Evolution and measurement of species diversity

SUMMARY
1According to ‘Gause's hypothesis’ a corollary of the process of evolution by natural selection is that in a community at equilibrium every species must occupy a different niche. Many botanists have found this idea improbable because they have ignored the processes of regeneration in plant communities.
2Most plant communities are longer-lived than their constituent individual plants. When an individual dies, it may or may not be replaced by an individual of the same species. It is this replacement stage which is all-important to the argument presented.
3Several mechanisms not involving regeneration also contribute to the maintenance of species-richness:


(a). differences in life-form coupled with the inability of larger plants to exhaust or cut off all resources, also the development of dependence-relationships,
(b) differences in phenology coupled with tolerance of suppression,
(c) fluctuations in the environment coupled with relatively small differences in competitive ability between many species,
(d) the ability of certain species-pairs to form stable mixtures because of a balance of intraspecific competition against interspecific competition,
(e) the production of substances more toxic to the producer-species than to the other species,
(f) differences in the primary limiting mineral nutrients or pore-sizes in the soil for neighbouring plants of different soecies, and
(g) differences in the competitive abilities of species dependent on their physiological age coupled with the uneven-age structure of many populations.


4The mechanisms listed above do not go far to explain the indefinite persistence in mixture of the many species in the most species-rich communities known.
5In contrast there seem to be almost limitless possibilities for differences between species in their requirements for regeneration, i.e. the replacement of the individual plants of one generation by those of the next. This idea is illustrated for tree species and it is emphasized that foresters were the first by a wide margin to appreciate its importance.
6The processes involved in the successful invasion of a gap by a given plant species and some characters of the gap that may be important are summarized in Table 2.
7The definition of a plant's niche requires recognition of four components:


(a) the habitat niche,
(b) the life-form niche,
(c) the phenological niche, and
(d) the regeneration niche.


8A brief account is given of the patterns of regeneration in different kinds of plant community to provide a background for studies of differentiation in the regeneration niche.
9All stages in the regeneration-cycle are potentially important and examples of differentiation between species are given for each of the following stages:


(a) Production of viable seed (including the sub-stages of flowering, pollination and seed-set),
(b) dispersal, in space and time,
(c) germination,
(d) establishment, and
(e) further development of the immature plant.


10In the concluding discussion emphasis is placed on the following themes:


(a) the kinds of work needed in future to prove or disprove that differentiation in the regeneration niche is the major explanation of the maintenance of species-richness in plant communities,
(b) the relation of the present thesis to published ideas on the origin of phenological spread,
(c) the relevance of the present thesis to the discussion on the presence of continua in vegetation,
(d) the co-incidence of the present thesis and the emerging ideas of evolutionists about differentiation of angiosperm taxa, and
(e) the importance of regeneration-studies for conservation.

The maintenance of species-richness in plant communities: the importance of the regeneration niche

Identification of the selective forces on plant dispersal engenders theoretical argument, empirical study, and speculation. We separate evidence, testable hypotheses, and conjecture surrounding two major questions in dispersal ecology. The first asks what ecological, and ultimately evolutionary, advantages exist in seed dispersal. Astonishingly little is known about the advantages to a parent plant that are actually conferred by investment in dispersal structures. Does dispersal enable seeds and ultimately seedlings to escape mortality near the parent? Is continual recolonization of unstable habitats the primary advantage? Must seeds find rare microhabitats suitable for reestablishment? Such issues are addressed through joint consideration of dispersal and establishment-those stages both mediated by parental provisioning and subject to the highest mortality in the life of a plant. The second broad question asks what general and explicit environmental forces influence the timing and mode of dispersal. Do climates or seasons favor one dispersal mode over another? Do differences in number, size, morphology, or nutritional quality of fruits influence frugivore choice, and consequently differential dispersal of species or individuals within species? Studies of dispersal process and mode should be intimately connected. A general objective of this paper is to explore the degree to which they are integrated and, in so doing, to catalyze their union. We emphasize topics most in need of critical attention: the evolutionary ecology of dispersal process and mode. Excellent recent reviews consider such related topics as dispersal mechanism (131, 184), seed dormancy (1, 30), phytogeography (11, 115, 146), masting and predator satiation (105, 156), and succession (68, 69, 189).

https://www.jstor.org/stable/info/2097067

Ecology of Seed Dispersal

The effects of initial nitrogen and lignin contents of six species of hardwood leaves on their decomposition dynamics were studied at the Hubbard Brook Experimental Forest. Rate con- stants (k) for annual leaf mass loss ranged from -0.08 to -0.47. The rate constants (k) had a negative linear correlation (r2 = .89) with the ratio of initial lignin concentration to initial nitrogen concentra- tion. Decomposition dynamics of the litter materials were described by inverse linear relationships between the percentage of original mass remaining and the nitrogen concentration in the residual material. Initial lignin concentration was highly correlated (r2 = .93) with the slope of the inverse linear relationship for each of the litter types.

Nitrogen and Lignin Control of Hardwood Leaf Litter Decomposition Dynamics

An understanding of community structure should be based on evidence that the growth and regulation of the component populations in the community are affected in a predictable manner by natural physical disturbances and by interactions with other species in the community. This study presents an experimental evaluation of the effects of such disturbances and competitive interactions on populations of sessile organisms in the rocky intertidal community, for which space can be demonstrated to be the most important limiting resource. This research was carried out at eight stations on the Washington coastline which have been ranked according to an exposure/desiccation gradient and subjected to comparable manipulation and observation. Physical variables such as wave exposure, battering by drift logs, and desiccation have important effects on the distribution and abundance of many of the sessile species in the community. In particular, wave exposure and desiccation have a major influence on the distribution patterns of all the algae and of the anemone Anthopleura elegantissima. The probability of damage from drift logs is very high in areas where logs have accumulated along the intertidal. Log damage and wave exposure have complementary effects in the provision of free space in a mussel bed, as wave shock enlarges a patch created by log damage by wrenching the mussels from the substratum at the periphery of the bare patch. Competition for primary space results in clear dominance hierarchies, in which barnacles are dominant over algae. Among the barnacles, Balanus cariosus is dominant over both B. glandula and Chthamalus dalli; B. glandula is dominant over C. dalli. The mussel Mytilus californianus requires secondary space (certain algae, barnacles, or byssal threads) for larval settlement, but is capable of growing over all other sessile species and potentially is the competitive dominant of space in the community.

Competition, Disturbance, and Community Organization: The Provision and Subsequent Utilization of Space in a Rocky Intertidal Community

Publisher Summary This chapter reviews the ecology of fire in grasslands. It describes several generalizations such as environmental alterations, effects on species of plants, effects on vegetation, and associated animals. The grassland essentially includes any herb-dominated vegetation, herb dominated layers of savanna or open forests. Lightning is considered a universal natural cause of vegetation fires. A wide variety of other non-anthropeic causes of fire include volcanic eruptions, spontaneous combustion, and the occurrence of sparks when boulders roll down a slope. It is noted that these natural fires have been occurring from the earliest appearance of land vegetation. Their effect is to injure a plant community to a greater or lesser degree, after which the community escapes similar disturbance for sufficient time to allow a restoration of its former structure and composition. Grass fires are characterized by a rather narrow zone of flames advancing across a finely divided and homogeneously dispersed fuel. The fine texture of fuel makes it more sensitive to weather change than is true of woody aerial stems.

Ecology of Fire in Grasslands

Steppe vegetation of Washington.

Plant communities: a textbook of plant synecology.

During the rainy season the semi-deciduous forest in tropical north-western Costa Rica bears a close rememblance to deciduous forests of temperate eastern U.S.A. and central Europe in their summer aspect. This physiognomic resemblance is ecologically striking since the former temperature varies but little through the year with rainfall differing markedly, whereas in the latter precipitation is abundant at all seasons but temperatures are strongly contrasted between summer and winter. During a year's residence in the Costa Rican forest, records were made of tree growth, flowering, and fruiting with the objective of characterizing such features of the vegetation more closely than has yet been done, and where feasible drawing comparisons with the temperate zone forest.

http://www.jstor.org/stable/pdfplus/2258048.pdf

Rexford F. Daubenmire

Papers

Ecology of Fire in Grasslands

Steppe vegetation of Washington.

Plant communities: a textbook of plant synecology.

Phenology and Other Characteristics of Tropical Semi-Deciduous Forest in North-Western Costa Rica

Plants and environment