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• W1986968973 abstract "In view of the paucity of information on the phenology of even the commonest species of mosses, a system is proposed to facilitate the accumulation of phenological data. The following essential developmental stages of the moss life cycle based on morphology and ecology are recognized and described: a) embryonic calyptra, b) seta with calyptra, c) 1 Grateful acknowledgement is made to Dr. H. A. Crum for examining many problem specimens and to Dr. H. E. Robinson for examining certain Sphagna. 2 Department of Biology, University of Wisconsin Center, Wausau, Wisconsin 54401. This content downloaded from 207.46.13.114 on Thu, 26 May 2016 06:39:52 UTC All use subject to http://about.jstor.org/terms 290 THE BRYOLOGIST [Volume 68 capsule green with calyptra, d) capsule operculate and postmeiotic, e) capsule de-operculate, f) spore wall bulging, g) protonema, h) bud on protonema, i) juvenile stem, j) juvenile gametangium, k) antheridium, and 1) archegonium. A method for coding and presenting the phenological data is described. As an example of the method, developmental stages and substages are recorded among the 103 species and one variety of mosses the flora of Rib Mountain, Marathon County, Wisconsin. In addition, 75 of the taxa are new records for Marathon County. Students of vascular plants have for many years recognized the importance of including phenological data their studies and collections. Some bryologists have included in fruit, c. fr. or similar notations on their packets, but this is done usually as an aid identification of the specimen rather than as recognition that the moss actively undergoes various stages of growth and that one or more of these stages is present the collection. Even if these stages of growth are recognized, c. fr. is an inadequate designation to record them. The ultimate goal of including phenological data is to explain the factors which permit the development of a given stage a species or another taxon. This can be done by studying an individual colony of a species throughout its life cycle. However, infraspecific variation is usually such that the species grows different sites and habitats a region. Conclusions based on a study of one colony cannot be applied to other colonies and cannot be correlated with regional climatic data. If one colony is studied, microclimatic data should also be collected. Another method with wider applicability is to study certain or all species of a region, disregarding the infraspecific variability. In many areas regional environmental data are collected, and general correlations may be made between environmental factors and phenological stages. In Grimme's (1903) monumental work the phenological stage of every collection of the 207 species collected the vicinity of Uppsala was recorded for many years. He described with varying degrees of completeness (depending upon the number and time of collections of each species) the phenology of the entire moss flora of the region. More detailed studies of individual species are published by Greene (1960) for Pylasia polyantha, Hypnum cupressiforme v. resupinatum, Brachythecium rutabulum, and Mnium hornum; Towle and Gilbert (1904) for Polytrichum commune and P. juniperinum; Towle (1905) for Atrichum; Towle (1906) for Mnium sylvaticum and M. affine; and Arnell (1875, 1905) for several species. The study of individual species is optimal and provides complete data for the species, while Grimme's approach provides data of varying completeness on the entire regional flora. However, the latter approach can be used by practically all bryologists the normal course of other studies. A few seconds only are required to scan a This content downloaded from 207.46.13.114 on Thu, 26 May 2016 06:39:52 UTC All use subject to http://about.jstor.org/terms 1965] FORMAN: MOSS PHENOLOGY 291 specimen and record the phenological stages present. If done for just a few years by bryologists over a wide geographic area, the general phenology of the majority of our species will be known. The purpose of this paper is (1) to present a system for the recognition and recording of the phenological stages of moss development and (2) to give an example of the use of this system. The need for phenological data is evident to the ecologist attempting to explain the distribution of any moss. For example, spore germination studies are done the laboratory for physiological purposes, but if the time of actual spore dispersal and spore germination are unknown, the factors governing these phenomena the field cannot be understood. The temperature necessary for meiosis can be delineated and checked after it is known when meiosis takes place. A recent study of Tetraphis pellucida (Forman 1964) defines the growth conditions for the species at several developmental stages, but the lack of general phenological data from different regions limits the ecological applicability of the data. Likewise phenology as a taxonomic tool is used by vascular plant systematists and recently by some bryologists (e.g., Greene & Greene 1960, Jones 1947). STAGES OF THE LIFE CYCLE The stages of a life cycle are merely divisions of a continuous process of growth and development. At one extreme time-lapse photography would provide the most complete series of stages, but the differences between stages would become inconsequential. The alternation of a sporophyte generation with a gametophyte generation at the other extreme is insufficiently detailed. The present system of 12 stages given below is devised to correspond to the major morphological and ecological differences the life cycle. The morphological basis is fundamental and no two stages are delimited unless morphologically distinct. In addition, the relative ease of recognition of stages is emphasized, such that macroscopic characteristics are used lieu of microscopic ones whenever practical. Certain environmental conditions are necessary for each stage to take place. The ecological basis is whether the environmental factors necessary for one part of the life cycle are or are not likely to be the same as those for the following part of the life cycle. For example, elongation of the seta is not subdivided into early and late stages since the ecological conditions necessary for each are likely to be essentially the same. But growth of the embryo a perichaetium is separated from growth of the seta since the ecological tolerances of the two stages are likely to be different. The 12 stages recognized, lettered a to 1, are described below and diagrammed Figure 1. The lettering arbitrarily begins with the diploid generation following fertilization. a) Embryonic calyptra. (This corresponds with the development of the embryo following fertilization.) This stage commences with This content downloaded from 207.46.13.114 on Thu, 26 May 2016 06:39:52 UTC All use subject to http://about.jstor.org/terms 292 THE BRYOLOGIST [Volume 68" @default.
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• W1986968973 title "A System for Studying Moss Phenology" @default.
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