SLJCCESSION

Communities are in ;i more or less continual process of change (Clements 1910). These changes result in
part from the reactions and coactions of the organisms themselves and in part from such external forces
as changing i)hysiogra|)liy, changing climate, and organic evolution. The habitat is usually affected as
well as the community, and as the habitat changes, new species invade it and become established, and
old species di.sappear. These changes are especially noticeable in dominant s])ecies, since these species
exert a controlling role over the composition and structure of the community as a whole. The
replacement of one comnnmity by another is succession, and succession continues until a climax or final
stage is reached. -Succession is a process. The series of steps or communities comjirising a successional
sequence lead- ing to the climax is the sere. Seres are sometimes classified according to the
predominant force that is bringing them about. These forces are biolic, cli matic, physiofnalhic. and
t/eoloc/ic and their resultant seres are commonly called hioscres. clisercs, eoseres, and geoscres.

A. Productivity

A characteristic of communities that has be- come of considerable importance in modern ecological
research is productivity. The number of individuals or biomass present in a community at any one time is
the standing crop. At the beginning of the year or reproductive season the standing crop is usually small,
but as reproduction and growth take place there is an increase in the amount of organic matter making
up the biomass of the community. The pro- duction of organic matter per unit of time and area is
productivity. Productivity is commonly indicated on a yearly basis, but it is also possible to measure
monthly, weekly, or daily production. Small standing crops may have a high productivity and large stand-
ing crops a low productivity, hence average bio- mass or standing crop differentials between different
communities is not comparative of the productivity of the habitats in which these communities occur.
The largest standing crop which a habitat can support without deterioration, or the maximum number of
biomass of animals that can survive least favorable yet tolerable environmental conditions during a
stated period of time, is the carrying capacity. Carrying capacity is determined not just by the amount of
food available, but also by shelter, social tolerance, and other factors (Edwards and Fowle 1955). A
variety of methods are being used to measure productivity of different kinds of organisms and of dif-
ferent habitats. It is desirable to indicate productivity as accurately as possible in descriptions or analyses
of community dynamics.

B. biotic succession

Biotic succession is brought about by forces inherent within the community and in the activities of the
plants and animals themselves. The most important of these activities are the organismal reactions and
coactions that produce modifications in the habitat and interrelations between species. Important
reactions involve filling in of ponds with plant and animal remains, the addition of organic nutrients to
sterile soil, and the reduction in light intensity by increasing density of plant growth. With progressive
improvement of the soil and changing light and moisture conditions, a series of new dominants come
into the area. When invasion of new species occurs, in- tense competition develops ; if the invaders are
successful, the old species disappear as a new community replaces the old one. Contributing factors that
may be involved are dif ferences in growth and dispersal rates, which are different for different species.
After a forest fire or logging operation, herbaceous plant growth is immediately stimulated ; since herbs
grow rapidly, they become dominant within a year or two. Shrubs begin to spread, and tree suckers or
seedlings also appear quickly, but because they require a longer time to reach maturity several years
may elapse before they gain control of the area. Succession is considerably influenced by the kinds of
propagules available in the vicinity. Seeds, spores, and the like are dispersed more or less readily,
depending on form. Some kinds of animals roam more widely or spread more readily into new areas
than do other kinds. The composition of the community that develops and the rapidity with which it
becomes established depends, in large part, on the rates at which different species invade. When the
volcanic island of Krakatoa in the East Indies blew up in 1883, all plant and animal life on it and on two
adjacent islands was destroyed. The following year, the only living animal reported was a single small
spider. In 1886, it was apparent that the pioneer plant stage consisted of a crust of blue-green algae
covering the lava. A few mosses were also present, as were many ferns, and a scattering of some 15
species of flowering plants including 4 species of grass. The vegetation stage following the algae
consisted predominantly of ferns, but the grasses had become dominant over most of the island. By
1906, woodland had appeared, which has since de veloped into an increasingly luxuriant mixed forest.
Dispersal of seeds, spores, and other propagules was effected by wind, sea, animals, and man, in that
descending order of importance. The order in which the propagules reached the islands greatly
influenced the succession that occurred ; as vegetation developed it reacted on the soil and habitat,
bringing about conditions amenable to the return of the tropical rain forest (Richards 1952). Very little
study of animal life was made until 1908, and then only for three or four days. At that time, 202 species
were found on Krakatoa and 29 on a nearby island. There were no earthworms, snakes, or mammals
present, but there were many spiders and centipedes, a number of insects, 2 species each of land snails
and lizards, and 16 species of birds. A more thorough survey in 1921 revealed 770 species of animals
including rats, apparently introduced in 1918, and bats, first noticed in 1920. In 1933, 1100 species were
found, including 3 species of earth worms, but true forest mammals had not y§t appeared and many
families of forest birds were unrepresented. As with plants, the invasion by animals de- pended on wind,
sea, and man and other animals, in that descending order of importance. Survival and establishment of
the animal species was correlated with the stage of vegetation that was reached. It is of interest,
however, that scavenger species appeared first, then omnivores, herbivores, and finally predators and
parasites. Succession of animals depended in large part on the speed with which they reached the island
and on their finding proper food and shelter (Dammerman 1948). Bioseres may be broadly grouped as
priseres and sttbseres, depending on whether they develop on pri- mary or secondary bare areas. A
primary bare area is a sterile habitat, such as rock, sand, clay, or water. A secondary bare area is a
denudation resulting from temporary flooding, fire, logging, cultivation, overgrazing, or other
phenomenon that does not produce an extreme disturbance of the soil or substratum. Since in the latter
the habitat has already sup- ported community life, and since the soil or sub- stratum is already in an
advanced stage of develop- ment, the resulting subsere progresses rapidly and the early pioneer stages
of the prisere do not usually occur. A subsere will develop following the destruction of any stage in a
prisere, and the species composition of the stages in the subsere will be influenced by the particular
priseral community that was destroyed. The early stages or communities that make up the prisere
depend largely on the type of bare area on which the prisere originates. As succession proceeds,
however, later stages in the various seres in any area having a relatively uniform and humid climate
come to be more and more alike. The successional devel- opment from widely diversified communities in
initially different habitats to closely similar or identical climax communities in habitats that have also
become much alike is called convergence.

C.Climatic succession

With changes in climate, environmental conditions often surpass the limits of tolerance of established
plants and animals. The result is the replace- ment of the existent community by another. A most
interesting clisere is the one that has oc- curred since the northward retreat of the continental glacier of
Pleistocene time (Sears 1948, Deevey 1949, Table 21-1 ). .Stages in this clisere may be detectedand even
the relative duration of each stage measured from an analysis of the number and persistence of different
kinds of pollen grains at various depths in peat bogs. To make such studies, a core of peat is obtained
from the deepest part of the bog by means of a special hand auger. The lowest portion of the core is the
oldest ; the most recently formed is at the top. Samples of the core at various depths are suit- ably
prepared, examined under a microscope, and the pollen grains identified and counted. The predominant
kind of pollen at any level of the core represents the probable prevailing species of plants in the vicinity
at the corresponding period of time, although the proportionality between all kinds of pollen and abun-
dance of the various species may not be exact (Davis and Goodlett 1960). Thus, during the last 20,000
years, the clisere in eastern North America is repre- sented in simplified form by the following climaxes
and climates, reading downward to present time :

spruce, fir cold, moist

pine cool, dry

hemlock, oak, beech warm, moist

oak, hickory warm, dry

beech, oak, hemlock cool, moist The climate is conjectured from the relation of similar flora to climate at
the present time. The complete clisere occurs only in regions near the southern limit of the reach of the
glacier. The later stages have not developed in more northern localities where the glacier has been gone
for a shorter time and where the climate has not warmed up sufficiently. Climatic succession actually
occurs at all levels in the biosere, as the serai stages leading to one climatic climax is replaced by the
corresponding stages leading to another climax (Table 7-(y). land and floodplain forests. Stages in the
erosion cycle, as it occurs in a stream, may be discerned by examination of habitat and animal life
progressively from headwaters to mouth.

D. Physiographi succession
Changes in the earth's surface bring a change of communities. The sea alternately inundated and
retreated from the Atlantic coastal plain during the Pleistocene as ice, in which large amounts of water
were tied up, alternately melted and formed. In earlier times, the sea inundated much of the conti-
nental interior, and on its recession the eosere of plant and animal communities which developed cov-
ered vast areas. Mountain-building brings the replacement of lowland communities with new ones that
invade at higher elevations. As mountains erode, the eosere progresses in the opposite direction, until
base-level or peneplanation is attained. The development that the eosere will undergo with continued
erosion is sometimes locally apparent in the difference between up land and floodplain forests. Stages in
the erosioncycle, as it occurs in a stream, may be discerned by examination of habitat and animal life
progressively from headwaters to mouth.

E. Geologic succession

The evolution of new forms of life and dis- persal of them through the world entails replacement of pre-
existing forms and gradual change in the composition and character of communities. The first or-
ganisms to appear on earth were unicellular forms confined to the sea (Table 3-1). During the Cambrian
and Ordovician periods, the marine animal life differentiated rapidly into a rich variety of inverte- brate
types and the anlage of vertebrates. The Silurian and Devonian periods are noteworthy for the invasion
of fresh water and land by both animals and plants. Fishes became predominant both in fresh water and
the sea. During the remainder of the Paleozoic era, a luxuriant flora evolved, especially in swampy areas.
Modern conifers, such as spruce, fir, juniper, tamarack, cypress, and yew made their appearance.
Amphibians became the predominant ad- vanced animal types, and a diversified invertebrate fauna
occurred in all habitats. In the Mesozoic era, the existing land flora of giant rushes, tree-ferns, and cycads
gave way to forests of hardwoods which then spread over the world. Conifers persisted. The earliest
woody angiosperms probably originated in the Jurassic and included sas- safras and poplar. The forests
soon contained elms, oaks, maples, and magnolias. Herbaceous angio- sperms, such as grasses and
sedges, appeared towards the end of the era but did not become important in North America until the
drying up of the interior oftiie continent in the middle Cenozoic. Although the Mesozoic is predominantly
the age of the giant reptiles that lived on the land and in the water and flew through the air, less
conspicuous types such as the toothed birds, archaic mammals, and insects were de- veloping rapidly. At
the beginning of the Cenozoic era, ancient types of animals, including the great reptiles and many types
of invertebrates, became extinct and mamals rose to predominance. It is significant that the rich
development of mammals, birds, and insects came after the worldwide establishment of the
angiosperms with their rich nutrient seeds, fruits, and grasses. Pleistocene glaciation brought a major
change in the habitat of these animals, and some large mammals disappeared. The last stage of the
geosere, the Recent epoch, brought the dominance of man. Only future ages will determine whetlier this
stage is climax, or whether new and different types of animal and plant life will someday evolve to
replace man andto continue the succession into the indefinite future. As higher types evolved in each
taxonomic group, primitive forms mostly died out. The first primitive wingless insects appeared in the
Devonian ; in the Pennsylvanian there were giant forms of primitive dragonflies, cockroaches, and
grasshoppers. One cockroach had a wingspread of nearly 12 cm. By the Permian, these giant forms
disappeared and were re- placed by many modern orders. During early Mesozoic, most modern families
of insects were established, and by the Upper Cretaceous many modern genera are recognizable. Of the
two great groups of warm-blooded ani- mals, the earliest mammals had originated by the Triassic.
Modern orders did not become well diflfer- entiated until very late Cretaceous or early Paleocene,
modern families by the Oligocene, and modern genera by the Pliocene (Simpson 1953).