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Vegetative Propagules Glime, J. M. 2017. Adaptive Strategies: Vegetative Propagules. Chapt. 4-10. In: Glime, J. M. Bryophyte Ecology. Volume 1. 4-10-1 Physiological Ecology. Ebook sponsored by Michigan Technological University and the International Association of Bryologists. Last updated 24 April 2021 and available at <http://digitalcommons.mtu.edu/bryophyte-ecology/>. CHAPTER 4-10 ADAPTIVE STRATEGIES: VEGETATIVE PROPAGULES TABLE OF CONTENTS Vegetative Reproduction ................................................................................................................................... 4-10-2 Adaptations ....................................................................................................................................................... 4-10-8 Fragmentation ................................................................................................................................................... 4-10-8 Leaves and Stems ..................................................................................................................................... 4-10-10 Regenerants .............................................................................................................................................. 4-10-14 Protonemata ............................................................................................................................................. 4-10-14 Perianths ................................................................................................................................................... 4-10-14 Calyptrae .................................................................................................................................................. 4-10-15 Involucres ................................................................................................................................................. 4-10-16 Paraphyses................................................................................................................................................ 4-10-16 Falling Epiphytes ..................................................................................................................................... 4-10-18 Moss Balls ................................................................................................................................................ 4-10-18 Animals – Breaking or Ingesting Bryophytes .......................................................................................... 4-10-19 Extreme Environments ............................................................................................................................. 4-10-19 Antarctic & Arctic ............................................................................................................................ 4-10-19 Alpine ............................................................................................................................................... 4-10-20 Vegetative Diaspores ...................................................................................................................................... 4-10-20 Stem and Leaf Gemmae ........................................................................................................................... 4-10-25 Rhizoidal Gemmae ................................................................................................................................... 4-10-27 Rhizoidal Tubers ...................................................................................................................................... 4-10-29 Protonemal Gemmae ................................................................................................................................ 4-10-30 Liberation Mechanisms ................................................................................................................................... 4-10-32 Protonemal Gemmae ................................................................................................................................ 4-10-32 Tmema ..................................................................................................................................................... 4-10-33 Diaspore Bank ................................................................................................................................................. 4-10-34 Propagule Dispersal Distances ........................................................................................................................ 4-10-35 Propagule Survival/Longevity ......................................................................................................................... 4-10-36 Propagule Establishment ................................................................................................................................. 4-10-42 Desiccation ............................................................................................................................................... 4-10-43 Inhibitors .................................................................................................................................................. 4-10-43 Establishment and Rarity ......................................................................................................................... 4-10-44 Reproductive Problems ............................................................................................................................ 4-10-45 Epiphytes ................................................................................................................................................. 4-10-45 Symbionts Needed ................................................................................................................................... 4-10-46 Tradeoffs ......................................................................................................................................................... 4-10-47 Summary ......................................................................................................................................................... 4-10-48 Acknowledgments ........................................................................................................................................... 4-10-48 Literature Cited ............................................................................................................................................... 4-10-48 4-10-2 Chapter 4-10: Adaptive Strategies: Vegetative Propagules CHAPTER 4-10 ADAPTIVE STRATEGIES: VEGETATIVE PROPAGULES Figure 1. Deciduous branch tips of Campylopus cryptopodioides. Photo by Michael Lüth, with permission. Vegetative Reproduction Vegetative reproduction comes in many forms. These We might assume that in disturbed habitats, such as range from breakage of leaves and stems that have become industrial areas, vegetative means to reproduce might offer brittle through desiccation to specialized gemmae that advantages over more fragile-seeming protonemata occur on leaves or special stems and are nurtured by the necessitated by spore dispersal. But this is not always true. parent plant, ready to go when finding a suitable substrate, Nordhorn-Richter (1982) found the extent of bryophyte to tubers that occur on roots and protonemata. distribution in an industrial area of Germany was not Such asexual structures seem to have evolved through improved by presence of asexual propagules except among heterochrony (developmental change in the timing of members of Plagiothecium (Figure 3-Figure 4). On the events, leading to changes in size and shape), including other hand, vegetative reproduction offers the advantage of neoteny (retention of juvenile characteristics in adults of being ready to start developing an adult plant, reaching species) Newton & Mishler 1994). Hence, we can see maturity much more quickly than a plant from a spore. gemmae that look like reduced leaves, as in Aulacomnium Furthermore, bryophytes produce asexual propagules at a (Figure 2). younger age than that needed for sexual reproduction. Chapter 4-10: Adaptive Strategies: Vegetative Propagules 4-10-3 2. decay of older gametophyte parts with the separation of younger parts 3. development of multiple shoots by rhizomes and stolons 4. development of gametophores from rhizoids 5. regeneration from fragments 6. production of specialized "propagula." Frey and Kürschner (2011) shortened and summarized this list, identifying three types of asexual reproduction that are recognized today: 1. asexual reproduction s. str. by regeneration from somewhat specialized caducous organs (leaves, leaf apices, shoots, branches, bulbils) and by production of specialized propagules (gemmae, protonemal brood cells, tubers) 2. fragmentation of plants or parts of plants into essentially unspecialized fragments 3. clonal reproduction (cloning from branches, stolons, Figure 2. Aulacomnium palustre gemmae that are modified and rhizomes). leaves. Photo by Zen Iwatsuki, with permission. Cloning results when a branch of a protonema or a plant becomes separated from its parent. This can occur by decay of the protonema or disintegration of plant modules. In gametophores, cloning requires the formation of ramets (individuals, "daughter plants"). Frey and Kürschner (2011) consider cloning to be a "keystone factor for asexual reproduction, habitat colonization, and habitat maintenance." While this cloning mechanism is not dispersal in the usual sense, it
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