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Topic 5: Biological Communities: The Biome Concept
Ecological Convergence (Fig. 5.1)
Cactaceae – New World Euphorbaceae – Old World
Convergence • Convergence is the process by which unrelated organisms evolve a resemblance to each other in response to common environmental conditions: – similar adaptive responses emerge in response to similar selective conditions – e.g. tree growth habit evolved independently in many plant families “battle for light with height” – e.g. mangroves - thick, leathery leaves, root projections, and viviparity “Ecological Equivalents” The Biome Concept • Character (plant and animal life) of natural terrestrial communities is determined by climate, topography, and soil. • Because of convergence, similar dominant plant forms occur under similar conditions. • Biomes are categories that group communities by dominant plant forms. Biomes - Key Points • Geographic distributions of biomes correspond closely to major climate zones. • biomes intergrade continuously and recognizing boundaries is difficult – Zones of overlap called ecotones • Matching of biomes and environment occurs because no single type of plant can endure the entire range of conditions on earth. Adaptations and Environment -- Not the Whole Story • Distributions of species are not solely a function of relationships to physical environment: – biotic interactions shape these distributions – chance and history play important roles Climate is the major determinant of plant distribution. • Climatic factors typically establish limits of plant distributions: – e.g. sugar maple, Acer saccharum, in eastern North America, is limited by… Ecological Tolerances • Several tree-sized maples in eastern North America have distributions that broadly overlap that of sugar maple: – because of different ecological tolerances, these other species exhibit distinctive environmental preferences, even when their ranges overlap… Figure 5.3
Dry sites Wet sites Moist sites
Variations in topography and soils influence plant distributions. • Topography in mountains creates a wide range of moisture conditions: – each species exhibits a local and distinctive optimum; in northern California… Environmental Optimum Form and function match the environment. • Adaptations match each species to the environment where it lives: – all species are to some extent specialized: • insect larvae from ditches and sloughs survive without oxygen longer than related species from well-aerated streams • marine snails from the upper intertidal tolerate desiccation better than their relatives from lower levels – we recognize both specialists and generalists Other Considerations • Certain species make their environments more favourable for themselves: – decaying foliage of evergreen species of poor soils produces organic acids, leaching minerals from soil – species such as beavers (Castor canadensis) are considered to be ecosystem engineers because they ‘construct’ habitats • Availability of moisture is the single most important climatic factor defining biomes: – because heat influences moisture stress, temperature and precipitation together are the determinants of boundaries of major biomes Walter’s Climate Classes (Fig. 5.4) Whittaker’s Classification of Biomes • Whittaker related major biomes to annual temperature and precipitation. • The biomes fall in a triangular area with corners representing following conditions: – warm-moist – warm-dry – cool-dry Whittaker’s Scheme (Fig. 5.5) Whittaker’s Scheme 2 • Equatorial and tropical climate zones (mean temperatures between 20oC and 30oC) – precipitation ranges from 0 to 400+ cm/yr • Temperate climate zones (mean temperatures between 5oC and 20oC) – precipitation ranges from 0 to 300+ cm/yr • Boreal and polar climate zones (mean temperatures less than 5oC) – precipitation typically below 200 cm/yr Whittaker’s Scheme - Other Considerations • Fire shapes vegetation toward drier end of spectrum within each temperature range: – typically in grassland and shrub biomes where: • moisture is intermediate (sufficient productivity for fuels to accumulate) • seasonal droughts occur (fuels dry out sufficiently to burn) – fire favours grasses and forbs over woody plants – species of these systems are adapted to or are specialized for frequent fires Major Terrestrial Biomes (Fig. 5.8) Temperate Climate Zones • Temperate zone is characterized by temperatures between 5o-20oC at low elevations, with frost throughout the zone: – found between 30oN and 45oN in North America and between 40oN and 60oN in Europe – biomes differentiated by: • total amounts and seasonality of precipitation • length of frost-free season or growing season Temperate Seasonal Forest (Fig. 5.9) Temperate Seasonal Forest (Fig. 5.9) Temperate Seasonal Forest Biome 1 • Develops under moderate climates with winter freezing: – growing season is 130-180 days – precipitation exceeds evapotranspiration • Found principally in eastern North America, Europe, and eastern Asia. • Vegetation is dominated by deciduous trees with understorey of small trees and shrubs, often abundant herbs. Temperate Seasonal Forest Biome 2 • Warmer and drier parts of the temperate seasonal forest biome are dominated by needle-leaved trees, typically pines: – found principally in North America along the Atlantic and Gulf coasts and at higher elevations in the western states – needle-leaved forests typically develop under conditions of drought and nutrient stress – fires may be frequent and species can resist fire damage Temperate Rain Forest (Fig. 5.10) Temperate Rain Forest (Fig. 5.10) Temperate Rain Forest Biome • Develops primarily in warm temperate climates: – mild winters, heavy winter rains, summer fogs common • Found principally in the northwestern US, adjacent British Columbia, southern Chile, New Zealand, Tasmania. • Vegetation is dominated by tall evergreen trees, such as Douglas fir and coastal redwood: – extensive during Mesozoic era – not as diverse as its tropical counterparts Temperate Grassland/Desert (Fig. 5.11) Temperate Grassland (Fig. 5.11) Temperate Grassland/Desert Biome 1 • Found in continental climate zones: – summers are hot and wet; winters are cold – growing season is 120-300 days – fires are a dominant influence • Extensive grasslands develop, called prairies in North America, steppes in central Asia. • Vegetation is dominated by grasses and forbs: – fire is frequent and most species have underground fire- resistant stems Temperate Grassland/Desert Biome 2 • Grasslands grade into deserts in arid continental climates: – winters are cold and summers hot – precipitation is 25-50 cm/yr – fires are infrequent because of low fuel accumulation – grazing can exert strong pressure on vegetation • Grasslands are widespread in the western US, from Great Basin southward. • Vegetation is dominated by shrubs, such as sagebrush, or small trees, such as piñon pine and juniper. Woodland/Shrubland (Fig. 5.12) Woodland/Shrubland (Fig. 5.12) Woodland/Shrubland Biome • Develops in Mediterranean-type climate (cool, wet winter, warm dry summer): – fires are frequent and most plants have adaptations to fire (resistant seeds or root crowns) • Typically found at 30-40o latitude, west coasts, common in southern Europe, southern California, central Chile, Cape region of South Africa. • Vegetation is dominated by sclerophyllous evergreen shrubs. Subtropical Desert (Fig. 5.13) Figure 5.13 Subtropical Desert Biome • These are highly variable systems found under extreme aridity: – develop at 20o-30o north and south latitude – rainfall is sparse (less than 25 mm) – creosote bush is common in subtropical American deserts, with associated cacti, shrubs, and small trees: • subtropical deserts typically have summer rainfall, with high species diversity, prominent annual flora Boreal and Polar Climate Zones • These zones have average temperatures below 5oC. • Boreal forest (taiga) develops between temperatures of 5oC and -5oC. • Tundra develops at temperatures below - 5oC. Boreal Forest (Fig. 5.14) Boreal Forest (Fig. 5.14) Boreal Forest Biome • Climate is extremely cold, with temperatures as low as -60oC in winter: – average annual temperature is below 5oC, precipitation 40-100 cm/yr – growing season is 50-100 days • Boreal forest is centered on a broad belt at 50-60 oN latitude across North America and Eurasia. • Also called taiga, vegetation of low diversity dominated by evergreen needle-leaved trees, typically spruce and fir (spruce-moose biome). Tundra (Fig. 5.15) Tundra (Fig. 5.15) Tundra Biome • Exceedingly cold climate, with brief, but active, growing season in summer: – soils are permanently frozen, thaw to depth of 0.5-1 m during brief summer growing season – precipitation is less than 60 cm/yr, but soils may be saturated because of impeded drainage • Found at high latitudes, north of boreal forest belt (but superficially similar systems occur in alpine zones). • Tundra is a treeless expanse of dwarf, prostrate woody shrubs. Equatorial and Tropical Climate Zones • Located within 20o of the equator. • Daily temperature variation exceeds monthly variation through the year. • Environments are largely distinguished by differences in the seasonal pattern of rainfall. • Frost is not a factor; plants and animals cannot tolerate freezing. Tropical Rain Forest (Fig. 5.16) Tropical Rain Forest (Fig. 5.16) Tropical Rain Forest Biome • Climate is continually warm and moist: – precipitation is in excess of 200 cm/yr, biseasonal, but never less than 10 cm in any month • Occupies three important regions, in South/Central America, West Africa, Indo-Malayan region. • These are exceedingly diverse forests, dominated by evergreen or seasonally deciduous broad- leaved trees, featuring diverse growth forms including lianas and epiphytes. Tropical Seasonal Forest/Savannah (Fig. 5.17) Tropical Seasonal Forest/Savannah (Fig. 5.17) Tropical Seasonal Forest/Savannah Biomes 1 • Found where climate is seasonally dry, but has sufficient moisture to support forest: – progressively drier tropical habitats support dry forests, thorn scrub, and true deserts • Occur worldwide within the tropics, but typically beyond 10oN or S of the equator. • Tropical seasonal forests have a preponderance of deciduous species. Tropical Seasonal Forest/Savannah Biomes 2 • Savannahs are grasslands with scattered trees. • These are typical of large areas of semiarid tropics, especially at high elevations in East Africa. • Rainfall is strongly seasonal: – 90-150 cm/yr but driest 3-4 months receive less than 5 cm each – fire and grazing play important roles in maintenance of these systems Modification of the Biome Concept for Aquatic Ecosystems • The biome concept in its strict sense does not exist for aquatic ecosystems: – independent classifications for aquatic systems, focused predominantly on physical factors… Aquatic Ecosystems - Streams • Lotic ecosystems have running water. • Streams form wherever precipitation exceeds evaporation, draining excess water. • Streams may be divided into principal habitats: – riffles (where water runs over rocky substrate) – runs (deeper stretches of slow-moving water) – riparian zone (floodplain – interface with land) • Streams exhibit continuous change in conditions from headwaters downstream, captured in river continuum concept. • Streams exhibit downstream drift of organisms/material. Streams (Fig. 5.18) Aquatic Ecosystems - Lakes • Lentic ecosystems have standing water. • Lakes form in any kind of depression (typically effects of glaciation or geological activity). • May be divided into principal habitats: – littoral zone (shallow zone with rooted vegetation) – limnetic zone (open water beyond littoral zone) – benthic zone (bottom sediments, habitat for burrowing animals and microorganisms) Lakes (Fig. 5.20) Aquatic Ecosystems - Estuaries • Are special environments at the mouths of rivers, especially where outflow is partially enclosed (such as barrier islands). • Unique because they are the interface between fresh and salt water habitats: – typically highly productive because of influx of nutrients and their rapid exchange between sediments and surface waters – often edged by extensive tidal marshes with emergent vegetation Estuaries (Fig. 5.21) Aquatic Ecosystems - Oceans • Oceans are enormously complex systems, with conditions varying with temperature, depth, current, substrate, tides. • Oceans are often classified according to depth: – littoral zone (between high and low tides, exhibits dramatic zonation) – neritic zone (beyond low tide to edge of continental shelf, often subdivided into photic and aphotic zones, typically productive) – oceanic zone (deeper waters, also divided into photic and aphotic zones, typically unproductive) Oceans (Fig. 5.22) Aquatic Ecosystems - Coral Reefs • Coral reefs are special oceanic systems: – develop in shallow waters of warm oceans – may develop around volcanoes, which may subside or erode, leaving a ring-like atoll – are typically highly productive: • nutrients released from erosion on landward side • nutrients forced up as deeper-water currents encounter the reef – are typically highly diverse Coral Reefs (Fig. 5.24) REEF and ATOLL FORMATION Summary 1 • Climate has profound effects on dominant plant forms. Each region has characteristic vegetation differing in growth form, leaf morphology, and seasonality of foliage. • Major vegetation types can be used to classify ecosystems into biomes associated with major climatic classes. • Walter’s approach classifies regions on the basis of climates having characteristic vegetation. Summary 2 • Whittaker’s approach classifies regions on the basis of vegetation having characteristic climates. • Principal climatic zones (tropical, temperate, boreal, polar) and their biomes are distinguished first on the basis of temperature, then precipitation, and its seasonality. • Aquatic systems are not classified into biomes, but we distinguish streams, lakes, estuaries, oceans, and coral reefs.