Chap.19 Landscape Dynamics

Chap.19 Landscape Dynamics
地景動態 Smith & Smith (1915) Elements of ecology. 9th. Ed. Pearson. 鄭先祐 (Ayo) 教授生態科學與技術學系國立臺南大學環境與生態學院

Chapter Opener Landscape showing the contrast between highly managed agricultural fields in the foreground and native forest communities occupying the hills in the background. Ayo NUTN 2017

Chapter 19 Landscape Dynamics
Ecological communities have a spatial boundary and a spatial context within the larger landscape. A mosaic (馬賽克) is the patchwork of different types of land cover. The landscape mosaic is defined by changes in the physical and biological structure of the distinct communities, called patches. Landscape ecology (地景生態學) is the study of the causes behind the formation of patches and boundaries and the ecological consequents of these spatial patterns on the landscape. Ayo NUTN 2017

Figure 19.1 Fig A view of a Virginia landscape showing a mosaic of patches consisting of different types of land cover: natural forest, plantations, fields, water, and rural development. Ayo NUTN 2017

Environmental processes create a variety of patches in the landscape. Transition zones (offer diverse conditions and habitats Patch size and shape are crucial to species diversity The theory of island biogeography(島嶼生物地理學理論) applies to landscape patches Landscape connectivity permits movement between patches Ayo NUTN 2017

Metapopulation (關聯族群) and metacommunity are central concepts in the study of landscape dynamics Frequency, intensity and scale determine the impact of disturbances (干擾) Various natural processes function as disturbances Human disturbance creates some of the most long-lasting effects The landscape represents a shifting mosaic of changing communities Ayo NUTN 2017

19.1 Environmental Processes Create a Variety of Patches in the Landscape
Ayo NUTN 2017 Patches are relatively homogenous community types that differ from their surroundings in structure (e.g., size, shape) and in species composition Patches making up the landscape mosaic result from the interactions of factors: geology, topography, soils, and climate Human activity makes its mark on the broad-scale distribution of communities Fragmentation of the natural landscape into isolated patches of forest, grassland, and shrubland

Fig. 19.3 Fragmentation and isolation of Poole Basin, Dorset, England.
Ayo NUTN 2017 Figure 19.3 Fig Fragmentation and isolation of Poole Basin, Dorset, England.

Ayo NUTN 2017 Fig Fragmentation and isolation of Poole Basin, Dorset, England. Between 1759 and 1978, the area lost 86% of its heathland (40,000 ha to 6,000 ha), changing from 10 large blocks separated by rivers to 1084 pieces – nearly half of these sites are less than 1 ha, and only 14 sites are larger than 100 ha. Many landscape patterns reflect early land-survey methods that divided land into sections This checkerboard pattern has a lasting impact on the landscape. (Fig. 19.4)

Ayo NUTN 2017 Figure 19.4 Fig Township boundaries in southwestern Wisconsin. Although the boundaries of some townships are defined by natural features on the landscape, most boundaries were established by the U.S. Rectangular Survey System used in the 18th and 19th centuries.

Ayo NUTN 2017 Landscape patches vary considerably in size and shape Determined by variations in geology and soil conditions and natural events (fire and grazing) The area, shape, and orientation of the landscape patches influence: Habitat suitability Wind flow Dispersal of seeds Movement of animals

19.2 Transition Zones Offer Diverse Conditions and Habitats
Ayo NUTN 2017 The edges of the landscape mark the perimeter of each patch Inherent edges are stable and permanent Induced edges are subject to successional changes over time A border is the place where the edge of one patch meets the edge of another Narrow and abrupt Wide with a transition zone or ecotone Perforated (穿孔的) Straight (直線的) Convoluted (迴旋的)

(a) Narrow, sharp, abrupt border (b) Wide border
Ayo NUTN 2017 Figure 19.5 Fig Types of borders (a) Narrow, sharp, abrupt border (b) Wide border (c) Convoluted (迴旋的) border (d) Perforated (穿孔的) border

Ayo NUTN 2017 Borders vary in length and have an associated vertical structure. Borders connect patches through fluxes of material, energy, and organisms. The height, width, and porosity (多孔性) of borders influence the gradients of wind flow, moisture, temperature, and solar radiation between adjoining patches.

Ayo NUTN 2017 Fig Microhabitat variation (temperature, light intensity, and relative humidity) across edges between oak woodland patches and two different matrix habitats, chaparral and grassland, in the Santa Cruz Mountains of central California. For the x-axis, positive values indicate the distance from an edge into the woodland habitat; negative values indicate the distance into the matrix habitat. Bars represent standard errors of the means. Figure 19.6

Ayo NUTN 2017 Environmental conditions in transition zones enable certain plant and animal species to colonize border environments. Plants tend to be more shade intolerant and can tolerate dry conditions. Animals usually require two or more habitat types within their home range. Edge species (邊緣物種) are those restricted exclusively to the edge environment.

Ayo NUTN 2017 Figure 19.7 Fig Map of territories of a true edge species, the indigo bunting, which inhabits woodland edges, hedgerows灌木樹籬, roadside thicket灌林叢, and large gaps in forests that create edge conditions.

Ayo NUTN 2017 Borders blend elements from all adjacent patches and offer unique habitats with relatively easy access to adjacent communities. The edge effect is the phenomenon where edge communities are often quite diverse. The edge effect can also create problems Attracts more predators Restricts dispersal

Ayo NUTN 2017 Figure 19.8 Fig Changes in vertical and horizontal structure of a border through time.

19.3 Patch Size and Shape Are Crucial to Species Diversity
Ayo NUTN 2017 Large habitat patches contain a greater number of individuals (population size) and species (species richness) than do small patches The increase in population size is a function of the increasing carrying capacity for the species, More area  more home ranges and territories Larger patches are more likely to contain variations in topography and soils Greater diversity of plant life  a wider array of habitats for animal species

Ayo NUTN 2017 Patch size and shape affect the relative abundance of edge and interior environments Only a larger patch can develop interior conditions.

Ayo NUTN 2017 Figure 19.9a Fig Relationship of habitat patch size to edge and interior conditions. All habitat patches are surrounded by edge. (a) assuming that the depth of the edge remains constant, the ratio of edge to interior decreases as the habitat size increases.

Ayo NUTN 2017 Figure 19.9b Fig (b) The general relationship between patch size and area of edge and interior. Below point A, the habitat is all edge. As size increases, interior area increases, and the ratio of edge to interior decreases.

Ayo NUTN 2017 Figure 19.9c Fig (c) This relationship of size to edge holds for a square or circular habitat patch. Long, narrow woodland islands whose widths do not exceed the depth of the edge would be edge communities, even though the area may be the same as that of square or circular ones.

Ayo NUTN 2017 Interior species (內部物種) require conditions characteristic of interior habitats and stay away from the abrupt changes associated with border environments. The probability of finding certain species may increase or decrease with patch size depending on whether they are edge or interior species.

(a) The catbird貓鵲and the robin知更鳥 are familiar edge species.
Ayo NUTN 2017 Figure 19.10a Fig Difference in habitat responses between edge species and area-sensitive or interior species. The graphs indicate the probability of detecting these species from a random point in patches of various size. Dashed lines indicate the 95% confidence intervals for the predicted probabilities. (a) The catbird貓鵲and the robin知更鳥 are familiar edge species.

Ayo NUTN 2017 Figure 19.10b Fig (b) The worm-eating warbler and ovenbird灶巢鳥are ground-nesting birds of the forest interior. The probability of finding them in small patches is low.

Ayo NUTN 2017 Figure 19.10c Fig (c) In contrast to edge and interior species, other species– such as the Carolina chickadee山雀 and Eastern wood pewee京燕– are insensitive to patch area.

Ayo NUTN 2017 The minimum habitat size needed to maintain interior species differs between plants and animals For plants, environmental conditions are more important to persistence than is patch size Several studies (e.g., R.F. Whitcomb) have revealed a pattern of increasing bird species diversity with patch size

Ayo NUTN 2017 Fig The number of bird species (species richness) plotted as a function of the area of (a) woodland or (b) grassland habitat. Area (x-axis) in both graphs is presented on a log scale. Different symbols in (a) refer to results from surveys conducted during three different time periods. Figure 19.11

19.4 The Theory of Island Biogeography Applies to Landscape Patches
Ayo NUTN 2017 Early explorers noted that large islands hold more species than do small islands J.R. Forster on Captain Cook’s voyage (1772–75) P. Darlington: on islands, a tenfold increase in land area leads to a doubling of the number of species The various patches that form the vegetation patterns across the landscape suggest islands of different sizes

Ayo NUTN 2017 Figure 19.12 Fig Number of bird species on various islands of the East Indies in relation to area (island size). The x- and y-axes are plotted on a log10 scale.

Ayo NUTN 2017 The theory of island biogeography has been applied to the study of terrestrial landscapes. R. MacArthur (Princeton University) and E.O. Wilson (Harvard University) developed this theory in 1963. The number of species established on an island represents a dynamic equilibrium between the immigration of new colonizing species and the extinction of previously established ones.

Ayo NUTN 2017 Figure 19.13 Fig According to the theory of island biogeography, immigration rate declines with increasing species richness while extinction rate increases.

Ayo NUTN 2017 Species on the mainland are the possible colonists to an uninhabited island. The species with the greatest dispersal ability will be the first to colonize the island. The immigration rate will decline as the number of species on the island increases. Few "new" species left to colonize. The immigration rate will be zero when all mainland species exist on the island.

Ayo NUTN 2017 The rate of species extinction on the island will increase with species number Based purely on chance Later immigrants will have less access to habitats and resources Competition will increase An equilibrium species richness (S) is achieved when immigration rate = extinction rate Equilibrium species richness (S) is affected by The distance of the island from the mainland Size of the island

Ayo NUTN 2017 Figure 19.14 Fig (a) Immigration rates are distance related. Islands near a mainland have a higher immigration rate and associated equilibrium

Ayo NUTN 2017 Figure 19.14a

Ayo NUTN 2017 Figure 19.14b

Figure 17.21 The Equilibrium Theory of Island Biogeography
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Regional Biogeography
Simberloff and Wilson worked with mangrove islands in Florida, where they were able to manipulate whole islands. Islands were sprayed with insecticides to remove all insects and spiders. After one year, species numbers were similar to numbers found before the experiment. Also, islands closest to a source of colonists had the most species, and the farthest island had the least. Ayo NUTN 2017

Figure 17.23 The Mangrove Experiment (Part 1)
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Figure 17.23 The Mangrove Experiment (Part 2)
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Ayo NUTN 2017 "Any patch of habitat isolated from similar habitat by different, relatively inhospitable terrain traversed only with difficulty by organisms of the habitat patch may be considered an island." (D. Simberloff) Mountaintops (山頂) Bogs (泥沼) Ponds (池塘) Dunes (沙丘) Areas fragmented by human land use Individual hosts of parasites

19.5 Landscape Connectivity Permits Movement between Patches
Ayo NUTN 2017 In some situations, corridors (廊道) connect patches of similar habitat. Strips of vegetation similar to the patches that they connect. Many corridors are of human origins (e.g., windbreaks, drainage ditches). Connectivity is the extent to which a species (or population) can move among patches within the matrix.

Fig. 19.15 Examples of corridors: (a) hedgerow灌木
Ayo NUTN 2017 Figure 19.15a Fig Examples of corridors: (a) hedgerow灌木

Fig. 19.15 Examples of corridors: (b) Riverine vegetation
Ayo NUTN 2017 Figure 19.15b Fig Examples of corridors: (b) Riverine vegetation

Ayo NUTN 2017 Corridors facilitate the movement among different patches and can encourage gene flow between subpopulations and help reestablish species in habitats that have experienced local extinction. Different-sized gaps in corridors allow certain organisms to cross while restricting others.

Ayo NUTN 2017 Negative impacts of corridors They offer scouting positions for predators Avenues for the spread of disease Provide a pathway for the invasion of exotic species If too narrow, they can inhibit the movement of social groups Corridors may provide habitats (especially in suburban and urban settings)

Ayo NUTN 2017 Roads dissect the landscape and have effects on adjacent patches of land. All types of roads affect roadside vegetation Salt spread during snow removal. Particulate matter from tires and exhaust. Chemical pollutants from automobiles. Where roads invade, people and development follow.

Field Studies: Nick M. Haddad
Ayo NUTN 2017 Corridors are thought to facilitate movement between connected patches of habitat. Increase gene flow. Promote reestablishment of locally extinct populations. Increase species diversity . Understanding the importance of corridors is useful to conservation biology.

Ayo NUTN 2017 N.M. Haddad (North Carolina State University) studies the influence of corridors on the dispersal and population dynamics of butterfly species. Savannah River Site, a National Environmental Research Park in South Carolina. Butterfly movement rates from a central patch to four surrounding peripheral patches were compared. A 25 m corridor connected the central patch to one of the peripheral patches but not to the other three patches.

Ayo NUTN 2017 Figure 1 Fig.1 Map of experimental landscape locations and aerial photograph of one landscape, sowing patch configuration.

Ayo NUTN 2017 Haddad tracked the movement patterns of the common buckeye (鹿眼蝶) and variegated (斑駁的) fritillary (豹紋蝴蝶) — both common to early successional habitats. The common buckeye was three to four times more likely to move from center patches to connected patches than to unconnected patches. The variegated fritillary was twice as likely to move down corridors than through forests when moving from the center patch. Neither was more likely to move to winged patches than to rectangular patches.

J. coenia (common buckeye) and E. Claudia (variegated fritillary)
Ayo NUTN 2017 Figure 2. Movement rates of butterfly species between connected and isolated patches. J. coenia (common buckeye) and E. Claudia (variegated fritillary) both moved between connected patches more often than between isolated patches. Figure 2

Ayo NUTN 2017 How might the corridor length influence butterfly dispersal? Interpatch distance and the presence or absence of a connecting corridor were varied. Distances: 64, 128, 256, or 384 m The movement patterns of the same species of butterfly were compared. Both species moved more frequently between patches connected by corridors. Interpatch movement was negatively related to interpatch distance.

Ayo NUTN 2017 Figure 3 Fig. 3. Mean proportion (+ standard error) of individuals (males ) Junonia (common buckeye) marked in a patch who moved one of four distances to an adjacent patch. Circles indicate mean proportions moving between patches connected by a corridor; squares indicate mean proportions moving between unconnected patches.

19.6 Metapopulation and Metacommunity
Ayo NUTN 2017 Separated populations interconnected by the movement of individuals are called metapopulations. The concept of metapopulations is central to a study of landscape dynamics because it provides a framework for examining dynamics of a discrete population on the larger landscape. The model of island biogeography can be viewed as a consequence of the metapopulation dynamics of the set of species that occupy the larger landscape.

Ayo NUTN 2017 Models of interacting metapopulations have been used to explain patterns in species succession, species richness and composition, and the food web structure of communities. Metapopulation models can be used to predict the incidence of a species on a given habitat patch as well as the expected number of species.

The set of local communities that are linked by the dispersal of multiple potentially interacting species define the metacommunity. The interaction among communities is influenced by size, shape, spatial arrangement of the habitat patches and the matrix in which they are imbedded. Habitat patches within the metacommunity are not static. Successional changes influence species composition of some patches relative to others. Ayo NUTN 2017

Metapopulations Concept 10.4: Many species have a metapopulation structure in which sets of spatially isolated populations are linked by dispersal. For many species, areas of suitable habitat exist as a series of favorable sites that are spatially isolated from one another. Metapopulations —spatially isolated populations that are linked by the dispersal of individuals or gametes. Metapopulations are characterized by repeated extinctions and colonization. Ayo NUTN 2017

Figure 10.16 The Metapopulation Concept
Members of the species occasionally disperse from one patch of suitable habitat to another. Figure The Metapopulation Concept Ayo NUTN 2017

Metapopulations Populations of some species are prone to extinction for two reasons: 1. The landscapes they live in are patchy (making dispersal between populations difficult). 2. Environmental conditions often change in a rapid and unpredictable manner. But the species persists because the metapopulation includes populations that are going extinct and new populations established by colonization. Ayo NUTN 2017

Metapopulations Extinction and colonization of habitat patches can be described by the following equation: p = Proportion of habitat patches that are occupied at time t c = Patch colonization rate e = Patch extinction rate Ayo NUTN 2017

Metapopulations The equation was derived by Richard Levins (1969, 1970), who made several assumptions: 1. There is an infinite number of identical habitat patches. 2. All patches have an equal chance of receiving colonists. 3. All patches have an equal chance of extinction. 4. Once a patch is colonized, its population increases to its carrying capacity more rapidly than the rates of colonization and extinction (allows population dynamics within patches to be ignored). Ayo NUTN 2017

Metapopulations This leads to a fundamental insight: For a metapopulation to persist for a long time, the ratio e/c must be less than 1. Some patches will be occupied as long as the colonization rate is greater than the extinction rate; otherwise, the metapopulation will collapse and all populations in it will become extinct. Ayo NUTN 2017

Metapopulations It led to research on key issues:
How to estimate factors that influence patch colonization and extinction. Importance of the spatial arrangement of suitable patches. Extent to which the landscape between habitat patches affects dispersal. How to determine whether empty patches are suitable habitat or not. Ayo NUTN 2017

Metapopulations Habitat fragmentation —large tracts of habitat are converted to spatially isolated habitat fragments by human activities, resulting in a metapopulation structure. Patches may become ever smaller and more isolated, reducing colonization rate and increasing extinction rate. The e/c ratio increases. If too much habitat is removed, e/c may shift to >1, and the metapopulation may go extinct, even if some suitable habitat remains. Ayo NUTN 2017

Metapopulations In studies of the northern spotted owl in old-growth forests in the Pacific Northwest, Lande (1988) estimated that the entire metapopulation would collapse if logging were to reduce the fraction of suitable patches to less than 19%. Ayo NUTN 2017

The northern spotted owl thrives in old-growth forests of the Pacific north-west, such forests include those that have never been cut, or have not been cut for 190 years or more. Figure The Northern Spotted Owl Ayo NUTN 2017

Metapopulations Research on the skipper butterfly in grazed calcareous grasslands in the U.K. highlighted two important features of many metapopulations: Isolation by distance. The effect of patch area (or population size—small patches tend to have small population sizes). Isolation by distance —patches that are located far from occupied patches are less like to be colonized than near patches. Patch area: Small patches may be harder to find, and also have higher extinction rates. Ayo NUTN 2017

Patches that had the largest area and were closes to occupied patches were most likely to be colonized. Figure Colonization in a Butterfly Metapopulation Ayo NUTN 2017

Metapopulations Isolation by distance can affect chance of extinction—a patch that is near an occupied patch may receive immigrants repeatedly, making extinction less likely. High rates of immigration to protect a population from extinction is known as the rescue effect. The pool frog is found in about 60 ponds along the Baltic coast in Sweden. Research to determine why pool frogs are not found in all ponds within its range included measurement of several environmental variables. Ayo NUTN 2017

Within the geographic range of the pool frog, different ponds are occupied by the species at different times. This map shows the results of three survey, in 1962, 1983, and 1987. Figure A Frog Metapopulation (Part 1) Ayo NUTN 2017

Figure 10.19 A Frog Metapopulation (Part 2)
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Metapopulations Several factors influenced the metapopulation:
Ponds far away from occupied ponds experienced low colonization rates and high extinction rates. Pond temperature —warmer ponds were more likely to be colonized successfully because breeding success was greater in them. Ayo NUTN 2017

19.7 Frequency, Intensity, and Scale Determine the Impact of Disturbances
Ayo NUTN 2017 A disturbance is any relatively discrete event — fire, windstorm, flood, extreme temperature, drought, or epidemic — that disrupts community structure and function. Disturbances create and are influenced by patterns on the landscape. Ecologists distinguish between disturbance events and disturbance patterns that characterize a landscape over time.

Ayo NUTN 2017 Disturbance regime Intensity (強度) is measured by the proportion of total biomass that is killed or eliminated — influenced by the magnitude of the physical force involved Scale (尺度) refers to the spatial extent of the impact relative to the size of the affected landscape Frequency (頻度) is the mean number of disturbances that occur within a particular time interval

Ayo NUTN 2017 Fig Map of North America showing generalized fire frequencies for major vegetation communities. Figure 19.16

Ayo NUTN 2017 Frequency is linked to a disturbance’s intensity and scale. Small-scale disturbances (the death of a tree in a forest) occur quite frequently; large-scale disturbances (fire) are rarer.

Ayo NUTN 2017 A gap is generated by a small-scale disturbance and becomes a site of localized regeneration and growth within the community Within a gap, the physical often differs substantially from conditions in the surrounding area Large-scale disturbances (e.g., fire, logging) result in substantial reduction or even elimination of local populations and significantly modify the physical environment Followed by a period of colonization

19.8 Various Natural Processes Function as Disturbances
Ayo NUTN 2017 Disturbance is a powerful force for change in the physical environment Many disturbances arise from natural causes Wind and ice storms Moving water (e.g., waves) Hurricanes Floods Lightning fires Grazing Animal activity Insect outbreaks

Ayo NUTN 2017 Figure 19.17 Fig Erosion of coastal dunes by storm surges creates breaks in the front dunes, resulting in areas of inundation (淹沒).

Ayo NUTN 2017 Fire is a major agent of disturbance and can be a major determinant of landscape patterns with the following effects Release of nutrients though pyromineralization (火礦化作用) Preparation of the seedbed (苗床) Can lead to an increase in light, water, and nutrient availability to the surviving and colonizing plants

Ayo NUTN 2017 Fire return rate is influenced by: Occurrence of droughts Biomass Burn intensity Human interference

Ayo NUTN 2017 Figure 19.18a Fig (a) Crown fire resulting in a landscape mosaic of patches of burned and unburned forest.

Ayo NUTN 2017 Figure 19.18b Fig (b) Fires of great intensity can profoundly influence ecosystems. Fire consumed the ground layer, exposing bedrock and mineral soil. The forest never recovered.

Ayo NUTN 2017 Grazing by domestic and native herbivores can cause disturbance Domestic cattle spread seeds of mesquite (豆科灌木) and other shrubs in grasslands of the southwestern United States Overpopulations of white-tailed deer (白尾鹿) have decimated (毀滅) the forest understory in eastern North America The African elephant is considered a major influence on the development of savanna communities

Ayo NUTN 2017 Figure 19.19 Fig A small beaver (水獺) dam about 2 m high along a stream in the Rocky Mountains. The reservoir of water behind the dam alters the stream flow.

Ayo NUTN 2017 Beavers modify many forested areas by damming streams — they alter the structure and dynamics of flowing water. Snow geese have affected brackish and freshwater marshes, resulting in erosion.

Ayo NUTN 2017 Outbreaks of insects (e.g., gypsy moths, spruce budworms) defoliate large areas of forest and cause the death or reduce the growth of affected trees. Human activity is ongoing and involves continuous management of an ecosystem  a more profound affect than natural disturbances Cultivating cropland for agriculture Timber harvesting

Ayo NUTN 2017 Figure 19.21 Fig Block clear-cutting in a coniferous forest in the western United States. Such cutting fragments the forest.

19.10 The Landscape Represents a Shifting Mosaic of Changing Communities
Ayo NUTN 2017 The community mosaic is ever changing due to large and small disturbances A shifting mosaic is composed of patches, each in a phase of successional development

Ayo NUTN 2017 Fig Representation of a forested landscape as a mosaic of patches in various stages of successional development. Figure 19.22 Although each patch is continuously changing, the average characteristic of the forest may remain relatively constant– in a steady state.

19.10 The Landscape Represents a Shifting Mosaic of Changing Communities
Ayo NUTN 2017 Shifting-mosaic steady state (F.H. Borman and G. Likens) Steady state is a statistical description and it describes the average state of the landscape The mosaic of patches is not static — each is continuously changing — but the average composition of the landscape may remain fairly constant. The current mosaic of land cover is maintained by active processes, many of which are forms of human-induced disturbance.

Some Burning Questions
Fire is the key to managing the longleaf pine system. Prescribed burning is used as a management tool for conserving species in many ecosystems where fire has historically been a regular disturbance. But prescribed burning can have unintended consequences. In some Florida longleaf pine forests, burning left openings that allowed cogongrass, an invasive plant from Asia, to become established. Ayo NUTN 2017

Figure 22.19 Prescribed Burning Is a Vital Management Tool in Some Ecosystems
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This grass causes fires to burn hotter, higher, and more evenly on a horizontal plane. Hotter fires cause increased mortality of longleaf pine seedlings and native wiregrass, favorable conditions for further infiltration (滲透)of cogongrass. Should managers burn or not? The timing and frequency of the prescribed burns can be crucial. Ayo NUTN 2017

People are also part of the landscape that is managed by burning. Education is necessary, as well as carefully controlled burning and safety measures. Ayo NUTN 2017

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