J Ha/lori Bot. Lab. No. 84: 29- 35 (July 1998)

ON THE AND PHYLOGENY OF FLEISCHEROBRYUM (MUSCI, )

1 1 TIMO KOPO EN AND V11vr VIRTA EN

ABSTRACT. According to " Index Muscorum" the Fleischerobryum Loeske has four species, F longicolle (Hampe) Loeske, F macrophyllum Broth., F eurybrochis (Ren. & Card.) Fleisch., and F wallisii (C. Miill.) Loeske. The two latter belong to the related genus Philonotis Brid. and have been synonymized. The capsule of Fleischerobryum shows several primitive characters: The position of the capsule is horizontal or slightly pendulous, the capsule is long cylindrical or slightly asymmetric, the neck is large, ea one third of the length of the capsule, and the stri ae of the exothecium of Fleischerobryum are less regularly organized than are the striae in Philonotis species with ovoid or gibbous and hori­ zontal capsules. Evolutionary lines fro m the primitive capsule of Fleischerobryum both to gibbous and regul arly striate capsules, and to upright smooth capsules with reduced peristomes are possible. Fleischerobryum macrophyllum and F longicolle have wide mammillose basal leaf cells and narrow­ er more distinctly mammillose/papillose cells in the apical leaf like some Philonotis species. In the basal cells the mammilla is proximal, and in the di stal cell s the papilla/mammilla is distal. In the mid­ leaf cells of Fleischerobryum, cells both with central papillae and with di stal papillae are present. In Philonotis, with a few exceptions, the site of the papillae/mamillae is fi xed either to the distal end or proximal end of the cell, on both cell ends, or papillae are central on the cell. A hypothesis is here presented that the ancestor of Philonotis, and possibly of all Bartramiaceae, was a having a cap­ sule similar to that of the Fleischerobryum, and wide and smooth leaf cells.

INTRODUCTION Loeske (1910) described the genus Fleischerobryum Loeske and based its separation from Philonotis Brid. on its capsule shape. Fleischerobryum has long cylindric or slightly asymmetric horizontal or pendulous capsules with a long neck, while the capsules of Philonotis are ovoid or gibbous and have a short neck. He included two species, Fleischer­ obryum longicolle (Hampe) Loeske and F wallisii (C. Miill.) Loeske. Fleischer (1923) added F eurybrochis (Ren. & Card.) Fleisch., and Brotherus ( 1926) described F macro­ phyllum Broth. The generic status of Fleischerobryum has been accepted by later revisors and monographers of the family Bartramiaceae (Kabiersch 1937, Ochi 1962, 1963, Griffin & Buck 1989, Koponen & Norris 1996), and by the major floras (Bartram 1939, Gangulee 1974, Chen et al. 1963, Noguchi 1989). F eurybrochis was synonymized with Philonotis vescoana (Besch.) Par. by Koponen & Norris ( 1996), and Fleischerobryum wallisii with Philonotis hastata (Duby) Wij k & Marg. by Bartram ( 1939), so that Fleischerobryum now has only two species, both of them Asiatic (Fig. 1). Koponen (1996a) discussed the chraracters useful in the taxonomy of Philonotis. He emphasized the significance of the distribution of mammillae/papillae on leaf cells in dif-

1 Department of Ecology and Systematics, Division of Systematic Biology, P. 0 . Box 7, FIN- 00014 University of Helsi nki , Fin land. 30 J. Hattori Bot. Lab. No. 84 I 9 9 8

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Fig. l. Distribution of Fleischerobryum longicolle (Hampe) Loeske. ferent parts of the leaves and the morphology of the exothecial cells of the capsule. In our studies of Bartramiaceae (Koponen l 996b, Koponen 1998, Koponen & Norris 1996, Virta­ nen 1996, 1997, Virtanen & Koponen 1998) we have paid much attention to these characters.

CAPSULE SHAPE AND STRUCTURE OF EXOTHECIUM The short, nearly spherical capsule with a short neck in most of the genera and species of the fami ly Bartramiaceae is the characteristic which holds the genera together. In the genus Philonotis the upright capsules tend to be ::+:: globose, and the horizontal capsules ovoid, ellipsoid, or gibbous. The horizontal capsules we have studied until now are striate. This means that in the exothecium two kinds of cells occur, thick-walled and thin-walled. The thin-walled cells are in longitudinal rows between broader areas of thick-walled cells. When the mature capsule dries up, the thin-walled cells shrink, and thick-walled cell areas remain intact. The taxonomists see these sunken areas of shrunken cells as furrows or stri­ ae. The function of these furrows is obvious: they push the spores out of the capsule. Great variability exists in the horizontal and gibbous capsules as to the size and position of their thin-walled cell areas. Rows of thin-walled cells may be several or only one cell wide; the cells may be elongate or isodiametric. These rows of thin-walled cells may be evenly dis­ tributed all over the exothecium, or be present only on the dorsal part of the capsule; the ventral part of the capsule then remains smooth or is wrinkled (Fig. 2). The striations and wrinkling of the upright capsules of the species in Philonotis sect. T. K oPONEN & V V IRTANEN: Taxonomy and phylogeny of Fleischerobryum 31

Bartramidula (Bruch & Schimp. in B.S.G.) Mitt. are more variable. Some species have fur­ rowed capsules, or furrows occurring only on the upper half of the capsule. In some of the species with upright capsules no striations appear at all. Then the capsules are wrinkled or remain smooth when the capsule dries up. In the exothecium of the wrinkled capsules the thin-walled cell areas are not in rows but irregularly situated. In the exothecium of the cap­ sules remaining smooth all the exothecial cells are similar; there are no size differences, nor differences in the cell wall thickness. We have found these characters useful in the tax­ onomy of Philonotis species. The capsule of Fleischerobryum differs in several characters from that of Philonotis. The position of the capsule is horizontal or slightly pendulous, the capsule is long cylindri­ cal or slightly asymmetric, and the neck is large, ea one third of the length of the capsule. The striae of the exothecium of Fleisherobryum are less regularly organized than the striae in Philonotis species, with ovoid or gibbous and horizontal capsules. The capsule of Fleis­ cherobryum much resembles the capsules of such families as Bryaceae and Mniaceae, s. lat. We consider these characters plesiomorphic and hypothesize that the direction of evolu­ tion was from the primitive capsule of Fleischerobryum to gibbous and regularly striate capsules, and to upright smooth capsules with reduced peristomes.

L EAF C ELL PAPILLOSITY The papillosity/mamillosity of the leaf cells in Philonotis has been used both as a spe­ cific and as a sectional character (Koponen l 996a), and in the floras it is one of the key characters (e.g. Smith 1978, Nyholm 1960). Both the size of mammillae/papillae and their sites on the cell show variation. In the species which have two papillae/mammillae on one cell these two papillae are on the opposite ends of the cell; ifthe distal papilla is on the dor­ sal leaf surface, the proximal one is on the ventral surface, and vice versa. One of these two papillae is usually higher or more pronounced, which gives the leaf areolation its character­ istic outlook. If there is only one papilla/mammilla on Philonotis leaf cell, it may be on the distal or on the proximal cell end, or be central on the cell. Species of Philonotis are report­ ed with no leaf cell papillae at all. Such must be classified by their other characters. The shape and size of the mammillae/papillae vary in relation to the width of the leaf cells. If the basal leaf cells are wide, they are only mammillose, but the narrow distal cells of the same leaf have papillae. In plants with narrow basal leaf cells, the cells are papillose down to the base. These differences in mammillosity/papillosity give a characteristic out­ look to the leaves under the compound microscope, because the mammillose part is more transparent than the narrow-celled papillose part (Koponen l 996a). The size and shape of the papillae differ, the papillae which under the compound microscope look simple, may actually be rather complicated structures (Koponen l 996a). In Fleischerobrym macrophyllum and F longicolle the combination of leaf cell areola­ tion and distribution ofmammillae/papillae differ from that of Philonotis: The area of wide thin-walled cells fills nearly all the basal halfof the leaf, basal cells are low mammillose in the proximal end of the cell; the leaf cells in the narrow apex are narrow and with firm walls, and are low mammillose/papillose at the distal end. In the mid-leaf there are several cells which have central papillae (Fig. 3). However, the size of and height of 32 J. Hattori Bot. Lab. No. 84 I 9 9 8

a

3mm 2mm

d e f I 3mm lmm

Fig. 2. Capsules shapes and striation types of some Bartramiaceae. - a. Fleischero­ bryum longicolle (Hampe) Loeske (from Wichura 2905b, H-BR). b. Philonotis falcata (Hook.) Mitt., section Philonotula (from Gamble 25423, H-BR). c. Philonotis fontana (Hedw.) Brid., section Philonotis (from Koponen 21215, H). d. Philonotis sikkimensis (Kab.) T. Kop., section Bartamidula (from holotype, Wichura, H-BR). e. Philonotis dispersa Card. & P. de la V., section Bartramidula (from Perrottet, as P macrocarpa, H-BR). f. Philonotis calomicra Broth., section Bartramidula (from Williams, H-BR). - Use the 2 mm scale for a, the 3 mm scale for b, c and d, and the 1 mm scale for e and f. mammilla/papilla in Fleischerobryum are especially variable, and we have seen specimens of with leaf cells practically smooth. F longicolle is often misidentified in herbaria as Philonotis fontana (Hedw.) Brid. which has wide, thin-walled basal leaf cells. Basal cells with proximal mammilla, mid-leaf cells with central papilla, and apical cells with distal papilla are unique character of Fleischerobryum, and separate it from any species of Philonotis. In the central leaf of Philonotis seriata Mitt. mid-leaf cells may have a central T. K OPONEN & V ViRTANEN: Taxonomy and phylogeny of Fleischerobryum 33

Fleischerobryum ~hilonotis

sect. Philonotis PhiloLtula\~amidula VJgqqqQQnov. ~Q 800@ uu QQO)Ql)QQ Fig. 3. The site of papillae/mamillae in leaf cells and capsules shapes of Fleischero­ bryum Loeske and of sections of Philonotis Brid. (The row just below capsules - apical leaf cells, the row in the middle - median leaf cells, the undermost row - basal leaf cells.) papilla, but the distal leaf cells in that species have a proximal papilla.

PHYLOGENY We hypothesize here on the phylogeny of Fleischerobryum and Philonotis. We base these preliminary thoughts on the morphology of the capsules and leaf cells and add the re­ duction of the peristome, which is an apomorphic character of section Bartramidula (Fig. 2). Both these character groups are in direct relation to survival and therefore under evolu­ tionary stress. The characters of the capsule are significant in spore dispersal, and leaf cell characters, such as basal leaf cell size and the papillosity, with water uptake and conduction. We believe that the ancestor of Philonotis, and possibly of all Bartramiaceae, had a capsule similar to that of the present Fleischerobryum. Similar capsules occur in the pre­ sent families Bryaceae and Mniaceae s. !at. An evolutionary line from primitive capsule of Fleischerobryum to gibbous and regularly striate capsules with short neck and further to upright smooth capsules with reduced peristomes is probable. It may be emphasized that the reduction of the peristome and the change of capsule position in Bartramidula (which may have taken place several times, making Bartramidula polyphyletic) may be simultane- 34 J. Hattori Bot. Lab. No. 84 I 9 9 8 ous with the diminishing size and change of habitats, and, accordingly, with the life strate­ gies of these plants. The species of Philonotis section Bartramidula are smaller than many other Philonotis species which occupy stable habitats, such as springs and moist rocks. The labels of herbarium specimens record Bartramidula from more ephemeral sites, such as eroded soil or road side banks. We assume that the leaf cells of the ancestor of Fleischerobryum and Philonotis were wide and smooth. Again, many species of the present families Bryaceae and Mniaceae s. !at., and also of the Funariaceae, have such cell characters. In them the leaf cells in all parts of the leaf are more or less of the same shape and size. The large and nearly smooth basal leaf cells in Fleischerobryum and some species of Philonotis resemble this ancestral type. However, in most of the Philonotis the apical leaf cells differ by being narrow and with firm walls, and many species have all their leaf cells narrow. In these advanced species of Philonotis, the papillosity/mammillosity of the leaf cells is fixed, as was discussed above. In Fleischerobryum the papillosity/mammillosity is not fixed, a character which we consid­ er plesiomorphic.

CONCLUSION The ancestor of Fleischerobryum and Philonotis possibly had a capsule similar to pre­ sent Fleischerobryum, and wide and smooth leaf cells without papillae/mammillae. Such plants are at present found in the families Bryaceae and Mniaceae s.lat., and in Funari­ aceae. However, since the present study is based on the Philonotis flora only of Europe and Asia, with the rest of the Bartramiaceae not included, we are not yet ready to present a de­ tailed cladogram.

SYNOPSIS OF FLE/SCHEROBRYUM Fleischerobryum longicolle (Hampe) Loeske Morph. Syst. Stud. Laubm. 127. 1910 .-Bartramia longicollis Hampe in C. Mill!., Syn. Muse. Frond. I: 478. 1848. - Philonotis longicollis (Hampe) Mitt., J. Linn. Soc. Bot. Suppl. 1: 64. 1859.­ Type : Indonesia. Java, Junghuhn, Hb. Gottscheanum (not seen). Fleischerobrym macrophyllum Broth. Philippine J. Sci. 31 : 285 . 1926. - Type: Philippines. Luzon, Bontoc Subprovince, Mount Polis, Ramos and Edano 38289 (H-BR). Philonotis turneriana (Schwaegr.) Mitt. var. robusta E. B. Bartr., Ann. Bryol. 8: 13. 1936, syn. nov. - Type. China. Kweichow Prov., Ma Isooho, alt. 800m, S. Y Chea 499 (FH).

EXCLUDED TAXA Fleischerobryum wallisii (C. Miill.) Loeske Stud. Morph. Syst. Stud. Laubm. 127. 1910.-Bartramia wallisii C. Mill!., Linnaea 38: 554. 1874 ("Wallisi").-Philonotis wallisii (C . Miill.) Jaeg., Ber. St. Gallischen Naturwiss. Ges. 1877- 78 : 437 (Gen. Sp. Muse. 1: 70 l ). 1880. - Type: Philippines. lnsulae Philippinae, Luzon, Mahahai, regione montosa, 1871 G. Wallis , "Hb. C. Milll ." (H-BR, lectotype, sel. by Koponen & Norris 1996); BM-Bescherelle, isolectotype, SM-Hampe, isolectotype ). - Synonymized with Philonotis hastata by Bartram ( 1939). T. KoroNEN & V YIRTANEN: Taxonomy and phylogeny of F/eischerobryum 35

Fleischerobryum eurybrochis (Ren. & Card.) Fleisch. Musci FI. Buitenzorg 4: 1663 . 1923. - Philonotis eurybrochis Ren. & Card., Rev. Bryol. 23 : 10 I. 1896. - Type: Indonesia. Java, Foret de Tji bodas, sur Jes pierres de la cascade de Tjibeurreum, 1894- 1895 M. 1 Massart 1234 (BM -Bescherelle, syntype).- Synonymized with Philonotio' vescoana by Koponen & Norris ( 1996).

A CKNOWLEDGEMENTS The Academy of Fin land Fellowships nos. 153706 and 10134229 to the author Kopo­ nen for the project "Biodiversity of bryoflora in tropical southeast Asia" are cordially ac­ knowledged.

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