad and Odyssey. Students will explore both the most famous versions of these myths in the canonical texts of classical literature and interpretations of myth in art, music, theater, and literature from the classical era to the modern day. This course will be especially beneficial for those students interested in art history, cultural anthropology, gender theory, ancient civilizations, and world religions.

Classical Mythology

Introduction to Art

Offered only to incoming students, Aurora eases the college transition through fun and accessible virtual expeditions. As part of the Sirius Aurora experience, tour the United States from your computer, explore natural spaces close to home, and connect with your group virtually — all students of all abilities welcome. Participants gain not only skills from trained upperclassmen, but insight into college life in a relaxed environment. The classroom component introduces students to health and wellness topics such as: eating well on campus, group dynamics, time management, and mental and emotional wellness. Students earn all 3 GHW credits required to graduate (KINES 89). No outdoor experience required! For more information call (814) 865-3890 or visit

There are four sections of the Aurora Pride. Lion and Nittany are paired with CAS 100A (Effective Speech). Roar and State are paired with ENGL 15 (Rhetoric and Composition). Students should register for one section.

AURORA Outdoor Orientation Programs - Sirius

Effective Speech

Offered only to incoming students, Aurora eases the college transition through fun and accessible virtual expeditions. As part of the Sirius Aurora experience, t our the United States from your computer, explore natural spaces close to home, and connect with your group virtually — all students of all abilities welcome. Participants gain not only skills from trained upperclassmen, but insight into college life in a relaxed environment. The classroom component introduces students to health and wellness topics such as: eating well on campus, group dynamics, time management, and mental and emotional wellness. Students earn all 3 GHW credits required to graduate (KINES 89). No outdoor experience required! For more information call (814) 865-3890 or visit

There are four sections of the Aurora Pride. Lion and Nittany are paired with CAS 100A (Effective Speech). Roar and State are paired with ENGL 15 (Rhetoric and Composition). Students should register for one section.

Aurora Outdoor Orientation Program - Sirius

Rhetoric and Composition

This pride explores health issues in an integrated way by taking into account the many different factors—biological, behavioral, social, cultural, and environmental—that influence health throughout the life span. Health will be emphasized as a state of physical, mental, and social well-being and not just the absence of disease. Students will learn to identify the many factors that contribute to differences in health and health risk for individuals and groups. You will apply health principles and research findings to health promotion, disease prevention strategies, and public health policies.

Biobehavioral Health

Rhetoric & Composition

Through this course, students will gain a well-rounded level of competency in the use of spreadsheet software as a tool. Students will develop problem-solving strategies while gaining insight on the tactical use of spreadsheets. Concepts are contextualized in a broader discussion of information systems management including data security, ethical issues, social media, distributed (cloud) services, and emerging trends. This is a required course for all majors in the Smeal College of Business.

There are five sections of the Business Problem Solving Pride: (1) Blue, (2) Lion, (3) Nittany, (4) Roar, and (5) State. The ENGL 15 course for the State section will meet in-person.

This pride is only open to students enrolled in the Smeal College of Business or the Division of Undergraduate Studies.

Business Problem Solving

Rhetoric and Composition

This pride is designed to both give students an introduction into their study in the Smeal College of Business through PSU 6 (1 credit) and BA 297 (1 credit) and to engage students in contemporary issues around race, ethnicity, and culture through SOC 119 (4 credits). Students will be able to understand the extent and pervasiveness of racial, ethnic , and cultural inequality and discrimination in the U.S. and explain how unequal systems impact different ancestry groups both in the U.S. and around the world. Students will be able to compare historical causes and consequences of sociological processes such as immigration, assimilation , and multiculturalism, and they will develop new ways of understanding how these processes are expressed in popular culture (e.g., art, music, literature) and intergroup dynamics. Students will be provided with intellectual tools to formulate more thoughtful questions and responses when they engage others in everyday discussions about the many racial, ethnic , and cultural factors and forces that pull people and groups together and push them apart. Students will also explore pathways to allow them to discover new ways to understand their own racial and ethnic place in the world and the history of their own families.

This pride is only open to students enrolled in the Smeal College of Business or the Division of Undergraduate Studies.

Race, Ethnicity, and Culture

Business Seminar

This course introduces basic statistical concepts and models within the framework of business problems and applications. Students learn about the usefulness of business statistics to decision making, how to perform basic statistical and analytical procedures, and how to interpret, critically evaluate, and analyze data. SCM 200 or STAT 200 is a n entrance to major requirement for Smeal students . SCM 200 is a 4-credit course. Students enrolling in this pride will be enrolling in 7 academic credits.

There are two sections of the Business Statistics Pride: (1) Lion and (2) Nittany. Students should enroll in one section.

Introduction to Statistics for Business

Effective Speech

This lab-based pride is designed to introduce you to the process and practice of chemical research. The research will focus on the knowledge and application of two core concepts central to understanding compound’s structure-reactivity relationship: polarity and pH. You will be required to think about structures of organic compounds as they relate to certain periodic trends and the connectiveness to these core concepts. You will be shown how these studies tie into the research projects of Dr. Xin Zhang, a chemical biologist at Penn State.

This pride is only available to students in the Eberly College of Science and the Division of Undergraduate Studies

Research Experience in Chemistry

Effective Speech

Many of the most pressing issues facing the world, such as climate change, racial inequality, food insecurity, and other social injustices , lie at the intersection of society, economy, and environment. These problems exist at multiple scales from the hyperlocal to the global. Successfully addressing these challenges requires residents, communities, governments, businesses, and other organizations and institutions to work collaboratively on solutions using skills in critical thinking, community leadership, research, problem-solving, and communication. This pride will introduce students to the basic concepts, skills, and practices for community development and engagement that are necessary for addressing such issues.

Community Development Concepts and Practice

Effective Speech

This pride provides an introduction to computer systems literacy. The history, architecture , and operation of computing systems and underlying computing theory are covered. The intent of this course is to ensure that students with diverse backgrounds can gain the information technology fundamental skills and understanding to succeed with subsequent in-depth courses in the Cybersecurity Analytics and Operations curriculum. At the same time , the general nature of the introduction may make it useful for other programs that involve education in concepts and skills relating to information and computing systems.

There are two sections of the Cybersecurity Pride: (1) Lion (in person) and (2) Nittany (virtual). Students should enroll in one section.


Rhetoric and Composition

This pride provides an introduction to computer systems literacy. The history, architecture , and operation of computing systems and underlying computing theory are covered. The intent of this course is to ensure that students with diverse backgrounds can gain the information technology fundamental skills and understanding to succeed with subsequent in-depth courses in the Cybersecurity Analytics and Operations curriculum. At the same time , the general nature of the introduction may make it useful for other programs that involve education in concepts and skills relating to information and computing systems.

There are two sections of the Cybersecurity Pride: (1) Lion (in person) and (2) Nittany (virtual). Students should enroll in one section.


Effective Speech

Th is pride introduces students to the basic historical contexts within which key democratic events arise engage s them in close readings of speeches, tracts, and polemical writing and develop s skills in critical thinking and writing. Students will attend to a broad spectrum of voices, including those of historically under-represented peoples as well as canonical figures.

Landmark Speeches on Democracy and Dissent

Effective Speech

The EMS First-Year Seminar is designed to encourage students to begin the important process of 'thinking for a living.' The primary focus of the seminar is to promote critical reading and thinking skills, and to help students develop effective written and oral arguments. Students learn the importance of reading, sharing ideas and collaborating, through independent and group research, discussion and debate. We want our students to think about some of the major issues facing the world today, the role that science and technology have played in defining and addressing these issues, the way in which present-day thinking has been shaped by the past, and the development of scientific thought. The Seminar's content focuses on communication skills, but these are addressed within the context of issues relevant to the disciplines represented in EMS. As such, the discussions range across topics such as the Earth and its resources scientific and technical aspects of global habitability development of the advanced materials necessary for sustaining and advancing civilization in the 21st century and the social, economic, and political factors that shape and constrain society's view of the Earth system. Students undertake three to five major writing projects throughout the semester, as well as several smaller one to two paragraph written assignments. Grades are determined from their performance on the written papers, oral presentations, and in-class participation. The Seminar is a required course for all EMS first-year students at the University Park campus and, together with English 015 and either English 202 or Speech Communications 100, will satisfy the Writing and Speaking requirements of General Education.

Sociological Perspectives on Sex and Sexuality

Sociologists representing all three major theoretical perspectives study the role sexuality plays in social life today. Scholars recognize that sexuality continues to be an important and defining social location and that the manner in which sexuality is constructed has a significant effect on perceptions, interactions, and outcomes.

Structural Functionalism

When it comes to sexuality, functionalists stress the importance of regulating sexual behavior to ensure marital cohesion and family stability. Since functionalists identify the family unit as the most integral component in society, they maintain a strict focus on it at all times and argue in favor of social arrangements that promote and ensure family preservation.

Functionalists such as Talcott Parsons (1955) have long argued that the regulation of sexual activity is an important function of the family. Social norms surrounding family life have, traditionally, encouraged sexual activity within the family unit (marriage) and have discouraged activity outside of it (premarital and extramarital sex). From a functionalist point of view, the purpose of encouraging sexual activity in the confines of marriage is to intensify the bond between spouses and to ensure that procreation occurs within a stable, legally recognized relationship. This structure gives offspring the best possible chance for appropriate socialization and the provision of basic resources.

From a functionalist standpoint, homosexuality cannot be promoted on a large-scale as an acceptable substitute for heterosexuality. If this occurred, procreation would eventually cease. Thus, homosexuality, if occurring predominantly within the population, is dysfunctional to society. This criticism does not take into account the increasing legal acceptance of same-sex marriage, or the rise in gay and lesbian couples who choose to bear and raise children through a variety of available resources.

Conflict Theory

From a conflict theory perspective, sexuality is another area in which power differentials are present and where dominant groups actively work to promote their worldview as well as their economic interests. Recently, we have seen the debate over the legalization of gay marriage intensify nationwide.

For conflict theorists, there are two key dimensions to the debate over same-sex marriage—one ideological and the other economic. Dominant groups (in this instance, heterosexuals) wish for their worldview—which embraces traditional marriage and the nuclear family—to win out over what they see as the intrusion of a secular, individually driven worldview. On the other hand, many gay and lesbian activists argue that legal marriage is a fundamental right that cannot be denied based on sexual orientation and that, historically, there already exists a precedent for changes to marriage laws: the 1960s legalization of formerly forbidden interracial marriages is one example.

From an economic perspective, activists in favor of same-sex marriage point out that legal marriage brings with it certain entitlements, many of which are financial in nature, like Social Security benefits and medical insurance (Solmonese 2008). Denial of these benefits to gay couples is wrong, they argue. Conflict theory suggests that as long as heterosexuals and homosexuals struggle over these social and financial resources, there will be some degree of conflict.

Symbolic Interactionism

Interactionists focus on the meanings associated with sexuality and with sexual orientation. Since femininity is devalued in U.S. society, those who adopt such traits are subject to ridicule this is especially true for boys or men. Just as masculinity is the symbolic norm, so too has heterosexuality come to signify normalcy. Prior to 1973, the American Psychological Association (APA) defined homosexuality as an abnormal or deviant disorder. Interactionist labeling theory recognizes the impact this has made. Before 1973, the APA was powerful in shaping social attitudes toward homosexuality by defining it as pathological. Today, the APA cites no association between sexual orientation and psychopathology and sees homosexuality as a normal aspect of human sexuality (APA 2008).

Interactionists are also interested in how discussions of homosexuals often focus almost exclusively on the sex lives of gays and lesbians homosexuals, especially men, may be assumed to be hypersexual and, in some cases, deviant. Interactionism might also focus on the slurs used to describe homosexuals. Labels such as “queen” and “fag” are often used to demean homosexual men by feminizing them. This subsequently affects how homosexuals perceive themselves. Recall Cooley’s “looking-glass self,” which suggests that self develops as a result of our interpretation and evaluation of the responses of others (Cooley 1902). Constant exposure to derogatory labels, jokes, and pervasive homophobia would lead to a negative self-image, or worse, self-hate. The CDC reports that homosexual youths who experience high levels of social rejection are six times more likely to have high levels of depression and eight times more likely to have attempted suicide (CDC 2011).

Queer Theory

Queer Theory is an interdisciplinary approach to sexuality studies that identifies Western society’s rigid splitting of gender into male and female roles and questions the manner in which we have been taught to think about sexual orientation. According to Jagose (1996), Queer [Theory] focuses on mismatches between anatomical sex, gender identity, and sexual orientation, not just division into male/female or homosexual/hetereosexual. By calling their discipline “queer,” scholars reject the effects of labeling instead, they embraced the word “queer” and reclaimed it for their own purposes. The perspective highlights the need for a more flexible and fluid conceptualization of sexuality—one that allows for change, negotiation, and freedom. The current schema used to classify individuals as either “heterosexual” or “homosexual” pits one orientation against the other. This mirrors other oppressive schemas in our culture, especially those surrounding gender and race (black versus white, male versus female).

Queer theorist Eve Kosofsky Sedgwick argued against U.S. society’s monolithic definition of sexuality and its reduction to a single factor: the sex of someone’s desired partner. Sedgwick identified dozens of other ways in which people’s sexualities were different, such as:

  • Even identical genital acts mean very different things to different people.
  • Sexuality makes up a large share of the self-perceived identity of some people, a small share of others’.
  • Some people spend a lot of time thinking about sex, others little.
  • Some people like to have a lot of sex, others little or none.
  • Many people have their richest mental/emotional involvement with sexual acts that they don’t do, or don’t even want to do.
  • Some people like spontaneous sexual scenes, others like highly scripted ones, others like spontaneous-sounding ones that are nonetheless totally predictable.
  • Some people, homo-, hetero-, and bisexual, experience their sexuality as deeply embedded in a matrix of gender meanings and gender differentials. Others of each sexuality do not (Sedgwick 1990).

Thus, theorists utilizing queer theory strive to question the ways society perceives and experiences sex, gender, and sexuality, opening the door to new scholarly understanding.

Throughout this chapter we have examined the complexities of gender, sex, and sexuality. Differentiating between sex, gender, and sexual orientation is an important first step to a deeper understanding and critical analysis of these issues. Understanding the sociology of sex, gender, and sexuality will help to build awareness of the inequalities experienced by subordinate categories such as women, homosexuals, and transgender individuals.


When studying sex and sexuality, sociologists focus their attention on sexual attitudes and practices, not on physiology or anatomy. Norms regarding gender and sexuality vary across cultures. In general, the United States tends to be fairly conservative in its sexual attitudes. As a result, homosexuals continue to face opposition and discrimination in most major social institutions.

Short Answer

  1. Identify three examples of how U.S. society is heteronormative.
  2. Consider the types of derogatory labeling that sociologists study and explain how these might apply to discrimination on the basis of sexual orientation.


Gene flow and immigration: genetic diversity and population structure of lions (Panthera leo) in Hwange National Park, Zimbabwe

The genetic diversity and population structure of a population of African lions in Hwange National Park, Zimbabwe, was studied using 17 microsatellite loci. Spatial genetic analysis using Bayesian methods suggested a weak genetic structure within the population and high levels of gene flow across the study area. We were able to identify a few individuals with aberrant or admixed ancestry, which we interpreted as either immigrants or as descendants thereof. This, together with relatively high genetic diversity, suggests that immigrants from beyond the study area have influenced the genetic structure within the park. We suggest that the levels of genetic diversity and the observed weak structure are indicative of the large and viable Okavango-Hwange population of which our study population is a part. Genetic patterns can also be attributed to still existing high levels of habitat connectivity between protected areas. Given expected increases in human populations and anthropogenic impacts, efforts to identify and maintain existing movement corridors between regional lion populations will be important in retaining the high genetic diversity status of this population. Our results show that understanding existing levels of genetic diversity and genetic connectivity has implications, not only for this lion population, but also for managing large wild populations of carnivores.

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4. Discussion

Our analyses demonstrate the epidemiological importance of variation in group size in social species. As the variance of the group size distribution increases, the model demonstrates that the epidemic threshold decreases and that both the mean and variance of small outbreak sizes should increase. Above the epidemic threshold, that is, for diseases capable of causing large epidemics, the effect of group size variability depends on the transmissibility of the disease. For mildly contagious diseases, more variable group sizes promote larger epidemics, whereas for more highly contagious diseases, the effect reverses and group size variability inhibits epidemics.

These findings have important implications for disease-control strategies, including vaccination, quarantine and culling, which often target social groups that are likely to decrease the epidemic threshold. Prior studies have highlighted the importance of targeting groups with high numbers of interacting neighbours or who occupy a central location in the network [41]. Our study suggests that factoring the group size distribution into epidemiological assessments can improve both understanding of disease dynamics and efforts to prevent and mitigate outbreaks. For example, vaccination efforts should perhaps be targeted to reduce the variation in the number of susceptible individuals per group (perhaps by vaccinating a fraction of individuals in large groups) rather than simply reducing the total number of susceptible individuals population-wide or immunizing all individuals in a few groups. For conservation biologists, this work also suggests that assessments of disease risk in endangered animal populations should carefully consider variation in both contact patterns and group sizes.

The effect of the hierarchical structure of a population on the transmission of infectious diseases can similarly apply to the transmission of information. In human populations, the transmission of rumours and cultural traits between families or communities is thus probably influenced by the variance in social group size. One can also predict that the transmission of computer viruses between small networks of interconnected computers (e.g. intranets) is more likely to become epidemic when the size of subnetworks is more variable.

We have introduced EEGS as a simple, intuitive metric for quantifying the epidemiological impact of group size variation and the epidemiological vulnerability of a population. As long as the network structure of the population is sufficiently random (lacks clustering or other local structure), EEGS can be estimated using only the mean and variance of the group size distribution. EEGS is independent of degree distribution and indicates which homogeneous network with same-sized groups has comparable epidemiological properties. The original network and the EEGS homogeneous network share the same epidemic threshold and thus are vulnerable to the same suite of pathogens. However, the two networks will not necessarily experience outbreaks of similar magnitude. Analogous metrics can also be computed by equating the mean outbreak size (as illustrated here with the Serengeti lion network) or the probability of an epidemic instead of the epidemic threshold. However, such metrics can generally only be computed numerically or using Monte Carlo simulations.

The model makes several simplifying assumptions. First, it assumes that between-group transmission is an increasing function of the size of the groups. This happens, for example, when all the individuals of two interacting groups come into contact, or when movements of potentially infected individuals between connected social groups are proportional to the size of the groups (as in the gravity model [42,43]). This may also be the case when disease vectors such as mosquitoes are more likely to detect large social groups than small social groups (e.g. malaria in primates [44,45]). It also assumes that when disease reaches a group, all individuals in the group become infected. This may not hold when the within-group disease transmission rate or the within-group connectivity is low. The model can be modified to handle such scenarios by replacing the group size distribution with the distribution of within-group outbreak sizes. This distribution can, for example, be inferred using stochastic SIR compartmental models for highly intra-connected groups [46] or network-based models for sparser groups. Within-group network structures may be sensitive to stochastic effects and yield highly variable within-group outbreak sizes, which will, in turn, affect the epidemiological vulnerability of the larger population. In these cases, the dynamics of infectious diseases will depend on the two-level within- and between-group contact network.

Second, the model assumes that group sizes are randomly distributed across the landscape. In reality, group sizes may depend on the quality of locally available resources, with large groups occurring around resource-rich areas and small groups found in more marginal areas [47]. If group sizes are spatially clustered, then the epidemiological impact of the group size variability may be different, depending on the size and spatial distribution of these clusters.

Finally, we assume that group size and degree are not correlated. If there is a positive correlation between the two, that is, larger groups have more inter-group contacts, then we expect the effects of group size variability to be amplified. The epidemic threshold would then be lower and epidemic size for high transmission rates would be smaller.

In conclusion, group size variability strongly impacts disease transmission in hierarchical populations. This pertains not only to group-living wild animals such as the Serengeti lions, but more generally to structured populations, including patch-structured wild plants, cultivated crops, herd-structured domestic livestock and community-structured humans.

Watch the video: Σκοτεινή Τριάδα u0026 Σκοτεινή Τετράδα: Ναρκισσισμός, Ψυχοπάθεια, Μακιαβελισμός, Σαδισμός (October 2022).