ANTH105 Quiz 1

What is Anthropology?

  • Anthropology is the broad study of humankind around the world and throughout time.

  • It is also concerned with both the biological and the cultural aspects of humans.

  • Anthropology is the study of humans– their origins, biological characteristics, cultural development, social relationships, and more– based in scientific methods

The four main subdivisions of Anthropology

  • Physical/Biological Anthropology

  • Archeology

  • Cultural Anthropology- The material aspects of society, the norms and values shared among people, transmitted and learned

  • Linguistic Anthropology- The study of language and how it varies across different societies

  • Fifth field: Applied Anthropology- Taking anthropology and applying it to the real world

Physical/Biological Anthropology

  • How the human species has changed physically over time (hundreds of millions of years)

    • Biological evolution

  • Subfields:

    • Primatology- the study of primates

    • Genetic inheritance- how physical characteristics are transmitted and evolving over time

    • Paleoanthropology- researching our human ancestors and looking at the fossil evidence for human evolution

    • Human biology- modern human health and disease

    • Bioarchaeology- human culture through human remains

    • Human behavioral ecology- how humans adapt to their local environment, mainly focusing on the physical environment

What is evolution?

  • Evolution is the change in inherited characteristics of biological populations over successive generations.

  • It is also the change in the genetic structure of a population

  • Evolution is NOT-

    • Moving toward some sense of perfection

    • Consistent

Definitions

  • Adaptation- a feature of an organism created by the process of natural selection.

  • Adaptive trait- a heritable trait that increases the probability of survival and/or reproduction in a given environment.

    • An adaptive trait in one population may be maladaptive in another.

    • Examples: thick fur, colored fur, altered eyesight

The History of Evolutionary Theory

  • Pre-Darwin thought

    • Species are fixed and unchanging

  • Aristotle (300ish BC): the Great Chain of Being

    • Unchangeable

  • Linnaeus (1700s)

    • Father of modern taxonomy 

    • Binomial nomenclature (Homo sapiens)

    • Fixity of species

  • James Hutton (1700s)

    • Father of modern geology

    • Uniformitarianism/gradualism- Earth changes very slowly over time

    • Influenced Darwin’s ideas

  • Jean Baptiste-Lamark (1744-1829)

    • Simple to complex evolution

    • Coined the term evolution- change over time

    • Main ideas

      • Animals inherit traits from previous generations

      • Change happens over time

      • Change has something to do with the environment

    • “Acquired traits”

    • Law of use/disuse

  • Thomas Malthus (1766-1834)

    • Studied population dynamics

    • Populations struggle when resources are limited

  • George Cuvier (1769-1832)

    • Paleontologist

    • Fossils can be used to assess time for life on Earth

    • Catastrophism- when animals go extinct, mass extinctions

  • Erasmus Darwin (1731-1802)

    • Charles Darwin’s grandfather

    • A proponent of the theory that species change over time

  • Alfred Russel Wallace (1823-1913)

    • Came up with the theory of natural selection independent of Darwin

  • Charles Lyell (1797-1875)

    • Geologist

    • Earth is changing

    • Earth changes gradually and in uniformed and predictable processes

Paving the way for Darwin

  • Theorists who cast doubt on Divine Creation

    • Georges Cuvier

    • Charles Lyell

    • Erasmus Darwin

  • Contributing to the theory of natural selection

    • James Hutton

    • Jean-Baptists Lamarck

    • Thomas Malthus

    • Alfred Russel Wallace

Charles Darwin

  • Medical student

  • Interest in natural history

  • Meant to become an Anglican parson

  • Invited to be on the voyage of the HMS Beagle

    • Allowed him to take extensive notes and learn about fossils and marine creatures

    • Served as a companion to the captain

  • Poor health

  • Published 3 geological books within 6 years

  • 1858: began writing On the Origin of Species

Darwin and Wallace

  • Alfred Russel Wallace

    • Wrote “On the Law which has Regulated the Introduction of New Species”

  • Darwin quickly wrote a short paper explaining natural selection

  • Later published On the Origin of Species

    • Nearly died

  • Darwin’s contribution: the mechanism- how the change happens

Eastern Evolutionary Thought

  • People in the Middle East had ideas on evolution before Darwin

    • Al-Jahiz, Iraq, (776-868 CE)

    • Ibn Miskawayh, Persia, (930-1030)

    • Nasir ad-Din Tusi, Persia, (1201-1274)

    • Ibn Khaldun, North Africa, (1322)

Darwin’s Postulates

  • There is a struggle for existence.

  • There is variation in features related to survival and reproduction.

  • This variation is passed from generation to generation (or from parent to offspring).

  • These three postulates make evolution by means of natural selection inevitable

Survival of the Fittest?

  • Survival of the fit enough.

  • NOT coined by Charles Darwin

  • Social Darwinism at its worst

More definitions

  • Fitness- An individual’s genetic representation in future generations

Social Darwinism

  • “Survival of the fittest” applied to:

    • Eugenics (Margaret Sanger)

    • Racism

  • Francis Galton

    • Darwin’s cousin

An Example of Adaptation by Natural Selection

  • Galapagos Islands, Ecuador

    • Darwin’s finches

    • Peter and Rosemary Grant, Princeton

  • Postulate 1: struggle for existence

    • 1200 birds reduced to 180

  • Postulate 2: variation

    • Beak depth varied

    • Influenced survival

  • Postulate 3: heritable

    • Beak depth passed from parent to offspring

Darwin’s Difficulties Explaining Variation

  • Questions?

    • Blending inheritance?

    • How is variation maintained if natural selection eliminates it?

    • Where do novel forms come from if not already present in the original population?

  • Natural selection produces variation.

    • Darwin thought that offspring were the blend of two parents. He was very wrong.

Issues with Darwin’s Theory

  • Blending inheritance could not work.

  • It would cause a reduction in the variation of traits over time.

Gregor Mendel

  • Monk

  • Discovered laws of inheritance

  • Bred garden pea plants

  • Worked with traits with two variants

Mendelian Genetics

  • Crossbreeding experiments

  • Independent assortment

    • Each of these particles is equally likely to be transmitted from eggs and sperm

  • Mendel’s work was “rediscovered” in early 1900s.

  • Chromosomes discovered

    • Chromosomes are the small living structures in cells that divide and replicate

  • Gene

    • Different versions are “alleles”

    • If alleles are the same: homozygous

    • If alleles are different: heterozygous

  • Genotype

    • Alleles carried by organism

  • Phenotype

    • Observable characteristics

  • Dominant

  • Recessive

Modern Synthesis: Key Concepts

  • Natural selection operates on the phenotype, not the genotype.

  • Natural selection can change gene frequencies in a population.

Mechanisms of Evolutionary Change

  • Natural selection results in genetic adaptation.

  • Genetic drift- random changes

  • Gene flow- exchange of genetic info between populations

  • Mutation- alters the bases in the DNA molecule, happens by chance

Types of Adaptation

  • Acclimatization- response to environmental pressures

  • Developmental- occurs during development

  • Behavioural- learned adaptations to environment

  • Psychological- a mental change to better navigate the world

Are we still evolving?

  • It is a never-ending process.

  • Our environment is still changing.

  • Evolution doesn’t care about health. It maximizes reproduction.


Human Evolution

  • 200,000 YA- the first humans appeared

  • 60,000-40,000 YA “The Great Leap Forward”

  • 10,000-now- from hunter-gather to frustrated commuter

Sociobiology

  • E. O. Wilson: “Genetic leash”

  • Sociobiology

    • Differential reproductive success shapes the evolution of social behaviors of ALL organisms

  • Genetic determinism: driven by genes

  • Humans were “vehicles” that genes used to reproduce themselves


E. O. Wilson and Richard Dawkins

  • Sociobiology: The New Synthesis by Wilson

  • Selfish Gene by Dawkins

  • How does one explain altruism?


Critiques of Sociobiology

  • Ignored culture and learning

  • The research was conducted on animals and plants, not humans

  • Assumed specific human behaviors could be linked to genes


Modern Approaches to Evolution and Human Behaviour

  • Human behavioral ecology

    • The study of the adaptive nature of human behavior

    • Mating, parenting, division of labor, cooperation

    • HBE examines how environmental and ecology factors influence human behaviour

    • How do we measure reproductive benefits?

      • Counting babies: not feasible

      • What about culture?

  • Evolutionary Psychology

    • The study of human adaptation from a psychological perspective

    • Reproductive success, genetics, phonetics adaptation

    • EEA- the environment of evolutionary adaptiveness

    • Pleistocene prior to cultural revolution

    • Environmental mismatch

    • The aim is to identify the selection pressures that shape the human psyche in our evolutionary history

      • Repeatedly encountered problems

    • The human brain - like any other organ - is comprised of specialized “domain-specific” modules that cope with a range of adaptive problems

    • W.E.I.R.D.

      • Pencil and paper methods: data is almost entirely self-report with a small sample of modern humans

  • Gene-culture coevolution/Cultural evolution


Anthropology and Evolution

  • The first anthropologists of the late 1800s utilized evolution but not Darwin's principles (natural selection)

  • Unilineal evolution- cultures change over time

  • Continuous change toward “perfection”


Herbert Spencer

  • Interested in evolution as a general phenomenon

  • It has little to do with variation and reproduction

  • Focused on Lamarkian concepts of evolution

    • Ignored natural selection


Henry Morgan (1818-1881) and Sir Edward Burnett Tylor (1832-1917)

  • Unilineal evolutionists

  • Morgan:

    • Savagery

    • Barbaris

    • Civilization

  • Tylor:

    • Animism

    • Polytheism

    • “Enlightened” monotheism

  • Ethnocentric- belief that your culture is superior to others

  • Cultural relativism- observing a culture in its own light (nonjudgemental)


Leslie White (1943)

  • Cultural evolution leads to increased complexity

  • Developed a non-ethnocentric way to measure cultural evolution

  • Culture advances as “the amount of energy per capita increases”


Julian Steward (1902-1972)

  • Environment impacts human behavior

  • Multilinear evolution: culture can evolve in any number of distinct  patterns depending on environmental circumstances


Key concepts:

  • Phenotypic flexibility- changing behavior to match the environment

  • Environmental conditions include: ecological dimensions, cognitive, social, and political

  • Intrinsic constraints are particular to the organism, such as physiological limitations

    • Height, dietary restrictions, allergies, reading level, mobility

  • Extrinsic constraints are due to the environment (man-made or not)

    • Famine, drought, food desert, flooding, prices, deforestation

Overview

  • Microevolution: changes within a species

  • Macroevolution: the process of how new species evolve

  • Ligers


What is a species?

  • Biological Species Concept

  • Species: life can be clustered into distinct types where the individuals can produce fertile offspring

    • Ernst Mayr

  • However, species are difficult to define

    • Often they do not follow the “rules”

    • Liges!

      • Liligers?

  • Biological species concept: a group of organisms that interbreed in nature and are reproductively isolated

  • Gene flow: members of the same species share genes

  • Offspring are infertile


The Ecological Species Concept

  • Reproductive isolation is not necessary

  • Natural selection keeps species distinct from one another

    • Intermediate cannot survive because of selection pressures

  • Support:

    • Almost half of all species are not reproductively isolated

    • Some species have little to no gene flow

  • Both reproductive isolation and natural selection are important

  • Offspring don’t survive


The Origin of Species

  • Three types:

    • Allopatric

    • Parapatric

    • Sympatric

  • Allopatric Speciation

    • Geographic isolation

    • Combination of reproductive isolation and natural selection

    • Reinforcement: mating behaviors may evolve differences, which ultimately leads to reproductive isolation

    • Don’t recognize each other as mates

  • Parapatric Speciation

    • “Next-door neighbors”

    • Distinct species with hybrid zones

    • Natural selection + partial reproductive isolation keeps species separate

    • Difficult to observe in nature

    • More common in marine species because total geographic isolation is not possible

  • Sympatric Speciation

    • Natural selection may favor two equally successful solutions to the same problem

    • Species stay in one ecosystem even though there is no geographic barrier

    • Only uses natural selection

Why do we study primates?

  • We share similarities in behavior and anatomy

  • They provide models of how adaptation shapes environments that shaped our ancestors

  • Homology: the existence of shared structure in genes between different species, there is a shared ancestor to create this

  • Analogy: the existence of similar traits in two different species, but the species are not related, you can understand what selection pressures were present to give rise to the trait


Conversion Evolution

  • Evolutionary process by which similar traits/adaptations emerge in unrelated species

  • Happens with analogy


Primates are diverse!

  • Body size

  • Social structures

  • Diet

  • Nocturnal vs Diurnal

  • Behaviors

  • Locomotion

  • Habitats


What are the characteristics that primates have in common?

  • Hind-limb dominant locomotion

  • Nails instead of claws

  • Grasping hands and feet

  • Reduced olfaction and enhanced vision

  • Foward-facing eyes encased in bone

  • Relatively large brain

  • Long gestation

  • Small litters

  • Long juvenile period

  • Long lifespan

  • Increased dependence on learning and behavioral flexibility


Primate Biogeography

  • Tropical animals

  • Mostly forest-dwelling, arboreal animals

  • Modern range

    • Americas, Africa, Asia

  • Ancient range

    • Included North America, Europe


Key Features of Primates

  • How does the diet of a primate impact its home range?

    • Availability

    • Seasonality of food

  • Some primates are quite territorial

  • Home ranges do not overlap

  • Why so territorial?

    • Lemurs

    • Gibbons

  • Access to males and resources

  • Some primates are not as territorial

    • Home ranges do overlap

    • Territorial or not?

      • Costs/Benefits


Primate Sociality

  • Why are primates social?

    • Shared resources

    • More mates

    • Impacts hormones

    • Predator avoidance

  • Costs

    • More competition

    • Disease transmission

  • Five basic types


Mating Strategies

  • Monogamy/pair-bonding

    • One male, one female

  • Polyandry

    • One female, multiple males

  • Polygyny

    • One male, multiple females

  • Polygynandry

    • Multiple males, multiple females


Primate Taxonomy

  • There are approximately 300 different species of primates in the world

  • They are not all equally related to one another

  • Using genetic and morphological data, scientists can split the primate order into two large suborders:

    • Strepsirrhini (strepsirrhines)

    • Haplorrhini (haplorrhines)


Strepsirrhines

  • Tooth comb

  • Slightly smaller brain

  • Tapetum (reflective layer in back of the eye)

  • Divided into to infraorders

    • Lemuriformes (lemurs)

    • Lorisformes (lorises and galagos)


Hominoids (apes)

  • Lesser apes

  • Orangutans

  • Great apes

  • Family Hominiade (great apes)

    • Orangutans (Pongo)

    • Gorillas (Gorilla)

    • Chimpanzees (Pan)

    • Bonobos (Pan)

    • Humans (Homo)

Overview

  • Humans are equally related to chimpanzees and bonobos

  • Humans shared a common ancestor with chimpanzees and bonobos 5 to 7 million years ago


History of Research: Chimpanzees

  • 1925-1940: studies on captive chimpanzees

    • The Mentality of Apes

  • 1960s-1970s: studies on wild chimpanzees

  • Jane Goodall: Gombe Stream National Park

  • Toshisada Nishida: Mahale Mountains


History of Research: Bonobos

  • 1928: Harold Coolidge and Ernst Schwartz discovered the “pygmy chimpanzee”

  • Studied in captivity due to the Congo civil war


Current Perspectives

  • Past research:

    • Wild chomps vs. captive bonobos

  • Current:

    • Larger perspective and test subject size


Physiological Differences

  • Bonobos: “Pygmy chimp”

    • Misnomer

  • Distinct body proportions

    • Shorter upper limbs

    • Longer lower limbs

  • Bonobos: slender build, bright pink lips, black face

  • Chimpanzees: robust build, face color changes with age, dark lips


Sexual Dimorphism

  • Inherited sex differences: size, shape, color


Chimpanzee Social Organization

  • Fission-fusion:

    • Band together and disperse (size of the group in flux)

    • Large groups that form smaller parties

    • Party size depending on resources

  • Male philopatry

    • Male dominance and hierarchical

  • Females disperse in adolescence

    • 9-14 years old

    • Transition can take 2 years

  • Female hierarchy in a new group:

    • Unrelated to females

    • Age-based


Chimpanzee Parental Care

  • “Promiscuous” mating patterns

    • Among other types

  • Mothers provide majority of care

  • Infanticide has been observed in:

    • Gombe, Mahale, and Kibale

    • Explained by sexual selection

    • Uncertain paternity


Chimpanzee Mating Patterns

  • Restricitive: The dominant male restricts access to fertile females

  • Consortship: adult pair leaves the community

  • Extra group: females leave


Chimpanzee Warfare

  • Chimpanzees and Humans:

    • Only primates to engage in coalitionary warfare

  • Chimpanzees:

    • Highly territorial

    • Kill and eat other primates

  • More likely to threaten

    • Killing is the last strategy


Bonobo Social Organization

  • Fission-fusion

    • Groups range from 1 to 20 bonobos

  • Male philopatry, female dispersal

    • Females live around 7-9 years old

  • Strong female-female bonds

    • As group size increases, male-female bonds increase

  • Males and females interact to

    • Mate

    • Kinship ties

    • Rank acquisitions

      • Mother-son pairs

  • Related mixed-sex pairs interact more frequently than unrelated mixed-sex pairs

  • Strong female bonds keep males away from food


Bonobo Sexual Behaviour

  • Most defining characteristic of Bonobos

  • Purpose of sex:

    • Reproduction

    • Dominance

    • Rank


Bonobo Parental Care

  • “Promiscuous” mating patterns

  • Paternal uncertainty

    • Fathers do not invest

  • Emphasis on maternal care

  • Infanticide is rare


How do humans compare to chimpanzees and bonobos?

  • Emotions

    • Chimpanzees display similar emotions as humans

  • Nonverbal communication

    • Kissing, embracing, holding hands, hurling rocks

  • Intelligence

    • Sophisticated cooperation

  • Childhood

    • Long childhood

  • Culture!


Culture in Chimpanzees

  • Chimpanzees tool use

    • Occupy many different habitats

    • Observed in more than 25 contexts

    • Considerable variation in type and purpose

    • Mainly used to acquire insects


Tool Use in Bonobos

  • Study conducted in Lomako

  • Examined cultural patterns found in chimpanzees

    • 14 of 65 traits found in Bonobos

    • Branch dragging, leaf sponge, branch clasp, branch shake, etc.


Sex Differences

  • Bonobos

    • Females are more avid tool users than males

    • Larger tool range behavior

    • More tools for play and food acquisition

  • Chimpanzees

    • Females are more avid tool users than males

Primate Mating Systems

  • Mating systems: the way animals find mates and care for their offspring

  • We are primates.

  • There is variation in primate mating systems

  • There are some basic rules in mammals


Life History Theory

  • Three tradeoffs that all species make:

    • Reproductive effort

    • Somatic effort

    • Growth

  • Example: Pregnancy

    • High investment in reproduction

    • Low investment in growth

    • Low investment in maintenance

      • The immune system down-regulates

      • The immune system sees the fetus as a parasite

  • Asymmetrical investment in offspring

Women

Men

Minimum Parental Investment

1200 days (40 months)

1 day

Reproductive Age Limit

~44 years

~60 years

Maximum Reproductive Potential

10ish

30-1000

  • Males' reproductive success is limited by access to mates

  • Female reproductive success in limited by access to resources


Males do not invest when…

  • They can easily use their resources to secure additional mates

  • When caring for offspring would not increase their offspring’s fitness


Trivers’ Parental Investment Theory

  • Inter-sexual selection

    • More investment = more selective

  • Intra-sexual selection

    • Less investment = more competitive


Intrasexual Selection

  • Males compete for territory, control of females, or opportunities to fertilize

    • Combat is a common form of pre-mating competition


Intersexual Selection

  • When males cannot monopolize resources, females may be able to “choose” among the males

  • Females choose males based on tail lengths, which is presumed to indicate the relative fitness of a male


Sex differences

  • Humans:

    • Sexual dimorphism

    • Ornamentation and appearance

      • Same-sex competition?

      • Secure mates?

      • Signal quality?

      • Signal resources?


Sexual vs Natural Selection

  • Guppies - sexual selection can favor traits that reduce survival

  • Natural selection favors traits that make the male blend in

  • Sexual selection favors traits that make the male stand out


Parent-Offspring Conflict

  • Trivers (1974)

    • Conflict over care

  • Parental care is costly:

    • Trade-off: current vs future reproduction

    • Time: Lost mating opportunities

    • Energy: energy invested in one offspring cannot be placed in another


Conflict over Care…

  • Parents distribute resources evenly

  • Parents and children want the same resources

  • Sibling rivalry

    • A small age difference between siblings leads to more conflict over resources


Breastfeeding & Weaning

  • Breastfeeding is costly

  • Cost increases over time

  • Results in mother-infant conflict

    • Increased birth spacing

    • Neglect


Comparisons with Other Primates

  • Baboons

    • Weaning conflict long and intense

    • Frequent turnover of the Alpha male

      • Siblings generally share 25% of genes

  • Gorillas

    • Less weaning conflict

    • New alpha male will kill infants

      • No relation to them


Infanticide

  • Parental investment is a limited resource

    • Allocation

  • The women who sacrifice their offspring tend to be young, poor, unmarried, and socially isolated

The Evolutionary Psychology of Attraction

  • Common, prevailing view for women’s preferences:

  • Males who can provide a range of resources for the female

    • Food

    • Shelter

    • Protection from other males

  • Females should have evolved preferences for males who:

    • Have good financial prospects

    • Are older than themselves

    • Have higher social status

    • Display hard-working and industrious character

  • Males are looking for women with a youthful appearance

  • Beauty:

    • Unwrinkled skin

    • Bright eyes

    • Full red lips

    • Glossy hair

    • Lack of facial blemishes and facial hair

    • Facial symmetry

    • Body size and shape


Waist-to-Hip ratio

  • WHR = measurement at smallest point of waist and widest point of hips

    • Before puberty = no sex differences

    • After puberty = female fat accumulation around hips

  • Western averages:

    • Healthy men = 0.85-0.95

    • Healthy (premenopausal) women = 0.65 - 0.80

  • Darwin: there are no universal signs of beauty

  • 1980’s: Evolutionary psychologists

    • Universal mate preferences

  • Body fat distribution

    • Hips and buttocks

    • Associated with menarche

  • “Honest Signal” of fertility

  • WHR of 0.7 is a signal of attractiveness

  • Low WHR:

    • Indicates that a woman is not pregnant

    • There women have higher circulating b-estradiol and progesterone

    • Might facilitate conception during in-vitro fertilization

  • Women with a high WHR and low BMI have a harder time becoming pregnant

  • Men with low testosterone have lower WHR

  • Hadza of Tanzania

    • WHR of 0.9 is considered most attractive

    • However, the profile of women mattered

      • When viewed from the side, men preferred a WHR of 0.6

  • Matsigenka of Peru living in the Amazon

    • WHR of 0.9 is considered most attractive


Problems with WHR Studies

  • Images are not realistic

  • What is being measured: BMI or WHR?

  • Inconsistent manipulators of the study stimuli:

    • Stimuli is often not culturally or ecologically relevant

  • Status of participant (Ex. hungry)


Tsimane Study

  • Tsimane men (N=66)

  • Controlled for BMI to look at preferences


Polygyny

  • Why would a woman put up with polygyny?

    • Resources

    • Genes

  • Examples: suppose it costs two units of stuff to produce a viable offspring

  • Male coercion

    • Only if men use force and will not lead to higher reproductive success

  • Polygyny Threshold

    • Females select males because of resources will lead to higher reproductive success

  • Dogon of Mali in West Africa (Strassman)

    • Patrilineal

    • Arranged marriage

    • Male dominated

      • 46% have more than one wife

    • IMR: high

      • 20% of babies die in the first year

      • 46% of children die by age five

    • Wealthy men:

      • More wives

      • Male Coercion Model


Polyandry

  • Non-classical polyandry

    • Mostly informal

    • No co-residence

    • Partible paternity: 41% of South American tribes studied

      • Offspring more likely to live to adolescence because the child had more fathers

    • Sex with secondary father may cease when child is born

    • Low frequency

  • Classical

    • Organized at the state level

      • Agricultural and Agropastoralism

    • Cohabitation

    • 9% to 50% of all marriages

    • Formal (residential) polyandry

    • Co-residence

  • Monomarital Principle

    • Land is scarce

      • Less land fragmentation

    • Marriage of brothers allows land to stay in family

    • Sri Lanka and India

  • Skewed Sex Ratio

    • Shortage of women from polygyny

    • “Junior husband” to have some chance at reproduction

  • Male Absenteeism

    • Fear of female abduction

    • Inuit

      • High male mortality

    • Fraternal polyandry to protect wife

  • Father Effect

    • Consequences for the survival and well-being of a child

      • Partible paternity

      • Leads to informal polyandry

    • Children with two fathers were more likely to survive to age 15

  • Evolutionary Biological Theories

    • Classical: Operational Sex Ratio (OSR)

      • Male-biased

    • The rarer sex (females) becomes more picky

  • Male-Cooperative polyandry

    • Joint male defense of a territory

    • When a female’s territory overlaps with territories of other males