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Campylopus, a Modern and Successful Genus !?

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Campylopus, a Modern and Successful Genus !? View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by Hochschulschriftenserver - Universität Frankfurt am Main 91 Tropical Bryology 2: 91- 101, 1990 Campylopus, a modern and successful genus !? Jan-Peter Frahm Universität Duisburg, Fachbereich 6, Botanik, Postfach 101503, 4100 Duisburg, W-Germany Abstract. Campylopus with nearly 200 revised species is one of the largest genera of mosses in the world and has extremely broad geographical, altitudinal and ecological ranges. Factors to be considered for the rich speciation are 1) the enormous anatomical plasticity of the structure of the costa, 2) the ability for vegetative propagation utilizing different methods, and 3) the special twist mechanism of the cygneous setae. Phytogeographical interpretations of present ranges of species of Campylopus lead to the conclusion that this genus is of Gondwanalandic origin. Most of the species seem to be cool temperate in origin. They have adapted to dry habitats in the Mesozoic and invaded the tropical mountains during the Tertiary, accompanied by rich speciation. This is supported by the physiology of these species. According to preliminary gas exchange measurements, tropical montane species do not differ from temperate species, and are not able to stand climatic conditions of the tropical lowland rainforests; a relatively young habitat for bryophytes. A cladistic analysis of the infrageneric categories of Campylopus using phytogeographical evaluations of apomorphic character states shows that the section Homalocarpus seems to be the most primitive and the subgenera Campylopidulum and Thysanomitrion the most derived. 1. Introduction given in textbooks, but probably no more The somewhat provocative title goes back than 10.000 (4.000 species of hepatics to Schuster (1984), who (as a and 6.000 species of mosses). There are hepaticologist) interpreted some only a few genera of mosses with numbers distribution maps of Campylopus and of species comparable to Campylopus, called this genus ‘modern and successful’. but these have not yet been revised critically So it would be interesting to prove whether on a worldwide basis, and therefore a moss genus such as Campylopus is Campylopus may be one of the largest really modern and successful. genera of mosses in the world. Based on In total, more than 1000 species of an estimated total of about 6000 species of Campylopus have been described. Twenty mosses, i of every 30. mosses (or 3.3% of years ago, Index Muscorum (Wijk et al. all mosses) belong to the genus 1959) listed more than 700 legitimate Campylopus. species. Since that time, the number has The genus Campylopus has a wide been reduced by worldwide revisions to latitudinal range between 650N (Alaska) less than 200. This is a common effect in and and 650S (South Sandwich Islands) bryophytes, especially in tropical ones, and an enormous altitudinal range between where rates of reduction owing to sealevel and more than 4500 m elevation. worldwide revisions, are about 60-70%. The species occur on a variety of different This indicates that there are not 25.000 substrates such as nutrient poor sand, humic species of bryophytes in the world as soil, peat, rocks, rotten and living wood, 93 2.1 Gametophyte seems to function only in wet habitats, 2.1.1 Costa since the cell walls of large hyalocysts are very delicate and can probably not stand The most conspicuous vegetative character desiccation. In species pairs the species of of the genus Campylopus is the broad wet habitats have larger hyalocysts than costa with extreme diversity of internal species in drier habitats, as exemplified by differentiation. There is no comparable Campylopus shawii Wils. growing in range of variation in the structure of the swamps and C. setifolius Wils. growing costa in any other moss genus. All the on wet cliffs (Frahm 1985). It seems, different expressions between types with however, to be paradox that species of wet two stereid bands and a median deuter habitats should depend on storing water, band (as commonly found in the Dicrana- where there is no dry daily or annual ceae and also other families) and costae season. Thus it may be that the function of with no stereids at all can be explained by the hyalocysts (as well in the Leucobrya- a few cell divisions (fig. 1). Thus the ceae) have been misinterpretated. As Ro- mechanism behind this broad variety of binson (1985) has observed, the hyalocysts structures is relatively simple. in the Leucobryaceae often show air What are the advantages of these different bubbles, and the major function may not costal structures? be water storage but gas exchange. This a. mechanical stability. Stereids with firm hypothesis is supported by the location of thickened walls, provide mechanical the chlorocysts, which are situated in the support either in layers on the adaxial middle of the leaf without any direct contact surface or in bundles in the abaxial part of to the atmosphere. the costa. They are supposed to be good Another method of water storage is protection against shrinking caused by between the dorsal lamellae of the costa. desiccation and are therefore are found In species pairs, the species of drier habitats frequently in dry adapted species. In have the longer lamellae, for example species pairs (often sympatric species Campylopus pilifer Brid. and C. introflexus differing mainly in one character of the (Hedw.) Brid. (Frahm 1985). costa and the habitat), the species of the d. Water conducting is probably possible wetter habitat has ventral hyalocysts, not only from cell to cell or through the whereas that of the drier habitat has ventral pectine lamellae of the walls but also more stereids. One of many examples is efficiently through bundles of stereids. In Campylopus comosus (Reinw. & some species the alar cells (functioning for Hornsch.) Bosch & Lac. growing on soil, water uptake from the water conducted with small ventral hyalocysts and C. along the stem) bulge into the costa, flagelliferus (C. Müll.) Jaeg. growing in indicating a conducting pathway from the the same range but epiphytic with ventral alar cells to the costa. stereids (Frahm 1985, Frahm et al. 1985) e. Gas exchange combined with photo- b. Photosynthesis. Assimilation is carried synthesis. Effective gas exchange requires out by a median band (the so called deuter a large surface. Lamellae (being cells) and a dorsal band of chlorocysts. chlorocysts) seem to function not only for Since the width of the costa is usually water storage but also for gas exchange as more than half of the leaf width, and the shown by Campylopus pilifer Brid. var. basal laminal cells are often lamellatus (Mont.) Gradstein & Sipman, nonchlorophyllose, the costa is the main which is not found in arid environments source for photosynthesis. like C. pilifer s.str. but in rainforests. In c. Water uptake and storage. Water is this kind of habitat, water storage is not easily taken up and stored by hyalocysts, required, but gas exchange in high especially the large ventral ones. This humidity may be a problem. 94 Thus the plasticity of different structures Campylopodiella (but not in other closely of the costa allows adaptations to a broad related genera such as Atractylocarpus or variety of habitats. The facility to modify Pilopogon). such structures has promoted the broad The sigmoid curve in the seta is caused by ecological and geographical ranges and its development (Frey & Frahm 1987 figs. enhanced the high number of species. 1-7). When the seta has reached half of its final length, it curves down, the capsule 2.1.2 Vegetative propagation developes, often leaves the calyptra between the perichaetial leaves and curves Compared with other genera of mosses, up again. vegetative propagation is relatively In this way the young sporophyte important in Campylopus. In this genus, developes protected in the perichaetial all different kinds of vegetative propagation leaves which is an advantage, especially from microphyllous branches and in exposed or dry habitats. deciduous leaves to special brood leaves, The twist mechanism of the seta causes broken leaves and stem tips are found. If wide movements of the capsule in a 40% of all species are known only sterile, moistened state at the same time as when the importance of vegetative propagation the peristome is open. The function of is evident. The vegetative propagation these circular movements of the capsule is allows a species to survive in a habitat as not clear. It may enhance spore dispersal a unisexual clone. It also allows clones to to all directions, but this is a relatively survive long unfavourate climatic small advantage compared with the conditions with no sexual reproduction. anatomical structures necessary for these At least it allows propagation over short movements. Spore dispersal in wet weather distances until clones of different sexes conditions also prohibits long distance meet and allow fertilization. Apparently dispersal. However, since Campylopus as most of the species (if not all) are able to many other genera of Dicranaceae are vegetative propagate by more than one dioicous, male and female spores need to kind of vegetative propagation. land near to one another in order to establish new male and female clones which are 2.1.3 Perichaetia situated close enough to allow fertilisation. The mechanism for these uncoiling Differentiated bud-like perichaetia function movements have not been known yet but for splash cup dispersal of spermatozoids could be explained by a study of the and also as protection for the young ultrastructure of the setae (Frey & Frahm sporophyte during the stage of 1987). The epidermal cell layer of the development when the seta is curved into setae possesses a three layered wall with the perichaetial leaves. Such perichaetia asymmetric lumens. SEM and TEM are found in all infrageneric categories of studies revealed a distinct lamination in Campylopus.
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