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Peristome Variations in the Genus Fissidens: an SEM Study Author(S): Bruce H Peristome Variations in the Genus Fissidens: An SEM Study Author(s): Bruce H. Allen Source: The Bryologist, Vol. 83, No. 3 (Autumn, 1980), pp. 314-327 Published by: American Bryological and Lichenological Society Stable URL: http://www.jstor.org/stable/3242441 Accessed: 09/08/2010 07:49 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/action/showPublisher?publisherCode=abls. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. American Bryological and Lichenological Society is collaborating with JSTOR to digitize, preserve and extend access to The Bryologist. http://www.jstor.org The Bryologist 83(3), 1980, pp. 314-327 Copyright @ 1980 by the American Bryological and Lichenological Society, Inc. Peristome Variations in the Genus Fissidens: An SEM Study BRUCEH. ALLEN Departmentof Biology, Buckhout Laboratory,The PennsylvaniaState University, University Park, PA 16802 Abstract. Peristomes of 19 species representing two subgenera of the genus Fissidens were examined using scanning electron microscopy. Seven types of peri- stomes were distinguished on the basis of variations in the trabeculae and the lamellae of the dorsal and ventral surfaces. These seven types are designated the scariosus-, bryoides-, excurrentinervis-, obtusifolius-, taxifolius-, subbasilaris- and manateensis-types of peristomes. In some instances, species having similar peristome types are gametophytically similar; in other instances species having the same peristome type have different gametophytes. In still other instances, species with dissimilar peristome types have similar gametophytes. The genus Fissidens is a homogeneous group of mosses distinguished by an unusual leaf structure. The leaves are equitant and each is composed of a ventral, boat-shaped proximal portion consisting of two vaginant laminae and an undivided distal ventral lamina; dorsally there is an undivided dorsal lamina. In addition, the leaves are distichous and complanate. In the Musci, the peristome is considered an important indicator of natural relationships (Philibert 1884-in Taylor 1962, Dixon 1932, Buck & Crum 1978). The peristome of Fis- sidens, usually described as dicranoid, consists of a single row of 16 peristome teeth that are haplolepideous (having a single row of dorsal "scales"). Thus, the genus Fissidens is a taxon apparently closely related to the Dicranaceae on the basis of the sporophyte but which possesses a distinctive gametophyte easily separating it from all other haplolepi- deous mosses. Conspectus of the genus Fissidens. The genus Fissidens was established by Hedwig (1801) to accommodate 14 species. Since that time the number of species ascribed to Fissidens has increased to approximately 876 (Wijk et al. 1962, 1969). Along with this increase in the number of species, a large number of subgeneric taxa has been described. Brotherus (1924) recognized 4 subgenera, viz., Polypodiopsis (C. M0ll.) Broth., Eufissi- dens Hedw. (=Fissidens), Pachyfissidens (C. M011.) Kindb. and Octodiceras (Brid.) Broth. Subgenus Fissidens was further divided into twelve sections, viz., Weberiopsis C. Mill., Reticularia Broth., Bryoidium C. Miill., Pachylomidium C. Mill., Pycnothallia C. Miill., Heterocaulon C. Miill., Semilimbidium C. MWll., Aloma C. Miill., Crenularia C. Mill., Crispidium C. Miill., Amblythallia C. Miill. and Serridium (C. Mill.). In the past, subgeneric taxa within Fissidens have been based principally on gameto- phytic characters. The peristome was never utilized in the interpretation of relationships within Fissidens. This is because differences seen in structure of the peristome with the light microscope appear minor. However, Florschiitz (1964) believed the current division of the genus was "highly artificial." Species he considered closely related were positioned into disparate sections. Furthermore, there has been disagreement among authors as to the number and rank of the subgeneric taxa within Fissidens, even though all treatments 007-2745/80/314-327$1.65/0 1980] ALLEN: PERISTOMEVARIATIONS 315 have utilized the same characters. Indeed, recent treatments of groups within the genus (Norkett 1969, Bruggeman-Nannenga 1978) have resulted in a reduction in the number of subgeneric taxa on the basis that variation of characters within the gametophyte may be environmentally induced. It would appear that a better understanding of relationships within Fissidens could be achieved if additional characters were utilized. The sporophyte should be considered a possible source of new characters in view of its conservative nature (Buck & Crum 1978) and the potential of recent techniques developed for observing spo- rophytic features, e.g., transmission electron microscopy, scanning electron microscopy (SEM), and chemical analysis (Mueller 1973). To date only three studies have been done on Fissidens using SEM: one by Berlin et al. (1974) who studied the leaf structure of nine species of Fissidens, and two by Mueller (1973, 1974) who observed the morphology of the peristome of F. limbatus Sull. and F. osmundoides Hedw. MATERIALS AND METHODS Peristomesof 22 specimensbelonging to 19 species in two subgeneraand nine sections of Fissidens (see Brotherus1924) were used. With the exception of three species, F. manateensis, F. scariosus and F. polypodioides,peristomes viewed were taken from single populations.An attemptwas made to view the peristomesof at least six plants per gathering.Except where otherwise stated, all spec- imens are deposited at PAC. Specimens examined: SUBGENUSFISSIDENS. Sect. Aloma C. Muill.: F. exilis Hedw., U.S.A., Pennsylvania, Centre Co., Allen 956; F. prionodes Mont., Mexico, Chiapas, Sharp et al. 4096; F. subulatus Mitt., Brazil, Rond6nia, Reese 13332. Sect. Amblyothallia C. Miill.: F. asplenioides Hedw., Mexico, Mexico, Delagadillo 2718; Sect. Bryoidium C. Miill.: F. bryoides Hedw. sensu lato, U.S.A., Pennsylvania,Fayette Co., Allen 270; Sect. CrenulariaC. Miill.: F. radicans Mont., U.S.A., Florida, Dade Co., Pursell 2915; Sect. Heterocaulon C. Miill.: F. excurrentinervis R. S. Williams, Peru, La Libertad, Hegewald 5112; F. milobakeri L. Koch, U.S.A., California, San Mateo Co., Koch 3394 (isotype);Sect. SemilimbidiumC. Mill.: F. obtusifoliusWils., U.S.A., Kansas, DouglasCo., Cridland 250; Sect. Serridium C. Miill.: F. adianthoides Hedw., U.S.A., Pennsylvania, Centre Co., Allen 364; F. bushii (Card. & Ther.) Card. & Ther., Canada, Quebec, Fabius 2207 (DUKE);F. cristatus Wils. ex Mitt., U.S.A., Tennessee, Greene Co., Sharp4528; F. osmundoidesHedw., U.S.A., Wisconsin, Douglas Co., Schroeder M183; F. polypodioides Hedw., U.S.A., Florida, Leon Co., Pursell 634, Mexico, Chiapas, Smith et al. 2873; F. subbasilaris Hedw., U.S.A., Tennessee, Greene Co., Sharp 4514; F. taxifolius Hedw., U.S.A., Pennsylvania, Centre Co., Allen 217; Sect. Weberiopsis C. Miill.: F. gladiolus Mitt., Tanzania, Morogoro, Crosby et al. 8749 (MO); F. scariosus Mitt., Brazil, Sio Paulo, Yano 211, Minas Geraes, Frahm 1543. SUBGENUSOCTODICERAS (BRID.) BROTH.F. man- ateensis Grout, U.S.A., Florida, WakullaCo., Redfearn2060, Louisiana, Lafayette Parish, Reese et al. 6410. In the selection of peristomes, an attemptwas made to use maturecapsules with intact opercula. Surface features of the teeth are easily eroded once the operculumis lost from a capsule. Mature sporophyteswere dissected and mountedwith double-stickadhesive tape onto aluminumspecimen mounts. In order to avoid a buildupof electrical charge on the specimens, a thin layer of gold was applied to the specimens using a HummerII sputterer.The specimens were viewed with a Super Mini-SEMat 25 kV. RESULTS Study reveals considerable variation in the surface features of the peristomes of Fis- sidens species. Among the representatives of 19 species studied, seven types of peristome can be distinguished on the basis of variation in the trabeculae and lamellae of the outer and inner surfaces of the teeth (see Table 1). These seven types are here designated as the scariosus-, bryoides-, excurrentinervis-, obtusifolius-, taxifolius-, subbasilaris- and man- ateensis-types of peristomes. Scariosus-type of peristome (Fig. 1-10). The scariosus-type of peristome is exhibited by F. scariosus, (Fig. 1-2), F. radicans (Fig. 3-4), F. prionodes (Fig. 5, 10), F. exilis (Fig. 6-7), and F. subulatus (Fig. 8-9). It is characterized by a dorsal surface on which the w TABLE1. A comparisonof the seven peristome-typesobserved in this study. Trabeculae Lamellae Outer surface Inner surface Outer surface Inner surface fimbriate smooth Smooth or papillose ON Scariosus-type Unthickened, Usually bearing High, closely
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