Journ. Hattor; Bot. Lab. No. 64: 29-36 (June 1988)

SYMPOSIUM 5-14: OF HIGHER CATEGORIES AND GENERIC CONCEPTS IN BRYOPHYTES (Organizers: R. M. SCHUSTER and D. H. VITT)

ANOTHER VIEW OF FAMILIAL DELIMITATION IN THE

WILLIAM R. BUCK'

ABSTRACT: The Hookeriales are divided into five families, the , , Adelotheciaceae, Leucomiaceae and Callicostaceae. The families are delimited primarily on gameto­ phytic characters, including several little-used ones, because the order is considered fairly uniform in peristome features except for a repeatable reduction of cross-slriolate to papillose exostome teeth. The order and each family are described and generic inclusions listed. Discussion is presented for controversal genera. A dendrogram is provided 10 illustrate tentative relationships between the families.

The Hookeriales, with strong representation at all elevations in tropical latitudes, are often considered exotic by north temperate bryologists due to the family'S scanty penetration into Europe and North America. The are among the most attractive of pleurocarpous , but notoriously variable, perhaps a result of their frequent association with humid habitats. As in other truly pleurocarpous orders (sensu Buck & Vitt 1986), some individual genera are widely disparate in morphological ex­ pression from others. This diverse morphology has led to several, very different familial schemes for the Hookeriales. The traditional treatment of the order (Brotherus 1925, following Fleischer 1923) recognizes six families. Miller (1971) proposed nine families and Crosby (1974) only two. Buck and Vitt (1986) removed some of the families (Symphyodontaceae and ) to other orders but basically delayed decision on familial delimita­ tion until further study. As a result of recent work on the Hookeriales for floristic purposes, I have proposed an alternative familial arrangement (Buck 1987) that will be expanded upon here. Previous classifications of the order have primarily stressed either gametophytic or sporophytic characters, but little attempt has been made to correlate the two gene­ rations. Not only has such an effort been made here, but several previously overlooked or little-used gametophytic characters have been utilized to construct the present classifi­ cation. Included among these characters are axillary hairs, pseudoparaphyllia, stem an­ atomy and calyptra anatomy. Many traditional characters such as costa type and leaf areolation (including borders) have also proved useful. However, peristome mor-

1 New York Botanical Garden, Bronx, NY 10458-5126, U.S.A. 30 Journ. Hattori Bot. Lab. No. 64 1 9 8 8

phology has only been useful in circumscribing the order. Exostome ornamentation has generally not been of value. Rather, it is proposed that papillose exostome teeth are polyphyletic and have been derived from cross-striate ones repeatedly in the order. This will be discussed in more detail below.

Hookeriales (Fleisch.) Fleisch., Hedwigia 61: 397. 1920. Plants small to robust, often pale-green when growing in shade, becoming golden to red-brown in sun, in mats or tufts. Stems creeping or erect, rarely pendent or floating, simple to regularly bipinnate, often complanate-foliate; paraphyllia none; pseudoparaphyllia absent, foliose or filamentous; rhizoids mostly smooth or lightly roughened; in cross-section stems undifferentiated throughout or with small, thick-walled cells (sometimes the outermost layer a hyalodermis) surrounding larger, thinner-walled cells, central strand absent or occasionally present; axillary hairs 2-many-celled, the basal one usually brown, the others (or rarely all) hyaline. Lateral and dorsal leaves often differentiated, lanceolate to suborbicuJar, sometimes concave, rarely plicate; margins usually plane, occasionally narrowly recurved or erect, entire to coarsely serrate, the teeth sometimes inflated, bordered or not; costa single, strong and double, short and double, or rarely absent; cells linear to isodiametric, usually smooth, some­ times uni- or pluripapiIJose or prorulose; alar cells not differentiated; propagula common, borne on stems, leaves, rhizoids or rarely protonemata. Autoicous or dioicous. Perichaetia often small, inconspicuous; leaves ± erect, pale, few. Setae short to long, often ornamented, especially above, straight or curved at apex; capsules exserted, mostly symmetric; exothecial cells usually collenchymatous; annulus present or absent, if present often adhering to oper­ culum, of thin-walled cells; operculum conic-rostrate, if annulus none the basal cells often differentiated; peristome double, exostome teeth 16, entire, often bordered, the inner surface usually wider than outer, not shouldered, attached at mouth, on front surface either furrowed (sometimes not furrowed) and cross-striolate or not furrowed and papillose; endostome with a high to low basal membrane, segments keeled, perforate or not, with baffle-like crosswalls, cilia usuall y absent, occasionally single, rarely to 3. Spores spherical, papillose, usually finely so. Calyptrae typically mitrate, rarely cucullate, lobed or fringed at base, often hairy, smooth or roughened. The Hookeriales, like most orders, are difficult to define. Among the characters that hold together most of the genera are short leaf cells, complete lack of alar differenti­ ation, exserted capsules, often on roughened setae, collenchymatous exothecial cells, rostrate opercula, baffle-like cross walls in endostomial segments and mitrate calyptrae. 1. Costa single, ending at or above midleaf, rarely absent; or leaves absent and only pro- tonemata present...... 2 2. Leaves bordered or absent; calyptra fringed; pseudoparaphyllia absent...... Daltoniaceae 2. Leaves un bordered ; calyptra hairy but not fringed; pseudoparaphyJlia filamentous ...... Adelotheciaceae 1 . Costa strong and double to absent ...... 3 3. Calyptra multistratose; axillary hairs mostly 3-5-celled; central strand usually present; leaf cells large and lax ...... Hookeriaceae 3 . Calyptra unistratose; axillary hairs 2-celled; central strand absent; leaf cells usually smaller and thicker-walled ...... , 4 W. R. BUCK: Another view of familial delimitation in Hookeriales 31

4. Calyptra cucullate; sterns without differentiated layers ...... Leucomiaceae 4. Calyptra mitrate; stem with small, thick-walled outer cells (or occasionally also with hyalodermis) and larger, thinner-walled inner cells .... Callicostaceae

Hookeriaceae Schimp., Coroll. Bryol. Eur. 101. 1855 [1856]. Cyathophoraceae Kindb., Bot. Centralbl. 76: 85. 1898; H. A. Mill., Phytologia 21 : 251. 1971 , nom. superf. Pterygophyllaceae Braithw., Brit. -PI. 3: 193. 1903, nom. illeg.; Rodway, Pap. & Proc. Roy. Soc. Tasmania 1912: 10. 1913. Plants relatively robust in soft to stiff, ± shiny, pale-green to golden, thin mats or tufts. Stems creeping or erect, little-branched, complanate-foliate; rhizoids tufted on stems, roughened; paraphyllia absent; pseudoparaphyllia filamentous or absent; in cross-section stems with cells all about the same size, the outer ones sometimes somewhat thicker-walled, central strand present, of small cells, rarely without central strand and epidermal cells small and thick-walled (Schimperobryum); axillary hairs (2-)3-5-celled, the basal one small and brown, the apical ones longer and hyaline. Leaves usually differentiated, the lateral ones larger, usually ovate, the dorsal ones smaller, often ± orbicular, rarely with the base modified into a small pouch, only slightly contorted when dry; margins usually plane, entire to serrate, not or only obscurely bordered; costa single and forked to absent; cells large, lax, smooth, hexagonal, thin-walled, usually ± porose; alar cells not differentiated; propagula present in leafaxils or on rhizoids or leaf apices. Autoicous or dioicous. Perichaetia along stems, some­ times ± bulbiform. Setae often short, sometimes elongate, smooth; capsules erect and short (on short setae) or inclined and cylindric (on long setae); exothecial cells usually strongly collenchymatous; annulus present; operculum conic-rostrate; peristome double, exostome teeth narrowly bordered, on front surface with a zig-zag center line or (in Achrophyllum) furrowed, cross-striolate or (in Cyathophorella) papillose, slightly projecting at back; endo­ stome with a fairly high (or rarely low) basal membrane, segments keeled, perforate, with baffle-like crosswalls, cilia none or rudimentary. Spores spherical, finely papillose. Calyptrae mitrate, smooth or slightly roughened, naked, mostly 4-6 cells thick. Genera: Achrophyl/um Vitt & Crosby (Pterygophyllum Broth., horn. illeg.), Cyatho­ phorella (Broth.) Fleisch., Cyathophorum P.-Beauv., Dendrocyathophorum Dix., J. E. Srn., Schimperobryum Marg. (Lamprophyllum Schimp. ex Broth., horn. illeg.). The Hookeriaceae are uniquely characterized by large, lax, often porose leaf cells, a weak costa, unbordered leaves, smooth setae, cross-striolate exostome teeth and multistratose calyptrae. The last character is truly remarkable, resulting in sturdy, almost rubbery calyptrae. Although it has long been described and even iIIustrated by morphologists (e.g., Hy 1884, p. 165, pl. 14, fig. 71; Janzen 1917, S. 214, Abb. 31), it has been ignored by systematists. I have verified it in Hookeria, Achrophyllum, Schim­ perobryum and Cyathophorum. It is this last genus and its allies which may appear most surprising in an alliance with Hookeria since it has long been associated with Hypoptery­ gium. Although Buck and Vitt (1986) transferred the Hypopterygiaceae to the Bryales, they (we) failed to evaluate the generic inclusions of the family. However, Cyatho­ phorum and Hypopterygium have little in common but dimorphic leaves. Cyatho­ phorum, unlike Hypopterygiaceae s. str., has little-branched plants, large, lax leaf cells, weak costa, unbordered leaves, erect capsules and multistratose calyptrae, among 32 Journ. Hattori Bot. Lab. No. 64 198 8 other differences. Although Cyathophorum and associated genera may deserve sub­ familial recognition within the Hookeriaceae, they are surely most similar to the other genera of the family and phylogenetic relationships are best represented by a single family.

Daltoniaceae Schimp., Syn. Musc. Eur. 478. 1860; H. A. Mill. , Phytologia 21 : 250. 1971, nom. superf. Nemataceae Broth., Nat. PR. 1(3): 916.1907, nom. inval. ; Ephemeropsidaceae Schultze-Motel, Taxon 19: 251. 1970. Distichophyllaceae H. A. Mill ., Phyto\ogia 21 : 249. 1971. Plants mostly small in mostly shiny, pale green to golden to reddish, tufts or mats, rarely reduced to a persistent protonemata. Stems erect to prostrate, mostly little branched, com­ planate-foliate or not; paraphyllia and pseudoparaphyJlia none; rhizoids at base of erect stems or all along prostrate ones, smooth or± roughened; in cross-section stems with all cells similar, the outer ones often somewhat thicker-walled and the inner ones somewhat larger, or with outer layer differentiated, central strand none; axillary hairs 3--4(- 12)-celled, the basal one short and brown or hyaline, the apical ones hyaline, short or elongate. Leaves differentiated or not, if so the lateral ones larger and broader, usually contorted when dry, lanceolate to ovate; margins plane, entire or less often serrulate, bordered by narrow, elongate cells ; costa single, mostly ± percurrent, sometimes short or absent (in Calyptrochaeta and Distichophyl­ lidium); cells short, ca. 1- 3: 1, firm-walled, smooth; alar cells not differentiated ; propagula sometimes in leafaxils or on rhizoids. Mostly autoicous or synoicous, less often dioicous. Perichaetia small, inconspicuous. Setae elongate, usually roughened (or rarely hirsute) above, rarely throughout, or smooth; capsules erect or rarely inclined, cylindrical, smooth or rarely pustulate; ,exothecial cells collenchymatous; annulus present or absent ; operculum conic­ rostrate; peristome double, exostome teeth narrowly bordered, on front surface with either median furrow and cross-striolate or papillose and unfurrowed, slightly projecting at back; endostome with a low basal membrane, segments keeled, with baffle-like crosswalls, perforate OI:not, cilia none or rarely single. Spores spherical, papillose. Calyptrae mitrate, fringed at base, o(ten somewhat roughened, usually naked, sometimes erect-hairy, unistratose. Genera: Calyptrochaeta Desv. (ErioplIs Brid., nom. illeg. ; Piloserioplls Sharp), Crosbya Vitt_(fi ellia J;Jroth., nom. illeg.), Daltonia Hook. & Tay!., Distichophyllidium Fleisch., Disticho­ phyl'lIf1l. Dozy & Molk., Goeb., Leskeodon Broth., Leskeodontopsis Zant. The Daltoniaceae are mostly characterized by small, little-branched plants with unic~state , bordered leaves ~ The elongate setae are often roughened above. The calyptra is mitrate, mostly naked but fringed .at the base. The:O.altoniaceae have~ been broadly interpreted by Crosby (1974) to include any genus in the order with papillose exostome teeth, thereby encompassing such diverse genera (1,& Daltoniq, Callicosta and Cyathophorella. However, generic pairs, truly almost inseparable when sterile, areJelega,ted to different families. For example, Daltonia and q;.osbJ!a ~ave identical growth habits, stem cross-sections, axillary hairs, leaf areola­ tion and borders, c,:ostal development, exothecia and calyptrae. They diffe~ only in e~9$tomeornamentation. Of a,·si!nilar nature is the the relationship.between Leskeodon r;OO:Pistichophyllun:z. There(ore"l ..c~J1 c>ply conclude.thatthere is a strong t~p.dency, W. R. BUCK: Another view of familial delimitation in Hookeriales 33

not only in the Daltoniaceae but in the whole order, to reduce cross-striolate exostome teeth to papillose ones, and that the latter condition is no indication of phylogenetic relationships. Ephemeropsis has long been considered worthy of familial distinction. Indeed, most classifications have given it subordinal status. However, it is an ancient genus, pre­ viously of wider dispersal, as evidenced by Eocene fossils from Europe (K6ck 1939). Although currently restricted to Southeast Asia and vicinity, the genus shows that at one time it had a distribution not too dissimilar to that of Daltonia today, that has mananged to survive in oceanic areas of Europe. Sporophytically Ephemeropsis is very close to Daltonia, and even Zielinski (1909) commented on the similarily of calyptrae of Daltonia and Ephemeropsis. Therefore, placement of Ephemeropsis in the Daltoniaceae best represents its relationships (and prevents the Daltoniaceae from being paraphyletic). The most questionable inclusion in the family, from my point of view, is Calyptro­ chaeta. The weak costa, well developed stem anatomy and curved seta are obviously out of place. However, the bordered leaves, 3-4-celled axillary hairs and fringed calyptra most strongly suggest an alliance with the Daltoniaceae rather than with any other family in the order. If a monotypic family were erected, one would still be left with a decision as to relationships. Therefore, I prefer to put Calyptrochaeta in the Daltoni­ aceae.

Adelotheciaceae Buck, Brittonia 39: 214. 1987. Plants medium-sized in green to brownish, usually smallish, subpendent tufts. Stem cre­ eping when in contact with substrate, ascending to dropping at apical end, complanate-foliate; paraphyllia none; pseudoparaphyllia filamentous, numerous ; rhizoids smooth or slightly roughened; in cross-section stems with smal1, thin-wal1ed cells surrounding larger, firm-walled cells, central strand none; axillary hairs 10-15-celled, al1 hyaline and of equal size. Lateral and dorsal leaves differentiated, ± ovate, decurrent; margins plane, denticulate-crenulate, not bordered; costa single; cells ± isodiametric, thick-walled, po rose, smooth or papillose ; alar cel1s not differentiated; propagula borne on microphyllous branches. Dioicous. Setae short, smooth or lightly roughened; capsules erect; exothecial cel1s ± col1enchymatous; annulus none; operculum conic-rostrate; peristome double, exostome teeth on front surface perforate along zig-zag center line, cross-striolate; endostome with a fairly high basal membrane, segments keeled, perforate, with baffle-like crosswalls, cilia rudimentary. Spores spherical, papillose. Calyptrae mitrate, plicate, not fringed, densely erect-hairy, smooth, unistratose. Genus: Adelothecium Mitt. (and possibly Bryobrothera Ther.). The Adelotheciaceae are characterized by a strong, single costa, isodiametric leaf cells and unbordered leaves. Highly specialized branches bear bifid/paired gemmae on gemmaphores. The calyptra, though densely erect-hairy, is not fringed at the base. Because of the large number of apomorphies, and the lack of similarity to other families in the order, I (Buck 1987) described it as a new family. I am not absolutely confident of its placement in the Hookeriales. Rather, it is somewhat possible that its affinities lie in the Leucodontales. Like that order, specifically the Pterobryaceae, it has numerous filamentous pseudoparaphyllia (Allen 1987), copious geIl)mae and many-celled axil- 34 Journ. Hattori Bot. Lab. No. 64 198 8 lary hairs. Also, densely hairy mitrate calyptrae are not incongruous in the Leucodon­ tales (e.g., Squamidium). However, the total lack of alar development and baffle-like crosswalls in the endostome have, at least for the time being, swayed me toward the Hookeriales. As additional data become available a possible relationship with the Leucodontales should be considered. It is very possible, despite phytogeographic incongruities, that Bryobrothera belongs in the Adelotheciaceae. I know no more now that I did when establishing the family (Buck 1987), but am increasingly impressed with the similarities between the leaves, especially costal development and areolation, of Adelothecium and Bryobrothera. I am most likely to feel more comfortable in placement of the genus if I could see it in the field.

Leucomiaceae Broth., Nat. Pfl. 1(3) : 1095. 1908. Plants slender to robust in mostly pale green, sometimes silvery or golden, often extensive, fiat mats. Stems creeping to scrambling, fragile, irregularly branched to subpinnate, corn pla­ nate-foliate; paraphyllia and pseudoparaphyllia none; rhizoids tufted on stems, mostly lightly roughened; in cross-section stems with all cells thin-walled and similar, central strand none; axillary hairs 2-celled, the basal one short and brown, the apical one elongate and hyaline. Leaves sometimes slightly contorted when dry, often ± falcate, lateral and dorsal leaves somewhat differentiated, lanceolate to ovate, ± symmetric; margins mostly plane, entire to serrulate, not bordered; costa none or very short and double ; cells long-rhomboidal to linear, thin-walled, lax, smooth; alar cells not differentiated; propagula none. Autoicous, synoicous or dioicous. Perichaetialleaves erect, pale, similar to vegetative leaves. Setae elongate, smooth throughout or slightly roughened at apex; capsules inclined to horizontal, mostly symmetric, cylindric, sometimes with a well defined neck; exothecial cells ± collenchymatous; annulus of thin-walled cells; operculum rostrate; peristome double, exostome teeth narrowly bordered, on front surface typically with a median furrow, cross-striolate below, coarsely papillose above, slightly projecting at back; endostome with a fairly high basal membrane, segments keeled, perforate, with baffle-like crosswalls, cilia 1- 3, ± nodose, sometimes short or rudi­ mentary. Spores spherical, finely papillose. Calyptrae cucul1ate, smooth or roughened above, naked or sparsely hairy, unistratose. Genera : Leucomium Mitt., Rhynchostegiopsis C. MliIl. The Leucomiaceae are characterized by flaccid, ecostate leaves with long, lax cells; the stems are composed of thin-walled cells throughout and are thus fragile. The capsule is perfectly hookerioid except for the presence of cilia in some taxa. However, were it not for the cucullate calyptra there would be little justification for familial segregation. The family has but two genera. Rhynchostegiopsis is a small, underdescrib­ ed, neotropical genus characterized by serrate leaf margins, often associated with a zig-zag leaf apex, and better developed endostomial cilia.

Callicostaceae Crum, Bryologist 87: 210. 1984. Pilotrichaceae Kindb., Bot. Centralbl. 77: 54. 1899, nom. iIIeg. Hookeriaceae subfam. Hookeriopsidoideae H. A. Mill., Phytologia 21: 248. 1971. Plants medium-sized to robust, sometimes small, in pale green to golden mats or tufts. W. R. BUCK: Another view of familial delimitation in Hookeriales 35

Stems mostly creeping, sometimes erect, scarcely branched to regularly bipinnate, often com­ planate-foliate; paraphyllia none; pseudoparaphyllia none or foliose; rhizoids usually all along stem, sometimes only at base, mostly somewhat roughened; in cross-section stems with or without a hyalodermis over small, thick-walled cells surrounding larger, thin-walled cells, central strand none; axillary hairs 2-ceJled, a short, basal brown one and an elongate hyaline apical one. Lateral and dorsal/ventral leaves often differentiated in shape, areolation and costal development, sometimes isophyllous, sometimes contorted when dry, lanceolate to broadly ovate; margins plane or less often erect or narrowly recurved, mostly serrulate to serrate, rarely entire, not or sometimes conspicuously bordered by narrow, elongate cells; costa mostly strong and double, sometimes short and double, rarely absent, never single; cells isodiametric to linear, thin- to thick-walled, smooth, prorulose, or papillose (with I-several papillae over each lumen); alar cells not differentiated; propagula common, especially in leaf axils. Autoicous or dioicous. Perichaetia mostly along primary stems, or if with creeping stems and erect primary branches, then along branches, small, inconspicuous, usually with pale leaves. Setae usually elongate, sometimes short, smooth or roughened; capsules usually inclined, if erect then seta short and/or peristome reduced; exothecial cells collenchymatous or not; annulus present or not; operculum conic-rostrate; peristome double, exostome teeth narrowly to broadly bordered, on front surface either with a furrow and cross-striolae or no furrow and papillae; endostome with a high to low basal membrane, segments keeled, per­ forate or not, with baffle-like crosswall s, cilia none or rarely present. Spores spherical, mostly finely papillose. Calyptrae mitrate, not or rarely (Crossomitrium) fringed, often hairy, the hairs usuaJly erect, often ± roughened, unistratose. Genera: Actinodontium Schwaegr., Amblytropis (Mitt.) Broth., Brymela Crosby & Alien, Callicosta C. MUll. (Pilotrichum Brid., horn. ilIeg.), Callicostellopsis Broth., Crossomitrium C. MUll., Cyc!odictyon Mitt., Diplonellron Bartr., Helicoblepharum (Mitt.) Broth., Hemiragis (Brid.) Brid. (Harpophyllum Spruce ex Ren. & Card.), Hookeriopsis (Besch.) Jaeg., Hypnella (c. MUll.) Jaeg., Lepidopi/idium (c. MUll.) Broth., Lepidopilum (Brid.) Brid., Philophyllllm C. MUll., Pilotrichidium Besch., Sauloma (Hook. f. & Wils.) Mitt. (Pulvinella Broth. & Herz.), Schizomitrium B.S.G. (Callicostella (c. MUll.) Mitt.), Stenodesmus (Mitt.) Jaeg., Stenodictyon (Mitt.) Jaeg., Tetrastichium (Mitt.) Card., Thamniopsis (Mitt.) Fleisch., Trachyxiphillm Buck, Vesicu/ariopsis Broth. (Achrohypnella Herz.). The Callicostaceae accommodate most of the genera in the Hookeriales, many of which are confined to relatively recent geological areas, thus giving the impression of great genetic vitality. Probably due to lack of adequate study, I am still unable to understand more than the basic evolutionary trends in the family. The strong, double costa is the primary claim to distinctiveness for the family but it has been reduced repeatedly, as has the cross-striolate ornamentation of the exostome teeth. Two large groups of genera perhaps indicate lines of evolution. One, including Callicosta and Schizomitrium, has a stem without a hyalodermis and when the exostome becomes reduced the teeth are only slightly bordered. The second lineage involves genera, such as Thamniopsis and Lepidopilum, with a hyalodermis and exostome teeth, when reduced, strongly bordered. In the most tentative way I have speculated on the relationships of the families of the Hookeriales (see dendrogram). I suspect that the common ancestor of the order 36 Journ. Hattori Bot. Lab. No. 64 1 988

Daltoniaceae Leucomiaceae Hookeriaceae \ Adelotheciaceae \' Callicostaceae

calyptra 'ringed / calyptra cucullate \ leaves bordered / leal cells large costa strong & double \ :r:e:,dooaraPhYllia lilamentose calyptfa multistratose central strand present y~"'~"~~'

leaf cens small costa single leaf cells elongate axillary hairs 2-celled leaf cells short plants freely branched adlary hairs 3-celled plants little-branched

Dendrogram representing postulated relationships between the families of the Hookeriales. had unbordered leaves with either a short and double or weak single costa, shortish laminal cells and undifferentiated alar cells. The seta was elongate, bearing an inclined, symmetric capsule with cross-striolate exostome teeth. The calyptra was probably mitrate. Perhaps the Hookeriales shared a common ancestor with the Leucodontales. The proposed classification seems the most parsimonious, with the fewest reversals. It is proposed primarily to stimulate work on the group and is not meant to be indelible.

LITERA TURE CITED ALLEN, B. 1987. On distinguishing Pterobryaceae and Meteoriaceae by means of pseudoparaphyllia. Bryol. Times 42: 1-3. BROTHERUS, V. F. 1925. Hookeriales. In: A. ENGLER & K. PRANTL (eds.), Die natiirlichen Pflanzen­ familien, ed. 2, 11 : 214- 278. BUCK, W. R. 1987. Taxonomic and nomenclatural rearrangement in the Hookeriales with notes on West Indian taxa. Brittonia 39: 210-224. BUCK , W. R. & D. H. VITT 1986. Suggestions for a new familial classification of pleurocarpous mosses. Taxon 35: 21 - 60. CROSBY, M. R. 1974. Toward a revised classification of the Hookeriales (Musci). J. Hattori Bot. Lab. 38: 129-141. FLElSCHER, M. 1923. Allgemeine Uebersicht des natiirlichen Systems der Laubmoose. In : Die Musci der Flora von Buitenzorg 4 : XI-XXUL Hy, F. C. 1884. Recherches sur l'archegone et le developpement du fruit des Muscinees. Ann. Sci. Nat. Bot. VI, 18 : 105-206, pis. 9-14. JANZEN, P. 1916-1917. Die Haube der Laubmoose. Hedwigia 58 : 156-192 (1916),193-280 (1917). KOCK, C. 1939. Fossile Kryptogamen aus der eoziinen Braunkohle des Geiseltales. Nova Acta Leop., n.s. 6 : 333-359, Taf. 37-45. MILLER, H. A. 1971. An overview of the Hookeriales. Phytologia 21 : 243-252. ZIELlNSKI, F. 1909. Beitriige zur Biologie des Archegoniums und der Haube der Laubmoose. Flora 100: 1-36.