Journ. HallOl'i BOI. Lab. No. 61 : 309- 332 (Dec. 1986)

SCIAROMIADELPHUS A. ABR. & 1. ABR. - THE RELATIONSHIP BETWEEN EXTANT AND FOSSIL SPECIMENS

R YSZARD O CHYRA

ABSTRACT. Sciaromiadelphus lon!!.ifolius A. Abr. & I. Abr., a fossil moss from the Pliocene deposits from Bashkiria in the Soviet Union, is redescribed and illustrated . It is congeneric with and closely related to the extant Sciaromium barllellii Crum & Steere from Hispaniola, which is also redescribed and ill ustrated. The new genus Sciaromiella Ochyra is proposed and two new combinations, S . longifolia (A. Abr. & I. Abr. ) Ochyra and S. barllellii (Crum & Steere ) Ochyra, are made. Sciaromiadelphus A. Abr. & I. Abr. , a fossil moss genus is congeneric with the extant Sciaromiella , the latter having a priori ty over the former. Sciaromiella is considered to be closely related to Sciaromiopsis Broth., a monotypic genus from China. Sciaromiopsis is redescribed and ill ustra ted. It includes only S. sinellSLI (Broth.) Broth., and S. brevifolia Broth. is synonymous wit h the former species. S. nipponensis Sak. from Japan is di scussed and this species is inseparable from Brachy rhecium populeum (Hedw.) B., S. & G . var. populeum. T he fa milial placement of Sciaromiella and SCiaromiopsis is di scussed and they are placed in the newl y described su bfamily Sciaromiopsoideae Ochyra of the Donrichardsiaceae Ochyra. A key for genera and spec ies of Donrichardsiaceae is gi ven and the geographical affiliation of the Sc iaromiopsoideae is considered.

INTRODUCTION Sciaromiade/phus A. Abr. & 1. Abr. is a fossil moss genus that includes only one species, S. longifolius A. Abr. & I. Abr., that have been found in the Pliocene moss assemblage of the K inelian deposits from Bashkiria in the Soviet Union (Abramova & Abramov (967). The moss was recovered from the unsurpassed well-preserved fossil remain s of considerable thickness (Abramov (965). Contrary to most fossilized moss specimens that have hitherto been reported, there is excellent preservation of in these Pliocene deposits and the Abramovs were able to examine cellular details of the leaves and to make transverse sections of the stems and leaves as with extant . This resulted in an excellent description of S. /ongifo/ius and therefore an evaluation of its possible affinities with both fossil and extant taxa of mosses was feasible. Abramova and Abramov (1967) included Sciaromiadelphus in the Amblystegi­ aceae. However, they did not discuss the problem of the taxonomic position of this newly described genus and stated only that the general aspect of S. longifolius is compatible to that of some species of Drepanocladus, for in stance D. aduncus var. kneifJi.i or D. jluitans, especia Jl y in habit and leaf shape. On the other hand, the generic name suggests its si milarity and relationship to Sciaromium (Mitt.) Mitt., a genus characterized by having limbate leaves. During the course of my on-going revision of Sciaromium and potentiaJly related

I Laboratory ot Bryology, Institute of Botany of the Polish Academy of Sciences, Lubicz 46, PL-31-512 Krak6w, Poland. 310 Journ. Hattori Bot. Lab. No. 61 I 9 8 6 genera with limbate leaves I critically re-examined the holotype of S. longifolius housed in the Komarov Botanical Institute in Leningrad (LE) and compared it to extant genera having limbate leaves, including Sciaromium and Sciaromiopsis Broth., genera that are currently placed in the Amblystegiaceae (Brotherus 1925; Vitt 1984). Special attention was given to Sciaromium bartlettii Crum & Steere, a species described from two collections from Hispaniola (Crum & Steere 1958). It soon became obvious that the fossil , Sciaromiadelphus longifolius, and the extant, Sciaromium bartlettii, shared many critical and taxonomically important characteristics, suggesting close relation­ ships and affinities between these two taxa, and their possible congeneric status.

DESCRIPTION OF SCIAROMIADELPHUS LONGlFOLIUS (Fig. 1) Plants rather robust, rigid and wiry in texture. Stems elongate, 5- 8 cm long, si mple or sparsely fastigiately branched, somewhat curved at tips, in transverse section composed of 2- 3 rows of smaller, thick-walled cortical cells surrounding 5- 6 rows of larger, rather firm-walled

FIG. I. Sciaromiadelphus longifolius A. Abr. & I. Abr. 1-3: leaves; 4: portion of side­ branch; 5: mid-leaf cells; 6: angular cells of the upper leaf; 7: upper marginal cells without border; 8: transverse section of the costa in the upper part; 9: transverse section of the costa in the median part; 10-11 : transverse sections of leaf borders; 12: transverse section of the stem (according to Abramova and Abramov 1967). R. OCHYRA: Sciaromiadelphus - relationship between extant and fossil mosses 3 11 inner cells, central strand lacking or inconspicuous. Leaves crowded and appressed at tips of stems and branches, rather loosely arranged below, erect to erect spreading, (3.0- )3.5- 4.0(- 5.0) mm long, (0.5-)0.8-\.0(- 1.3) mm wide, ovate-lanceolate and gradually long to filiform acuminate, straight or slightly secund, somewhat rounded to the insertion, not auriculate and non-decurrent; margins plane, entire to sli ghtly and distantly serrulate, bordered by 6--12 rows of narrow, elongate, thick -walled, brown cells in 2 layers; borders well-developed and conspicuous in older, lower leaves, reaching usually the apex and confluent with the costa or ending somewhat below the apex, in uppermost stem and branch leaves narrower and indistinct or lacking; costa single, terete, stout, 70-80/lm wide at base, sli ghtly tapered, percurrent to long excurrent as a stout, cuspidate point, in transverse section bi-convex, composed of uniform, small, thick-walled cells, and only sometimes with some ventral outer cells at the base larger than inner cells; lamina cells unistratose, frequently with bistratose, longi tudinal streaks extending as spurs from costa and margins or developing independently; upper median cells flexuose, somewhat prorate, linear-rhomboidal, becoming longer downwards, (26- )40-55(-60)/lm long, 8- 9/lm wide; basal cells wider and shorter in several rows, short-oblong to oblong-hexagonal, rather thin-walled, more lax, 15- 25 ~lm long, 10-13 /lm wide. Sexual organs and sporophytes unknown. The distinctive features of the monotypic fossil genus Sciaromiadelphus are erect to erect-spreading, ovate-lanceolate to lanceolate leaves that gradually taper into a long acumen and in general aspect resemble those found in a variety of Drepanoc/adus species; distinct, bistratose borders of the leaves composed of 6-- 12 rows of narrow, thick-walled, linear cells that are usually confluent with the percurrent to long excurrent costa; and linear-rhomboidal, flexuose and prorulose lamina cells.

DESCRIPTION OF SCIAROMJUM BARTLETTlI (Fig. 2) Plants stiff, yellow-green or golden-green to brownish, glossy, in loose mats. Stems up to 6 cm long, prostrate, sparsely fastigiately branched, brown, naked, in transverse section composed of 2-3 rows of small cortical cells with incrassate walls and 5-7 rows of larger, rather firm-walled cells in an inner area, central strand absent or inconspicuous and obscure; paraphyllia none, pseudoparaphyllia not abundant, foliose to suborbicular, entire to irregularly serrulate. Leaves erect, densely set, sl ightly flexuose and twisted in the upper part of stem and branches when dry, \.9- 2.1 mm long, 0.8- 0.9 mm wide at the base, ovate-lanceolate, gradually tapering into a rather short acumen, flat or somewhat concave; margins plane, entire or si nuate to distantly indistinctly serrulate, bordered all around by 5- 10 rows of narrow, elongate, linear­ flexuose, thick-walled, yellowish-brown to brownish cells, not sharply demarcating from lamina cells, in transverse section bistratose, consisting of thick-walled, homogeneous cells; costa single, yellow, 95-110/lm wide at the base, gradually narrowed to the apex, percurrent to short excurrent, confluent with bistratose margins and forming bistratose, cuspidate point, in transverse section composed of uniform, thick-walled cells, bi-convex; lamina cells unistratose, occasionally with bistratose strands radiating from costa and margins as spurs, rather obscure, green to yellowish-green, Iinear-flexuose throughout the lamina, 60-110 /lm long, 8-9/lm wide, with moderately incrassate walls, papillose by projections of the upper ends; basal cells noticeably wider and shorter than upper cells, short rectangular to oblong-hexagonal, greenish to yellowish-green, ll- 14/lm wide, 40-55/lm long; cells at the insertion similar in shape, not coloured and not forming auricles. Sex apparently dioicous. Male gametangia bud-like, 312 Journ. Hattori Bot. Lab. No. 61 I 9 8 6

, . 2 01 22 19 j16 1\ ." J 20 18 00-0\ , \ F IG. 2. Sciaromium barllettii Crum & Steere. I: habit; 2: portion of branch when dry; 3--6: leaves; 7- 9: pseudoparaphyllia; 10: transverse sections of leaves; 11 : transverse section of the stem; 12: angular cells of the leaf base; 13: median leaf cells; 14: median leaf cells at margin; 15: perigonium; 16-18: outer perigonial bracts; 19-22: inner perigonial leaves [1-4, 7- 14 drawn from isotype (KRAM); 5- 6, 15- 22 drawn from paratype (MICH)] . R. OCHYRA: Sciaromiadelphus - re lationship between extant and fossil mosses 313

0.7- 0.8 mm long; outer perigonial bracts lingu late, truncate or broadly rounded at the apex, entire and irregul arly si nuose, 0.2- 0.4 mm long, 0.2-0. 3 mm wide; inner perigonial leaves broadly ovate-lanceolate , deeply concave, graduall y narrowed into a broad, short apex, 0.5- 0.6 mm long; antheridia few, claviform, short-stalked, about 250 JLm long; paraphyses few, hyaline, filiform, reach ing beyond the antheridia. Perichae ti a and sporophytes unknown .

G ENERIC POSITION OF S CIAROMIUM BARTLETTlI The generic position of S. bart/ellii was one of the most problematic and difficult tasks of my revisionary study of Sciaromium. This sterile moss does not seem to fi t the concept of any extant genus of pleurocarpous mosses. Therefore, it is only possible to speculate on affinities of this unique moss, exclusive of sporophyte characters. It is obvious that the problem of the systematic disposition of mosses should be judged upon as many sporophytic and gametophytic characters as possible. However, many examples ind icate convincingly that strict adherence to the principle of placing a genus within one or the other moss famil y or order based exclusively on peristome morphology may yield polyphyletic groups. In fact, many families and genera of hypnobryalean mosses are established mainly on the basis of gametophyte characters, because the peristome does not offer any help in this matter, being perfectly hypnaceous in structure. Hence, gametophyte characters may not be disregarded in a discussion of the phylogenetic position of taxa in question. It should, however, be remembered that in all such speculations it is necessary to be careful because of the deceptiveness of vegetative characters and convergence showed by organisms of remote phylogenetic position that live in similar t:nvironmental niches. Crum and Steere (1958) compared Sciaromium bart/eltii to Sciaromium fryei R. S. Williams [= Limbellafryei (R. S. Williams) Ochyra, comb. no v.; basionym: Sciaromium fryei R. S. Williams, The Bryologist 35: 52, fig . 5 (1- 8), 1933] from Oregon in western North America, a species closely related to the Hawaiian Limbella tricostala (Sull.) Broth. These two taxa, however, are only similar in having limbate leaves while all other character states, including leaf areolation and branching pattern, are totally different. Limbel/afryei has short, smooth, irregular, 4-6-angular, quadrate, rhombic, oblong-rhomboidal or shortly oblong upper lamina cells, whereas those of S. bartfellii are elongate, linear-fiexuose and prorate. The branching pattern of the stem is also a significant character since it gives the moss its growth habit. Alas, this important character is frequently neglected in moss taxonomy (Koponen 1982). Species of Limbella are typically dendroid mosses and this aspect of the plants, combined with leaf areolation, the presence of characteristic stipe-Ieaves and the leaf shape, indicate a close affinity of this genus to Thamnobryum and Porothamnium of the Thamnobryaceae (Ochyra 1987a). On the other hand, S. bart/ettii has typically prostrate aspect of the plants and this character precludes any close affinity of this species with Limbella. Similarly, the placement of S. bart/eltii by Crum and Steere (1958) in the section Limbidium of Sciaromium appears to be unwarranted. This section consists of 15 species that are widely distributed in the warmer and cool temperate zone of the 314 Journ. Hattori Bol. Lab. No. 61 I 986

Southern Hemisphere, from Juan Fernandez Islands to South Africa and Kerguelen Islands (Brotherus 1925). After careful taxonomic study of this complex, I (Ochyra 1987c) found that all these species are inseparable from one other and actually they represent a highly variable, amphibious moss, Vittia pachyloma (Mont.) Ochyra, which is the only representative of the Vittiaceae Ochyra, a monotypic fami.Jy having a typical austral distribution pattern. S. bartlettii is evidently anomalous in the section Limbidium of Sciaromium. There has been an increasing tendency in past years to reunite many species characterized by having limbate leaves in one, all-encompassing genus Sciaromium; this arises, surely, from stressing superficial features, rather than structural features in the definition of this genus. If defined in terms of the character states enumerated by Brotherus (1925), Sciaromium forms a heterogeneous taxon encompassing many different genera of rather distant affinity (Ochyra 1987b, c, d , e). The only real character, which was used by Crum and Steere (1958) to include S. bartlettii in the section Limbidium ( = Vittia Ochyra) is the presence of bordered lea ves. However, Vittia is a well-defined genus warranted by several structural character states including strong, polystratose leaf margins consisting of four to many layers of cells and short, rhombgidal to hexagonal, smooth to very slightly prorulose lamina cells. Contrary to Vittia, S. bartlettii has only bistratose, relatively weak leaf borders that are not sharply set off from the lamina cells; the latter are elongate, linear-flexuose and distinctly prorate. These characters seems to be sufficient to preclude any closer affinity of both taxa. Thus, S. bartlettii otherwise does not fit the concept of the section Limbidium of Sciaromium. Sciaromium bartlettii is also absolutely different from species of the other sections of Sciaromium. Species of the section Aloma Dusen have totally elimbate leaves and do not show even a superficial similarity to S. bartlettii (Ochyra 1985a, 1986a, 1987b). Finally, S. bartlettii is entirely different from the eastern North American S. lescurii (Sull.) Broth. [= Platylomella lescurii (Sull.) Andrews], which is the only species of the section Pia tyloma Broth. of Sciaromium. This taxon is described as having bistratose leaf borders, rhombic to oblong-hexagonal or shortly oblong, smooth lamina cells and the presence of filiform paraphyllia on the surfaces of the stem and branches. The combination of these character states corroborates the generic distinctiveness of this taxon as it was earlier suggested by Kindberg (1897) and Andrews (1945, 1950). Platylomella lescurii is very similar (or identical?) to the fossil Sciaromium laxirete A. Abr. & I. Abr. from the Pliocene deposits from Bashkiria in the Soviet Union (Abramova & Abramov 1967) and obviously these two taxa are totally different from S. bartlettii. Thus, the latter species appears to be different from all species included by Brotherus (1925) in Sciaromium, and it is deemed necessary to seek similarities of this species with other moss genera. Limbate leaves, a basic diagnostic character of S. bartle ttii, are frequently observed in a variety of both pleurocarps and acrocarps. Among mosses with pleurocarpous growth form, pluristratose or otherwise differentiated leaf borders are most often found in taxa growing in rheophilous or semi-aquatic habitats and have evidently an adaptative nature (Vitt & Glime 1984; Ochyra 1987a), and can be found in R. OCHYRA: Sciaromiadelphus - relationship between extant and fossil mosses 315 both genera of isobryalean and hypnobryalean mosses. Bordered leaves also occur frequently in hookeriacean mosses, but such a leaf condition in most genera of this group is simultaneously combined with double or no costa and obviously precludes any affinity of S. bartlettii with Hookeriales. Perhaps the only exception here is Cros@ya Vitt, in which leaves external1y resemble very much those of S. bartlettii. They are bordered by several rows of firm-wal1ed, linear cel1s and the costa is single, percurrent to short excurrent and confluent with borders. The borders, however, are unistratose throughout and only slightly swollen and the leaf margins are recurved. In addition, species of Crosbya can be readily distinguished by smooth, shortly rhombic to elongate-rhomboidal lamina cells with more or less incrassate walls, and the plants are predominantly epiphytic and occasionally produce fusiform gemmae at the base of the outer side of the leaves (Vitt 1977). There are some rheophytes with limbate leaves among isobryalean mosses that seemingly might be related to S. bartlettii. These are some species of Neckeropsis Reichdt. including N. submarginata Card. ex Touw, N. moutieri (Broth. & Par. in Par.) Fleisch., N. boniana (Besch.) Touw & Ochyra and N. negrosense (Bartr.) Touw & Ochyra from the eckeraceae (Touw & Ochyra 1987). However, all these species have leaf areolation of rounded or rhombic cells above and elongate-rectangular below, and smooth throughout the lamina and this leaf cell pattern indicate that there is no closer relationship between S. bartlettii and neckeraceous mosses. There is a great resemblance, at least by the overall aspect of the leaves, of S. bartlettii with some species of Pterobryella (c. Muell.) Jaeg., expecial1y with P. rigida (Mitt.) Touw, a species known from New Caledonia and the New Hebrides (Miller et al. 1978). This species has leaves distinctly bordered above by several rows of linear, firm-walled cells, forming a partly bistratose border that is not sharply set off from lamina cells and is confluent with the percurrent to long excurrent costa. The lamina cells are linear to Iinear-flexuose, distinctly papillose by projections of the upper ends, and have strongly incrassate, occasionally slightly porose, walls. Traditionally, Pterobryella was included in the Pterobryaceae as a separate subfamily and exhibits all the typical features of this family as circumscribed by Brotherus (1925) and Argent (1973). From the bulk of genera and species of this tropical family, Pterobryella differs in a typically dendroid habit of the plants and for this reason Buck and Vitt (\986) raised Pterobryelloideae to the family status and placed it together with other dendroid moss families in the suborder Hypnodendrineae of (= Hypnobryales). Otherwise, Pterobryella has basal cells smal1, isodiametric and with incrassate walls and characteristic bright orange-brown tinge. All aforementioned characters are entirely lacking in S. bartlertii and this definitely excludes any alliance of this species with Pterobryella. There are only two genera of hypnobryalean mosses that appear to be closely related to S. bartlellii. These are the fossil Sciarorniadelphus and the extant Sciaromiopsis. Plants in both genera are prostrate in habit, have bordered leaves, and lamina cel1s that are elongate, Iinear-flexuose and prorate; these character states are exactly the same in S. bartlettii. The only differences between Sciaromiopsis and 316 Journ. Hattori Bot. Lab. No. 61 1 986

Sciaromiadelphus are that the former has multistratose leaf borders, leaf margins that are sharply serrate above and basal lamina cells that are differentiated into distinct, sizable auricles. These character states seems to be sufficient to consider Sciaromiopsis and Sciaromiadelphus as distinct, but closely related, genera. Taxonomic problems and a description of Sciaromiopsis are presented below. A detailled comparison, primarily of leaf shape, border and areolation as well as anatomical details, has revealed that S. bartleltii can be placed in the concept of Sciaromiadefphus, and shares with it critical and taxonomically important characteris­ tics. These include prostrate habit of the plants; ovate-lanceolate, gradually tapering into a short or long acumen, leaves with plane, entire to slightly sinuate or minutely serrulate margins that are bordered with 5- 12 rows of elongate, thick-walled, yellowish to brown cells disposed in two layers; single and stout costa, which is confluent with the borders; and elongate, oblong-hexagonal to linear-f1exuose, prorate lamina cells with occasional bistratose streaks. There are only a few examples in the kingdom of living genera that were earlier described from fossil material. One of these is the coniferous genus Metasequoia, which was described for the first time by Miki (1941) for the fossil Sequoia disticha Heer. Subsequenly, a number of species of Metasequoia were described from Cretaceous and Tertiary fossils in North America (Chaney 1951) and Eurasia (Sveshnikova 1975). However, an extant representative of this genus, M. gfyptrostroboides, was discovered and described in China by Hu and Cheng (1948). The discovery of the extant species of Sciaromiadelphus has serious nomenclatural consequences. According to Article 58.1 of the Sydney Code (Voss 1983) when an extant taxon of plants, with the exception of Algae, and a fossil (or subfossil) taxon of the same rank are united, the name of the extant taxon is treated as having priority. The case of Sciaromiadelphus fongi/o/ius and Sciaromium bartleltii are indeed parallel to that of Melasequoia which is cited as example 2 in Article 58.1 of the Code, but there are some subtle differences. After the discovery of living M. glyptrostrobides, conservation of Metasequoia Hu & Cheng 1948 against Metasequoia Miki 1941 was accepted since this name was in common use in palaeobotany before the discovery of the soon famous extant tree several years later. Contrary to Metasequoia, Sciaromiadelphus is known to very few people. In addition, the decission to unite both taxa of mosses is based entirely upon my study of different plants, all of which are sterile. Therefore, it seems to me that intentional publication of a later homonym for a little known name and a proposal of its conservation at the same time is rather strange. It appears more reasonable to propose a new generic name for the extant species, which according to Article 58.1 of the ICBN would have preference, even if the fossil Sciaromiadelphus is included. This solution appears to have the advantage of being independent from an assumption about the congeneric status of both taxa in question, which may be right or wrong. Consequently, I propose the name Sciaromiella to replace Sciaromiadelphus. The' name is a derivative of the commonly known name Sciaromium that, unfortunately, proved to be synonymous with Echinodium Jur. (Churchill 1986; Ochyra 1987b). For R. OCHYRA : Sciaromiadelphus - relationship between extant and fossil mosses 317 reasons, given below, the genus is placed in the Donrichardsiaceae.

Sciaromiella Ochyra, gen. novo Donrichardsiacearum Plantae aquaticae, sat robustae, fusco-virides vel fuscae, nitentes. Caulis elongatus, prostratus, sat rigidus, irregulariter fastigiate ramificatus, 6 vel pluria cm longus, paraphylliis nullis, pseudoparaphylliis fo liosis ve l suborbicularibus, irregulariter serrulatis vel integerrimis, in sectione transversa ovalis. e cell~lis extern is 2- 3-stratosis, minoribus, fuscis, parietibus crassis, internis 5-7-stratosis, magnis, parietibus sat firmis, fasciculo centrali indistincto vel nullo. Folia caulinaria et ramealia similia, 2.0--4.0 x 0.8- 0.9 mm magna, dense vel remote disposita, ovato-Ianceolata, breviter vellonge acuminata, stricta velleviter falcato­ secunda. Margines folii plani, integerrimi vel distanter parce serrulati, ubique bistratoso limbati vel limbo infra folii apicem evanido, e cellulis elongatis, 5- 12 seriatis, parietibus crassis, fuscis. Costa solitaria, lutea, crassa, 70--110 Jlm latitudine, teres, percurrens vel longe excurrens, cum marginibus crassis confluens, in sectione transversa e cellulis incrassatis, minoribus, plus minusve aequali bus constructa. Lamina unistratosa, saepe radiis bistratosis, liberis vel e \:osta et marginibus egredientibus, praedita. Cellulae laminae 8- 9 Jlm latae, 50--1 10 Jlm longae, oblongo-hexagonales vellineari-flexuosae, minute papillosae, commissuris crassis. Perichaetia ut sporophyta ignota. Plants rather robust, prostrate, stiff, in loose or dense, yellowish-green to brownish, usually glossy mats. Stems elongate, sparsely fasciculately branched, without tomentum of rhizoids, in older parts frequently appearing bristly because of erosion of laminae except only for strong costae; branches erect-spreading, elongate; in transverse section with 2- 3 rows of smaller, brown, strongly incrassate cortical cells surrounding 5- 7 rows of larger cells with moderately incrassate walls in an inner area, central strand absent or indistinct; paraphyllia none, pseudoparaphyllia foliose or suborbicular, entire or serrulate. Stem and branch leaves similar, erect, sli ghtly contorted and twisted when dry, close to distant, straight or with curved tips, flat to sl ightly concave, 2.0-4.0mm long, 0.8- 0.9mm wide at the base, ovate-lanceolate, gradually narrowed to a short or long acumen, non-decurrent; margins plane, entire to slightly sinuate or minutely and indistinctly, distantly serrulate, bordered with 5-12 rows of elongate, thick-walled, yellow to brownish cells, not sharply demarcated from la mina cells, disposed in 2 layers or sometimes border indistinct above in upper leaves; costa single, strong, terete, yellow, subpercurrent to long excurrent, confluent with borders, 70- 110 flm wide at the base, gradually tapered, in transverse section consisting of small, incrassate, homogeneous cells; lamina cells oblong-hexagonal to linear-flexuose, 50- 110 flm long, 8- 9 flm wide, prorate, short-oblong in 2-3 rows at the leaf base, not differentiated in the basal angles. Sporophytes and perichaetia unknown. Type species: Sciaromiella bartlellii (Crum & Steere) Ochyra, comb. novo [Basionym: Sciaromium (Limbidium) bartlellii Crum & Steere, Am. Midland Natur. 60( I): 40, fig . 95- 103, 1958]. Holotype: Haiti, Beaumont to Morne Geffrard, 2000--2600 ft alt., arrondissement de Jeremie (north of divide on road from Les Roseaux to Les Cayes), southern peninsula, H. H. Bartlett 17665, May 13, I 945-MICH'; isotypes: KRAM!, NY!, TNS! Paratype: Haiti, attached to stones in swift, cold, clear stream Riviere Glace between Camp Perrin and Jeremie, Faith P. Mackeness 29 , Oct. I, 1943- MICH!; isoparatypes: CANM! , NY! Sciaromiadelphus A. Abr. & I. Abr. , Nov. Sist. Niz. Rast. 4: 334. 1967, syn. novo Type species: Sciaromiadelphus longifolius A. Abr. & I. Abr., Nov. Sist. iz. Rast. 4: 334, fig . 2 (1 - 12), pis 3--4, 1967 == Sciaromiella longifolia (A. A br. & I. Abr.) Ochyra, comb. novo Holotype: Iz Pliotsenovykh otlozheniy u dier. Monchady Bashk. ASSR. sky. No. 16, glubina 30m, prieparaty I- IV i kiern ["From the Pliocene deposits near the village of Monchady, Bashkirian Soviet Autonomous Socialist Republic, bore­ hole No. 16, depth 30m, slides I- IV and drill-core"] - LE!; thereafter Fig. 2 (1 - 12) and plates 3- 4 in Abramova and Abramov (1967) and Fig. I in the present paper. 318 Journ. Hattori Bot. Lab. No. 61 1 9 8 6

ON SCIAROMJOPSIS

The moss genus Sciaromiopsis was established by Brotherus (1924) to accom­ modate two species. One of them, S. brevi/olia, was described as a new species, whereas the second one Brotherus (1922) described as Sciaromium (Limbidium) sinense and subsequently transferred it to the newly created Sciaromiopsis: Later, Sakurai (1951) added to this genus a third species, S. nipponensis, from Japan. Brotherus (1924 1929) considered Sciaromiopsis to be closely related to Sciaromium, mainly because of the presence of limbate leaves, and also to Leptodictyum (Schimp.) Warnst. because of its leaf areolation of elongate cells and more or less differentiated auricles. Therefore, the genus has become firmly established as a member of the Amblystegiaceae (Brotherus 1925). While discussing taxonomic problems in Sciaromium, Andrews (1945) also gave his attention to Sciaromiopsis. Unfortunately, his entire discussion on the relationships of this genus was based on comparison of descriptions and illustrations provided by Brotherus (1924, 1925, 1929), and he did not examine any authentic material of the taxa. This approach has led Andrews (1945) to the erroneous conclusion that S. brevifolia is presumably congeneric with Handeliobryum Broth., a genus known from the Himalayan region including Assam, Nepal, Sikkim and Tibet. The lack of agreement as to Andrews' (1945) conclusion was expressed by Chopra (1975) and Gangulee (1976). Finally, Noguchi (1978), having examined original collections of Sciaromiopsis species and Handeliobryum, stated definitely that both genera are not related, excepting for the superficial similarity of Iimbate leaves. Also, I (Ochyra 1986b) came to the same conclusion in a discussion of the taxonomy and geographical distribution of Handeliobryum sikkimense (Par.) Ochyra. In addition, Noguchi (1978) found that Sciaromiopsis sinensis and S. brevi/olia are conspecific, whereas S. nipponensis is inseparable from Brachythecium populeum (Hedw.) B. , S. & G . var. quelpaertense (Card.) Tak. Unfortunately, Noguchi's (1978) paper is written entirely in Japanese and is not available for a large part of the bryological community. Moreover, he did not discuss the relationships of Sciaromiopsis and the problem of its familial placement. During the course of the present study, I critically examined all available collections of species described under Sciaromiopsis. S. nipponensis was described by Sakurai (1951) from a single collection from the Akaishi Range on Honshu, Japan. I share the opinion of Noguchi (1978) that this taxon is con specific with Brachythecium populeum. The plants of the type of S. nipponensis are prostrate and the stem leaves are concave with margins recurved or revolute below (Fig. 3). The revoluteness of the margins gives the leaves an illusion of a border as it was misinterpreted by Sakurai (1951). Noguchi (1978) included S. nipponensis in the synonymy of B. populeum var. quelpaertense. According to Takaki (1955) this variety differs from the type variety of B. populeum only in having broad and very plicate leaves with numerous quadrate basal cells. It seems that the taxonomic value of this variety is doubtful, as these characters vary greatly in B. populeum and all possible integradations are easily R. OCHYRA: Sciaromiadelphus - relationship between extant and fossil mosses 319

FIG. 3. BrachYlheciwn populeum (Hedw.) 8., S. & G. I: habit; 2: portion of the stem; 3- 5: stem leaves; 6--7: branch leaves; 8: apex of the stem leaf; 9: upper marginal cells; 10: median marginal cells; 11: angular cells of the stem leaf (all drawn from the holotype of Sciaromiopsis nipponensis Sak., MAK). 320 Journ. Hattori Bot. Lab. No. 61 I 986

FIG. 4. Sciaromiopsis sinensis (Broth.) Broth. I : habit; 2: portion of branch when wet; 3: portion of branch when dry; 4-9: leaves; 10-12: pseudoparaphyllia (1-7, 10-12 drawn from the holotype of Sciaromium sinense Broth., H-Broth; 8- 9 drawn from the holotype of Sciaromiopsis brevi/olia Broth., H-Broth). R.OCHYRA: Sciaromiadelphus - relationship between extant and foss il mosses 321 observed. Type material of S. nipponensis fall s within the range of variation of the type plants of B. populeum. Brachythecium popu{eum (Hedw.) B., S. & G ., Bryo l. Eur. 6: 7.535. 1853. var. popu{eum. Sciaromiopsis nipponensis Sak., J. Jap. Bot. 26: 201 . 2. 1951 . Type: Honshu, Prov. Kai Mt. Akaishi , ad rupes in rivulo Koshibu (Leg. N. Takak i - Typus in Herb. K . Sakurai No. 19505, 17-J ul y- 1949) (Holotype: MAK !; isotype: TNS!), syn. novo Having examined type specimens of Sciaromiopsis sinensis and S. brevifolia, I can confirm Noguchi's (1978) conclusion that these species are identical. Additionally, it appears that Sciaromiopsis shares many essential character states with Sciaromiella that are of importance in discussion of the familial placement of these genera that are evidently anomalous in the Amblystegiaceae.

Sciaromiopsis Broth., Sitzungsber. Ak. Wiss. Wien Math. Naturw. Kl. Abt. I, 133: 580. 1924. (Figs 4-5) Relatively robust plants, in wide, loose, rather rigid, somewhat glossy mats, brown below, sordid-yellow, golden or yellowish-brown to golden-brown above. Stems elongate and creeping, up to 12 cm long, not radiculous, freely and irregularly to fasciculately branc hed; branches elongate, sparsely pinnate. ascending, flexuose. usually curved at tips; stems and branches brown below, greenish-brown to yellowish-brown above. densely foliate above, commonly entirely denuded or bristly below because of erosion of leaf laminae excepting stout midribs and borders; cortical cells in 2-3 layers, small , thick -walled, surronding 3-6 layers of larger, rather firm­ walled, inner cells, central strand conspic uous; paraphyllia lacking, pseudoparaphyllia scattered, fo liose, suborbicular, distinctly crenulate at the broadly rounded apex. Leaves loosely set, erect. usually homomallous and shrinked when dry, erect-spreading when moist, (1.6-)1. 7- 2.1( - 2.3 ) mm long, 0. 7-0.8 mm wide, narrowly to broadly ovate-lanceolate, rarely oblong­ lanceolate, gradually acuminate, straight or somewhat falcate and asymmetric, plane, rounded at the insertion, auriculate and decurrent; margins plane, sharply serrate above, remotely and sharply serrulate to subentire below, bordered by about 7- 12 rows of thick-walled, linear, brown cells usually in 4-5 layers, not sharply demarcated from lamina cells; costa single, stout, terete, 80-92 pm wide near the base, yellow-brown, gradually tapered, in transverse section consisting of incrassate, nearly homogeneous cells, disappearing below the apex or excurrent into the apex and confluent, frequently asymmetrically, with borders; lamina cells shortly oblong or linear­ rhomboidal to linear-flexuose, thick-walled or seldom thin-walled, (35- )45-85(- 95) pm long, (5.5- )7.0- 9.0(- 13.0) pm wide, unistratose, frequently with conspicuous longitudinal, bi-to polystratose streaks usually extending as spurs from the margins; cell s at the basal angles clearly differentiated in somewhat excavate, sizable groups extending nearly to the costa, hyaline or yellowish-brown, rather firm-walled, forming distinctly decurrent auricles. Inflorescences and sporophytes unknown. Lectotype species (selected here): Sciaromiopsis sinensis (Broth.) Broth., Sitzungsber. Ak. Wiss. Wien Math. Naturw. Kl. Abt. I, 133: 580. 1924 [= Sciaromium (Limbidium) sinense Broth., Sitzungsber. Ak. Wiss. Wien Math. Naturw. K I. Abt. I, 131 : 218. 1922]. Type: Prov. Setschwan: In montium Daliang-schan (territorio Lolo) ad orientem urbis Ningyiien regione temperata, in rivis utrinque in juga Dsiliba; ca. 2900- 3300m (Nr. 1542) (Holotype: " Handel-Mazzetti, Iter Sinense 1914-1918 su mptibus Academiae scientiarum Vindobonensis suspectum. Nr. 1542. Sciaromium sinense Broth. n. sp. = Sciaromiopsis sinensis (Broth.) Prov. Setschwan austro-occid.: In montium Daliang-schan (territorii Lolo) ad oerientem urbis Ningyiien regione temperata, in rivi utrinque in juga Dsiliba; all. S. m. ca. 2900-3300 m. Leg. 21. IV . 1914 Dr. Heinr. Frh. v. Handel-Mazzetti (Diar. Nr. 372)" - H-Broth !; isotypes: BP!, BR!, c l, F!, FH', FH- 322 Journ. Hattori Bot. Lab. No. 61 I 986

FIG. 5. Sciaromiopsis sinensis (Broth.) Broth. 1- 2: angular cells; 3- 4: leaf apices; 5: marginal cells in mid-leaf; 6: transverse section of the stem; 7: transverse sections of the leaves (1, 4,6 drawn from the holotype of Sciaromium sinense Broth., H-Broth; 2- 3, 5, 7 drawn from the holotype of Sciaromiopsis brevifolia Broth., H-Broth).

Fleisch!, KRAM! , L! , M' , MO !, NY! , PR !, TNS! , w!). Synonyms: SCiaromiopsis brevi/olia Broth., Sitzungsber. Ak. Wiss. Wien Math. Naturw. Kl. Abt. I, 133: 580. 1924. Type: Prov. Setschwan austro-occid.: In montis Lungdschu-schan prope urbem Huili regione temperata ad rupes diabasicas in rivis; ca. 3000 m (Nr. 926) [Holotype: "Handel-Mazzetti Iter Sinense 1914-1918 sumptibus Academiae scientiarum Vindobonensis suspectum. Nr. 926. Pseudolimbella laxirelis Broth. n. sp. Sciaromiopsis brevifolia Broth. Prov. Setschwan austro-occid.: In montis Lungdschu-schan R. OCHY RA: Sciaromiadelphus - relationship between extant and fossil mosses 323 prope urbem Huili regione temperata ad rupes in rivis, substr. melaphyrico, all. s. m. ca 3000 m. Leg. 25. Ill. 1914 Dr. Heinr. Frh. v. Handel-Mazzetti (Diar. Nr. 236)" - H-Broth!; isotypes: FH-Fleisch!, w!]. Sciaromiopsis acuminata Broth. n. nudo in sehed. Chirotype: " 1542. Sciaromiopsis acuminata Broth. n. sp . Setschwan, terril. Lolo, Daliang-schan, in rivis, 2900-3300 m, 4/ 1914, leg. H. v. Handel-Mazzetti, comm. Brotherus" - FH-Fleisch l , syn. novo Additional collection seen : China, Prov. Yunnan bor.-occid., trans vicum Gandhaidse ad urbem Lidjiang ("Likiang"), in regionis temperate lapidibus rivo irrigatis supra vic. Akalii, subsr. calceo, all. S. m. ca 3000 m, 19. VI. 19 15, leg. H. Frh. V. Handel-Mazzetti 6836 (Diar. Nr. 1217) (H-Broth, w). Sciaromiopsis is a distinct, monotypic genus endemic to Sichuan and Yunnan, China. It is distinguished by a unique combi nation of several structural character states, some of which are critical in determining its relationships. These are sum­ marized as fol1ows: (1) plants aquatic, prostrate, without paraphyl1 ia.and with foliose, suborbicular pseudoparaphyl1 ia; (2) leaves bordered with 7-12 rows of narrow, linear, thick-walled cells in 4-5 layers; (3) costa si ngle, strong below, usually confluent with thickened borders, seldom disappearing below the apex; (4) lamina cells li near­ rhomboid al to linear-flexuose, firm-walled, usually distinctly prorate, unistratose with frequent bi- to polystratose streaks extending as spurs from the borders; and (5) basal cells with a well-developed alar region forming decurrencies. From discussion of the taxa with limbate leaves presented earlier in this paper, it is evident that the only genus, which shares many important characters with Sciaromiopsis is Sciaromiella. These include unicostate, limbate leaves; elongate, linear-rhomboidal to linear-flexuose and prorate lamina cells; and foliose, suborbicular pseudoparaphyllia. In addition, the plants are aquatic, rather stiff, freely and irregularly branched, and the leaves are of comparable size and shape. However, leaf borders in Sciaromiella are bistratose, whilst those in SCiaromiopsis are 4-5-stratose; the leaf margins are sharply serrate above to remotely serrulate below in Sciaromiopsis, whereas they are entire to slightly distantly serrulate in Sciaromiella; basal cells form sizable, pellucid, decurrent auricles in Sciaromiopsis, while they are not auriculate and are non-decurrent in Sciaromiella.

THE FAMILIAL PLACEMENT OF SCIAROMIELLA AND SCIAROMIOPSIS The familial placement of Sciaromiella and SCiaromiopsis is difficult owing to the sterile condition of these plants. Therefore, it is only possible to speculate on their relationships. It is hard to deny that the principles of classification of higher taxa of mosses should be based primarily upon peristome morphology and many recent studies in different groups of mosses strongly support this view (Buck 1980a, b, 1981 ; Buck & Crum 1978; Vitt 1984; Buck & Vitt 1986). However, it cannot be neglected that in some groups of mosses, for instance in H ypnales (= H ypnobryales), most of the taxa are characterized by having perfect hypnaceous peristomes, which do not otTer good characters useful in phylogenetic considerations. It is rather unquestionable that the evolution of hypnobryalean mosses has led to diversification of gametophytes, whereas sporophytes, and especially peristomes, have remained similar, with only slight modifications. Hence, most genera and families within the Hypnales are established mainly on the basis of a combination of gametophyte characters. This 324 lourn. Hattori Bot. Lab. No. 61 1 986 principle is highly justified with the only one reservation. It is necessary to recognize characters of minor taxonomic value, sometimes of evidently adaptative nature, from those that are significant and fundamental in all phylogenetic speculations. Unfortunately, the strategy of earlier bryologists dealing with higher taxa of pleu­ rocarps has been to emphasize the value of superficial similarities of gametophytes in grouping mosses in families and genera. This approach has led to the production of artificial assemblages, including taxa of remote phylogenetic affinity but externally similar due to convergence exhibited by plants growing in identical habitats. There are much evidence to support this point-of-view, e.g. Entodonaceae (Buck 1980a), Plagiotheciaceae (Buck & Ireland 1985), Sematophyllaceae (Seki 1968) and (Nishimura et al. 1984). Species that are currently included in Sciaromiella and Sciaromiopsis have been invariably placed in the Amblystegiaceae (Brotherus 1922, 1924, 1925, 1929; Crum & Steere 1958; Abramova & Abramov 1967). The concept of this family dates back to the end of the last century when Roth (1899) segregated it from the large and all­ encompassing Hypnaceae. Originally, Amblystegiaceae included only genera characte­ rized by having unicostate leaves, rhombic to linear-flexuose, smooth lamina cells, smooth setae and arcuate capsules strongly contracted below the mouth with perfect hypnaceous peristomes, viz. Amblystegium B., S. & G., Cratoneuron (S ull .) Spruce, Drepanocladus (c. Muell.) G. Roth, Calliergon (Sull.) Kindb. and Campylium (Sull.) Mitt. (only a part of the species presently known as Campyliadelphus (Kindb.) Kanda). In Roth's understanding, Amblystegiaceae were a consistent grouping of genera, excepting perhaps Cratoneuron. However, it seems that this author considered paraphyllia as taxonomically unessential character. The concept of Amblystegiaceae that is now in common use dates from Brotherus (1908, 1925) and Fleischer (1915- 1922). Unfortunately, modifications and additions made by these and many subsequent authors have considerably garbled the original idea of Roth. The Amblystegiaceae have become rather an ecological assemblage of a variety of taxa mostly related to moist habitats. Hence, a re-interpretation of this family in terms of modern moss taxonomy is needed. The Amblystegiaceae, as I understand them, are defined by the following set of characters: (1) unicostate perichaetial and vegetative leaves (in the latter midribs may be sometimes spurred or forked above but never double or none); (2) lack of paraphyllia; (3) foliose pseudoparaphyllia; (4) unistratose, smooth, rhombic to linear lamina cells; (5) smooth setae; (6) cylindric to oblong-cylindric, arcuate capsules strongly constricted below the orifice; (7) perfect hypnaceous peristome with yellow­ brown exostome teeth and 1- 3 cilia. If the above definition of this family is accepted, then many genera and species that are presently there are anomalous in it and these should be shifted to other families of hypnobryalean mosses or even placed in families of their own. This re­ fers especially to three groups of taxa: (1) bi- or ecostate taxa (Scorpidium, Ortholimnobium, Calliergonella, Pseudohygrohypnum, some species of Campylium and Hygrohypnum); (2) taxa having paraphyllia (Cratoneuron, Cratoneuropsis, Sasaokaea, R. OCHYRA: Sciaromiadelphus. - relationship between extant and fossil mosses 325

Platylome/la, Crassicosta); and (3) taxa with variously polystratose leaf laminae and papillose lamina cells (Platylomella, Donrichardsia, Sciaromiopsis, Sciaromiella, Sciaromium ). The problem of the familial pl acement of some taxa of the latter group has been discussed in length elsewhere, including Donrichardsia (Ochyra 1985b), Sciaromium (Ochyra 1985a, I 986a, 1987a, b, c) and Platylomella (Ochyra 1987d). Discussion presented below is focused on relationships and affinities of Sciaromiella and Sciaromiopsis. These genera appear to be the closest relatives of species of the Donrichardsiaceae. This family was segregated from the Amblystegiaceae for Donrichardsia macroneuron (Grout) Crum & Anderson, a monotypic, endemic genus from Texas (Ochyra 1985b). Later, I added to this family Richardsiopsis lacustris (Herz. & Rich. in Rich.) Ochyra, a monotypic genus from Peru (Ochyra 1986a). This family is circumscribed as including rheophytes with strong, single costae; elongate, linear-flexuose, distinctly prorate lamina cells that are variously polystratose; a lack of paraphyllia; and presence of foliose, suborbicular pseudoparaphyllia. The same set of characteristics is also observed in Sciaromiella and Sciaromiopsis. Multistratosity of leaf laminae is variously expressed in those four genera. In Donrichardsia, there exist polystratose streaks extending as spurs from the costa or being independent; sometimes they are situeated near the leaf margins giving the leaves an appearance as those found in taxa with perfectly limbate leaves. In Richardsiopsis, bistratose streaks extend also as spurs from the costa or are independent, and the lamina is frequently totally bistratose in the upper part. In addition, the marginal cells of the leaves have a tendency to be narrower and somewhat swollen, and give the impression of a hint of a border. What has not been accomplished in either Donrichardsia or Richardsiopsis appears to become a reality in both Sciaromiella and Sciaromiopsis. Both of the latter have perfectly developed leaf borders and, additionally, leaf laminae are variously poly­ stratose by spurs extending from the border and costa. Thus, the most reasonable solution, until sporophytes are found in those four genera, is the placement of them in the Donrichardsiaceae, since the combination of their gametophytic characters is unknown in any other family of pleurocarps. Because these genera form two dis­ tinct complexes, well diagnosed by the presence or the lack of leaf borders, I pro­ pose to include Sciaromiella and Sciaromiopsis into a subfamily of their own, Sciaromiopsoideae, and to retain Donrichardsia and Richardsiopsis in the type subfamily, Donrichardsioideae. Donrichardsiaceae Ochyra subfamilia Sciaromiopsoideae Ochyra, subfam. novo Plantae aquaticae, sat robustae, rigidae, paraphylliis nullis, pseudoparaphylliis foliosis et sub­ orbicularibus, laminis ubique multistratose limbatis et radiis multistratosis frequenter praeditis, cellulis oblongo-hexagonalis vel lineari-flexuosis et papillosis et costis 80--1 IO I'm latis iam dignoscenda. Typus: Sciaromiopsis Broth. As circumscribed herein, Sciaromiopsoideae are pleurocarpous rheophilous mosses that are characterized by plants that are mostly rigid; lack of paraphyllia; having foliose, sub­ orbicular, variously serrulate or crenulate pseudoparaphyllia; leaves that have a single costa, 326 Journ. Hattori Bot. Lab. No. 61 1 9 8 6

80--110 pm wide and are bordered with 5~12 rows of linear, thick-walled cells arranged in 2- 5 layers and usually confluent with the costa; leaf cells that are distinctly prorate, oblo.ng­ hexagonal to linear-flexuose; and basal cells that are differentiated in sizable, decurrent auricles or only slightly differentiated from the upper lamina cells. Relationships of the Donrichardsiaceae are rather obscure, mostly due to the sterile condition of the plants. If we assume that they have the standard hypnoid peristome (what is highly probable), than we must seek their relatives among families grouped in the superfamily Brachytheciacanae as defined by Buck and Vitt (1986). Prorate and firm-walled lamina cells as well as rigid texture of the plants are characteristic of all taxa included in this family and suggest its affinity to the Rhytidiaceae. On the other hand, there is a strong similarity of the Donrichardsiaceae to the Amblystegiaceae. However, it is hard to accept a view that such apomorphic character states as prorulose and thick-walled lamina cells may have evolved from rather thin-walled and smooth lamina cells. Judging from the gametophyte characters it seems to be more reasonable to seek relatives of the Donrichardsiaceae in the Stereophyllaceae-Fabroniaceae complex of families. The Stereophyllaceae were recently circumscribed by Buck and Ireland (1985) to accommodate six genera centered around Stereophyllum. The family is recognized by unicostate leaves and by large groups of quadrate to rectangular alar cells as well as by perfect hypnalean peristome and by the presence of filamentous or less commonly foliose pseudoparaphyllia. Leaf areolation of many species of this family, especially Entodontopsis, luratzkea and Eulacophyllum, resemble very much that of species of Donrichardsiaceae. Lamina cells in these genera are linear to rhomboidal, thin- or thick-walled and frequently dorsally prorulose. Moreover, in some species of Entodontopsis and Stenocarpidium, there is a strong tendency fo r narrowing marginal cells of the lamina that form an indistinct border like that found in Richardsiopsis lacustris. It appears that perfectly limbate leaves occur sometimes in the Stereophyllaceae-Fabroniaceae complex. Unistratose borders are known in Anacamptodon marginatus (Dixon) Buck from South Africa, and bistratose margins occur in A.fortunei Mitt. of China and Japan (Buck 1980b). The Fabroniaceae are an anomalous family, having many very specialized features, but Buck and Vitt (1986) consider it as a derivative of the Stereophyllaceae. Considering the great similarity of vegetative features, especially leaf areolation, it can be suggested that the Donrichardsiaceae and Stereophyllaceae are closely related and probably have evolved from common ancestors. Contrary to the Stereophyllaceae that occupy dry, terrestrial environmental niches, Donrichardsiaceae have evolved in wet habitats and therefore exhibit typical features of rheophytes. This is expressed in different shape of the basal cells of the leaves. They are either weakly differentiated (Donrichardsia, Sciaromiella bartlettii) or form a distinct band of larger cells (Sciaromiella longifolia, Richardsiopsis), and in the extreme case of Sciaromiopsis form distinct, sizable, pellucid auricles that are typical of numerous, unrelated rheophytes. Species currently included in the Donrichardsiaceae may be keyed as follows: I. Leaves elimbate or indistinctly bordered by unistratose, linear, narrow cells (subfamily R. OCHYRA: Sciaromiadelphus - relationship between extant and fossil mosses 327

Donrichardsioideae) ...... 2 2. Leaves oblong-ovate, bluntly acute to obtuse or rounded-obtuse; margins se rrulate above; costa 160- 200).lm wide, su bpercurrent to short excurrent; lamina cells wit h numerous polystratose strands ...... Donrichardsia macroneuron 2. Leaves narrowly lanceolate, tapered into a long acumen; margins entire; costa (80-)IOO- 130(- 140)/lm wide, long excurrent; lamina cells irr~gu l arly bistratose below the excurrency of the costa with longitudinal, bi stratose strands below ...... Richardsiopsis lacustris I. Leaves bordered by 5- 12 rows of narrow, linear cells arranged in 2-5 layers (subfamily Sciaromiopsoideae) ...... 3 3. Borders of lamina 4-5-stratose; leaves sharply serrate above, serrulate to subentire below; basal lamina cells enlarged, pellucid forming sizable, decurrent auricles ...... Sciaromiopsis sinensis 3. Borders of lamina bistratose throughout; margins entire or sinuate to remotely, minutely serrulate above; basal cells larger from the lamina cells, opaque, forming non- decurrent band (Sciaromie/la) ...... 4 4. Leaves gradually short acuminate, up to 2.1 mm long; lamina cells 60-110 llm long Iinear-flexuose throughout; costa 95- 110 /lm wide...... Sciaromiella bartlettii 4. Leaves gradually long acuminate, (3.0- )3 .5- 4.0(- 5.0) mm long; lamina cells oblong-hexagonal above to Iinear-flexuose below, (26-)40- 55(- 60)/lm long; costa 70-80/lm wide ...... Sciaromiella longifolia

ON THE GEOGRAPHICAL DISTRIBUTION OF THE DONRICHARDSIACEAE Donrichardsiaceae seems to be a relictual group of mosses, since particular species of this family have very restricted geographical ranges (Fig. 6). Extinction of Sciaromiella longifolia clearly confirms the retreat of this highly specialized moss family. The lack of any modes of reproduction, neither sexual nor asexual doubtless affects this situation. However, this is not exceptional among rheophilous mosses, many of which exhibits simi lar reproductive performances (Ochyra 1985a). Donrichardsia macroneuron is endemic to the Edwards Plateau in Texas (Crum & Anderson 1979; Wyatt & Stone burner 1980). Richardsiopsis lacustris is restricted to two lakes in Peru (Richards 1984), whereas Sciaromiopsis sinensis has a very limited geographical range in southwestern China (Sichuan, Yunnan), an area that is particularly rich in monotypic and stenotypic moss genera (Ochyra 1986b). Similarly, Sciaromiella bartlettii is known only from two stations from Haiti (Crum & Steere, 1958; Steere 1985). As I was kindly informed by William R. Buck (in lilt. 18 December 1985) this species had not been recollected at the type locality. The distribution of the Donrichardsiaceae, and especially Sciaromiella, is anom­ alous and difficult to categorize into floristic groups. Taking into account the range of the only living representative of Sciaromiella, S. bartlettii, it should be considered as an endemic genus to the West Indies. The moss flora of this area, especially of the Greater Antilles, is relatively rich in endemic species (Crosby 1969), but at the genus level endemism is very low. Apart from Sciaromiella and Teniolophora, there are no strict endemic genera of mosses, though some of them, fo r instance Pilotrichidium or W IV 00

.... o c :3 :t ~ 5 ::l. o:l ~ r Ol ".. Z ? a-

FrG. 6. Total range of the Donrichardsiaceae. I: Donrichardsia macro neuron (Grout) Crum & Anderson; 2: Richardsiopsis lacustris (Herz. & Rich. in Rich.) Ochyra; 3: Sciaromiella bart/ell;; (Crum & Steere) Ochyra; 4: S. longijolia (A. Abr. & I. Abr.) Ochyra; 5: Sciaromiopsis sinensis (Broth.) Broth.

>0 00 a- R. OCHYRA: Sciaromiadeiplius - relationship between extant and fossil mosses 329

Eucamptodontopsis, appear to have here the main occurrence centre. On the other hand, the moss flora of the West I ndies has strong continental affiliations (S teere 1984, 198 5). The problem of the geological hi story of the West lndies is still debatable and unsolved. If we accept an idea that the Greater Antilles are of volcanic origin and not truly continental (Khudoley & Meyerhoff 1971), then an assumption can be advanced that S. bartlettii migrated to Hispaniola from the ancient plateaux of Central America via long range dispersal. On the other hand, Schuchert (1935, 1955) postulates that during the Eocene time there was a land connection between the West Indies and Central America. Thus, plant migration could have occurred from Central American plateaux to the West Indies before the permanent separation, probably in the Miocene, and certainly in the Pli ocene, time. Acceptance of this view implies that the present occurrence of S. bartlettii in Hispaniola may be interpreted as a fragment of presumably broader original range. Sciaromiella probably represents an ancient Laurasian taxon and its geographical range reflects the geological and climatic history of the supercontinent Laurasia in Tertiary and Pleistocene times. Extensive glaciations that occurred on this vast land mass by Oligocene time or late in the Tertiary, and especially in the Pleistocene time, caused mass extermination of Bryophyta taxa or at least their regional obliteration or reduction to relictual status (Dickson 1973; Schuster 1983; Miller 1984). As a result, a considerable number of bryophytes that had been distributed more or less con­ tinuously in the Arcto-Tertiary region show presently highly disjunct distribution patterns. This is especially true for western Eurasia, where such west-east barriers as the Alps, the Carpathians, the Caucasus and the Tian-Shan Mountains exist. These precluded southward migration of species and in consequence the possibility of their survival in more southerly refugia as in North America or eastern Asia where there predominate meridional mountain chains that are conducive for such migrants. Therefore, the present bryophyte flora of western Eurasia is very impoverished in comparison to that of eastern Asia or North America. The case of S. longifolia is one of many examples, which support this situation. Other examples include Platylomella lescurii (Sull .) Andrews, presently endemic to eastern North America on the ancient Appalachian Plateau, but its close relative (or identical taxon?) was descri bed from the Pliocene deposits of Bashkiria as Sciaromium laxirete A. Abr. & I. Abr. (Ochyra 1987d); Andoa berthelotiana (Mont.) Ochyra, an endemic genus from Macaronesia, was found in a rich Neogene assemblage of mosses from Duab at the western end of the Caucasus (Abramova & Abramov 1959). From the same deposits also was described Echinodium savicziae A. Abr. & I. Abr., a species of the genus that presently exhibits an unusual Australasian-Macaronesian distri­ bution (Churchill 1986). The general assumption is that the entire Tertiary was a period of progressive regional extinction of taxa of the Donrichardsi aceae. These were probably never an important constituent of the flora of the Arcto-Tertiary Region . Decimated remnants of this family have survived presently in southerly refugia in southwestern China (Sciaromiopsis) or on ancient plateaux in Central America (Donrichardsia, 330 Journ. Hattori Bot. Lab. No. 61 I 9 8 6

Sciaromiella), and Richardsiopsis might have migrated along the Cordilliera to Peruvian Andes. Donrichardsiaceae, in the Northern Hemisphere, exhibits a classic example of the eastern Asian-eastern North American distribution pattern. I watsuki and Sharp (1967) listed more than 30 moss species showing this type of range. Further examples, parallel to the Sciaromiella case, are some species of Drummonidia (Vitt 1972) and a pair of species of Astomiopsis, namely A . sinensis Broth. and A . radiculosa (Herz.) Buck & Zand. (Buck 1979; Buck & Zander 1980).

ACKNOWLEDGMENTS. I wish to thank the Directors and Keepers of the following herbaria for kindly allowing me to borrow specimens during the course of this work without which this study would not have been possible: BP, BR, C, CAN M, F, FH, H, L, M, MAK, MICH, MO, NY , PR, S, TNS and w. Special thanks are due to Drs A. L. Abramova and I. I. Abramov for their kindness when I visited the Komarov Botanical Institute in Leningrad, Dr H. Crum for making me a present of the type material of Sciaromium bartlellii, and to D r William R. Buck for information on his attempt to recollect this species on Haiti. I a lso greatly appreciate suggestions of Drs Dale H. Vitt and Paul L. Redfearn, J r. for improving the manuscript. I owe a particular word of gratitude to Dr Ridef Grolle for his important aid in nomenclatural questions and to Dr Hisatsugu Ando for his kind translation of Dr Noguchi's (1978) paper. At last, but not least of all, I am very grateful to Dr T . Tacik for his kind correction of my Latin and to my wife, Halina, for her illustrations.

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