Environmental Biology I: Ecology as an Interdisciplinary Science, the Origin of Life and Speciation, Human Ecology, Ecosystem Structure and Function, Energy Flow, Succession, Ecotones, Niches, Stability and Services, the Types of Ecosystem and the Biomes

Unit III opens with the ecosystem. This chapter takes the unit's first five items: ecology as an interdisciplinary science, the origin of life and the modes of speciation, and human ecology and settlement; the structure and functions of ecosystems, the energy flow models, food chains and webs, ecological pyramids and succession; ecotones, edge effects, habitats and niches, stability and ecosystem services; the basis of ecosystem classification with the desert, forest, rangeland, wetland, lotic, lentic, estuarine and oceanic types; and the eight biomes the syllabus names. The ecology of energy flow and the biogeochemical cycles also appears in the geography course on this platform; here it is taught as the working ecology of an environmental scientist, with the ten per cent rule and net productivity worked.

1. Ecology as an interdisciplinary science, the origin of life, speciation and human ecology

Ernst Haeckel coined the word ecology in 1866 for the study of organisms in relation to their environment. It is interdisciplinary by necessity: energy flow is physics, nutrient cycling is chemistry, soils and climate are earth science, population dynamics is mathematics, and human ecology draws on sociology and economics. It is studied at nested levels, from the individual through the population, community and ecosystem to the landscape and the biosphere, with new properties emerging at each level.

The Oparin-Haldane hypothesis of the 1920s proposed that life arose by chemical evolution in a reducing early atmosphere without free oxygen, organic molecules accumulating as a "primordial soup". Miller and Urey (1953) passed electric sparks through methane, ammonia, hydrogen and water and obtained amino acids, showing that the building blocks can form abiotically. Later ideas include the RNA world, in which RNA served both as genetic material and catalyst, and the possible role of deep-sea hydrothermal vents. The earliest widely accepted fossils, microbial mats called stromatolites, are about 3.5 billion years old, and oxygenic photosynthesis by cyanobacteria later oxygenated the atmosphere in the Great Oxidation Event, about 2.4 billion years ago.

Mode of speciationHow populations become isolatedExample
AllopatricA geographical barrier splits a population, and the parts divergeSpecies on either side of a mountain range or on different islands
PeripatricA small group colonises a new area at the edge of the rangeDarwin's finches on the Galápagos
ParapatricAdjacent populations diverge across an environmental gradient with limited gene flowGrasses tolerant of heavy metals on mine spoil beside normal pasture
SympatricNew species arise within the same area, often by polyploidy or a shift to a new hostPolyploid wheat; the apple maggot fly shifting from hawthorn to apple

Mayr's biological species concept defines a species as a group of interbreeding natural populations reproductively isolated from other such groups; isolation may be prezygotic (habitat, timing, behaviour, mechanical incompatibility) or postzygotic (hybrid inviability or sterility, as in the mule). Human ecology, developed by the Chicago school of Park and Burgess in the 1920s, applied ecological ideas such as competition, invasion and succession to cities. An urban settlement is a heterotrophic ecosystem: it imports food, water, energy and materials from a hinterland many times its own area and exports wastes, which is why its ecological footprint far exceeds its boundaries.

2. Ecosystem structure and function: energy flow, food chains and webs, pyramids and cycles

An ecosystem, a term introduced by Tansley in 1935, has abiotic components (energy, climate, water, inorganic substances and organic compounds such as humus) and biotic components: producers (autotrophs), consumers (herbivores, carnivores, omnivores) and decomposers (bacteria and fungi, with detritivores). Its two basic functions are the one-way flow of energy and the cycling of matter. Gross primary productivity (GPP) is the total rate of photosynthesis; net primary productivity (NPP) is what remains after the producers' own respiration, NPP = GPP − R, and it is the energy actually available to consumers. Tropical rainforests, estuaries, coral reefs and swamps have the highest NPP per unit area, and the open ocean and deserts the lowest, though the ocean's great area makes it a large total producer.

Odum's energy flow models show the same principle at increasing detail. The single-channel model traces energy from producers through herbivores to carnivores, with losses to respiration at every step. The Y-shaped model splits the flow into a grazing food chain (plants to herbivores to carnivores) and a detritus food chain (dead organic matter to decomposers and detritivores to their predators); in most forests and many aquatic systems far more energy passes through the detritus chain. The universal model applies to any living component, showing ingested energy divided into assimilation and egestion, and assimilation into respiration and production. Food chains interlock into food webs, which give consumers alternative prey and so stabilise the community. Because only about 10% of energy passes from one level to the next (Lindeman, 1942), if producers fix 20,000 kJ, herbivores receive about 2,000 kJ and carnivores about 200 kJ, which is why food chains rarely exceed four or five links.

Ecological pyramidAlways upright?Inverted example
NumbersNoA single large tree supporting thousands of insects; a host with many parasites
Biomass (standing crop)NoOpen ocean, where a small standing crop of fast-dividing phytoplankton supports a larger biomass of zooplankton
Energy (productivity)YesNone, because energy is lost at every transfer

Matter, unlike energy, cycles. Biogeochemical cycles are of two broad types: gaseous cycles, such as those of carbon, nitrogen and oxygen, have their main reservoir in the atmosphere or ocean and are relatively well buffered; sedimentary cycles, such as that of phosphorus, have their reservoir in rocks and sediments, turn over slowly and are easily disrupted, which is why phosphorus lost to the sea is effectively lost to the land for geological time. Each cycle has a large, slow reservoir pool and a smaller, active exchange pool between organisms and their surroundings.

3. Ecological succession

Succession is the orderly, directional change in the species composition of a community over time. Primary succession begins on a surface never before colonised, such as bare rock, a lava flow or a new sand dune, and is slow because soil must first form; secondary succession follows disturbance of an existing community, such as an abandoned field or a burnt forest, where soil remains, and is faster. On bare rock (a lithosere, a kind of xerosere) the sequence runs from crustose lichens, which begin weathering, to foliose lichens, mosses, herbs, shrubs and finally forest; in water (a hydrosere) from phytoplankton to submerged plants, floating plants, reed swamp, sedge meadow, woodland and forest, as sediment and organic matter fill the basin. Changes caused by the organisms themselves are autogenic; those caused by outside forces such as siltation or climate are allogenic.

The end point is the climax community, in equilibrium with its environment. Clements's monoclimax theory held that each climatic region has a single climax; Tansley's polyclimax theory allowed several climaxes set by soil, topography and fire; and Whittaker's climax-pattern hypothesis sees a continuous pattern of climax communities tracking environmental gradients. Connell and Slatyer (1977) proposed three mechanisms by which early species affect later ones: facilitation, tolerance and inhibition. Odum (1969) summarised the trends of a developing ecosystem: biomass and organic matter increase, species diversity and structural complexity increase, food chains become web-like and detritus-based, nutrient cycles become more closed, K-selected species replace r-selected ones, and the ratio of gross production to community respiration (P/R) moves towards one.

⚠️ Pioneers on rock are crustose lichens, not mosses
Mosses need some soil and moisture, which the lichens create by secreting acids and trapping dust. In a hydrosere the pioneers are phytoplankton. An option naming mosses or grasses as the first colonists of bare rock is wrong.

4. Ecotones, edge effects, habitats and niches, ecosystem stability and ecosystem services

An ecotone is the transition zone between two communities, such as a forest edge meeting grassland, or a mangrove between land and sea. It often holds species of both communities plus some found only there, and the resulting tendency for greater diversity and density at the junction is the edge effect; species that prefer such zones are edge species. Fragmentation multiplies edges, which benefits edge species but harms interior species that need large blocks of undisturbed habitat. The habitat is where an organism lives, its address; the niche is its role, its profession. Grinnell (1917) used niche mainly in the spatial sense, Elton (1927) for an organism's functional place in the community, and Hutchinson (1957) defined it as an n-dimensional hypervolume of all the conditions and resources in which a species can persist. The fundamental niche is that hypervolume without competitors; the realised niche is the narrower part actually occupied in their presence. By Gause's competitive exclusion principle, two species with identical niches cannot coexist indefinitely.

Stability has two components: resistance, the ability to withstand a disturbance without changing, and resilience, the ability to return to the original state after one. The diversity-stability hypothesis of MacArthur (1955) and Elton (1958) holds that more diverse communities, with more pathways for energy, are more stable. Robert May (1973) showed mathematically that randomly assembled complex systems tend to be less stable, and the modern view is that real communities are stabilised not by complexity as such but by particular structures, such as many weak interactions, functional redundancy among species and the insurance that different species respond differently to change. Other factors affecting stability are the presence of keystone species, the frequency and intensity of disturbance, and the connectivity of habitat.

Ecosystem service (Millennium Ecosystem Assessment, 2005)Examples
ProvisioningFood, fresh water, timber, fibre, fuel, genetic resources, medicines
RegulatingClimate regulation, flood control, water purification, pollination, pest and disease control
CulturalRecreation, tourism, spiritual and religious values, aesthetic inspiration, education
SupportingSoil formation, nutrient cycling, primary production, which underpin all the others

Costanza and colleagues (1997) estimated the value of the world's ecosystem services at about 33 trillion US dollars a year, more than the global economic output of the time, a figure intended less as a price than as a demonstration that services treated as free are not.

5. The basis of classification and the types of ecosystem

Ecosystems are classified by habitat (terrestrial or aquatic, and within aquatic, freshwater, estuarine or marine), by origin (natural or human-made, such as croplands, plantations and reservoirs), by the dominant vegetation or physiognomy, and, in Odum's energetic scheme, by the source of energy: unsubsidised natural solar-powered systems such as the open ocean, naturally subsidised ones such as estuaries enriched by tides, human-subsidised solar-powered ones such as farms, and fuel-powered urban-industrial systems.

Ecosystem typeDefining featuresIndian example
Hot desertRainfall usually below 250 mm, high temperature range; plants with succulence, CAM photosynthesis and deep rootsThar Desert, Rajasthan
Cold desertHigh altitude, very low temperature and precipitation, strong radiationLadakh, Lahaul-Spiti
ForestTree-dominated; stratified canopy; the largest store of terrestrial carbonTropical evergreen forests of the Western Ghats; sal forests of central India
RangelandGrasslands, shrublands and savannas used for grazing; productivity limited by rainfall and grazing pressureBanni grassland, Kachchh; shola grasslands of the Nilgiris
WetlandMarsh, fen, peatland or water, natural or artificial, permanent or temporary, fresh, brackish or salt, including marine water not deeper than six metres at low tide (Ramsar definition)Chilika, Keoladeo, Loktak, the East Kolkata Wetlands
Lotic (running water)Current, high oxygen, organisms with attachment and streamlining; changes from source to mouth (river continuum concept, Vannote and colleagues, 1980)Himalayan torrents to the lower Ganga
Lentic (standing water)Littoral, limnetic, profundal and benthic zones; temperate lakes stratify into epilimnion, thermocline and hypolimnion, overturning in spring and autumnDal Lake, Wular Lake, village ponds
Estuarine (mangrove)Mixing of fresh and sea water, fluctuating salinity, very high productivity; mangroves with pneumatophores and viviparyThe Sundarbans; Bhitarkanika
OceanicNeritic and oceanic zones; a sunlit photic zone over a dark aphotic zone; productivity concentrated in upwelling and coastal watersThe Arabian Sea and Bay of Bengal, with seasonal upwelling off the south-west coast

6. Biomes: concept, classification and distribution

A biome is a large regional community of plants and animals with a characteristic physiognomy, determined above all by climate: Whittaker's diagram places the world's biomes on axes of mean annual temperature and precipitation. Biomes grade into each other along latitude, and a similar zonation repeats with altitude on high mountains. The syllabus spells chaparral as "Chapparal"; the correct spelling has one p.

BiomeClimateVegetation, soil and fauna
TundraVery cold, short growing season of about two months, low precipitation; permafrostMosses, lichens, sedges, dwarf shrubs; no trees; caribou, arctic fox, migratory birds
Taiga (boreal forest)Long cold winters, short summers, moderate precipitationEvergreen conifers such as spruce, fir and pine; acidic podzols; moose, lynx, wolves
Temperate grasslandHot summers, cold winters, 250 to 750 mm rainGrasses on deep, fertile chernozems; the prairies, steppes, pampas and veld; bison and grazing herds
Temperate deciduous forestFour distinct seasons, rain spread through the yearBroad-leaved trees such as oak, maple and beech that shed leaves in autumn; fertile brown forest soils
Highland icy alpineAbove the tree line on high mountains; cold, windy, intense ultravioletCushion plants, alpine meadows, rhododendron scrub; snow leopard, ibex, yak in the Himalaya
Chaparral (Mediterranean scrub)Hot, dry summers and mild, wet wintersDense evergreen sclerophyllous shrubs adapted to fire; California, the Mediterranean basin, central Chile, the Cape of South Africa, south-west Australia
SavannaWarm all year with distinct wet and dry seasonsGrassland with scattered trees, maintained by fire and grazing; the great herds and predators of East Africa
Tropical rainforestHot and wet all year, often over 2,000 mm of rainMulti-layered evergreen forest with the highest species richness on land; most nutrients are held in the vegetation, and the leached soils are poor once it is cleared

Key takeaways

  • Life probably arose by chemical evolution (Oparin-Haldane, Miller-Urey); species form by allopatric, peripatric, parapatric and sympatric routes; cities are heterotrophic ecosystems.
  • NPP = GPP − R; energy flows one way through grazing and detritus chains, about 10% passing each level, so the energy pyramid is always upright while those of number and biomass can invert.
  • Succession runs from pioneers (crustose lichens on rock, phytoplankton in water) to a climax, with biomass, diversity and closed nutrient cycles increasing and P/R approaching one.
  • Ecotones show the edge effect; Hutchinson's niche is an n-dimensional hypervolume, fundamental or realised; stability combines resistance and resilience; services are provisioning, regulating, cultural and supporting.
  • Ecosystem types run from hot and cold deserts to lotic, lentic, estuarine and oceanic waters, and biomes from tundra and taiga to chaparral (the syllabus's "Chapparal"), savanna and tropical rainforest, each set chiefly by temperature and rainfall.

Practice questions (10)

Attempt each one before opening the answer. Every explanation names the tempting wrong option as well as the right one, because that is where marks are lost.

  1. Which ecological pyramid is always upright?

    1. The pyramid of numbers
    2. None of the three pyramids, since all can be inverted in some ecosystems
    3. The pyramid of biomass
    4. The pyramid of energy
    Show answer

    Answer: D — The pyramid of energy

    Energy is lost as heat at every transfer, so each level must have less energy flow than the one below it. The pyramid of numbers inverts under a single large tree and that of biomass in the open ocean, where fast-turning phytoplankton support a larger standing crop of zooplankton.
  2. Producers in a grassland fix 20,000 kJ of energy. Assuming 10% transfer between successive trophic levels, how many kJ reach the secondary consumers?

    Numerical answer — type the value.

    Show answer

    Answer: 200

    Primary consumers (herbivores) receive 10% of 20,000 = 2,000 kJ, and secondary consumers (the carnivores that eat them) receive 10% of that, 200 kJ. Stopping at 2,000 kJ confuses primary with secondary consumers.
  3. Hutchinson's concept of the ecological niche is best described as

    1. an n-dimensional hypervolume of the conditions and resources within which a species can persist
    2. the position of an organism in a food chain only
    3. the physical place where an organism lives
    4. the transition zone between two communities
    Show answer

    Answer: A — an n-dimensional hypervolume of the conditions and resources within which a species can persist

    Hutchinson (1957) defined the niche as a multidimensional space of all the environmental variables a species tolerates and the resources it uses. The physical place is the habitat, the transition zone is an ecotone, and a purely trophic position is closer to Elton's functional niche.
  4. Which of the following are trends expected as an ecosystem develops towards climax, according to Odum? Select all that apply.

    1. Total biomass and organic matter increase
    2. Nutrient cycles become more open and leaky
    3. The ratio of gross production to community respiration approaches one
    4. r-selected species increasingly dominate
    Show answer

    Answer: A — Total biomass and organic matter increase; C — The ratio of gross production to community respiration approaches one

    In a maturing ecosystem biomass accumulates and production comes into balance with respiration, so P/R moves towards one. Nutrient cycles become more closed, conserving nutrients, and K-selected species replace the r-selected pioneers.
  5. A biome with hot, dry summers, mild, wet winters and dense, fire-adapted evergreen sclerophyllous shrubs is the

    1. taiga
    2. chaparral
    3. savanna
    4. tundra
    Show answer

    Answer: B — chaparral

    That is the Mediterranean climate of the chaparral, found in California, the Mediterranean basin, central Chile, the Cape and south-west Australia; the syllabus spells it "Chapparal". Savanna has a warm wet-and-dry climate with grass and scattered trees, and taiga and tundra are cold biomes.
  6. Assertion (A): In the open ocean the pyramid of biomass can be inverted. Reason (R): Phytoplankton reproduce so fast that a small standing crop can support a larger biomass of zooplankton. Choose the correct option.

    1. Both A and R are true, and R is the correct explanation of A
    2. Both A and R are true, but R is not the correct explanation of A
    3. A is true, but R is false
    4. A is false, but R is true
    Show answer

    Answer: A — Both A and R are true, and R is the correct explanation of A

    Both are true and R explains A. Biomass is a snapshot of standing crop, not a rate; phytoplankton turn over in days, so their small standing stock produces enough to feed a larger, slower-turning mass of consumers. The pyramid of energy, which measures rates, stays upright.
  7. In the Millennium Ecosystem Assessment, soil formation, nutrient cycling and primary production are classed as

    1. cultural services
    2. supporting services
    3. provisioning services
    4. regulating services
    Show answer

    Answer: B — supporting services

    Supporting services are the underlying processes on which all others depend. Provisioning services are products such as food and timber, regulating services include climate regulation and pollination, and cultural services are non-material benefits such as recreation and spiritual value.
  8. Which statements about thermal stratification in a temperate lake are correct? Select all that apply.

    1. The hypolimnion may become depleted of oxygen during summer stratification
    2. The thermocline is the layer in which temperature falls rapidly with depth
    3. The epilimnion is the cold, dense bottom layer
    4. Spring and autumn overturns mix oxygen and nutrients through the water column
    Show answer

    Answer: A — The hypolimnion may become depleted of oxygen during summer stratification; B — The thermocline is the layer in which temperature falls rapidly with depth; D — Spring and autumn overturns mix oxygen and nutrients through the water column

    In summer a warm, light epilimnion floats over a cold, dense hypolimnion, separated by the thermocline, and decomposition can strip the isolated hypolimnion of oxygen. When surface water cools or warms to about 4 °C in autumn and spring, density differences vanish and the lake mixes. The epilimnion is the warm upper layer, not the bottom one.
  9. Speciation that occurs when a population is split by a geographical barrier such as a mountain range is

    1. allopatric speciation
    2. parapatric speciation
    3. sympatric speciation
    4. polyploid speciation
    Show answer

    Answer: A — allopatric speciation

    Allopatric means "other homeland": the barrier stops gene flow and the isolated populations diverge. Sympatric speciation happens within one area, often through polyploidy, and parapatric speciation happens across a gradient between adjacent populations still in contact.
  10. A forest has a gross primary productivity of 25,000 kcal/m²/yr, and its plants use 60% of this in respiration. If primary consumers obtain 10% of the net primary productivity, how many kcal/m²/yr do they obtain?

    Numerical answer — type the value.

    Show answer

    Answer: 1000

    NPP = GPP − R = 25,000 − 0.6 × 25,000 = 10,000 kcal/m²/yr, and 10% of that is 1,000 kcal/m²/yr. Taking 10% of the GPP (2,500) ignores the plants' own respiration, which is exactly the part consumers can never obtain.