Volume 1, Chapter 4-6: Adaptive Strategies: Life Cycles
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Glime, J. M. 2017. Adaptive Strategies: Life Cycles. Chapt. 4-6. In: Glime, J. M. Bryophyte Ecology. Volume 1. Physiological 4-6-1 Ecology Ebook sponsored by Michigan Technological University and the International Association of Bryologists. Last updated 5 June 2020 and available at <http://digitalcommons.mtu.edu/bryophyte-ecology/>. CHAPTER 4-6 ADAPTIVE STRATEGIES: LIFE CYCLES TABLE OF CONTENTS Life Strategies .............................................................................................................................................. 4-6-2 Clonal Growth .............................................................................................................................................. 4-6-3 Foraging and Sharing ............................................................................................................................. 4-6-4 Implications for Reproduction ................................................................................................................ 4-6-5 Density Effects ...................................................................................................................................... 4-6-5 Tradeoffs ............................................................................................................................................... 4-6-6 r & K Strategies ............................................................................................................................................ 4-6-7 Bet Hedgers ........................................................................................................................................... 4-6-7 Dedifferentiation Issues ......................................................................................................................... 4-6-8 The r Strategist ...................................................................................................................................... 4-6-8 The K Strategist ..................................................................................................................................... 4-6-8 Life Cycle Strategies .................................................................................................................................... 4-6-9 Diaspore Banks.................................................................................................................................... 4-6-10 Tradeoffs ............................................................................................................................................. 4-6-10 Life Cycle Strategies based on During (1979, 1992) ............................................................................. 4-6-11 Fugitives .................................................................................................................................................... 4-6-11 Fugitives – species that live in unpredictable environments................................................................... 4-6-11 Colonists .................................................................................................................................................... 4-6-11 Colonists (sensu stricto) – species that live where habitat start is unpredictable ..................................... 4-6-11 Colonists .................................................................................................................................................... 4-6-11 Colonists (ephemerals) – gap-dependment species ................................................................................ 4-6-11 Colonists (pioneers) – species that live where habitat start is unpredictable ........................................... 4-6-11 Shuttles ...................................................................................................................................................... 4-6-11 Annual Shuttle – species that require small disturbances that last 1-2 years ........................................... 4-6-11 Short-lived Shuttle – species that don't avoid periods of severe stress .................................................... 4-6-12 Perennial (long-lived) Shuttle – species that require stable environments .............................................. 4-6-12 Perennial Stayers ........................................................................................................................................ 4-6-12 Perennial stayers (competitive) – forest floor ........................................................................................ 4-6-12 Perennial stayers (stress-tolerant) – fens, bogs, desert ........................................................................... 4-6-12 Dominants – bogs ....................................................................................................................................... 4-6-12 Generation Time ......................................................................................................................................... 4-6-13 Habitat Studies ........................................................................................................................................... 4-6-14 Summary .................................................................................................................................................... 4-6-15 Acknowledgments ...................................................................................................................................... 4-6-16 Literature Cited .......................................................................................................................................... 4-6-16 4-6-2 Chapter 4-6: Adaptive Strategies: Life Cycles CHAPTER 4-6 ADAPTIVE STRATEGIES: LIFE CYCLES Figure 1. Strap-shaped leaves of males and capsules with perichaetial leaves of females of the dioicous moss Diphyscium foliosum. Photo by Michael Lüth, with permission. Life Strategies By now it must be obvious to you that to understand growth, which implies the increase in size of a the life strategies of bryophytes, one must first understand physiological individual (Söderström 1994). On the other the life cycle (e.g. Figure 1). Frahm and Klaus (2001) state hand, growth can ultimately result in reproduction, as is the that bryophytes are able to react quickly to such events as case when the plant branches and is physiologically climatic fluctuations because of their short life cycle and connected, but later the branches separate and become ease of dispersal by spores. It is likely that when all other physiologically independent. As you can see in Figure 2, plant life has perished from some Earth catastrophe, it will the degree of reproduction by propagules depends on be the bryophytes that persist, surviving as spores or other habitat. Among British habitats, short-lived habitats propagules until conditions return to safety and once again (wood, bark, farmland, dung) have the highest degree of surrounding the earth due to their ability to travel great propagular reproduction (that is, by vegetative means) distances as "dust." (Herben 1994). On the other hand, the habitats with the By definition, the life cycle includes reproduction. greatest percent of the species are in the middle of the However, even such a widely used term has ambiguities. range of propagules. In bryophytes, we shall use this term to refer not only to the Using the principle that extreme conditions might sexual reproduction that results from union of sperm and provide the best test of the limits of an organism, we learn egg (ultimately resulting in spores produced by meiosis), that in the maritime Antarctic, bryophytes seem to have but also to the multitude of asexual (vegetative) means by enhanced sexual reproduction (Lewis Smith & Convey which bryophytes are able to make more, physiologically 2002). This is contrary to the generally accepted belief that independent plants (Mishler 1988). This definition bryophyte fertility decreases toward the poles. Rather, separates reproduction, which can permit relocation, from 43% of the bryophytes (19 species) in Marguerite Bay and Chapter 4-6: Adaptive Strategies: Life Cycles 4-6-3 47% of those on Alexander Island are known to produce of disadvantaged ramets that might later survive in the face capsules. But Lewis Smith and Convey attribute this to of adversity; increased precision with the sequestering of favorable microclimatic conditions. Nevertheless, in this space and dispersal of ramets; ability to monopolize extreme environment, the large majority of mosses with resources for the benefit of the genotype. They capsules were short, monoicous, acrocarpous taxa, furthermore include mobility, but I question whether this is suggesting that the predominance of dioicous taxa in more much of a gain when compared to the alternative of wider temperate climates may be possible because the distribution of propagules away from the parent. environment is less stressful. In the more extreme One possibility that has barely been explored is the environments of the Antarctic continent, the numbers of increase of genetic variability through production of these species producing capsules at similar latitudes (68-72°S) haploid genets. We had long assumed that the limited are much less (33%). morphological development of the Bryophyta and Marchantiophyta reflected a limited genetic diversity, a case to be expected when the dominant generation is haploid and asexual reproduction is common. However,