Evolution, Biodiversity and

Population Ecology

By
Moe Thazin Shwe, PhD
Evolution: The Source of Earth’s Biodiversity

• The honeycreepers and the other native animals and plants of Hawai‘i help reveal
how our world became populated with the remarkable diversity of life we see today.

• Scientific study shows us that our planet has progressed from a stark world
inhabited solely by microbes to a lush cornucopia of millions of species (FIGURE
3.1).

• A species is a particular type of organism or, more precisely, a population or group

of populations whose members share characteristics and can freely breed with one
another and produce fertile offspring.

• A population is a group of individuals of a particular species that live in a particular

area.

• Over vast spans of time, the process of biological evolution has shaped populations
and species, giving us the vibrant abundance of life that enriches Earth today.
• Evolution in the broad sense means change over time, and biological evolution
consists of change in populations of organisms across generations.

• Changes in genes (p. 47) often lead to modifications in the appearance or behavior of
organisms from generation to generation.

• Biological evolution results from random genetic changes and may be directed by
natural selection.

• Natural selection is the process by which inherited characteristics that enhance

survival and reproduction are passed on more frequently to future generations than
those that do not, thus altering the genetic makeup of populations through time.
• Evolution is one of the best-supported and most illuminating concepts in all of
science, and it is the very foundation of modern biology.

• Perceiving how species adapt to their environments and change over time is crucial
for comprehending ecology and learning the history of life.

• Evolutionary processes influence many aspects of environmental science, including

pesticide resistance, agriculture, medicine, and environmental health.
Natural selection shapes organisms and diversity

• In 1858, Charles Darwin and Alfred Russel Wallace each independently proposed
the concept of natural selection as a mechanism for evolution and as a way to explain
the great variety of living things.

• Both Darwin and Wallace were exceptionally keen naturalists from England who had
studied plants and animals in such exotic locales as the Galápagos Islands (Darwin)
and the Malay Archipelago (Wallace).

• In the century and a half since then, many thousands of scientists have refined our
understanding of natural selection and evolution.
• Natural selection is a simple concept that offers a powerful explanation for patterns
evident in nature.

• The idea of natural selection follows logically from a few straightforward premises
that are readily apparent to anyone who observes the life around us:
• Organisms face a constant struggle to survive and reproduce
• Organisms tend to produce more offspring than can survive
• Individuals of a species vary in their characteristics
• Variation is due to differences in genes, the environments in which genes are
expressed, and the interactions between genes and environment.

• As a result of this variation, some individuals of a species will be better suited to

their environment than others and will be better able to reproduce.
• Many characteristics are passed from parent to offspring through the genes, and a
parent that produces many offspring will pass on more genes to the next generation
than a parent that produces few or no offspring.

• In the next generation, therefore, the genes of better-adapted individuals will out-
number those of individuals that are less well adapted.

• From one generation to another through time, characteristics, or traits, that lead to
better and better reproductive success in a given environment will evolve in the
population.

• This process is termed adaptation, and a trait that promotes success is also called
an adaptation or an adaptive trait.
Selection acts on genetic variation

• For an organism to pass a trait along to future generations, genes in the organism’s
DNA (p. 47) must code for the trait.

• In an organism’s lifetime, its DNA will be copied millions of times by millions of cells.

• In all this copying and recopying, sometimes a mistake is made.

• Accidental changes in DNA, called mutations, give rise to genetic variation among
individuals.
• If a mutation occurs in a sperm or egg cell, it may be passed on to the next
generation.

• Most mutations have little effect, but some can be deadly, whereas others can be
beneficial.

• Those that are not lethal provide the genetic variation on which natural selection
acts.
• Genetic variation is also generated as organisms mix their genetic material through
recombination during sexual reproduction.

• When organisms reproduce sexually, a portion of each parent’s genes contributes to

the genes of the offspring.

• This process produces novel combinations of genes, generating variation among

individuals.
• When natural selection acts on genetic variation by favoring certain variants, it can
drive a feature in a particular direction (FIGURE 3.2).

• Because such evolutionary change generally requires a great deal of time, a species
cannot always adapt to environmental conditions that change quickly.

• For instance, the warming of our global climate today (Chapter 18) is occurring too
rapidly for most species to adapt, and we may lose many species to extinction as a
result.
• However, genetic variation can sometimes help pro- tect a population against novel
challenges. One of the honeycreeper species of the Hakalau

• Forest, the ‘amakihi, has in recent years been discovered in ‘o ̄ hi‘a trees at very low
elevations, well within the zone where avian malaria has killed off all other
honeycreepers.

• Researchers studying this population have determined that some of the ‘amakihis
living here when malaria arrived had genes that by chance gave them a natural
resistance to the disease.
• These resistant birds survived malaria’s onslaught, and their descendants
reestablished a population that continues to grow today.

• Similarly, the ‘apapane (another Hawaiian honeycreeper) and the ‘o ̄ ma‘o (a native
Hawaiian thrush) also are showing some degree of resistance to malaria.

• Scientists hope that perhaps some individuals of the rarer native birds of Hakalau
might also harbor resistance genes that may help them persist in the face of
malaria.
Selective pressures from the environment influence adaptation

• Environmental conditions determine what pressures natural selection will exert, and
these selective pressures affect which members of a population will survive and
reproduce.

• Over many generations, this results in the evolution of traits that enable success within
the environment in question.

• Closely related species that live in very different environments and thus experience
very different selective pressures tend to diverge in their traits as the differing
pressures drive the evolution of different adaptations (FIGURE 3.3a).

• Conversely, sometimes very unrelated species may acquire similar traits as they adapt
to selective pressures from similar environments; this is called convergent evolution
(FIGURE 3.3b).
• However, environments change over time, and organisms may move to new locations
and encounter new conditions.

• In either case, a trait that promotes success at one time or place may not do so at
another.

• Hawaiian honeycreepers such as the ‘apapane and the ‘i‘iwi fly long distances in
search of flowering trees.

• This behavior had long helped them to find the best resources across a diverse
landscape.
• However, once malaria arrived, the strategy backfired, as birds from malaria-free
areas would sometimes fly into death zones.

• Today, thousands of honeycreepers from mountain forests die each year when they
fly downslope and are bitten by malarial mosquitoes.

• As environmental conditions vary in time and space, adaptation becomes a moving

target.
Evidence of selection is all around us
• The results of natural selection are all around us, visible in every adaptation of every
organism.

• In addition, scientists have demonstrated the rapid evolution of traits by selection

in countless lab experiments, mostly with fast-reproducing organisms such as
bacteria, yeast, and fruit flies.

• The evidence for selection that may be most familiar to us is that which Darwin
himself cited prominently in his work 150 years ago: our breeding of domesticated
animals.

• In domesticated dogs, cats, and livestock, we have conducted selection under our
own direction.

• We have chosen animals with traits we like and bred them together, while not
breeding those with variants we do not like.
• Consider the great diversity of dog breeds (FIGURE 3.4a).

• People generated every type of dog alive today by starting with a single ancestral
species and selecting for particular desired traits as individuals were bred together.

• From Great Dane to Chihuahua, all dogs are able to interbreed and produce viable
offspring, yet breeders maintain striking differences among them by allowing only
like individuals to breed with like.

• This process of selection conducted under human direction is termed artificial

selection.
• Artificial selection has given us the many crop plants we depend on for food, all of
which people domesticated from wild ancestors and carefully bred over years,
centuries, or millennia (FIGURE 3.4b).

• Through selective breeding, we have created corn with bigger, sweeter kernels; wheat
and rice with larger and more numerous grains; and apples, pears, and oranges with
better taste.

• We have diversified single types into many—for instance, breeding variants of the
plant Brassica oleracea to create broccoli, cauliflower, cabbage, and brussels sprouts.

• Our entire agricultural system is based on artificial selection. We depend on a working

understanding of evolution for the very food we eat.
Evolution generates biodiversity

• Just as artificial selection helps us create new types of pets, farm animals, and crop
plants, natural selection serves to elaborate and diversify traits in wild organisms.

• Over the long term, natural selection helps lead to the formation of new species and
whole new types of organisms.

• Life’s complexity can be expressed as biological diversity, or biodiversity for short,

which refers to the variety of life across all levels of biological organization,
including the diversity of species, genes, populations, and communities (we will
introduce communities shortly: p. 78 and Chapter 4).
• Scientists have described about 1.8 million species, but many more remain undiscovered
or unnamed.

• Estimates for the total number of species in the world vary, but they range from 3
million up to 100 million.

• Hawaii’s insect fauna provides one example of how much we have yet to learn.

• Scientists studying fruit flies in the Hawaiian Islands have described over 500 species
of them, but they have also identified about 500 others that have not yet been
formally named and described.

• Still more fruit fly species probably exist but have not yet been found.
• Subtropical islands such as Hawai‘i are by no means the only places rich in biodiversity,
however.

• Step outside anywhere, and you will find many species within close reach.

• They may not always be large and conspicuous like Yellowstone’s bears or the Serengeti’s
elephants, but they will be there.

• Plants poke up from cracks in asphalt in every city in the world, and even Antarctic ice
harbors microbes.

• A handful of backyard soil may contain an entire miniature world of life, including insects,
mites, millipedes, nematode worms, plant seeds, fungi, and millions upon millions of bacteria.
Speciation produces new types of organisms

• which new species are generated is termed speciation.

• Speciation can occur in a number of ways, but the main mode is generally thought to
be allopatric speciation, whereby species form from populations that become
physically separated over some geographic distance.

• To understand allopatric speciation, begin by picturing a population of organisms.

• Individuals within the population possess many similarities that unify them as a
species because they are able to breed with one another and share genetic
information.

• However, if the population is broken up into two or more isolated areas, individuals
from one area cannot reproduce with individuals from the others.
• When a mutation arises in the DNA of an organism in one of these newly isolated
populations, it cannot spread to the other populations.

• Over time, each population will independently accumulate its own set of mutations.

• Eventually, the populations may diverge, growing so different that their members
can no longer mate with one another.

• Once this has happened, there is no turning back; the two populations can no longer
share genetic information, and they will embark on their own independent
evolutionary trajectories as separate species (FIGURE 3.5).

• The populations will continue diverging in their characteristics as chance mutations

accumulate that cause them to differ more and more.

• If environmental conditions happen to differ for the two populations, then natural
selection may accelerate the divergence
 Populations can be separated in many ways
• various ways. Lava flows can destroy forest, leaving small isolated patches intact (as
shown in Figure 3.5).

• Glacial ice sheets may move across continents during ice ages and split populations in
two. Major rivers may change course and do the same.

• Mountain ranges may be uplifted and divide regions and their organisms.
• Sea level may rise, flooding low-lying regions and isolating areas of higher ground as
islands.

• Drying climate may partially evaporate lakes, subdividing them into smaller bodies of
water.

• Warming or cooling temperatures may cause plant communities to shift northward or

south- ward, or upslope or downslope, creating new patterns of plant and animal
distribution.
• Alternatively, sometimes new areas are created and organ- isms colonize them,
establishing isolated populations.

• Hawai‘i provides an example. As shown in Figure 2.22 (Chapter 2, p. 59), the Pacific
tectonic plate moves over a volcanic “hotspot” that extrudes magma into the ocean,
building volcanoes that form islands once they break the water’s surface.

• The plate inches northwest, dragging each island with it, while new islands are
formed at the hotspot.

• The result, over millions of years, is a long string of islands, an archipelago.

• As each new island is formed, plants and animals that colonize it may undergo
allopatric speci- ation if they are isolated enough from their source population (see
THE SCIENCE BEHIND THE STORY, pp. 74–75).
• For speciation to occur, populations must remain isolated for a very long time, generally
thousands of generations.

• Then, if the geologic or climatic process that has isolated populations reverses itself—if the
glacier recedes, or the river returns to its old course, or warm temperatures turn cool again—
then the populations may come back together.

• This is the moment of truth for speciation.

• If the populations have not diverged enough, their members will begin interbreeding and
reestablish gene flow, mixing the mutations that each population accrued while isolated.

• However, if the populations have diverged sufficiently, they will not interbreed, and two species
will have been formed, each destined to continue on its own evolutionary path.
We can infer the history of life’s diversification by comparing
organisms
• Innumerable speciation events have generated complex patterns of diversity at
levels above the species level.

• Evolutionary biologists study such patterns, examining how groups of organisms

arose and how they evolved the characteristics they show.

• For instance, how did we end up with plants as different as mosses, palm trees,
daisies, and redwoods?

• Why do fish swim, snakes slither, and sparrows sing?

• How and why did birds, bats, and insects each independently evolve the ability to
fly?

• To address such questions, we need to know how the major groups diverged from one
another over time.
• Scientists represent this history of divergence by using branching, tree- like
diagrams called phylogenetic trees.

• Similar to family genealogies, these diagrams illustrate scientists’ hypotheses as to

how divergence took place (FIGURE 3.6).

• Phylogenetic trees can show relationships among species, groups of species,

populations, or genes.

• Scientists construct these trees by analyzing patterns of similarity among the genes
or external traits of present-day organisms and by inferring which groups share
similarities because they are related.
• Once we have a phylogenetic tree, we can map traits onto the tree according to
which organisms possess them, and we can thereby trace how the traits have
evolved.

• For instance, phylogenetic research shows that birds, bats, and insects are distantly
related, with many flightless groups between them. (Note in Figure 3.6 that birds
and mammals are separated on the tree and that insects are outside the tree.)

• Therefore, it is far simpler to conclude that these three very different groups
evolved flight independently than it would be to conclude that the many flightless
groups between them each lost an ancestral ability to fly.

• Because phylogenetic trees help biologists make such inferences about so many
traits, they have become one of the modern biologist’s most powerful tools.
• Knowing how organisms are related to one another also helps scientists to classify
them and name them, so that we can make sense of the life around us and
communicate effectively.

• Taxonomists use an organism’s physical appearance and genetic makeup to determine

its species.

• These scientists then group species by their similarity into a hierarchy of categories
meant to reflect evolutionary relationships.

• Related species are grouped together into genera (singular, genus), related genera
are grouped into families, and so on (FIGURE 3.7).

• Each species is given a two-part Latin or Latinized scientific name denoting its genus
and species.
• For instance, the ‘akiapo ̄la ̄‘au, Hemignathus munroi, is similar to other Hawaiian
honeycreepers in the genus Hemignathus.

• These species are closely related in evolutionary terms, as indicated by the genus
name they share.

• They are more distantly related to honeycreepers in other genera, but all
honeycreepers are classified together in the family Fringillidae. This system of
naming and classification was devised by Swedish botanist Carl Linnaeus (1707–1778)
long before Darwin’s work on evolution.

• Today biologists use evolutionary information from phylogenetic trees to help

classify organisms under the Linnaean system’s rules.
The fossil record teaches us about life’s long history

• Scientists also decipher life’s history by studying fossils.

• As organisms die, some are buried by sediment.

• Under certain conditions, the hard parts of their bodies—such as bones, shells, and teeth
—may be preserved, as sediments are com- pressed into rock (pp. 55–56).

• Minerals replace the organic material, leaving behind a fossil, an imprint in stone of the
dead organism (FIGURE 3.8, p. 76).

• In countless locations throughout the world, geologic processes across millions of years
have buried sediments and later brought sedimentary rock layers to the surface, revealing
assemblages of fossilized plants and animals from different time periods.
• By dating the rock layers that contain fossils, paleontologists (scientists who study
the history of Earth’s life) can learn when particu- lar organisms lived. The
cumulative body of fossils worldwide is known as the fossil record.
• The fossil record shows that:
• Across life’s 3.5 billion years on Earth, complex structures have evolved from simple
ones, and large sizes from small ones.

• However, simplicity and small size have also evolved when favored by natural selection; it
is easy to argue that Earth still belongs to the bacteria and other microbes, some of
them little changed over eons.

• Even fans of microbes, however, must marvel at some of the exquisite adaptations of
animals, plants, and fungi: the heart that beats so reliably for an animal’s entire lifetime
that we take it for granted; the complex organ system to which the heart belongs; the
stunning plumage of a peacock in full display; the ability of each and every plant on the
planet to lift water and nutrients from the soil, gather light from the sun, and turn it
into food; the staggering diversity of beetles and other insects; and the human brain and
its ability to reason. All these adaptations and more have resulted as the process of
evolution has generated new species and whole new branches on the tree of life.
Speciation and extinction together determine Earth’s biodiversity

• Although speciation generates Earth’s biodiversity, it is only part of the equation,

because the fossil record teaches us that the vast majority of species that once
lived are now gone.

• The disappearance of a species from Earth is called extinction.

• From studying the fossil record, paleontologists calculate that the average time a
species spends on Earth is 1–10 million years.

• The number of species in existence at any one time is equal to the number added
through speciation minus the number removed by extinction.
• Extinction occurs naturally, but human impact can profoundly affect the rate at
which it occurs (FIGURE 3.9).

• As we will see in Chapter 11, our planet’s biological diversity is being lost at a
frightening pace.

• This loss affects people directly, because other organisms provide us with life’s
necessities—food, fiber, medicine, and vital ecosystem services (pp. 21, 134–135,
170, 290).

• Species extinction brought about by human impact may well be the single biggest
problem we face, because the loss of a species is irreversible.
Some species are especially vulnerable to extinction

• In general, extinction occurs when environmental conditions change rapidly or

severely enough that a species cannot adapt genetically to the change; the slow
process of natural selection simply does not have enough time to work.

• All manner of events can cause extinction—climate change, the arrival of new
species, severe weather events, and more.

• In general, small populations are vulnerable to extinction because fluctuations in

their size could, by chance, bring the population size to zero.

• Species narrowly specialized to some particular resource or way of life are also
vulnerable, because environmental changes that make that resource or way of life
unavailable can doom them.
• Island-dwelling species are frequently vulnerable.

• Because islands are smaller than mainland areas and are isolated by water, only some
species reach islands, whereas many others never do.

• As a result, some of the pressures and challenges faced daily by organisms on the
mainland simply don’t exist on islands.
• For instance, only one land mammal—a bat—ever reached Hawai‘i naturally, so
Hawaii’s birds evolved for millions of years without having to defend against the
threat of predation by mammals.

• Likewise, Hawaii’s plants did not need to protect themselves against plant-eating
mammals.

• Because defenses are costly to invest in, most island birds and plants lost any
defenses their ancestors may have had.
• Eventually, people—first Polynesians and then Europeans— arrived in Hawai‘i,
bringing cattle, goats, pigs, rats, dogs, cats, and mongooses.

• Hawaii’s native organisms were completely unprepared.

• Rats, cats, and mongooses preyed on ground- nesting seabirds, ducks, geese, an ibis,
and flightless rails, driving a number of these birds extinct.

• Livestock ate through the vegetation, turning lush forests into desolate grasslands.

• All in all, half of Hawaii’s native birds were driven extinct within just decades after
human arrival.
• On a mainland, “islands” of habitat (such as forested mountaintops) can host endemic
species that are vulnerable to extinction (FIGURE 3.10).

• In the United States, many amphibians are limited to very small ranges.

• The Yosemite toad is restricted to a small region of the Sierra Nevada in California,
the Hou-ston toad occupies just a few areas of Texas woodland, and the Florida bog
frog lives in a tiny region of Florida wetland.

• Fully 40 salamander species in the United States are restricted to areas the size of
a typical county, and some of these live atop single mountains.
Earth has seen several episodes of mass extinction

• Most extinction occurs gradually, one species at a time.

• The rate at which this type of extinction occurs is referred to as the background
extinction rate.

• However, Earth has seen five events of staggering proportions that killed off
massive numbers of species at once.

• These episodes, called mass extinction events, have occurred at widely spaced
intervals in Earth’s history and have wiped out 50–95% of our planet’s species each
time.
• On a mainland, “islands” of habitat (such as forested mountaintops) can
host endemic species that are vulnerable to extinction (FIGURE 3.10).
• In the United States, many amphibians are limited to very small ranges.

• The Yosemite toad is restricted to a small region of the Sierra Nevada in

California, the Houston toad occupies just a few areas of Texas woodland,
and the Florida bog frog lives in a tiny region of Florida wetland.

• Fully 40 salamander species in the United States are restricted to areas

the size of a typical county, and some of these live atop single mountains.
The sixth mass extinction is upon us

• Many biologists have concluded that Earth is currently entering its sixth mass extinction
event—and that we are the cause.

• Indeed, the Millennium Ecosystem Assessment (p. 33) estimated that today’s extinction
rate is 100–1000 times higher than the background rate, and rising (p.301).

• Changes to Earth’s natural systems set in motion by human population growth,

development, and resource depletion have driven many species extinct and are threatening
countless more.

• The alteration and outright destruction of natural habitats, the hunting and harvesting of
species, and the introduction of species from one place to another where they can harm
native species—these processes and more have combined to threaten Earth’s biodiversity
(pp. 301–308).
• When we look around us, it may not appear as though a human version of an asteroid
impact is taking place, but we can- not judge such things on the timescale of a human
lifetime.

• On the geologic timescale, extinction over 100 years or even 10,000 years appears
instantaneous.

• In contrast, speciation is a slow enough process that it will take life millions of
years to recover— by which time our own species will most likely not be around.
Levels of Ecological Organization

• Extinction, speciation, and other evolutionary mechanisms and patterns play key
roles in ecology.

• Ecology is the scientific study of the distribution and abundance of organisms, the
interactions among organisms, and the relationships between organisms and their
environments.

• It is often said that ecology provides the stage on which the play of evolution
unfolds. The two are intertwined in many ways.
We study ecology at several levels

• Life exists in a hierarchy of levels, from atoms, molecules, and cells (pp. 41–47) up
through the biosphere, which is the cumulative total of living things on Earth and
the areas they inhabit.

• Ecologists are scientists who study relationships at the higher levels of this
hierarchy (FIGURE 3.11), namely at the levels of the organism, population,
community, ecosystem, and biosphere.

• At the level of the organism, the science of ecology describes relationships between
an organism and its physical environment.
• Organismal ecology helps us understand, for example, what aspects of a Hawaiian
honeycreeper’s environment are important to it, and why.

• In contrast, population ecology examines the dynamics of population change and the
factors that affect the distribution and abundance of members of a population.

• It helps us understand why populations of some species (such as endangered

honeycreepers) decline while populations of others (such as ourselves) increase.
• In ecology, a community consists of an assemblage of populations of interacting
species that live in the same area.

• A population of ‘akiapo ̄la ̄‘au, a population of koa trees, a population of wood-boring

grubs, and a population of ferns, together with all the other interacting plant,
animal, fungal, and microbial populations in the Hakalau Forest, would be considered a
community.

• Community ecology focuses on pat- terns of species diversity and on interactions

among species, ranging from one-to-one interactions to complex interrelationships
involving the entire community.
• Ecosystems encompass communities and the abiotic (nonliving) material and forces
with which community members interact.

• Hakalau’s cloud-forest ecosystem consists of its community plus the air, water, soil,
nutrients, and energy used by the community’s organisms.

•
• Ecosystems encompass communities and the abiotic (nonliving) material and forces
with which community members interact.

• Hakalau’s cloud-forest ecosystem consists of its community plus the air, water, soil,
nutrients, and energy used by the community’s organisms.

• Ecosystem ecology reveals patterns, such as the flow of energy and nutrients, by
studying living and nonliving components of systems in conjunction.

• Today’s warming climate (Chapter 18) is having ecosystem- level consequences as it

exerts impacts on the organisms of Hakalau and many other ecosystems throughout
the world.
• As new technologies allow scientists to learn more about the complex dynamics of
natural systems at a global scale, ecologists are increasingly expanding their
horizons beyond eco- systems to the biosphere as a whole.

• In the remainder of this chapter we explore ecology up through the population level.

• In Chapter 4 we examine com- munity ecology, and in Chapter 5 we consider ecology

at the levels of the ecosystem and biosphere.
Each organism has habitat needs

• At the level of the organism, each individual relates to its environment in ways that
tend to maximize its survival and reproduction.

• One key relationship involves the specific environment in which an organism lives, its
habitat.

• A species’ habitat consists of the living and nonliving elements around it, including
rock, soil, leaf litter, humidity, plant life, and more.

• The ‘akiapo ̄ la ̄ ‘au (FIGURE 3.12) lives in a habitat of cool, moist, montane forest of
native koa and ‘o ̄ hi‘a trees, where it is high enough in elevation to be safe from avian
malaria.
• Each organism thrives in certain habitats and not in oth- ers, leading to nonrandom
patterns of habitat use.

• Mobile organisms actively select habitats in which to live from among the range of
options they encounter, a process called habitat selection. I

• n the case of plants and of rooted animals (such as sea anemones in the ocean),
whose young disperse and settle passively, patterns of habitat use result from
success in some habitats and failure in others.
• Habitats are scale dependent.

• A tiny soil mite may use less than a square meter of soil in its lifetime.

• A vulture, elephant, or whale, in contrast, may traverse miles upon miles of air, land,
or water in just a day.

• Species also may have different habitat needs in different seasons; many migratory
birds use distinct breeding, wintering, and migratory habitats.
• The criteria by which organisms favor some habitats over others can vary greatly.

• The soil mite may assess available habitats in terms of the chemistry, moisture, and
texture of the soil and the percentage and type of organic matter.

• The vulture may ignore not only soil but also topography and vegetation, focusing
solely on the abundance of dead animals in the area that it scavenges for food.

• For a whale, water temperature, salinity, and the density of marine microorganisms
might be critical characteristics.

• Each species assesses habitats differently because each species has different needs.
• Habitat use is important in environmental science because the availability and quality
of habitat are crucial to an organism’s well-being.

• Indeed, because habitats provide everything an organism needs, including nutrition,

shelter, breeding sites, and mates, the organism’s very survival depends on the avail-
ability of suitable habitats.

• Often this need results in conflict with people who want to alter or develop a habitat
for their own purposes.
Niche and specialization are key
concepts in ecology

• Another way in which an organism relates to its environment is through its niche.

• A species’ niche reflects its use of resources and its functional role in a community.

• This includes its consumption of certain foods, its role in the flow of energy and
matter, and its interactions with other organisms.

• The niche is a multidimensional concept, a kind of summary of everything an organism

does.

• The pioneering ecologist Eugene Odum once wrote that “habitat is the organism’s
address, and the niche is its profession.”
• Organisms vary in the breadth of their niches.

• Species with narrow breadth, and thus very specific requirements, are said to be
specialists.

• Those with broad tolerances, able to use a wide array of resources, are generalists.

• A native Hawaiian honeycreeper like the ‘akiapo ̄la ̄‘au (see Figure 3.12) is a specialist,
because its unique bill is exquisitely adapted for feeding on grubs that tunnel
through the wood of native trees.
• In contrast, the common myna (a bird introduced to Hawai‘i from Asia) is a
generalist; its unremarkable bill allows it to eat many types of foods in many types
of habitats.

• As a result, the common myna has spread through virtually all areas of the Hawaiian
Islands where native birds have disappeared and where human development has
altered the landscape.

• Today it is one of Hawaii’s most numerous birds.

• Specialists succeed over evolutionary time by being extremely good at the things
they do, but they are vulnerable when conditions change and threaten the habitat or
resource on which they have specialized.

• Generalists succeed by being able to live in many different places and to withstand
variable conditions, but they may not thrive in any one situation as much as a
specialist would.

• An organism’s habitat preferences, niche, and degree of specialization each reflect

adaptations of the species and are products of natural selection.
Population Ecology

• Individuals of a species that inhabit a particular area make up a population. Species

may consist of multiple populations that are geographically isolated from one
another.

• This is the case with Hawaii’s state bird, the ne ̄ ne ̄ , a goose that grazes in open
grassy areas (see Figure 3.1d).

• Originally common throughout the Hawaiian Islands, the ne ̄ne ̄ (pronounced “nay-
nay”) was nearly driven to extinction by human hunting; livestock and alien plants
that destroyed and displaced the vegetation it fed on; and rats, cats, dogs, pigs, and
mongooses that preyed on its eggs and young.
• Ne ̄ne ̄s disappeared from all islands except the island of Hawai‘i, and in recent
decades biologists and wildlife managers have labored to breed it in captivity and
reintroduce it to protected areas on other islands.

• These efforts have met with success, and today ne ̄ne ̄s live in at least seven
populations on four islands.

• In contrast, the human species faces few threats from other animals, is a
consummate generalist, and has spread into nearly every corner of the planet.

• As a result, it is difficult to define a distinct human population on anything less than

the global scale. In the ecological sense of the word, all 7 billion of us comprise one
population.
Populations show characteristics that help predict their dynamics

• All populations—from humans to ne ̄ne ̄s—exhibit characteris- tics that help

population ecologists predict the future dynamics of the population.

• Attributes such as density, distribution, sex ratio, age structure, and birth and
death rates all help the ecologist understand how a population may grow or decline.

• The ability to predict growth or decline is useful in monitor- ing and managing
threatened and endangered species (see THE SCIENCE BEHIND THE STORY, pp.
82–83).

• It is also useful in studying human populations (Chapter 8). Understanding human

population dynamics is a central element of environmental science and is one of the
prime challenges for our society today.
• Population size Expressed as the number of individual organisms present at a given
time, population size may increase, decrease, undergo cyclical change, or remain
stable over time.

• Populations generally grow when resources are abundant and natural enemies are few.

• Populations can decline in response to loss of resources, negative impacts from other
species, or natural disasters that kill large numbers of individuals.

• Researchers estimate that the ne ̄ne ̄ population surpassed 25,000 birds before
Europeans reached the Hawaiian Islands.

• By the 1950s, after two centuries of impacts from hunting, agriculture, non-native
mammals, and invasive plants, the population was down to just 30 individuals.

• Since then, intensive conservation efforts have turned this decline around, and now
over 2000 ne ̄ne ̄s live on the Hawaiian Islands.
• The passenger pigeon, now extinct, illustrates the extremes of population size
(FIGURE 3.13).

• Not long ago it was the most abundant bird in North America; flocks of passenger
pigeons literally darkened the skies.

• In the early 1800s, ornithologist Alexander Wilson watched a flock of 2 billion birds
390 km (240 mi) long that took 5 hours to fly over and sounded like a tornado.

• Passenger pigeons nested in gigantic colonies in the forests of the upper Midwest
and southern Canada.

• Once settlers arrived and began cutting the forests, however, the birds made easy
targets for market hunters, who gunned down thousands at a time and shipped them
to market by the wagonload.

• By the end of the 19th century, the passenger pigeon population had declined to such
a low number that the birds could not form the large colonies they apparently
needed in order to breed.
• Population density The flocks and breeding colonies of passenger pigeons showed
high population density, another attribute that ecologists assess to understand
populations.

• Population density describes the number of individuals in a population per unit area.

• High population density makes it easier for organisms to group together and find
mates, but it can also lead to competition and conflict if space, food, or mates are in
limited supply.
• Overcrowded organisms may become vulnerable to the predators that feed on them,
and close contact among individuals can increase the transmission of infectious
disease.

• For these reasons, organisms sometimes leave an area when densities become too
high.

• In contrast, at low population densities, organisms benefit from more space and
resources but may find it harder to locate mates and companions.
• Population distribution Population distribution describes the spatial arrangement of
organisms in an area.
• Ecologists define three distribution types: random, uniform, and clumped (FIGURE
3.14).

• In a random distribution, individuals are located haphazardly in no particular pattern.

• This type of distribution can occur when the resources an organism needs are
plentiful throughout an area and other organisms do not strongly influence where
members of a population settle.

• A uniform distribution is one in which individuals are evenly spaced.

• This can occur when individuals hold territories or compete for space.

• In a desert where water is scarce, each plant needs space for its roots to gather
moisture. Plants may even poison one another’s roots as a means of competing for
space.
• In a clumped distribution, the pattern most common in nature, organisms arrange
themselves according to the availability of the resources they need to survive.

• Many Hawaiian honeycreepers tend to cluster near actively flower- ing trees that
offer nectar.

• Many desert plants grow in patches around isolated springs or along streambeds that
flow with water after rains.

• Human beings also exhibit clumped distribu- tion: People frequently aggregate in
villages, towns, or cities.
• Clumped distributions often indicate that species are seeking certain habitats or
resources that are themselves clumped.

• Distributions can depend on the scale at which one meas- ures them.

• At small scales a population may be distributed uniformly, yet this may occur within
one patch of a larger, clumped distribution.

• At very large scales, all organisms show clumped or patchy distributions, because
some parts of the total area they inhabit are bound to be more hospitable than
others.
• Sex ratio A population’s sex ratio is its proportion of males to females, and this can
influence whether the population will increase or decrease in size over time.

• In monogamous species (in which each sex takes a single mate), a 1:1 sex ratio
maximizes population growth, whereas an unbalanced ratio leaves many individuals of
one sex without mates.

• Most species are not monogamous, however, so sex ratios may vary from one species
to another.
• Age structure Populations generally consist of indi- viduals of different ages.

• Age distribution, or age structure, describes the relative numbers of organisms of

each age within a population.

• By combining this information with data on the reproductive potential of individuals

in each age class, a population ecologist can predict how the population may grow or
shrink.
• For many plants and animals that continue growing in size as they age, older
individuals reproduce more: A tree that is large because it is old can produce more
seeds, and a fish that is large because it is old may produce more eggs.

• In some animals, such as birds, the experience they gain with age often makes older
individuals better breeders.
• Human beings are unusual because we often survive
past our reproductive years.

• A human population made up largely of older (post-

reproductive) individuals will tend to decline over time,
whereas one with many young people (of reproductive or pre-
reproductive age) will tend to increase.

• We will use diagrams to explore these ideas further in

Chapter 8 (pp. 215–217) as we study human population growth.
• Birth and death rates All the preceding factors can influence the rates at which
individuals within a population are born and die.

• Just as individuals of differing ages have different reproductive capacities,

individuals of differing ages show different probabilities of dying.

• For instance, people are more likely to die at old ages than young ages. However, this
pattern does not hold for all organisms.

• An insect, a fish, or a toad produces large numbers of young, which suffer high
death rates. For such animals, death is less likely (and survival more likely) at an
older age than at a very young age.
• To show how the likelihood of survival varies with age, ecologists use graphs called
survivorship curves (FIGURE 3.15).

• There are three fundamental types of survivorship curves.

• Humans, with higher death rates at older ages, show a type I survivorship curve.

• Toads, with highest death rates at young ages, show a type III survivorship curve.

• A type II survivorship curve is intermediate and indicates equal rates of death at all
ages. Many birds are thought to show type II curves.
Populations may grow, shrink, or remain stable

• Now that we have outlined some key attributes of populations, we are ready to take
a quantitative view of population change by examining some simple mathematical
concepts used by population ecologists and by demographers (scientists who study
human populations).
• Population growth, or decline, is determined by four factors:

• Births and immigration add individuals to a population, whereas deaths and

emigration remove individuals.
• A convenient way to express rates of birth and death is to measure the number of
births and deaths per 1000 individuals per year.
• These rates are termed the crude birth rate and the crude death rate.
• If we are not interested in the effects of migration, we can measure the rate of
natural increase by subtracting the crude death rate from the crude birth rate:

• The rate of natural increase reflects the degree to which a population is growing or
shrinking as a result of its own internal factors.
• To obtain an overall population growth rate, the total rate of change in a
population’s size per unit time, we must also take into account the effects of
migration.
• Thus, we include terms for immigration and emigration (each expressed per 1000
individuals per year) in the formula, as follows:
• The resulting number tells us the net change in a popula- tion’s size per 1000
individuals per year. For example, a popu- lation with a crude birth rate of 18 per
1000/yr, a crude death rate of 10 per 1000/yr, an immigration rate of 5 per 1000/yr,
and an emigration rate of 7 per 1000/yr would have a popula- tion growth rate of 6 per
1000/yr:

• Thus, a population of 1000 in one year will reach 1006 in the next. If the population is
1,000,000, it will reach 1,006,000 the next year. Such population increases are often
expressed as percentages, which we can calculate using the following formula:
• By measuring population growth in terms of percentages, scientists can compare
increases and decreases in species that have far different population sizes.

• They can also project changes that will occur in the population over longer periods,
much like you might calculate the amount of interest your savings account will earn
over time.
Unregulated populations increase by exponential growth

• When a population increases by a fixed percentage each year, it is said to undergo

exponential growth.

• Imagine you put money in a savings account at a fixed interest rate and leave it
untouched for years.

• As the principal accrues interest and grows larger, you earn still more interest, and
the sum grows by escalating amounts each year.

• The reason is that a fixed percentage of a small number makes for a small increase,
but the same percentage of a large number produces a large increase.

• Thus, as savings accounts (or populations) become larger, each incremental increase
likewise gets larger. Such acceleration is a characteristic of exponential growth.
• We can visualize changes in population size by using population growth curves.

• The J-shaped curve in FIGURE 3.16 shows exponential growth.

• Populations of organisms increase exponentially unless they meet constraints.

• Each organism reproduces by a certain amount, and as populations get larger, there
are more individuals reproducing by that amount.

• If there are adequate resources and no external limits, ecologists theoretically

expect exponential growth.
• Normally, exponential growth occurs in nature only when a population is small,
competition is minimal, and environmental conditions are ideal for the organism in
question.

• Most often, these conditions occur when the organism is introduced to a new
environment that contains abundant resources to exploit.

• Mold growing on a piece of fruit, or bacteria colonizing a recently dead animal, are cases
in point.

• Plants colonizing regions during primary succession (p. 103) after glaciers recede or vol-
canoes erupt may also grow exponentially.

• In Hawai‘i, many of the species that colonized the islands from other locations
underwent exponential growth for a time after their arrival.

• One current example of exponential growth in the mainland United States is the
Eurasian collared dove (see Figure 3.16).
Limiting factors restrain population growth

• Exponential growth rarely lasts long.

• If even a single species were to increase exponentially for very many generations, it
would blanket the planet’s surface!

• Instead, every population eventually is constrained by limiting factors—physical,

chemical, and biological attributes of the environment that restrain population
growth.

• These limiting factors determine the carrying capacity, the maximum population
size of a species that a given environment can sustain.
• Ecologists use the S-shaped curve in FIGURE 3.17 to show how an initial
exponential increase is slowed and eventually brought to a standstill by limiting
factors.

• Called the logistic growth curve, it rises sharply at first but then begins to level
off as the effects of limiting factors become stronger.

• Eventually the collective force of these factors stabilizes the population size at its
carrying capacity.
• We can witness this process by taking a closer look at the data for the Eurasian
collared dove population, as gathered by thousands of volunteer birders and analyzed
by government biologists through the Breeding Bird Survey, a long-running citizen
science project.

• The dove first reached North America in Florida a few decades ago and
subsequently spread north and west.

• Today its numbers are growing fastest in western areas it has recently reached, and
slower in eastern areas where it has been present for longer.

• In Florida, it has apparently eached carrying capacity (FIGURE 3.18).

• Populations of other European birds that spread across North America in the past,
such as the house sparrow and European starling, have peaked and are today
beginning to decline.
• Many factors influence a population’s growth rate and carrying capacity.

• For animals in terrestrial environments, limiting factors include temperature

extremes; prevalence of disease; abundance of predators; and the availability of
food, water, mates, shelter, and suitable breeding sites.

• Plants are often limited by amounts of sunlight and moisture and the type of soil
chemistry, in addition to disease and attack from plant-eating animals.

• In aquatic systems, limiting factors include salinity, sunlight, temperature, dissolved

oxygen, fertilizers, and pollutants.

• To determine limiting factors, ecologists may conduct experiments in which they

increase or decrease a hypothesized limiting factor and observe its effects on
population size.
The influence of some factors depends on population density

• A population’s density can enhance or diminish the impact of certain limiting factors.

• Recall that high population density can help organisms find mates but can also
increase competition and the risk of predation and disease.

• Such factors are said to be density-dependent factors, because their influence

rises and falls with population density.

• The logistic growth curve in Figure 3.17 represents the effects of density depend-
ence.

• The larger the population size, the stronger the effects of the limiting factors.
• Density-independent factors are limiting factors whose influence is not affected
by population density.

• Temperature extremes and catastrophic events such as floods, fires, and landslides
are examples of density-independent factors, because they can eliminate large
numbers of individuals with- out regard to their density.

• The logistic curve is a simplified model, and real populations in nature can behave
differently.

• Some may cycle above and below the carrying capacity.

• Others may overshoot the carrying capacity and then crash, destined either for
extinction or recovery (FIGURE 3.19).
Carrying capacities can change

• Because environments are complex and ever-changing, carrying capacity can vary.

• If a fire destroys a forest, for example, the carrying capacities for most forest
animals will decline, whereas carrying capacities for species that benefit from fire
will increase.

• Our own species has proved capable of intentionally altering our environment so as to
raise our carrying capacity.

• When our ancestors began to build shelters and use fire for heating and cooking,
they eased the limiting factors of cold climates and were able to expand into new
territory.
• As human civilization has developed, we have overcome limiting factors time and
again through the development of new technologies and cultural institutions.

• People have managed so far to increase the planet’s carrying capacity for our
species, but we have done so by appropriating immense proportions of the planet’s
resources.

• In the process, we have reduced carrying capacities for countless other organisms
that rely on those same resources.
Reproductive strategies vary among species

• Limiting factors from an organism’s environment help to regulate its population size,
but the attributes of the organism also matter.

• For example, organisms differ in their biotic potential, or capacity to produce

offspring.

• A fish with a short gestation period that lays thousands of eggs at a time has high
biotic potential, whereas a whale with a long gestation period that gives birth to a
single calf at a time has low biotic potential.
• Giraffes, elephants, humans, and other large animals with low biotic potential
produce relatively few offspring during their lifetimes.

• Species that take this approach to reproduction require a long time to gestate and
raise their young, but the considerable energy and resources they devote to caring
for and protecting them helps give these few offspring a high likelihood of survival.

• Such species are said to be K-selected (because their populations tend to stabilize
over time near carrying capacity, commonly abbreviated K).

• Because their populations stay close to carrying capacity, these organisms must
compete to hold their own in a crowded world.

• Thus, natural selection favors investing in high-quality offspring that can be good
competitors.
• In contrast, species that are r-selected have high biotic potential and devote their energy
and resources to producing many offspring in a short time.

• Their offspring do not require parental care after birth, so r-strategists simply leave their
survival to chance.

• The abbreviation r denotes the per capita rate at which a population increases in the absence
of limiting factors.

• Population sizes of r-selected species fluctuate greatly, such that they are often well below
carrying capacity.

• This is why natural selection in these species favors traits that lead to rapid population
growth. Many fish, plants, frogs, insects, and others are r-selected.
• It is important to note, however, that these are two extremes on a continuum and
that most species fall somewhere between the extremes of r-selected and K-
selected species.

• Moreover, many organisms show combinations of traits that do not correspond to a

place on the continuum. A red- wood tree, for instance, is large and long-lived, yet it
produces many small seeds and offers no parental care.
 Conserving Biodiversity

• Environmental changes that affect populations have been taking place as long as life has
existed, but today human development, resource extraction, and population pressure are
speeding the rate of change and altering the types of change. Science is crucial in
helping us understand how we modify our environment.

• However, the threats to biodiversity have complex social, economic, and political roots,
so environmental scientists recognize that we must also understand these aspects if we
are to develop sustainable solutions.

• Fortunately, millions of people around the world are taking action to safeguard
biodiversity and to preserve and restore Earth’s ecological and evolutionary processes.

• We will explore these efforts more fully in our discussion of biodiversity and
conservation biology in Chapter 11.

• For now, let us see how Hawaiians have been confronting the challenges to their
biodiversity.
Introduced species pose challenges for native populations and
communities

• On top of our direct effects on populations, communities, and ecosystems, human

beings exert many indirect effects—for instance, by introducing species into areas
where they do not occur naturally.

• Some such introduced species thrive in their new surroundings, killing or displacing
native species
(pp. 106–107, 304–307).

• Island species are particularly vulnerable to introduced species: They have evolved in
isolation
in small areas with a limited community of other species, and so they lack defenses
against mainland species that are well adapted to deal with a broad array of enemies.
• The Hawaiian Islands have been utterly transformed by impacts from introduced
species.

• Cattle, goats, sheep, and pigs eat native vegetation, endangering plant populations and
altering entire landscapes.

• Alien grasses, shrubs, and trees spread across the landscapes that livestock have
altered.

• Rat, cats, dogs, and mongooses eat the eggs and young of ground- nesting birds with
impunity, and have driven a number of them extinct.

• Forest birds suffering already from predation and habitat loss now also struggle
against diseases like pox and malaria.

• Pigs have made the malaria problem worse, because they dig holes in the forest floor,
where rainwater forms shallow pools in which mosquitoes breed.
• As a result, biologists and land managers have found that trying to help a species in
trouble often means trying to eradicate or control populations of another that is
doing too well.

• For instance, in many areas pigs are being hunted and pig-free areas are being
fenced off.

• However, pigs are clever animals, and some usually escape.

• Moreover, native Hawaiian people who hunt pigs have no incentive to get rid of every
last pig; otherwise, they could no longer hunt them.
Innovative solutions are working

• Amid all the challenges of Hawaii’s extinction crisis, hard work is resulting in some
inspirational success stories, and several species have been saved from imminent
extinction already.

• At Hakalau Forest, ranchland is being restored to forest, invasive plants are being
removed and native ones are being planted, and ne ̄ne ̄ are being protected while new
populations of them are being established.
• Elsewhere across Hawai‘i, tracts of public land are being managed with similar goals
and techniques, and some private landholders have joined in conservation efforts.

• Early work at Hawai‘i Volcanoes National Park inspired the work at Hakalau, as well as
efforts by managers and volunteers from the Hawai‘i Division of Forestry and
Wildlife, The Nature Conservancy of Hawai‘i, Kamehameha Schools, and local
watershed protection groups.

• People are protecting land, removing alien mammals and weeds, and restoring native
habitats.

• Offshore, conservation efforts are gaining steam as well, as Hawaiians strive to

protect their fabulous coral reefs, seagrass beds, and beaches from pollution and
overfishing.

• The northwesternmost Hawaiian Islands are now part of the largest federally
declared marine reserve (p. 461) in the world.
• Hawai‘i and its citizens are reaping benefits from their conservation efforts—
economic benefits as well as ecological ones.

• The islands’ wildlife and natural areas draw tourists from around the world, a
phenomenon called ecotourism (FIGURE 3.20).

• A large percentage of Hawaii’s tourism is eco- tourism, and tourism as a whole draws
more than 7 million visitors to Hawai‘i each year, provides thousands of jobs to
Hawaiians, and pumps $12 billion annually into the state’s economy.
Climate change now poses an extra challenge

• Traditionally, people sought to conserve populations of threatened species by

preserving and managing tracts of land (or reas of ocean) designated as protected
areas.

• However, global climate change (Chapter 18) threatens this strategy. As

temperatures climb and rainfall patterns shift, conditions within protected areas
may turn unsuitable for the species they were meant to protect.

• Hawaii’s systems are especially vulnerable.

• At Hakalau Forest on the slopes of Mauna Kea, mosquitoes and malaria are expected
to move upslope into the refuge as temperatures rise, exposing more and more birds
to disease (FIGURE 3.21).

• Meanwhile, some researchers maintain that climate change will lower the cloud layer
atop Mauna Kea, reducing rainfall at high elevations and pushing the upper limit of
the forest downward.

• If they are correct, Hakalau’s honeycreepers may become trapped within a shrinking
band of forest by malaria from below and drought from above.
• On the Hawaiian island of Kaua‘i, the outlook is worse: Forests there are closer to
the mountaintops, so climate warming is expected to shift the forests upward until
they vanish, leaving their inhabitants nowhere to go.

• Already mosquitoes have moved upslope and bird populations are diminishing.

• Two honeycreeper species, the ‘akeke‘e and the ‘akikiki, were recently added to the
Endangered Species List (p. 314).
• The challenges of climate change mean that scientists and managers need to come up
with new ways to help save declining populations.

• We will learn about the many efforts being made across the world in our exploration
of biodiversity and conservation biology in Chapter 11.

• In Hawai‘i, it remains to be seen how effectively management and ecotourism can

stem the tide of challenges and help preserve natural systems in the long term.

• Resources and efforts to preserve habitat and protect endangered species will likely
need to be stepped up.

• Programs to restore altered communities to their former condition—as is being done

at Hakalau Forest—will also be necessary.

• The restoration of ecological communities is one phenomenon we will examine in our

next chapter, as we shift from populations to communities.
Conclusion

• The honeycreepers of Hakalau Forest National Wildlife Refuge, along with many
other Hawaiian species, have helped to illuminate the fundamentals of evolution and
population ecology that are integral to environmental science.

• The evolutionary processes of natural selection, speciation, and extinction help

determine Earth’s biodiversity.

• Understanding how ecological processes function at the population level is crucial to

protecting biodiversity threatened by the mass extinction event that many
biologists maintain is now underway.

• Population ecology also informs the study of human populations (Chapter 8), another
key endeavor in environmental science.