Preface to the Princeton Landmarks in Biology Edition vii Preface xi Symbols Used xiii 1. The Importance of Islands 3 2. Area and Number of Speicies 8 3. Further Explanations of the Area-Diversity Pattern 19 4. The Strategy of Colonization 68 5. Invasibility and the Variable Niche 94 6. Stepping Stones and Biotic Exchange 123 7. Evolutionary Changes Following Colonization 145 8. Prospect 181 Glossary 185 References 193 Index 201

The Theory of Island Biogeography

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

Seed dormancy is an innate seed property that defines the environmental conditions in which the seed is able to germinate. It is determined by genetics with a substantial environmental influence which is mediated, at least in part, by the plant hormones abscisic acid and gibberellins. Not only is the dormancy status influenced by the seed maturation environment, it is also continuously changing with time following shedding in a manner determined by the ambient environment. As dormancy is present throughout the higher plants in all major climatic regions, adaptation has resulted in divergent responses to the environment. Through this adaptation, germination is timed to avoid unfavourable weather for subsequent plant establishment and reproductive growth. In this review, we present an integrated view of the evolution, molecular genetics, physiology, biochemistry, ecology and modelling of seed dormancy mechanisms and their control of germination. We argue that adaptation has taken place on a theme rather than via fundamentally different paths and identify similarities underlying the extensive diversity in the dormancy response to the environment that controls germination.

/pdf/seed-dormancy-and-the-control-of-germination-4tdhm2985s.pdf

Seed dormancy and the control of germination

The proposal is made that seed scientists need an internationally acceptable hierarchical system of classification for seed dormancy. Further, we suggest that a modified version of the scheme of the Russian seed physiologist Marianna G. Nikolaeva be adopted. The modified system includes three hierarchical layers – class, level and type; thus, a class may contain levels and types, and a level may contain only types. The system includes five classes of dormancy: physiological dormancy (PD), morphological dormancy (MD), morphophysiological dormancy (MPD), physical dormancy (PY) and combinational dormancy (PY + PD). The most extensive classification schemes are for PD, which contains three levels and five types (in the non-deep level), and MPD, which contains eight levels but no types. PY is not subdivided at all but probably should be, for reasons given. Justifications are presented for not including mechanical dormancy or chemical dormancy in the modified scheme. PD (non-deep level) is the most common kind of dormancy, and occurs in gymnosperms (Coniferales, Gnetales) and in all major clades of angiosperms. Since, first, this is the class and level of dormancy in seeds of wild populations of Arabidopsis thaliana and, secondly, Type 1 (to which seeds of A. thaliana belong) is also common, and geographically and phylogenetically widespread, it seems that biochemical, molecular and genetic studies on seed dormancy in this model species might have rather broad application in explaining the basic mechanism(s) of physiological dormancy in seeds.

/pdf/a-classification-system-for-seed-dormancy-33n2carxeu.pdf

A classification system for seed dormancy

Maternal effects in plants were recognized as long ago as 1909 (32). Recent evidence, primarily over the last 15-20 years, shows that maternal effects can contribute substantially to the phenotype of an individual, and as we show, this has important consequences for the interpretation and' design of both ecofogical and genetic studies. Following a discussion of the consequences of maternal effects and an analysis of the different ways these effects can be estimated, we review the evidence for maternal effects from the fields of physiological ecology, crop science, and quantitative genetics. We do not review all of the literature because that would be a monumental task; rather, we focus on representative studies from each of these fields. It is our contention that despite evidence that maternal effects can have a large in­ fluence on offspring phenotype, few detailed studies have identified the specific causes of maternal effects, particularly in natural populations.

Maternal effects in plants

Seed dormancy in isanthus brachiatus (labiatae)

Intermediate complex morphophysiological dormancy in seeds of the cold desert sand dune geophyte Eremurus anisopterus (Xanthorrhoeaceae; Liliaceae s.l.).

of the dispersal distance-seed density curve was concave. Number of seeds retained on dispersing plants declined exponentially with increase in time after beginning of dispersal. The possible ecological advantages of an extended period of seed dispersal are discussed.

Ecological life history of the facultative woodland biennial Arabis laevigata variety laevigata (Brassicaceae): seed dispersal1

This chapter discusses the inheritance of seed dormancy, dormancy breaking, and germination requirements. It also covers variations in germination responses of seeds from different populations of the same species. Environmental factors are known to influence the mother plant, thereby causing changes in the germination characteristics of the progeny (seeds). However, the kinds of changes that can occur in seeds, as they develop under varying environmental conditions, also need to be considered. The chapter illustrates seed polymorphism, and cleistogamous or chasmogamous seeds representing interplay of genetics, environment, and genetics X environmental factors along with a discussion of their consequences on seedling recruitment. It explores the reasons for variation in seed dormancy breaking and germination requirements within a species. Genetics and the environment of the mother plant during the time of seed maturations are two most important factors controlling this variation. Thus, research is often required to determine the relative importance of each of these two factors in the variation in seed dormancy and germination of a particular species.

Causes of Within-Species Variations in Seed Dormancy and Germination Characteristics

Many studies have compared diaspore dispersal ability and degree of dormancy in the two diaspores of dimorphic plant species. A primary goal of these studies was to determine if germination and dispersal characteristics of the two morphs fit within a high risk–low risk (bet-hedging) life-history strategy, i.e. high dispersal/low dormancy in one morph versus low dispersal/high dormancy in the other one. In a survey of 26 papers on 28 diaspore dimorphic species, we found that in 12 of the studies, which were published between 1978 and 2008, seeds were stored, and thus possibly afterripened, before they were tested for germination. The 14 papers that tested fresh seeds were published between 1963 and 2010. Failure to test fresh seeds likely resulted in misinterpretation of the diaspore dispersal/dormancy strategy in some of the species investigated. We conclude that it is imperative that fresh seeds be tested for germination in order to be certain that the correct relationship between dispersal and dormancy is elucidated, and thus that the correct interpretation is made concerning life-history strategy and bet-hedging, in dimorphic species.

Jerry M. Baskin

Papers

Seed dormancy in isanthus brachiatus (labiatae)

Intermediate complex morphophysiological dormancy in seeds of the cold desert sand dune geophyte Eremurus anisopterus (Xanthorrhoeaceae; Liliaceae s.l.).

Ecological life history of the facultative woodland biennial Arabis laevigata variety laevigata (Brassicaceae): seed dispersal1

Causes of Within-Species Variations in Seed Dormancy and Germination Characteristics

The necessity for testing germination of fresh seeds in studies on diaspore heteromorphism as a life-history strategy