Misplaced Pages

#139860

220-509: One of the most important discoveries that was learned by observing the Kasekela chimpanzee community was the use of tools. On November 4, 1960, Goodall observed a chimpanzee that she had named David Greybeard using a grass stalk as a tool to extract termites from a termite hill. Later, she observed David Greybeard and another chimpanzee named Goliath stripping leaves off twigs to create termite fishing tools. Previously, tool use in chimpanzees

440-449: A miscarriage in 1970. Pom (1965–1983) was Passion's oldest known offspring to survive to adulthood. Pom was instrumental in helping her mother capture and kill community infants. Pom gave birth herself to Pan in 1978, and was very cautious with the infant around her mother. Pan died of injuries from an accident in 1981. Pom eventually disappeared about a year after her mother's death. She may have transferred to another community, but she

660-514: A 14-month-old human child that the niece of a member of the research team had carried into his territory. As a result, the Tanzanian National Parks Department considered killing Frodo. In 1988, he attacked visiting Far Side cartoonist Gary Larson , leaving him bruised and scratched. Frodo had a history of attacking the researchers observing him; Goodall was attacked by Frodo on multiple occasions and, in 1989,

880-462: A Kasekela infant named Tofiki in 2004. In his old age, his younger brothers sometimes provided him support, such as sharing meat and honey with him. The documentary Chimpanzee Family Fortunes ends with a scene of the orphaned young female Flirt finding solace after several lonely months by getting groomed by her older brother Freud. He fathered at least one infant, the female Cocoa to Candy, born in 2004 Freud's approach to maintaining his alpha status

1100-417: A baby of unknown sex in 1988. She was killed by males from the rival Mitumba Chimpanzee Community in 2005. A female chimpanzee leaving the safety of her community territory poses risks, and studies on chimpanzee intercommunity relations revealed that chimpanzees are likely to be attacked by the territorial males of a rival community, and the intensity of the attacks depends on the gender of the intruder, and if

1320-419: A brief amount of time, so they also rely on their legs. The appearance of the legs is similar in each caste, but the soldiers have larger and heavier legs. The structure of the legs is consistent with other insects: the parts of a leg include a coxa , trochanter , femur , tibia and the tarsus . The number of tibial spurs on an individual's leg varies. Some species of termite have an arolium, located between

1540-542: A carcass. In toque monkeys subordinates are often displaced from feeding sites by dominant males. Additionally, they are excluded from sleeping sites, and they suffer reduced growth and increased mortality. Subordinate individuals often demonstrate a huge reproductive disadvantage in dominance hierarchies. Among brown hyenas, subordinate females have less opportunity to rear young in the communal den, and thus have fewer surviving offspring than do high-ranking individuals. Subordinate males copulate far less with females than do

1760-447: A certain developmental path. Workers may also be fertile or sterile, however the term "worker" is normally reserved for the latter, having evolved in taxa that exhibit a bifurcated developmental pathway. As a result, sterile workers like in the family Termitidae are termed true workers and are the most derived, while those that are undifferentiated and fertile as in the wood-nesting Archotermopsidae are termed pseudergates , which are

1980-606: A close alliance against the other community females. Although males are perceived as the more aggressive sex in chimpanzees, studies at Gombe have revealed that there is substantial aggression among females with each other. One of two contexts for female-female aggressive encounters in Gombe was the presence of newborn infants. Fifi and Fanni were a formidable force and they carried out infanticidal attacks on other, lower-ranking females in her community. Such attacks were observed on fellow Kasekela female, Gremlin, on three different occasions;

2200-432: A coalition of males, but remained the top-ranked male. By 1977, he had regained his alpha status by forging alliances with other males, including his predecessor Humphrey. In 1979, he was toppled from his alpha status by 15-year-old Goblin, whom he had previously supported, but regained his alpha status once again by forging alliances with other males. Although a powerful male, Figan's ability to make alliances with other males

2420-401: A coalition of rival males formed that included his nephews Fudge and Fundi, and their father and uncle Sheldon and Sampson, and another nephew Zeus. In October 2016 the coalition attacked Ferdinand when he made one of his occasional appearances to try to assert his continuing dominance. Fudge and Fundi led the attack, with some support from their mother (and Ferdinand's sister) Fanni. Ferdinand

SECTION 10

#1732788100140

2640-492: A colony can reproduce, whereas the other colony members have their reproductive capabilities suppressed. This conflict over reproduction in some cases results in a dominance hierarchy. Dominant individuals in this case are known as queens and have the obvious advantage of performing reproduction and benefiting from all the tasks performed by their subordinates, the worker caste (foraging, nest maintenance, nest defense, brood care and thermal regulation). According to Hamilton's rule ,

2860-400: A colony of H. glaber was correlated with the individual's ranking position within a dominance hierarchy, but aggression between potential reproductives only started after the queen was removed. The social insects mentioned above, excluding termites, are haplodiploid . Queen and workers are diploid, but males develop from haploid genotypes. In some species, suppression of ovary development

3080-472: A developmental process called incomplete metamorphosis , going through multiple gradual pre-adult molts that are highly developmentally plastic before becoming an adult. Unlike in other hemimetabolous insects, nymphs are more strictly defined in termites as immature young with visible wing buds, which often invariably go through a series of moults to become winged adults . Larvae , which are defined as early nymph instars with absent wing buds, exhibit

3300-485: A dominant, authoritative manner in a group, this behaviour tends to prompt submissive responses from other group members. Similarly, when group members display submissive behaviour, others feel inclined to display dominant behaviours in return. Tiedens and Fragale (2003) found that hierarchical differentiation plays a significant role in liking behaviour in groups. Individuals prefer to interact with other group members whose power, or status behaviour complements their own. That

3520-425: A few hundred individuals to enormous societies with several million individuals. Most species are rarely seen, having a cryptic life-history where they remain hidden within the galleries and tunnels of their nests for most of their lives. Termites' success as a group has led to them colonizing almost every global landmass, with the highest diversity occurring in the tropics where they are estimated to constitute 10% of

3740-442: A fight for a female, and they father 44% of the lambs born in the population. These sheep live in large flocks, and dominance hierarchies are often restructured each breeding season. Burying beetles , which have a social order involving one dominant male controlling most access to mates, display a behavior known as sneak copulation. While one male at a carcass has a 5:1 mating advantage, subordinate males will tempt females away from

3960-412: A fight. Ferdinand maintained his alpha male position primarily through intimidation and threats and even surprising attacking males. He would bite rivals on the back to leave a scar. By the last two years of his reign Ferdinand's aggressiveness had made him so unpopular with other group males that he spent most of his time apart from them, returning to assert his dominance every so often. During that time

4180-513: A fontanelle, which appears as a circular pore or series of pores in a depressed region within the middle of the head. The fontanelle connects to the frontal gland, a novel organ unique to Neoisopteran termites which evolved to excrete an array of defensive chemicals and secretions, and so is typically most developed in the soldier caste. Cellulose digestion in the family Termitidae has co-evolved with bacterial gut microbiota and many taxa have evolved additional symbiotic relationships such as with

4400-584: A hierarchy is to prolong the colony lifespan. The top ranked individuals may die or lose fertility and "extra queens" may benefit from starting a colony in the same site or nest. This advantage is critical in some ecological contexts, such as in situations where nesting sites are limited or dispersal of individuals is risky due to high rates of predation. This polygynous behavior has also been observed in some eusocial bees such as Schwarziana quadripunctata . In this species, multiple queens of varying sizes are present. The larger, physogastric , queens typically control

4620-419: A hierarchy of potential reproductives. In the honey bee Apis mellifera , a pheromone produced by the queen mandibular glands is responsible for inhibiting ovary development in the worker caste . " Worker policing " is an additional mechanism that prevents reproduction by workers, found in bees and ants. Policing may involve oophagy and immobilization of workers who lay eggs. In some ant species such as

SECTION 20

#1732788100140

4840-437: A high rank. These include whether or not high rank gives them access to valuable resources such as mates and food. Age, intelligence, experience, and physical fitness can influence whether or not an individual deems it worthwhile to pursue a higher ranking in the hierarchy, which often comes at the expense of conflict. Hierarchy results from interactions, group dynamics, and sharing of resources, so group size and composition affect

5060-416: A larger role in establishing dominance (Roseler et al. , 1984). Subsequent research however, suggests that juvenile hormone is implicated, though only on certain individuals. When injected with juvenile hormone, larger foundresses showed more mounting behaviors than smaller ones, and more oocytes in their ovaries . Naked mole-rats ( Heterocephalus glaber ) similarly have a dominance hierarchy dependent on

5280-450: A palm tree by a strong wind and died four days later. Patti (c.1961–2005) was not born in the Kasekela community but joined the community in 1973. She was an assertive female who spent much time on her own. Her first two infants died before reaching adulthood, but she has successfully raised several chimps since. Besides Tapit, Tita, Tanga, Titan and Tarzan, she had two offspring who died in infancy, including her first born Pibi (1978) and

5500-464: A particularly difficult weaning conflict . This may account for Faustino being more skittish as a juvenile than his younger brother. Faustino grew into a large but mellow adult who reminded observers of Freud. He had worked his way up to become the beta, or second highest-ranking, male in the community until struck by a near fatal disease in 2005. Although Faustino survived, he lost his high rank. After recovering, Faustino slowly climbed his way back up

5720-454: A pedicel (typically shorter than the scape), and the flagellum (all segments beyond the scape and pedicel). The mouth parts contain a maxillae , a labium, and a set of mandibles . The maxillae and labium have palps that help termites sense food and handling. The cuticle of most castes is soft and flexible due to a resulting lack of sclerotization, particularly of the abdomen which often appears translucent. Pigmentation and sclerotization of

5940-405: A rather domineering and aggressive personality; according to Goodall, Passion was an indifferent mother to her three surviving offspring, but she became a very high-ranking female. She became more solitary later in life and died of some type of wasting disease in 1982. In addition to Pom, Prof and Pax, she gave birth to a baby in 1962 which died in infancy. She also had a stillborn baby, or possibly

6160-486: A region of the female wasp brain responsible for the synthesis and secretion of juvenile hormone, are naturally more dominant. A follow-up experiment utilized 20-hydroxyecdysone , an ecdysone known to enhance maturation and size of oocytes . The size of the oocytes plays a significant role in establishing dominance in the paper wasp. Foundresses treated with 20-hydroxyecdysone showed more dominance than did those treated with juvenile hormone, so 20-hydroxyecdysone may play

6380-550: A role in suppressing reproductive function. Glucocorticoids , signaling molecules which stimulate the fight or flight response , may be implicated in dominance hierarchies. Higher ranking individuals tend to have much higher levels of circulating glucocorticoids than subdominant individuals, the opposite of what had been expected. Two core hypotheses attempt to explain this. The first suggests that higher ranking individuals exert more energy and thus need higher levels of glucocorticoids to mobilize glycogen for energy use. This

6600-416: A rough life, being sickly as an infant, and he died as a juvenile. Tita (b. January 1984) is Patti's oldest daughter. She gave birth to her first child, Tibi, in 1995; he died in infancy. Another son, Tofiki (fathered by Sheldon), was born in 2000, and killed in 2004 when Freud attacked him while grooming Tita. Termite Termites are a group of detritophagous eusocial insects which consume

6820-575: A social order is created that is subject to change each time a dominant animal is challenged by a subordinate one. Dominance is an individual's preferential access to resources over another based on coercive capacity based on strength, threat, and intimidation, compared to prestige (persuasive capacity based on skills, abilities, and knowledge). A dominant animal is one whose sexual , feeding, aggressive, and other behaviour patterns subsequently occur with relatively little influence from other group members. Subordinate animals are opposite; their behaviour

Kasakela chimpanzee community - Misplaced Pages Continue

7040-416: A son named Ghurubu in 2016. In 2012, Golden gave birth to a daughter named Glamour. Golden also gave birth to a son named Gombe in 2017. She had another baby in 2022. Golden and Glitter have remained extremely close through adulthood. Passion (c.1951–1982) was a central female of the Kasekela community. She, along with her daughter Pom, captured, killed, and ate about ten newborns at Gombe. Passion possessed

7260-420: A stable hierarchy than did subdominant individuals, but the reverse was true at unstable times. Several areas of the brain contribute to hierarchical behavior in animals. One of the areas that has been linked with this behavior is the prefrontal cortex , a region involved with decision making and social behavior. High social rank in a hierarchical group of mice has been associated with increased excitability in

7480-404: A termite is in flight, its wings remain at a right angle, and when the termite is at rest, its wings remain parallel to the body. Due to termites being hemimetabolous insects , where the young go through multiple and gradual adultoid molts before becoming an adult, the advent of eusociality has significantly altered the developmental patterns of this group of insects of which, although similar,

7700-457: A thoracic sclerite in Diacamma ants inhibits ovary development; the only reproductive individual of this naturally queenless genus is the one that retains its sclerite intact. This individual is called a gamergate , and is responsible for mutilating all the newly emerged females, to maintain its social status. Gamergates of Harpegnathos saltator arise from aggressive interactions, forming

7920-484: A twin, he grew into a healthy but small adult. He reached a relatively high rank in the late 1990s but fell to a low-ranking male by 2004. He fathered at least two children, Mambo by Malaika, born in 2004, and Zinda (2006 – October 2012) by Trezia. He disappeared in early 2007. Galahad (1988-2000) was Gremlin's third son. His father was a low-ranking male, Atlas. Galahad was described by researchers at Gombe as mischievous and playful, often observed terrorizing baboons. In

8140-400: A unique white plumage; the higher the percentage of the crown that consists of white feathers, the higher the status of the individual. For other animals, the time spent in the group serves as a determinant of dominance status. Rank may also be acquired from maternal dominance rank. In rhesus monkeys , offspring gain dominance status based on the rank of the mother—the higher ranked the mother,

8360-426: A variety of decaying plant material , generally in the form of wood , leaf litter , and soil humus . They are distinguished by their moniliform antennae and the soft-bodied and often unpigmented worker caste for which they have been commonly termed " white ants "; however, they are not ants , to which they are only distantly related. About 2,972 extant species are currently described, 2,105 of which are members of

8580-466: A wide variety of predators. The queens can be particularly long-lived for insects, with some reportedly living as long as 30 or 50 years. In both the linear and bifurcated developmental pathways, the primary reproductives only develop from winged immatures (nymphs). These winged immatures are capable of regressively molting into a form known as brachypterous neotenics ( nymphoids ), which retain juvenile and adult characteristics. BN 's can be found in both

8800-429: Is submissive , and can be relatively easily influenced or inhibited by other group members. For many animal societies, an individual's position in the dominance hierarchy corresponds with their opportunities to reproduce. In hierarchically social animals, dominant individuals may exert control over others. For example, in a herd of feral goats it is a large male that is dominant and maintains discipline and coherence of

9020-592: Is Fanni's oldest surviving son. His father is Sheldon, a former alpha male. He is the fifth male of the F-family to reach alpha male status, overthrowing his own maternal uncle Ferdinand in October 2016. Fudge had joined a coalition that included his brother Fundi, his father Sheldon, his uncle Sampson and his cousin Zeus in opposition to Ferdinand. In October 2016 Fudge led an attack by the rival males on Ferdinand. Fundi

Kasakela chimpanzee community - Misplaced Pages Continue

9240-454: Is Fifi's fourth oldest son. His father was his uncle Figan's rival Evered, who was 41 years old when Ferdinand was conceived. He became the first wild great ape to have his birth recorded. By 2007, he had grown into a large enough male to wound the alpha male Kris, and researchers believed he would eventually become the third of Fifi's sons to become alpha male. In 2008, he succeeded in supplanting Kris as alpha male, after wounding Kris badly in

9460-416: Is Fifi's third oldest son. His father was brother Freud's rival Wilkie. At the beginning of the documentary Chimps: So Like Us , the infant Faustino is shown with mother Fifi and sister Flossi trying out facial expressions. Fifi became pregnant less than three years after Faustino was born, an unusually short time for a chimpanzee. As a result, Faustino had to be weaned at an unusually young age, leading to

9680-528: Is Gremlin's oldest daughter. Her father was Wilkie. As a juvenile, she helped her mother raise the twins Glitter and Golden, and this is shown in a number of documentaries about the community. Gaia gave birth to her first baby in April 2006, but the baby's grandmother Gremlin took the infant from her daughter, presumably in fear of it being attacked. Although Gremlin's instincts were justified from her own experiences of other community females attacking her newborns,

9900-617: Is abundant food and females will mate promiscuously. Because of this, males gain very little in fighting over females, who are, in turn, too large and strong for males to monopolize or control, so males do not appear to form especially prominent ranks between them, with several males mating with the same female in view of each other. This type of mating style is also present in manatees, removing their need to engage in serious fighting. Among female elephants, leadership roles are not acquired by sheer brute force, but instead through seniority, and other females can collectively show preferences for where

10120-730: Is believed to be from the Upper Cretaceous in West Texas , where the oldest known faecal pellets were also discovered. Claims that termites emerged earlier have faced controversy. For example, F. M. Weesner indicated that the Mastotermitidae termites may go back to the Late Permian , 251 million years ago, and fossil wings that have a close resemblance to the wings of Mastotermes of the Mastotermitidae,

10340-399: Is common in farming, the dominance hierarchy becomes less stable and aggression increases. Dominance hierarchies are found in many species of bird. For example, the blue-footed booby brood of two chicks always has a dominance hierarchy due to the asynchronous hatching of the eggs. One egg is laid four days before the other, and incubation starts immediately after laying, so the elder chick

10560-468: Is consistent with all four-winged insects. The mesothorax and metathorax have well-developed exoskeletal plates; the prothorax has smaller plates. Termites have a ten-segmented abdomen with two plates, the tergites and the sternites . The tenth abdominal segment has a pair of short cerci . There are ten tergites, of which nine are wide and one is elongated. The reproductive organs are similar to those in cockroaches but are more simplified. For example,

10780-707: Is correlated with reproductive success . Although a high rank is an advantage for females, clear linear hierarchies in female chimpanzees have not been detected. In "masculinized" female mammals like the spotted hyena ( Crocuta crocuta ), androgens (i.e. specifically, androstenedione and testosterone) are "implicated in the organization and activation of...nonreproductive behavioral traits, including aggression, social dominance, rough-and-tumble play, and scent marking". For aggressively dominant female meerkats ( Suricata suricatta ), they have "exceptionally high concentrations" of androgens, "particularly during gestation". The concept of dominance, originally called "pecking order",

11000-559: Is divided into the following clade and family groups, showing the subfamilies in their respective classification: The Neoisoptera , literally meaning "newer termites" (in an evolutionary sense), are a recently coined clade that include families such as the Rhinotermitidae and Termitidae . Neoisopterans have a bifurcated caste development with true workers, and so notably lack pseudergates (except in some basal taxa such as Serritermitidae : see below ). All Neoisopterans have

11220-546: Is expected to increase over time as a consequence of climate change . Increased urbanization and connectivity is also predicted to expand the range of some pest termites. The infraorder name Isoptera is derived from the Greek words iso (equal) and ptera (winged), which refers to the nearly equal size of the fore and hind wings. "Termite" derives from the Latin and Late Latin word termes ("woodworm, white ant"), altered by

SECTION 50

#1732788100140

11440-494: Is found mainly in the derived taxa (i.e. Neoisoptera ), and is believed to have evolved in tandem with the sterile worker caste as species moved to foraging for food beyond their nests, as opposed to the nest also being the food (such as in obligate wood-dwellers). There are three main castes which are discussed below: Worker termites undertake the most labor within the colony, being responsible for foraging, food storage, and brood and nest maintenance. Workers are tasked with

11660-565: Is hatched four days before the younger chick and has a four-day head start on growth. The elder, stronger chick almost always becomes the dominant chick. During times of food shortage, the dominant chick often kills the subordinate chick by either repeatedly pecking or by ousting the younger chick from the nest. The brood hierarchy makes it easier for the subordinate chick to die quietly in times of food scarcity, which provides an efficient system for booby parents to maximize their investment. In insect societies , only one to few individuals members of

11880-404: Is leader and which one is not. Sometimes dominant animals must maintain alliances with subordinates and grant them favours to receive their support in order to retain their dominant rank. In chimpanzees, the alpha male may need to tolerate lower-ranking group members hovering near fertile females or taking portions of his meals. Other examples can include Muriqui monkeys. Within their groups, there

12100-422: Is not homologous to that of the eusocial Hymenoptera . Unlike ants, bees, and wasps which undergo a complete metamorphosis and as a result only exhibit developmental plasticity at the immobile larval stage , the mobile adultoid instars of termites remain developmentally flexible throughout all life stages up to the final molt , which has uniquely allowed for the evolution of distinct yet flexible castes amongst

12320-411: Is not totally achieved in the worker caste, which opens the possibility of reproduction by workers. Since nuptial flights are seasonal and workers are wingless, workers are almost always non-breeders, and (as gamergate ants or laying worker bees ) can only lay unfertilised eggs. These eggs are in general viable, developing into males. A worker that performs reproduction is considered a "cheater" within

12540-624: Is observed consistently in hyenas , lemurs and the bonobo . The ring-tailed lemur is observed to be the most prominent model of female dominance. There are three basic proposals for the evolution of female dominance: In lemurs, no single hypothesis fully explains female social dominance at this time and all three are likely to play a role. Adult female lemurs have increased concentrations of androgens when they transition from non-breeding to breeding seasons, increasing female aggression. Androgens are greater in pregnant female lemurs, which suggests that organizational androgens might influence

12760-414: Is often what leads to the size differences that result in dominant-subordinate position rankings. Therefore, if during the winter aggregate, the female is able to obtain greater access to food, the female could thus reach a dominant position. In some species, especially in ants, more than one queen can be found in the same colony, a condition called polygyny . In this case, another advantage of maintaining

12980-581: Is supported by the fact that when food availability is low, cortisol levels increase within the dominant male. The second suggests that elevated stress hormones are a result of social factors, particularly when the hierarchy is in transition, perhaps resulting in increased aggression and confrontation. As a result, the dominant individual fights more and has elevated glucocorticoids during this period. Field studies of olive baboons in Kenya seem to support this, as dominant individuals had lower cortisol levels in

13200-442: Is the beta males that gain the most fitness, avoiding stress but receiving some of the benefits of moderate rank. The mating tactics of savanna baboons are correlated with their age. Older, subordinate males form alliances to combat higher-ranking males and get access to females. Fighting with dominant males is a risky behavior that may result in defeat, injury or even death. In bighorn sheep , however, subordinates occasionally win

13420-453: Is the environment. In populations of Kenyan vervet monkeys , high-ranking females have higher foraging success when the food resources are clumped, but when food is distributed throughout an area they lose their advantage, because subordinate females can acquire food with less risk of encountering a dominant female. A benefit to high-ranking individuals is increased foraging success and access to food resources. During times of water shortage

SECTION 60

#1732788100140

13640-406: Is to say, group members who behave submissively when talking to someone who appears to be in control are better liked, and similarly individuals who display dominant behaviours (e.g., taking charge, issuing orders) are more liked when interacting with docile, subservient individuals. Being subordinate offers a number of benefits. Subordination is beneficial in agonistic conflicts where rank predicts

13860-411: Is typical for female chimpanzees to emigrate from their natal group upon reaching maturity, within the Kasekela community only about half the females do so. Flossi may have chosen to emigrate to avoid mating attempts from her brother Frodo. Before emigrating from the Kasekela community, Flossi may have innovated the practice of fishing for ants to eat using a stick, which had not been observed before within

14080-521: The nymphal (winged) and apterous (wingless) lines. The bifurcation occurs early, either at the egg or the first two instars, and represents an irreversible and committed development to either the reproductive or non-reproductive lifestyles. As such, the apterous lineage consists mostly of wingless and truly altruistic sterile individuals (true workers, soldiers), whereas the nymphal lineage consists mainly of fertile individuals destined to become winged reproductives. The bifurcated developmental pathway

14300-477: The Archotermopsidae , which are able to retain high developmental plasticity even up to the late instars. In these basal taxa, the immatures are able to go through progressive ( nymph-to-imago ), regressive ( winged-to-wingless ) and stationary ( size increase, remains wingless ) molts, which typically indicates the developmental trajectory an individual follows. There is significant variation of

14520-457: The IMAX film Jane Goodall's Wild Chimpanzees . Later she brought her sons Figan and Faben, who would later become prominent members of the community to the camp, and when she came into estrus in 1963 she attracted the community's males to Goodall's camp. Flo was a high-ranking female who had at least five known offspring, three sons (Faben, Figan and Flint) and two daughters (Fifi and Flame). Flo

14740-674: The Nasutes , which have a horn-like nozzle projection ( nasus ) on the head. These unique soldiers are able to spray noxious, sticky secretions containing diterpenes at their enemies. Nitrogen fixation plays an important role in Nasute nutrition. Soldiers are normally a committed sterile caste and so do not molt into anything else, but in certain basal taxa like the Archotermopsidae they are known to rarely molt into neotenic forms that develop functional sexual organs. In species with

14960-872: The amygdala through lesion studies in rats and primates which led to disruption in hierarchy, and can affect the individual negatively or positively depending on the subnuclei that is targeted. Additionally, the dorsal medial PFC-medial dorsal thalamus connection has been linked with maintenance of rank in mice. Another area that has been associated is the dorsal raphe nucleus , the primary serotonergic nuclei (a neurotransmitter involved with many behaviors including reward and learning). In manipulation studies of this region, there were changes in fighting and affiliative behavior in primates and crustaceans. Female-biased dominance occurs rarely in mammals. It occurs when all adult males exhibit submissive behavior to adult females in social settings. These social settings are usually related to feeding, grooming, and sleeping site priority. It

15180-462: The anterior teats produce a greater quantity of milk. Once established, this teat order remains stable with each piglet tending to feed from a particular teat or group of teats. Dominance–subordination relationships can vary markedly between breeds of the same species. Studies on Merinos and Border Leicesters sheep revealed an almost linear hierarchy in the Merinos but a less rigid structure in

15400-455: The carpenter ant Camponotus floridanus , eggs from queens have a peculiar chemical profile that workers can distinguish from worker laid eggs. When worker-laid eggs are found, they are eaten. In some species, such as Pachycondyla obscuricornis , workers may try to escape policing by shuffling their eggs within the egg pile laid by the queen. Modulation of hormone levels after hibernation may be associated with dominance hierarchies in

15620-403: The claws , which is present in species that climb on smooth surfaces but is absent in most termites. Unlike in ants, the hind-wings and fore-wings are of equal length. Most of the time, the alates are poor flyers; their technique is to launch themselves in the air and fly in a random direction. Studies show that in comparison to larger termites, smaller termites cannot fly long distances. When

15840-416: The eusocial termite Reticulitermes speratus are capable of a long lifespan without sacrificing fecundity . These long-lived queens have a significantly lower level of oxidative damage, including oxidative DNA damage , than workers, soldiers and nymphs. The lower levels of damage appear to be due to increased catalase , an enzyme that protects against oxidative stress . Beta (biology) In

16060-503: The intromittent organ is not present in male alates, and the sperm is either immotile or aflagellate. However, Mastotermitidae termites have multiflagellate sperm with limited motility . The genitals in females are also simplified. Unlike in other termites, Mastotermitidae females have an ovipositor , a feature strikingly similar to that in female cockroaches. The non-reproductive castes of termites are wingless and rely exclusively on their six legs for locomotion. The alates fly only for

16280-437: The zoological field of ethology , a dominance hierarchy (formerly and colloquially called a pecking order ) is a type of social hierarchy that arises when members of animal social groups interact, creating a ranking system. A dominant higher-ranking individual is sometimes called an alpha , and a submissive lower-ranking individual is called a beta . Different types of interactions can result in dominance depending on

16500-588: The "egalitarian hypothesis", which predicts that status would affect reproductive success more amongst foragers than amongst nonforagers. High-ranking bonnet macaque males have more access to fertile females and consequently partake in most of the matings within the group; in one population, three males were responsible for over 75% of matings. In this population, males often vary in rank. As their rank improves, they gain more exclusive time with fertile females; when their rank decreases, they get less time. In many primates, including bonnet macaques and rhesus monkeys ,

16720-436: The "king" and "queen" that together form a lifelong monogamous pair. Also unlike ants, which undergo a complete metamorphosis , termites undergo an incomplete metamorphosis that proceeds through egg, nymph , and adult stages. Termite colonies are commonly described as superorganisms due to the collective behaviors of the individuals which form a self-governing entity: the colony itself. Their colonies range in size from

16940-539: The "relational model" created by the zoologist Frans De Waal . In systems where competition between and within the sexes is low, social behaviour gravitates towards tolerance and egalitarianism, such as that found in woolley spider monkeys . In despotic systems where competition is high, one or two members are dominant while all other members of the living group are equally submissive, as seen in Japanese and rhesus macaques, leopard geckos , dwarf hamsters , gorillas ,

17160-453: The 1970s, which proved that cannibalism, though deemed rare, does occur among both male and female chimpanzees. Clever females like Gremlin, who bore the brunt of infanticidal females, employed two counter strategies to avoid attacks on them and newborns. One strategy is being in close proximity to adult males for protection, as Gremlin initially did by sticking close to Frodo (the father of her twins) in 1998. The second strategy used by Gremlin

17380-594: The 2,972 extant termite species currently classified, 1,000 are found in Africa , where mounds are extremely abundant in certain regions. Approximately 1.1 million active termite mounds can be found in the northern Kruger National Park alone. In Asia , there are 435 species of termites, which are mainly distributed in China . Within China, termite species are restricted to mild tropical and subtropical habitats south of

17600-547: The BBC television program Chimpanzee Diary 1998 Galahad was depicted being supportive of his mother Gremlin, staying near by even though he could not intervene during her attack by other community females trying to take her newborn twins. He was developing into a favourite character for the human observers in Gombe, but was unfortunately struck down by the flu epidemic in 2000 and died in February of that year Gaia (b. February 1993)

17820-516: The Border Leicesters when a competitive feeding situation was created. Although many group-living animal species have a hierarchy of some form, some species have more fluid and flexible social groupings, where rank does not need to be rigidly enforced, and low-ranking group members may enjoy a wider degree of social flexibility. Some animal societies are "democratic", with low-ranking group members being able to influence which group member

18040-472: The Kasekela community (although fishing for termites had been) but soon after the practice spread throughout the community. Flossi has given birth to four offspring, the males Forest (1997– ) and Fansi (2001– ) and the females Flower (2005– ), and Falida (2009– ). Flossi became a high ranking female within the Mitumba community; more typically immigrating females remain at a low rank. Faustino (b. May 8, 1989)

18260-446: The Kasekela territory, further away from the other community females. Flossi (b. February 8, 1985) is Fifi's second-oldest daughter. Her father was Goblin. Unlike most other female chimpanzees in the F-family, upon reaching adulthood she emigrated from the Kasekela community to join the neighbouring Mitumba Chimpanzee Community in 1996, although her younger sister Flirt (who was born after Flossi emigrated) later joined her. Although it

18480-686: The Paleozoic and Triassic insects tentatively classified as termites are in fact unrelated to termites and should be excluded from the Isoptera. Other studies suggest that the origin of termites is more recent, having diverged from Cryptocercus sometime during the Early Cretaceous . The primitive giant northern termite ( Mastotermes darwiniensis ) exhibits numerous cockroach-like characteristics that are not shared with other termites, such as laying its eggs in rafts and having anal lobes on

18700-796: The Yangtze River. In Australia , all ecological groups of termites (dampwood, drywood, subterranean) are endemic to the country, with over 360 classified species. Because termites are highly social and abundant, they represent a disproportionate amount of the world's insect biomass . Termites and ants comprise about 1% of insect species, but represent more than 50% of insect biomass. Due to their soft cuticles, termites do not inhabit cool or cold habitats. There are three ecological groups of termites: dampwood, drywood and subterranean. Dampwood termites are found only in coniferous forests, and drywood termites are found in hardwood forests; subterranean termites live in widely diverse areas. One species in

18920-458: The age of almost 50, she gave birth to a son, Goodali. Although males are perceived as the more aggressive sex in chimpanzees, studies at Gombe have revealed that aggression between females does occur. Such attacks were observed of Fifi and her adult daughter, Fanni, aggressively harassing fellow Kasekela female, Gremlin, on three different occasions; the first being in 1993 following the birth of her daughter Gaia, and then five years later following

19140-558: The alpha male Ferdinand. Goodall has described Gremlin as one of her favourite chimpanzes, due to Gremlin’s patience as a mother and expertise in fishing for termites. She has raised twins, Golden and Glitter, who are the oldest known wild twin chimpanzees. She had male offspring in Getty (1982–1986), Galahad (1988–2000), Gimli (2004– ), Gizmo (2010–2016) and a female offspring in Gaia (1993– ). In 2015, she gave birth to Grendel and in 2019, at

19360-401: The alpha male position. Researchers speculated that Frodo was unaffected by the mange because he interacted less with the other apes, giving the parasite less opportunity to infect him. After taking over the alpha male position, Frodo demonstrated his new power by protecting his vanquished brother from the other males; typically the former alpha is attacked by the other males and forced to leave

19580-426: The alpha male was never the one to sire the most offspring, with that instead being a high-ranking but not top male. The complex relationship between rank and reproduction in this species is likely explained by the fact that rhesus macaques queue, rather than fight, for dominance, meaning that the alpha male is not necessarily the strongest or most attractive male. In rodents, the highest-ranking male frequently sires

19800-631: The alpha position results in individuals maintaining high rank for shorter periods of time and having an overall reduced health and longevity from the physical strain and costs of the position. The interpersonal complementarity hypothesis suggests that obedience and authority are reciprocal, complementary processes. That is, it predicts that one group member's behaviours will elicit a predictable set of actions from other group members. Friendly behaviours are predicted to be met with friendly behaviours, and hostile behaviours are predicted to be reciprocated with similar, hostile behaviours. When an individual acts in

20020-528: The animal biomass , particularly in Africa which has the richest diversity with more than 1000 described species. They are important decomposers of decaying plant matter in the subtropical and tropical regions of the world, and their recycling of wood and plant matter is of considerable ecological importance. Many species are ecosystem engineers capable of altering soil characteristics such as hydrology , decomposition, nutrient cycling , vegetative growth, and consequently surrounding biodiversity through

20240-710: The ape beat her head so violently her neck was nearly broken. Frodo fathered at least eight infants, second only to group male Wilkie, who fathered ten. Infants he fathered included: Fanni (b. March 19, 1981) is Fifi's oldest daughter. Her father was Goblin. She had her first infant, Fax, at just 11 years old, making her one of the youngest mothers seen in the community, although Fax did not survive to adulthood. She has also had five other children, males Fudge (1996– ), Fundi (2000– ), Fifty (2010– ) and Ficma (2015–) and females Familia (2004– ) and Fadhila (2007– ). Fudge, Fundi and Familia were fathered by Sheldon, who would later become alpha male. Familia disappeared in 2012. Fadhila

20460-464: The best position to detect and avoid predators, as well as to display their dominance to other members of their own species. It has been suggested that decision-taking about the actions of the group is commonly dissociated from social dominance. Given the benefits and costs of possessing a high rank within a hierarchical group, there are certain characteristics of individuals, groups, and environments that determine whether an individual will benefit from

20680-476: The birth of her twins. This was followed by another attack on Gremlin six years later following the birth of her next baby. Researchers could only speculate that the attacks on infants of other females had to do with food competition, as each female strives to secure resources for her own group of offspring and thus the expansion of another female's family unit could deplete resources. The attacks by Fifi and Fanni mirror those of previous perpetrators Passion and Pom in

20900-454: The carcass with pheromones and attempt to copulate before the dominant male can drive them forcefully away. In flat lizards , young males take advantage of their underdeveloped secondary sex characteristics to engage in sneak copulations. These young males mimic all the visual signs of a female lizard in order to successfully approach a female and copulate without detection by the dominant male. This strategy does not work at close range because

21120-554: The chemical signals given off by the sneaky males reveal their true nature, and they are chased out by the dominant. Subordinate individuals suffer a range of costs from dominance hierarchies, one of the most notable being reduced access to food sources. When a resource is obtained, dominant individuals are first to feed as well as taking the longest time. Subordinates also lose out in shelter and nesting sites. Brown hyenas , which display defined linear dominance in both sexes, allow subordinate males and females decreased time of feeding at

21340-463: The cichlid Neolamprologus pulcher , and African wild dog . Linear ranking systems, or "pecking orders", which tend to fall in between egalitarianism and despotism, follow a structure where every member of the group is recognized as either dominant or submissive relative to every other member. This results in a linear distribution of rank, as seen in spotted hyenas and brown hyenas. Dominance and its organisation can be highly variable depending on

21560-555: The colony, because it leaves disproportionately more descendants than do its sisters and mother. The advantage of remaining functionally sterile is only accomplished if every worker assume this "compromise". When one or more workers start reproducing, the "social contract" is destroyed and the colony cohesion is dissolved. Aggressive behavior derived from this conflict may result in the formation of hierarchies, and attempts of reproduction by workers are actively suppressed. In some wasps, such as Polistes fuscatus , instead of not laying eggs,

21780-458: The colony. Workers remain wingless and across many taxa become developmentally arrested, appearing to not change into any other caste until death. In some basal taxa, there is no distinction, with the "workers" (pseudergates) essentially being late instar larvae that retain the ability to change into all other castes. The development of larvae into adults can take months; the time period depends on food availability and nutrition, temperature, and

22000-423: The community for a period of time. As alpha male, Frodo maintained his position largely through intimidation. He rarely groomed other males, and often demanded that other males groom him. Frodo remained alpha male until becoming ill himself in 2002; he was then defeated by a coalition of several males and spent most of the next two years alone, recovering from his wounds and illness. He received antibiotics from

22220-461: The community, and the matriarch, Flo, played a particularly important role in acknowledging Goodall's acceptance as a human observer by the community. The G-family has produced at least one alpha male, and also the birth of several twins, which are rare among chimpanzees. There are other families as well which include the T-family and S-family (which has produced one alpha male). Flo (c.1919 – 1972)

22440-554: The conflicts with the rival Kahama community (which was formed by former Kasekela members) that occurred between 1973 and 1975, and was a leader in the killing of several Kahama males, including former Kasekela alpha male Goliath. Faben himself died in 1975, possibly as a result of further conflicts with the Kahama community. Following Faben's death, Figan had difficulty maintaining his alpha male position for several months and eventually lost his alpha status to Goblin. Figan (c.1953 – 1982)

22660-468: The contest. Larger stags have also been known to make lower-frequency threat signals, acting as indicators of body size, strength, and dominance. Engaging in agonistic behavior can be very costly and thus there are many examples in nature of animals who achieve dominance in more passive ways. In some, the dominance status of an individual is clearly visible, eliminating the need for agonistic behavior. In wintering bird flocks, white-crowned sparrows display

22880-514: The context or individuals involved. In European badgers , dominance relationships may vary with time as individuals age, gain or lose social status, or change their reproductive condition. Dominance may also vary across space in territorial animals as territory owners are often dominant over all others in their own territory but submissive elsewhere, or dependent on the resource. Even with these factors held constant, perfect dominance hierarchies are rarely found in groups of any great size, at least in

23100-431: The cuticle correlates with life history , with species that spend more time in the surface in the open tending to have a more sclerotized and pigmented exoskeleton. Consistent with all insects, the anatomy of the termite thorax consists of three segments: the prothorax , the mesothorax and the metathorax . Each segment contains a pair of legs . On alates, the wings are located at the mesothorax and metathorax, which

23320-483: The death of his mother Flo in 1972, Flint became depressed and ill; he died about three weeks after her. Flame (1968–1969) was Flo's daughter. Flo's son Flint, at age four, threw violent fits in order to suckle his mother's breast while newborn Flame was in Flo's arms. Flame disappeared shortly after she was born; there has been speculation she may have been killed by her jealous brother Flint. Freud (May 1971 – March 2014)

23540-604: The defeated, which include loss of reproductive opportunities and quality food, can hinder the individual's fitness. In order to minimize these losses, animals generally retreat from fighting or displaying fighting ability unless there are obvious cues indicating victory. These often involve characteristics that provide an advantage during agonistic behavior, such as size of body, displays, etc. Red stags , for example, engage in exhausting roaring contests to exhibit their strength. However, such an activity would impose more costs than benefits for unfit stags, and compel them to retreat from

23760-439: The derived and basal termite taxa, and generally serve as supplementary reproductives. Termites are often compared with the social Hymenoptera (ants and various species of bees and wasps), but their differing evolutionary origins result in major differences in life cycle. In the eusocial Hymenoptera, the workers are exclusively female. Males (drones) are haploid and develop from unfertilised eggs, while females (both workers and

23980-533: The descendants of the genus Cryptocercus . Some researchers have suggested a more conservative measure of retaining the termites as the Termitoidae, an epifamily within the cockroach order, which preserves the classification of termites at family level and below. Termites have long been accepted to be closely related to cockroaches and mantids , and they are classified in the same superorder ( Dictyoptera ). The oldest unambiguous termite fossils date to

24200-546: The developing offspring. Organizational androgens play a role in "explaining female social dominance" in ring-tailed lemurs , as androgens are associated with aggressive behavior in young females. Females that were "exposed to greater concentrations of maternal [androstenedione] late in fetal development were less likely to be aggressed against postnatally, whereas females that were...exposed to greater concentrations of maternal [testosterone]...were more likely to receive aggression postnatally". Dominance rank in female chimpanzees

24420-496: The developmental patterns in termites even across closely related taxa, but can typically be generalized into the following two patterns: The first is the linear developmental pathway , where all immatures are capable of developing into winged adults ( Alates ), exhibit high phenotypic potency, and where there exists no true sterile caste other than the soldier. The second is the bifurcated developmental pathway , where immatures diverge into two distinct developmental lineages known as

24640-418: The digestion of cellulose in food and are thus the most likely caste to be found in infested wood. The process of worker termites feeding other nestmates is known as trophallaxis . Trophallaxis is an effective nutritional tactic to convert and recycle nitrogenous components. It frees the parents from feeding all but the first generation of offspring, allowing for the group to grow much larger and ensuring that

24860-491: The dominance decisions of high-ranking individuals. For example, in a large group with many males, it may be difficult for the highest-ranking male to dominate all the mating opportunities, so some mate sharing is likely to exist. These opportunities available to subordinates reduce the likelihood of a challenge to the dominant male: mating is no longer an all-or-nothing game and the sharing is enough to placate most subordinates. Another aspect that can determine dominance hierarchies

25080-544: The drywood group is the West Indian drywood termite ( Cryptotermes brevis ) , which is an invasive species in Australia. Termites are usually small, measuring between 4 and 15 millimetres ( 3 ⁄ 16 and 9 ⁄ 16  in) in length. The largest of all extant termites are the queens of the species Macrotermes bellicosus , measuring up to over 10 centimetres (4 in) in length. Another giant termite,

25300-625: The early Cretaceous , but given the diversity of Cretaceous termites and early fossil records showing mutualism between microorganisms and these insects, they possibly originated earlier in the Jurassic or Triassic. Possible evidence of a Jurassic origin is the assumption that the extinct mammaliaform Fruitafossor from Morrison Formation consumed termites, judging from its morphological similarity to modern termite-eating mammals. Morrison Formation also yields social insect nest fossils close to that of termites. The oldest termite nest discovered

25520-429: The early 1960s was that male chimpanzees perform a " rain dance ," charging, calling, slapping the ground and trees and dragging branches in the rain. In the early 1970s the chimpanzees of the community were observed to engage in ongoing coordinated attacks against the chimpanzees of the neighbouring Kahama Chimpanzee Community, ultimately wiping it out. According to historian Ian Morris , this " Four Year War " represented

25740-409: The expansion of another female's family unit could deplete resources. The attacks by Fifi and Fanni mirror those of previous perpetrators Passion and Pom in the 1970s, which proved that cannibalism, although deemed rare, though deemed rare, does occur among both male and female chimpanzees. The loss of her mother in 2004 greatly reduced Fanni's dominance and she shifted her range to the southern part of

25960-602: The expansion of another female's family unit could deplete resources. The attacks by Fifi and Fanni mirror those of previous perpetrators Passion and Pom in the 1970s, which proved that cannibalism, though deemed rare, does occur among both male and female chimpanzees. In 2004, Fifi, who was by then 46 years old which made her the oldest living chimpanzee in the Kasekela community at that point in time, disappeared in September with her youngest and ninth offspring, two year old daughter Furaha. During that year she shifted her range to

26180-681: The extinct Gyatermes styriensis , flourished in Austria during the Miocene and had a wingspan of 76 millimetres (3 in) and a body length of 25 millimetres (1 in). Most worker and soldier termites are completely blind as they do not have a pair of eyes. However, some species, such as Hodotermes mossambicus , have compound eyes which they use for orientation and to distinguish sunlight from moonlight. The alates (winged males and females) have eyes along with lateral ocelli . Lateral ocelli, however, are not found in all termites, absent in

26400-409: The families Hodotermitidae , Termopsidae , and Archotermopsidae . Like other insects, termites have a small tongue-shaped labrum and a clypeus ; the clypeus is divided into a postclypeus and anteclypeus. Termite antennae have a number of functions such as the sensing of touch, taste, odours (including pheromones), heat and vibration. The three basic segments of a termite antenna include a scape ,

26620-424: The family Termitidae . Termites comprise the infraorder Isoptera , or alternatively the epifamily Termitoidae , within the order Blattodea (along with cockroaches ). Termites were once classified in a separate order from cockroaches, but recent phylogenetic studies indicate that they evolved from cockroaches, as they are deeply nested within the group, and the sister group to wood eating cockroaches of

26840-504: The far north of the Kasekela community territory, and the simultaneous advancement of the neighbouring Mitumba community further south nearing the Kasekela territory border suggests that Fifi and her youngster may have been fatally attacked by the Mitumba territorial males. A female chimpanzee leaving the safety of her community territory poses risks, particularly if she has a vulnerable youngster. Studies on chimpanzee intercommunity relations revealed that chimpanzees are likely to be attacked by

27060-414: The female workers begin being able to reproduce, but once being under the presence of dominant females, the subordinate female workers can no longer reproduce. In some wasp species such as Liostenogaster flavolineata there are many possible queens that inhabit a nest, but only one can be queen at a time. When a queen dies the next queen is selected by an age-based dominance hierarchy. This is also true in

27280-420: The first being in 1993 following the birth of her daughter Gaia, and then five years later following the birth of her twins. This was followed by another attack on Gremlin six years later following the birth of her next baby. Researchers could only speculate that the attacks on infants of other females had to do with food competition, as each female strives to secure resources for her own group of offspring and thus

27500-420: The first being in 1993 following the birth of her daughter Gaia, and then five years later following the birth of her twins. This was followed by another attack on Gremlin six years later following the birth of her next baby. Researchers could only speculate that the attacks on infants of other females had to do with food competition, as each female strives to secure resources for her own group of offspring and thus

27720-1011: The first fully sequenced termite genome, of Zootermopsis nevadensis , which was published in the journal Nature Communications , consists of roughly 500Mb, while two subsequently published genomes, Macrotermes natalensis and Cryptotermes secundus , are considerably larger at around 1.3Gb. External phylogeny showing relationship of termites with other insect groups:  (Mantises) Blaberoidea Corydiodea Blattoidae Lamproblattidae Cryptocercidae (brown-hooded cockroaches) Isoptera (Termites) Internal phylogeny showing relationship of extant termite families: Mastotermitidae Stolotermitidae Hodotermopsidae Hodotermitidae Archotermopsidae Kalotermitidae Stylotermitidae Serritermitidae Rhinotermitidae Termitidae There are currently 3,173 living and fossil termite species recognised, classified in 12 families; reproductive and/or soldier castes are usually required for identification. The infraorder Isoptera

27940-399: The first time scientists had observed chimpanzees "deliberately seek out, attack and leave for dead" chimps from another community, and it has been described as "the first record of lasting 'warfare' among [non-human] primates." Several families within the Kasekela chimpanzee community have been particularly prominent in books and documentaries. The F-family has produced five alpha males for

28160-485: The first wild chimpanzee to have his life documented from birth to death. Flint had difficulty weaning , and became distraught and angry when the birth of his sister Flame displaced him as Flo's baby. After Flame disappeared, possibly by the hand of Flint, he regained his enthusiasm, but resumed his needy, infantlike behaviour. After his nephew Freud was born, Flint became fascinated with the new infant and became an important influence of Freud's first year of life. Following

28380-518: The flock. He leads the group but shares leadership on a foraging expedition with a mature she-goat who will normally outlast a succession of dominant males. However, earlier work showed that leadership orders in goats were not related to age or dominance. In sheep, position in a moving flock is highly correlated with social dominance, but there is no definite study to show consistent voluntary leadership by an individual. In birds, dominant individuals preferentially select higher perches to put themselves in

28600-557: The forehead that lead to a gland which exudes defensive secretions, are a feature of the clade Neoisoptera and are present in all extant taxa such as Rhinotermitidae. The majority of termite species have mandibulate soldiers which are easily identified by the disproportionately large sclerotized head and mandibles. Among certain termites, the soldier caste has evolved globular (phragmotic) heads to block their narrow tunnels such as seen in Cryptotermes . Amongst mandibulate soldiers,

28820-702: The fungus Termitomyces ; in contrast, basal Neoisopterans and all other Euisoptera have flagellates and prokaryotes in their hindguts. Extant families and subfamilies are organized as follows: Termites are found on all continents except Antarctica . The diversity of termite species is low in North America and Europe (10 species known in Europe and 50 in North America), but is high in South America , where over 400 species are known. Of

29040-696: The genus Cryptocercus . Previous estimates suggested the divergence took place during the Jurassic or Triassic . More recent estimates suggest that they have an origin during the Late Jurassic , with the first fossil records in the Early Cretaceous . Similarly to ants and some bees and wasps from the separate order Hymenoptera , most termites have an analogous "worker" and "soldier" caste system consisting of mostly sterile individuals which are physically and behaviorally distinct. Unlike ants, most colonies begin from sexually mature individuals known as

29260-422: The group. In the red fox it has been shown that subordinate individuals, given the opportunity to desert, often do not due to the risk of death and the low possibility that they would establish themselves as dominant members in a new group. Animal decisions regarding involvement in conflict are defined by the interplay between the costs and benefits of agonistic behaviors. When initially developed, game theory ,

29480-573: The herd can travel. In hamadryas baboons, several high-ranking males will share a similar rank, with no single male being an absolute leader. Female bats also have a somewhat fluid social structure, in which rank is not strongly enforced. Bonobos are matriarchal, yet their social groups are also generally quite flexible, and serious aggression is quite rare between them. In olive baboons, certain animals are dominant in certain contexts, but not in others. Prime age male olive baboons claim feeding priority, yet baboons of any age or sex can initiate and govern

29700-419: The hierarchy, but it appears he will never become alpha male. However, he did help his younger brother Ferdinand attain alpha status in 2008, and continued to provide important support for Ferdinand in his brother's role as alpha, although as of early 2014 he would sometimes ally himself with Ferdinand's potential rival Titan. He is the father of Gaia's son Google, born in 2010. Ferdinand (b. August 19, 1992)

29920-416: The high-ranking males. In African wild dogs which live in social packs separated into male and female hierarchies, top-ranking alpha females have been observed to produce 76–81% of all litters. Subordinate animals engage in a number of behaviors in order to outweigh the costs of low rank. Dispersal is often associated with increased mortality and subordination may decrease the potential benefits of leaving

30140-565: The higher ranked the offspring will be (Yahner). Similarly, the status of a male Canada goose is determined by the rank of his family. Although dominance is determined differently in each case, it is influenced by the relationships between members of social groups. Individuals with greater hierarchical status tend to displace those ranked lower from access to space, to food and to mating opportunities. Thus, individuals with higher social status tend to have greater reproductive success by mating more often and having more resources to invest in

30360-598: The highest developmental potentiality and are able to molt into Alates , Soldiers , Neotenics , or Workers . Workers are believed to have evolved from larvae, sharing many similarities to the extent that workers can be regarded as "larval", in that both lack wings, eyes, and functional reproductive organs while maintaining varying levels of developmental flexibility, although usually to a much lesser extent in workers. The main distinction being that while larvae are wholly dependent on other nestmates to survive, workers are independent and are able to feed themselves and contribute to

30580-639: The highest phenotypic potency and can be described as totipotent ( able to molt into all alternative phenotypes ), whereas following instars can be pluripotent ( able to molt into reproductives and non-reproductives but cannot molt into at least one phenotype ), to multipotent ( able to molt into either reproductive or non-reproductive phenotypes ), to unipotent ( able to molt into developmentally close phenotypes ), and then finally committed ( no longer able to change phenotype, functionally an adult.) In most termites, phenotypic potency decreases with every successive molt. Notable exceptions are basal taxa such as

30800-415: The highest ranking female (queen) and her ability to suppress critically important reproductive hormones in male and female sub-dominants. In sub-dominant males, it appears that luteinizing hormone and testosterone are suppressed, while in females it appears that the suppression involves the entire suppression of the ovarian cycle . This suppression reduces sexual virility and behavior and thus redirects

31020-594: The highest-ranking vervet females have greater access than subordinates females to water in tree holes. In chacma baboons , the high-ranking males have the first access to vertebrate prey that has been caught by the group, and in yellow baboons the dominant males feed for longer without being interrupted. In many bird species, the dominant individuals have higher rates of food intake. Such species include dark-eyed juncos and oystercatchers . The dominant individuals in these groups fill themselves up first and fill up more quickly, so they spend less time foraging, which reduces

31240-436: The immatures. As a result the caste system of termites consists mostly of neotenous or juvenile individuals that undertake the most labor in the colony, which is in contrast to the eusocial Hymenoptera where work is strictly undertaken by the adults. The developmental plasticity in termites can be described similarly to cell potency , where each molt offers a varying level of phenotypic potency. Early instars typically exhibit

31460-447: The infant died just five months later. Gaia's second infant was stillborn, but her third pregnancy resulted in twins. However, once again her mother took over the babies, and one of the twins died at a few days old. She finally gave birth to a son named Google in 2010, fathered by Faustino. She gave birth to a son named Gabo in 2015. Glitter and Golden (b. July 1998) are the twin daughters of Gremlin, fathered by Frodo. Gaia contributed to

31680-548: The influence of Latin terere ("to rub, wear, erode") from the earlier word tarmes . A termite nest is also known as a termitary or termitarium (plural termitaria or termitariums ). The word was first used in English in 1781. Earlier attested designations were "wood ants" or "white ants", though these may never have been in wide use as termites do not exist in the British Isles . Termites were formerly placed in

31900-466: The interaction may walk over the subordinated, that in turn assumes a prostrated posture. To be effective, these regulatory mechanisms must include traits that make an individual rank position readily recognizable by its nestmates. The composition of the lipid layer on the cuticle of social insects is the clue used by nestmates to recognize each other in the colony, and to discover each insect's reproductive status (and rank). Visual cues may also transmit

32120-399: The king will continue to mate with the queen throughout their lives. In some species, the abdomen of the queen swells up dramatically to increase fecundity , a characteristic known as physogastrism . Depending on the species, the queen starts producing reproductive alates at a certain time of the year, and huge swarms emerge from the colony when nuptial flight begins. These swarms attract

32340-580: The large mounds constructed by certain species. Termites have several impacts on humans. They are a delicacy in the diet of some human cultures such as the Makiritare in the Alto Orinoco province of Venezuela , where they are commonly used as a spice. They are also used in traditional medicinal treatments of various diseases and ailments, such as influenza, asthma, bronchitis, etc. Termites are most famous for being structural pests; however,

32560-829: The level of developmental plasticity even between closely related taxa, with many species having true workers that can molt into the other apterous castes such as ergatoids (worker reproductive; apterous neotenics), soldiers, or the other worker castes. Pseudergates sensu stricto are individuals which arise from the linear developmental pathway that have regressively molted and lost their wing buds, and are regarded as totipotent immatures. They are capable of performing work but are overall less involved in labor and considered more cooperative than truly altruistic. Pseudergates sensu lato , otherwise known as false workers , are most represented in basal lineages ( Kalotermitidae , Archotermopsidae , Hodotermopsidae , Serritermitidae ) and closely resemble true workers in which they also perform most of

32780-439: The linear developmental pathway, soldiers develop from apterous immatures and constitute the only true sterile caste in these taxa. The primary reproductive caste of a colony consists of the fertile adult ( imago ) female and male individuals, colloquially known as the queen and king. The queen of the colony is responsible for egg production of the colony. Unlike in ants, the male and female reproductives form lifelong pairs where

33000-565: The longer they spend partaking in these high-energy activities, and they lose rank as a function of age. In wild male baboons, the highest-ranking male, also known as the alpha, experiences high levels of both testosterone and glucocorticoid, which indicates that high-ranking males undergo higher levels of stress which reduces fitness. Reduced health and longevity occurs because these two hormones have immunosuppressant activity, which reduces survival and presents opportunities for parasitic infestation and other health risks. This reduced fitness due to

33220-452: The males, which results in lethal aggression and fatality. The bodies of Fifi and Furaha were never found, but the confirmed death of fellow Kasekela female, Patti, in 2005, by neighbouring Mitumba community males provides a near definitive conclusion of the fate that befell Fifi and her youngster. Flint (1964–1972) was Flo's third son, and her first infant born after Goodall began observing the community. Goodall had intended that Flint become

33440-444: The mandibles have been adapted for a variety of defensive strategies: Biting/crushing ( Incisitermes ), slashing ( Cubitermes ), slashing/snapping ( Dentispicotermes ), symmetrical snapping ( Termes ), asymmetrical snapping ( Neocapritermes ), and piercing ( Armitermes ). In the more derived termite taxa, the soldier caste can be polymorphic and include minor and major forms. Other morphologically specialized soldiers includes

33660-412: The mating season. In many monogamous bird species, the dominant pairs tend to get the best territories, which in turn promote offspring survival and adult health. In dunnocks, a species of bird that experiences many mating systems, sometimes individuals will form a group that will have one dominant male who achieves all of the mating in the group. In the monogynous bee species Melipona subnitida ,

33880-463: The medial prefrontal cortex of pyramidal neurons , the primary excitatory cell type of the brain. High ranking macaques have a larger rostral prefrontal cortex in large social groups. Neuroimaging studies with computer stimulated hierarchal conditions showed increased activity in the ventral and dorsolateral prefrontal cortex, one processing judgment cues and the other processing status of an individual. Other studies have determined that lesions to

34100-412: The most basal. True workers are individuals which irreversibly develop from the apterous lineage and have completely forgo development into a winged adult. They display altruistic behaviors and either have terminal molts or exhibit a low level of phenotypical potency. True workers across different termite taxa ( Mastotermitidae , Hodotermitidae , Rhinotermitidae & Termitidae) can widely vary in

34320-497: The most offspring. The same pattern is found in most carnivores, such as the dwarf mongoose . The dwarf mongoose lives in a social system with one dominant pair. The dominant female produces all or almost all of the offspring in the living group, and the dominant male has first access to her during her oestrus period. In red deer, the males who experienced winter dominance, resulting from greater access to preferred foraging sites, had higher ability to get and maintain larger harems during

34540-663: The most primitive living termite, have been discovered in the Permian layers in Kansas. It is even possible that the first termites emerged during the Carboniferous . The folded wings of the fossil wood roach Pycnoblattina , arranged in a convex pattern between segments 1a and 2a, resemble those seen in Mastotermes , the only living insect with the same pattern. Kumar Krishna et al. , though, consider that all of

34760-461: The necessary gut symbionts are transferred from one generation to another. Workers are believed to have evolved from older wingless immatures ( Larvae ) that evolved cooperative behaviors; and indeed in some basal taxa the late instar larvae are known to undertake the role of workers without differentiating as a true separate caste. Workers can either be male or female, although in some species with polymorphic workers either sex may be restricted to

34980-625: The nest, though a "dwarf" queen will take its place in the case of a premature death. Dominance hierarchies emerge as a result of intersexual and intrasexual selection within groups, where competition between individuals results in differential access to resources and mating opportunities. This can be mapped across a spectrum of social organization ranging from egalitarian to despotic, varying across multiple dimensions of cooperation and competition in between. Conflict can be resolved in multiple ways, including aggression, tolerance, and avoidance. These are produced by social decision-making, described in

35200-565: The offspring of high-ranking individuals have better fitness and thus an increased rate of survival. This is most likely a function of two factors: The first is that high-ranking males mate with high-ranking females. Assuming their high rank is correlated with higher fitness and fighting ability, this trait will be conferred to their offspring. The second factor is that higher-ranking parents probably provide better protection to their offspring and thus ensure higher survival rates. Amongst rhesus macaques, higher-ranking males sired more offspring, though

35420-470: The order Isoptera. As early as 1934 suggestions were made that they were closely related to wood-eating cockroaches (genus Cryptocercus , the woodroach) based on the similarity of their symbiotic gut flagellates. In the 1960s additional evidence supporting that hypothesis emerged when F. A. McKittrick noted similar morphological characteristics between some termites and Cryptocercus nymphs . In 2008 DNA analysis from 16S rRNA sequences supported

35640-527: The outcome of a fight. Less injury will occur if subordinate individuals avoid fighting with higher-ranking individuals who would win a large percentage of the time — knowledge of the pecking order keeps both parties from incurring the costs of a prolonged fight. In hens, it has been observed that both dominants and subordinates benefit from a stable hierarchical environment, because fewer challenges means more resources can be dedicated to laying eggs. In groups of highly related individuals, kin selection may influence

35860-583: The position of termites being nested within the evolutionary tree containing the order Blattodea , which included the cockroaches. The cockroach genus Cryptocercus shares the strongest phylogenetical similarity with termites and is considered to be a sister-group to termites. Termites and Cryptocercus share similar morphological and social features: for example, most cockroaches do not exhibit social characteristics, but Cryptocercus takes care of its young and exhibits other social behaviour such as trophallaxis and allogrooming . Termites are thought to be

36080-446: The potential reproductive females. In eusocial insects, aggressive interactions are common determinants of reproductive status, such as in the bumblebee Bombus bifarius , the paper wasp Polistes annularis and in the ants Dinoponera australis and D. quadriceps . In general, aggressive interactions are ritualistic and involve antennation (drumming), abdomen curling and very rarely mandible bouts and stinging. The winner of

36300-406: The prefrontal cortex (when the area is severed to disrupt functioning to observe its role in behavior) led to deficits in processing social hierarchy cues, suggesting this area is important in regulating this information. Although the prefrontal cortex has been implicated, there are other downstream targets of the prefrontal cortex that have also been linked in maintaining this behavior. This includes

36520-433: The previous alpha male, which allowed Fudge to attain his alpha status. In 2019 Fundi helped Ferdinand rejoin the core group. Melissa (c.1950–1986) was a high-ranking female and mother of long-term alpha male Goblin. Researchers also suspected that she was related to alpha male Humphrey and another male, Mr McGregor. She was afflicted by polio in the 1966 epidemic, and for a while was paralyzed in her neck and shoulders, and

36740-642: The process. Although males are perceived as the more aggressive sex in chimpanzees, studies at Gombe have revealed that aggression among females with each other does indeed occur. One of two contexts for female-female aggressive encounters in Gombe was the presence of newborn infants. Fifi grew to be a formidable female who, although a very successful mother herself, was observed to have carried out ferocious and infanticidal attacks on other, lower-ranking females in her community. Such attacks were observed of Fifi and her adult daughter, Fanni, aggressively harassing fellow Kasekela female, Gremlin, on three different occasions;

36960-402: The queen from the colony but did not remove her bedding. They reasoned that if a primer pheromones were on the bedding then the sub-dominant's reproductive function should continue to be suppressed. Instead however, they found that the sub-dominants quickly regained reproductive function even in the presence of the queen's bedding and thus it was concluded that primer pheromones do not seem to play

37180-971: The queen seeks to maintain reproductive success by preventing workers from caring for their cells, pushing or hitting them using her antennae. Workers display aggression towards males, claiming priority over the cells when males try to use them to place eggs. There are costs to being of a high rank in a hierarchical group which offset the benefits. The most common costs to high-ranking individuals are higher metabolic rates and higher levels of stress hormones. In great tits and pied flycatchers , high-ranking individuals experience higher resting metabolic rates and therefore need to consume more food in order to maintain fitness and activity levels than do subordinates in their groups. The energetic costs of defending territory, mates, and other resources can be very consuming and cause high-ranking individuals, who spend more time in these activities, to lose body mass over long periods of dominance. Therefore, their physical condition decreases

37400-454: The queen's extremely high levels of circulating testosterone, which cause her to exert intense dominance and aggressiveness on the colony and thus "scare" the other mole-rats into submission. Research has shown that removal of the queen from the colony allows the reestablishment of reproductive function in sub-dominant individuals. To see if a priming pheromone secreted by the queen was indeed causing reproductive suppression, researchers removed

37620-415: The queen) are diploid and develop from fertilised eggs. In contrast, worker termites, which constitute the majority in a colony, are diploid individuals of both sexes and develop from fertilised eggs. Depending on species, male and female workers may have different roles in a termite colony. The life cycle of a termite begins with an egg , but is different from that of a bee or ant in that it goes through

37840-427: The raising of Gremlin’s twins, helping them to become the only known pair of wild chimpanzee twins known that have survived to the age of sexual maturity. Each of the pair exhibits a different personality. Glitter is shy and cautious, while Golden is more adventurous and willing to engage in rough and tumble activities. In 2011, Glitter gave birth to daughter named Gossamer, who was fathered by Sheldon. She gave birth to

38060-409: The reproduction costs of the worker caste are compensated by the contribution of workers to the queen's reproductive success, with which they share genes. This is true not only for the popular social insects ( ants , termites , some bees and wasps ), but also for the naked mole-rat Heterocephalus glaber . In a laboratory experiment, Clarke and Faulkes (1997) demonstrated that reproductive status in

38280-456: The researchers, which likely helped his recovery. In 2004, he once again began travelling with the rest of the community, but was unable to re-establish his alpha status. In his later years, Frodo mellowed. He became less aggressive and spent much time by himself, although he could still intimidate other chimpanzees on occasion. Frodo died in 2013. Frodo's aggression was not limited to colobus monkeys and other chimpanzees. In May 2002, he killed

38500-443: The risk of predation. Thus they have increased survival because of increased nutrition and decreased predation. In primates, a well-studied group, high rank brings reproductive success, as seen in a 1991 meta-analysis of 32 studies. A 2016 study determined that higher status increased reproductive success amongst men, and that this did not vary by type of subsistence (foraging, horticulture, pastoralism, agriculture). This contradicts

38720-474: The role. While Freud was alpha Goblin was able to be a wedge between Freud and Frodo, who had previously been close companions, but when Frodo overtook Freud, Goblin switched his allegiance to Frodo. Goblin died of disease in August 2004. After recovering from his injuries, Goblin maintained a high-ranking status within the community by forming alliances with the alpha males, especially Freud and Frodo. While he

38940-471: The same information. Paper wasps Polistes dominulus have individual "facial badges" that permit them to recognize each other and to identify the status of each individual. Individuals whose badges were modified by painting were aggressively treated by their nestmates; this makes advertising a false ranking status costly, and may help to suppress such advertising. Other behaviors are involved in maintaining reproductive status in social insects. The removal of

39160-430: The size of the colony. Since larvae and nymphs are unable to feed themselves, workers must feed them, but workers also take part in the social life of the colony and have certain other tasks to accomplish such as foraging, building or maintaining the nest or tending to the queen. Pheromones regulate the caste system in termite colonies, preventing all but a very few of the termites from becoming fertile queens. Queens of

39380-415: The social order of the paper wasp ( Polistes dominulus ). This depends on the queen (or foundress), possibly involving specific hormones. Laboratory experiments have shown that when foundresses are injected with juvenile hormone , responsible for regulating growth and development in insects including wasps, the foundresses exhibit an increase in dominance. Further, foundresses with larger corpora allata ,

39600-516: The species Polistes instabilis , where the next queen is selected based on age rather than size. Polistes exclamans also exhibits this type of hierarchy. Within the dominance hierarchies of the Polistes versicolor , however, the dominant-subordinate context in the yellow paper wasps is directly related to the exchange of food. Future foundresses within the nest compete over the shared resources of nourishment, such as protein. Unequal nourishment

39820-414: The species, including ritualized displays of aggression or direct physical violence. In social living groups, members are likely to compete for access to limited resources and mating opportunities. Rather than fighting each time they meet, individuals of the same sex establish a relative rank, with higher-ranking individuals often gaining more access to resources and mates. Based on repetitive interactions,

40040-414: The stability of hierarchical dominance. A subordinate individual closely related to the dominant individual may benefit more genetically by assisting the dominant individual to pass on their genes. Alpha male savanna baboons have high levels of testosterone and stress; over a long period of time, this can lead to decreased fitness. The lowest-ranking males also had high stress levels, suggesting that it

40260-409: The stranger is female, it depends on whether or not they are of any reproductive benefit to the males. Young, oestrus females are often spared an attack because they are deemed to be potential new mates, but an anestrus female (non-cycling or lactating) is considered of very little use for the males, which results in lethal aggression and fatality. Tapit (1979–1983) was Patti's second-oldest son. He had

40480-419: The strategies of individuals engaged in conflict has proven integral to establishing social hierarchies reflective of dominant-subordinate interactions. The asymmetries between individuals have been categorized into three types of interactions: As expected, the individual who emerges triumphant is rewarded with the dominant status, having demonstrated their physical superiority. However, the costs incurred to

40700-529: The study of optimal strategies during pair-wise conflict, was grounded in the false assumption that animals engaged in conflict were of equal fighting ability. Modifications, however, have provided increased focus on the differences between the fighting capabilities of animals and raised questions about their evolutionary development. These differences are believed to determine the outcomes of fights, their intensity, and animal decisions to submit or continue fighting. The influence of aggression, threats, and fighting on

40920-416: The sub-dominant's behavior into helping the queen with her offspring, though the mechanisms of how this is accomplished are debated. Former research suggests that primer pheromones secreted by the queen cause direct suppression of these vital reproductive hormones and functions however current evidence suggests that it is not the secretion of pheromones which act to suppress reproductive function but rather

41140-445: The survival of offspring . Hence, hierarchy serves as an intrinsic factor for population control, ensuring adequate resources for the dominant individuals and thus preventing widespread starvation. Territorial behavior enhances this effect. The suppression of reproduction by dominant individuals is the most common mechanism that maintains the hierarchy. In eusocial mammals this is mainly achieved by aggressive interactions between

41360-407: The territorial males of a rival community, and the intensity of the attacks depends on the gender of the intruder, and if the stranger is female, it depends on whether or not they are of any reproductive benefit to the males. Young, cycling females are often spared an attack because they are deemed to be potential new mates, but a lactating mother with a young infant is considered of very little use for

41580-450: The top of the female hierarchy in her community. Fifi had nine infants, seven of whom survived to independence. Her two oldest sons, Freud and Frodo, both have become the community's alpha male, as did another son, Ferdinand. One of her other sons, Faustino, also attained high rank. Her three surviving daughters were Fanni, Flossi and Flirt. She also had one son, Fred (1996–1997) and one daughter, Furaha (2002–2004) who died in infancy. Fred

41800-630: The top-ranked position in 1982, and was the unambiguous alpha male, able to control even coalitions of rival males, by 1984. Goblin maintained his alpha status until 1989, when he was badly beaten and injured by a coalition of males led by Wilkie. He was sterilized by a bite to his testicles in the process. He spent time away from the rest of the community recovering from his injuries, during which time Goodall administered medication to help him return to health. He later became Freud's key ally in becoming and retaining his position as alpha male, apparently going so far as to encourage Freud to challenge Wilkie for

42020-466: The vast majority of termite species are innocuous, with the regional numbers of economically significant species being: North America , 9; Australia , 16; Indian subcontinent , 26; tropical Africa , 24; Central America and the West Indies , 17. Of known pest species, 28 of the most invasive and structurally damaging belong to the genus Coptotermes . The distribution of most known pest species

42240-415: The wild. Dominance hierarchies in small herds of domestic horses are generally linear hierarchies whereas in large herds the relationships are triangular. Dominance hierarchies can be formed at a very early age. Domestic piglets are highly precocious and, within minutes of being born, or sometimes seconds, will attempt to suckle. The piglets are born with sharp teeth and fight to develop a teat order as

42460-447: The wings. It has been proposed that the Isoptera and Cryptocercidae be grouped in the clade " Xylophagodea ". Termites are sometimes called "white ants", but the only resemblance to the ants is due to their sociality which is due to convergent evolution with termites being the first social insects to evolve a caste system more than 100 million years ago. Termite genomes are generally relatively large compared to those of other insects;

42680-575: The work and are similarly altruistic, however differ in developing from the linear developmental pathway where they exist in a stationary molt; i.e they have halted development before the growth of wing buds, and are regarded as pluripotent immatures. The soldier caste is the most anatomically and behaviorally specialized, and their sole purpose is to defend the colony. Many soldiers have large heads with highly modified powerful jaws so enlarged that they cannot feed themselves. Instead, like juveniles, they are fed by workers. Fontanelles , simple holes in

42900-485: Was 28 years old when she gave birth to Faben. Most female chimpanzees have their first infant when 12–14 years of age, so it is possible that Flo had at least one infant prior to Faben that either died or, if it was a female, would have transferred to one of the other communities prior to Goodall's arrival to Gombe in 1960. When Flo died in 1972, she was given an obituary in Britain's Sunday Times . Faben (c.1947 – 1975)

43120-405: Was Fifi's oldest son. His father was a male from outside the Kasekela community. As a juvenile, he benefited from a high-ranking mother and an uncle, Figan, who was the alpha male. In 1993, Freud became the alpha male by unseating his childhood playmate Wilkie. While the alpha male, his younger but larger brother Frodo became the second highest-ranking male in the community. In 1997, while Freud

43340-457: Was Flo's oldest known offspring. He was a powerful male, but lost the use of one arm in a polio epidemic that ravaged the community in 1966. Despite his disability, he learned to create spectacular bipedal charging displays, which allowed him to regain a high rank within the community. His support was critical to Humphrey's ascension to alpha male rank in 1969, and to his brother Figan's ascension to alpha male in 1972. Faben also participated in

43560-597: Was Flo's second son. Goodall considered him to be Gombe's most intelligent chimpanzee. Although younger than Faben, he was able to dominate his older brother after Faben's arm was paralyzed from polio in 1966. With Faben's support, Figan was able to become alpha male in 1972 by defeating the prior alpha Humphrey and an older competitor Evered. At this time, Goodall described him as the most powerful alpha male in Gombe’s recorded history. After Faben's death in 1975, Figan lost his unquestioned alpha status, in that he could be dominated by

43780-421: Was Melissa's oldest son. He was discovered by Goodall when he was just a few hours old, with a twisted face that led her to name him Goblin. Goblin was protected by Figan as an adolescent, but at the young age of 15 in 1979 he challenged and defeated Figan to become the top-ranking male. He was defeated and brutally beaten by a coalition of older males later in the year, giving up his top ranking. Goblin regained

44000-456: Was able to intimidate even the larger Frodo with his displays. Freud died in March 2014, at more than 42 years old, which made him the oldest male whose age was known in the history of the community. Freud was one of the first wild chimpanzees whose entire life from birth to old age was documented by researchers. Frodo (June 30, 1976 – November 10, 2013) was Fifi's second-oldest son. His father

44220-472: Was able to regain alpha status. Ferdinand has been successful in siring offspring. Infants he fathered included Diaz by Dilly fathered at very young age of 12 in 2005, Tabora (2006– ) by Tanga, and Gizmo (2010–2016) by Gremlin. Flirt (b. July 1998) is Fifi's youngest surviving daughter. Her father was Kris, who would later become alpha male of the troop. She was a large baby and grew rapidly. This probably allowed her to survive when her mother died before she

44440-427: Was afflicted by sarcoptic mange , Frodo was able to take over the alpha status. After losing his alpha status, Freud settled into a role as a middle-ranking male, although could still threaten higher-ranking males when aroused. He was also able to serve as an important ally to other males. Although less aggressive than his brother Frodo, Freud is known to have engaged in at least one instance of infanticide , killing

44660-465: Was also able to intimidate other chimpanzees into sharing their kills with him if he was unsuccessful. From 1990 to 1995, Frodo killed 10% of the red colobus monkeys within his home range. His large size and aggressive nature allowed him to attain high status in the group, while his brother Freud was alpha male. On October 2, 1997, while Freud and other community members were suffering from sarcoptic mange , Frodo attacked and defeated Freud, thus attaining

44880-457: Was away from the group, perhaps with a fertile female, and Ferdinand aligned with Faustino and Fundi to rejoin the core group. Fudge was temporarily excluded from the group by this coalition but was eventually able to regain alpha status. Fundi (b. 2000) is Fanni's second oldest surviving son. His father is the former alpha male Sheldon; Fudge, the alpha male as of 2016, is his full brother. In 2016 Fundi helped Fudge attack their uncle Ferdinand,

45100-420: Was badly wounded in the attack and had to leave the group to recover. In his absence, Fudge was able to assert his dominance and attain the alpha position. After falling from alpha status, Ferdinand was generally excluded from the core group of male chimpanzees. But in 2019, while Fudge was away from the group, Ferdinand aligned with his brother Faustino and nephew Fundi to rejoin the core group. However, Fudge

45320-679: Was described in birds by Thorleif Schjelderup-Ebbe in 1921 under the German terms Hackordnung or Hackliste and introduced into English in 1927. In his 1924 German-language article, he noted that "defense and aggression in the hen is accomplished with the beak ". This emphasis on pecking led many subsequent studies on fowl behaviour to use it as a primary observation; however, it has been noted that roosters tend to leap and use their claws in conflicts. Wild and feral chickens form relatively small groups, usually including no more than 10 to 20 individuals. It has been shown that in larger groups, which

45540-413: Was different from his smaller predecessor Wilkie and his larger successor Frodo. At 99 pounds (45 kg), Freud was larger than the 81-pound (37-kilogram) Wilkie and smaller than the 113-pound (51-kilogram) Frodo. Freud was less aggressive than and groomed other males to help maintain alliances more often than Frodo, but engaged in more contact aggression and less grooming than Wilkie. While alpha male he

45760-427: Was establishing himself as a contender for the alpha position, he would occasionally beat up Goodall. In 1983, when his mother Melissa was in estrus, he forced himself upon her despite her efforts to resist. Fanni and Flossi, by Fifi, and Tanga (1989– ), by Patti, are his daughters. Gremlin (b. 1970) is Melissa's only daughter to survive to adulthood. Her father was Evered, who, more than 20 years later, also fathered

45980-411: Was fathered by Wilkie, another former alpha male. Fifty was also sired by Wilkie. As of 2021, Fudge is the alpha male of the community. Whilst most female chimpanzees leave their natal communities upon reaching sexual maturity, Fanni has remained in her natal Kasekela community alongside her mother (in contrast to her younger sister Flossi). Fanni remained close to her mother Fifi, with whom she formed

46200-645: Was fathered by one of Fifi's other sons, Frodo. Five-year-old Fifi was the focal point of the documentary People of the Forest: The Chimps of Gombe , in which her mother and siblings also featured, attempting to gain access to her infant brother Flint. All four of Fifi's surviving sons had all held the alpha and/or beta positions in the community. Grandson Fudge (by Fanni) also attained alpha male status. Late in Frodo's reign as alpha male, Fifi helped defend him against attacks and she suffered some injuries in

46420-476: Was forced to walk bipedally . She would regain use of her arms, although she never fully regained movement of her neck. She gave birth to two sets of twins, one set in 1976 of which neither survived the year, and another set in 1977, of which only one, Gimble, survived to adulthood. Her other offspring who survived to adulthood was daughter Gremlin. Her daughter Gremlin and her granddaughter Gaia have also given birth to twins. Goblin (September 1964 – August 2004)

46640-404: Was his primary supporter in the attack and they received some support from their mother Fanni, who is Ferdinand's sister. After seriously wounding Ferdinand in the attack, Fudge was able to assert his dominance over the remaining group males to attain alpha male status. Ferdinand was excluded from the core group of male chimpanzees after Fudge established himself as alpha male. But in 2019, Fudge

46860-482: Was instrumental to allow him to hold on to his alpha status. In 1982, Goblin unseated him again, and Figan disappeared and presumably died a few months later, although when last seen he appeared to be in good health. Although Figan was the alpha male for several years, he is not known for certain to have sired any infants, although he is likely the father of a few. Fifi (c.1958 – 2004) was Flo's oldest daughter. Fifi benefited from her high-ranking family, and ascended to

47080-480: Was never seen again. Prof (October 27, 1971 – 198?) was a male that survived to adulthood. He and Pom offered their little brother comfort following Passion's death. Prof and Pax associated closely for a number of years. Prof is deceased. Pax (b. December 16, 1977) was Passion's last offspring. He was only four years old when he was orphaned. He had the support of an elder brother and sister, and had very few behavioural issues following his mother's death. When Pax

47300-422: Was only rarely observed, and tool creation by non-human animals had never been observed. Until then, tool making was considered one of the defining characteristics of being human. Another important observation occurred a few days earlier, on October 30, 1960. On that day Goodall observed the community's chimpanzees eating meat, dispelling the notion that chimpanzees are vegetarians . A third observation by Goodall in

47520-495: Was seven years old. Although she spent a few lonely months after Fifi's death, she would begin to travel with her brothers. The documentary Chimpanzee Family Fortunes ends with Flirt finding solace after months of loneliness by being groomed by eldest brother Freud. She followed in the footsteps of her sister Flossi in eventually emigrating to the Mitumba Chimpanzee Community in 2013. Fudge (b. 1996)

47740-402: Was taking a newborn baby from her daughter, Gaia, in 2006, presumably in fear of it being attacked. Although Gremlin's instincts were justified from her own experiences, the latter case ended with the infant dying just five months later. Gimble (b. October 1977) is one of two of Melissa's sons to survive to adulthood. He was one of a set of twins; however, his twin brother Gyre died in 1978. As

47960-399: Was the matriarch of the F-family, so named because she and all her matrilineal descendants were given names beginning with the letter "F". In 1962, Flo was one of the first chimpanzees to approach Goodall's camp, along with her infant daughter Fifi. Video of Flo approaching Goodall and allowing Fifi to reach out to touch Goodall's forehead, letting Goodall know she had been accepted, is shown in

48180-483: Was the relatively low-ranking male Sherry. Even from a young age, Frodo was unusually large and aggressive. He learned to throw rocks as a juvenile and became very accurate with his aim; he sometimes threw them at his human observers, often hitting and bruising them. As an adult, he was one of the largest chimpanzees ever observed in the community, at about 113 pounds (51 kg) and remained quite aggressive. He also became an excellent hunter of red colobus monkeys, and

48400-423: Was very young, his mother was wounded in some type of conflict. Pax lost his testicles as a result of the conflict, which has affected his personality greatly. He is small for a male, doesn't compete in the male hierarchy, and is unable to reproduce. He also has been chosen by a couple of orphans as a companion. Pan (October 17, 1978 – 1981) was the only known offspring of Pom. He was injured after being blown from

#139860