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On the Peristome Types Found in the Fissidentaceae and Their Importance for the Classification

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On the Peristome Types Found in the Fissidentaceae and Their Importance for the Classification Journ . Hal/ori BOI. Lab. No. 68: 193- 234 (June 1990) ON THE PERISTOME TYPES FOUND IN THE FISSIDENTACEAE AND THEIR IMPORTANCE FOR THE CLASSIFICATION M. A. BRUGGEMAN-NANNENGA 1 AND W. B ERENDSEN 2 ABSTRACT Based on a study of peristomes of about 200 species of all subgenera and sections of the Fissidentaceae, 5 basic types of peristome are recognized: the bryoides-type, the scariosus-type, the zippelianus-type, the simi!irelis-type and the laxifolius-type. The scariosus-type is correlated with 28--40, but mostly circa 32 exothecial cells on the capsule periphery. The number of exothecial cells found with the other types is typically more than 40. The bryoides-type of peristome is characteristic of F. sect. Fissidens, F. su bg. Sarawakia and the genus Nanobryum . It also appears that these taxa arc gametophytically more similar to each other than to any other taxon, except that F. subg. Sarawakia is intermediate between F. subg. Octodiceras and sect. Fissidens. The scariosus-type of peristome is characteristic of F. subg. Aneuron, and the F. subg. Fissidens sections Areofissidens, Aloma, PYCllo lhallia, Semilimbidium and Crenularia. Within this group Aneuron and Areofissidens are gametophyticall y very si milar, and so are Pycnothallia, Semilimbidium and Crenularia. The zippe/ianus-typc of peristome is characteristic of F. sect. Crispidium; the similiretis-type of peristome is characteristic of F. sect. Amblyolhallia and the taxifolius-type of peristome is characteristic of F. sect Serridium. Several other peristome forms were also found; these, however, occur in one or just a very few species and are not characteristic of any supraspecific taxon. CONTE TS Introduction . .. .. ... .. .. ....... ...... ... .... .. .. ...... • .... • ...... .... ..... .. 194 Material and Methods ......... .. .. .. .. .. .. .......... .. .. , . .. .. ....... ... .. ...... ... 195 Key to the Main Peristome Types .......... .. .. ... ...... ......... ... .. ........ 197 Description of the Peristome Types .... .. .... .. .. .. ...... .. .. ...... ... ..... ... 197 Species with Deviating Peristomes ................... ..... ... .. .... .... .. ... ...... .... 216 Distribution of the Peristome Types over the Fissidentaceae .. ...... .. .. ....... .. ... .... .... 221 List of the New Names, New Combinations, New Synonyms and New Records Published in Bruggeman-Nannenga and Berendsen (1988) ... .. .. ............ ... .... ........ .. .. 226 Acknowledgements ..... .... .. .................. ... • . ... ..... .. ... • ........ 227 References ................. ... ... .. .. ...... .• .. ... .... .... .. .... ...... ... ..... 227 Index to the Species ... ....... ..... .. ..... .... ... .. .. .. .. .. ... .. ... 229 I Institute of Systematic Botany, State University Utrecht, Heidelberglaan 2, 3508 TC Utrecht, the Netherlands. 2 Department of Molecular Cell Bi ology, State Un i v~rsity Utrecht, Padualaan 8, 3584 CM Utrecht, the Netherlands. 194 Journ. Hattori Bot. Lab. No. 68 1 990 INTRODUCTION Philibert (1884), in his study on the peristomes of the mosses, considered the Fissidentaceae to have a dicranum-type of peristome. He remarked that the peristomes of some Fissidens species differ from the typical dicranum-type in having spirally arranged papillae on the filaments. Several genera of Fissidentaceae have been described on the basis of the peristome teeth. For instance Octodiceras Brid. was segregated from Fissidens Hedw. by Bridel (1806) by having 8 instead of 16 peristome teeth. Later it was recognized that Octodiceras too has 16 peristome teeth (Montagne 1837). Muller (1901) described Moenkemey((ra which is distinct from Fissidens in its undivided teeth. The generic status of this taxon was questioned by Grout (1938), who found undivided teeth in gametophytically disparate species. Although Dixon (1941) and Potier de la Varde (1951) continued to describe species in this genus it is now generally considered a synonym of Fissidens. In 1909 two more genera with undivided teeth were described, Fissidentella Card. and Simplicidens Herz. Dury (unpublished thesis, 1974) considered Fissidentella perpusi/la Card., the only species in the genus, to be identical with Fissidens Jouta-djallonii. Pursell (1982) reduced Simplicidens to Fissidens. Sainsburia, characterized by undivided, rimose teeth, was described by Dixon (1941 A) . Its only species, S. novae-zealandiae Dix., is now reduced to a synonym of Fissidens taylorii C. Muell. (Dixon in Steere, 1947; Sainsbury, 1955). Pursell (1982) in an overview of the Fissidentaceae concluded that none of the genera characterized by undivided teeth are gametophytically different from Fissidens, and proposed to consider all of them as synonyms of Fissidens. Fleischer (1904) divided the Fissidentaceae into species with spirally thickened peristome filaments and species with annularly thickened filaments. Brotherus (1909, 1924) in his subdivision of the Fissidentaceae used this character along with several others to distinguish sections. M ueller (1973) studied the ultrastructure of the peristome of F. limbatus Sull. with the scanning and transmission electron microscope. He was the first to show that the spiral orpamentation on the peristome filaments consists of an inner and outer layer each with oblique riblets, which together give the impression of a spiral. Allen (1980) in a SEM study of the Fissidens peristome compared peristomes of 20 species from 9 different sections. He distinguished 7 peristome types, which are in part correlated with gametophytic types. With light microscopy, however, it appears that some of the characters used by Alien to diagnose his groups are not constant, though his groups are generally sound. The aim of the present work has been to describe the types of peristome found in the Fissidentaceae, to determine whether they support the existing classification and to determine their usefulness in elucidating the natural relationships among taxa. The Fissidentaceae have a haplolepidous peristome. This type of peristome, as discussed by Edwards (1984), consists of eight completely divided pairs of teeth. The teeth, in most species, are divided into two filaments or prongs. Each tooth is formed M. A. BRUGGEMAN-NANNENGA & W. BERENDSEN: Peristome types in Fissidentaceae 195 vertical wall lamellae I-----<=+.__ trabeculae median wall DIAGRAM I: outer surface (left) and inner surface (right) of a pair of peristome teeth. by two cohering layers of cells, an outer one or primary peristomiallayer (PPL), which in this paper is indicated as the OP and an inner one or inner peristomial layer (IPL), which herein is indicated as JP. Typically, the outer layer of each pair of teeth consists of two vertical rows of cells (diagram I and PI. 4c), while the inner layer consists of three rows (diagram I and PI. 4d). The outer layer of each tooth consists of one row of vertical cells, the inner one of one and a half row, hence the basic unit of the haplolepidous peristome is the teeth-pair rather than the individual tooth. The main body of the teeth (diagram I) is formed by the cohering median OP and JP cell walls, the lamellae. The cohering horizontal walls of adjacent cells together form a trabecula. Occasionally the vertical walls connecting the trabeculae and the lamellae are persistent. There are marginal vertical walls both on the OP and JP sides as well as approximately median ones. The latter occur only on the JP side and are often oblique rather than vertical. The two filaments are slightly unequal since one is composed of a vertical row of complete cells and the other is composed of a vertical row of cell halves (the splitting follows the median walls of the inner layer). Three different zones of ornamentation can be recognized in most teeth. In the lowermost one, the anchor zone, the teeth form a continuous ring, which may protrude beyond the theca or be completely inserted. This zone usually consists of smooth, complete cells, ie cells with all walls persistent. The zone above it is the basal, undivided part, usually with an elaborate, often taxonomically important, ornamentation. At the bifurcation the ornamentation usually changes rather abruptly into that which is characteristic of the lower part of the filaments. This ornamentation, too, is often taxonomically important. Species with undivided teeth often have a similar abrupt change in ornamentation at approximately the height where one would expect the bifurcation. In the filaments there usually is a gradual change in ornamentation distally. MATERIAL AND METHODS The peristomes of circa 200 species were studied with the LM (light microscope) and those of 2S species with the SEM (scanning electron microscope). With a few exceptions, only dried herbarium material was used. 196 Journ. Hattori Bot. Lab. No. 68 1 9 90 The herbaria are abbreviated according to the Index Herbariorum (Holmgren & Keuken 1974). The collectors herbarium is indicated as follows: PC-B = herbarium Bizot, PC-F = herbarium of France, PC-g = general herbarium, PC-PV = herbarium Potier de le Varde, PC-Th = herbarium Theriot, H-Br = herbarium Brotherus and H-SOL = herbarium S.O. Lindberg. A few collecti ons are from private herbaria: hb. Brugg.-Nann. (herbarium of the first author) and hb. Melick (herbarium of H. van Melick, Eindhoven). As a rule only one collection per species is cited in the li sts of investigated material. Author citations are given only on the first occasion when a species is mentioned. It is possible that some of the investigated species li sted in this paper have
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