Altitudinal Zonation of Bryophytes on the Huon Peninsula, Papua New Guinea. a Floristic Approach, with Phytogeogra- Phic Considerations

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61 Tropical Bryology 2: 61-90, 1990

Altitudinal zonation of Bryophytes on the Huon Peninsula, Papua New Guinea. A floristic approach, with phytogeographic considerations

Johannes Enroth Department of Botany, University of Helsinki, Unioninkatu 44, SF-00170, Helsinki, Finland.

Abstract. The study is based on the major part of the bryophyte material collected during the Koponen-Norris expedition on the Huon Peninsula, Papua New Guinea, in 1981. Only taxa which were collected at least twice are included. Five altitudinal zones, the boundaries of which are indicated by discontinuities in the bryophyte flora, are distinguished: 0 - 300 m, 300 - 1200 m, 1200 - 2200(-2300) m, 2200(-2300) - 2800(- 2900) m, and 2800(-2900) -3400 m. These zones, each characterized by a typical species assemblage, are well in accordance with some earlier New Guinean zonation schemes based on the phanerogamic flora and vegetation. The most obvious correlations between bryophytes’ altitudinal ranges on the Huon Peninsula and their general phytogeography are: New Guinean or Western Melanesian endemics, as well as Malesian endemics, are concentrated at relatively high altitudes (zones III-V); Asian - Oceanian and Asian - Oceanian - Australian taxa, notably mosses, are relatively strongly represented at low to moderate altitudes (zones I-III); species which have their main distribution in the northern hemisphere occur at high altitudes; 'cosmopolitan' species either have wide vertical ranges or are restricted to high altitudes.

Introduction Nadkarni (1978), and Johns (1982). Robbins (1959) and Royen (1980) mainly The altitudinal zonation of vegetation in dealt with the New Guinean mountain the tropics is distinct and, in many regions, vegetation. The treatment of Australasian rather well documented (see, for example, alpine bryophytes by Ramsay et al. (1986) Richards 1957: 346-368). The zones are must be mentioned in this connection, mainly distinguished by changes in the since it includes a list of the alpine mosses phanerogam flora and the gross structure of New Guinea. Two studies by Seifriz of the vegetation relative to altitude. Often (1923, 1924) on Mt. Gedeh, Java, have a the zone characterizations also include bearing on the present study. In the former general observations on the abundance work he defined five vegetational and species- richness of lower crypto- 'subzones' above 4600 ft (1400 m) based gams, notably epiphytic ones, at different on phanerogams, and in the latter reported altitudes. Such general accounts of the that floristic and quantitative differences zonation of the vegetation in New Guinea regarding mosses and lichens at different have been presented by Lane-Poole altitudes are also very profound. Despite (1925), Paijmans (1976), Gressitt & certain lack of profoundness in Seifriz’s 62 latter study - he was neither a bryologist Santa Marta, Colombia. Based on relevée nor a lichenologist - it clearly demonstrates analysis concerning species presence, that the altitudinal zonation of lower substrate preference, and percentage cover cryptogams closely follows that of higher of bryophytes, they distinguished five plants and vegetation, and it can be altitudinal bryophyte zones between 500- considered a pioneer work in this respect. 4100 m a.s.l. Their results are discussed in more detail in the Results and Discussion It is currently a well-known fact that the section below. Gradstein & Frahm (1987) bryophyte flora and vegetation in tropical presented a floristic study on the altitudinal lowland forests are poor, become richer zonation of mosses in NE-Peru and came with increasing altitude and reach their up with 4 main zones two of which they culmination in the so- called mossy forests further subdivided. Their results will be - the terms cloud forests and mist forests considered further below. have also been employed - which occur in the lower and upper montane zones (e.g. The present study surveys the altitudinal Richards 1984). The reasons for the zonation of bryophytes on the Huon bryological poverty of tropical lowland Peninsula, Papua New Guinea, mainly on forests have been subject to some debate a floristic basis. The reasons for such an and disagreement, but it was quite approach to the problem are given below convincingly pointed out by Frahm (1987), (see Materials and Methods). Some through measurements in the field and observed correlations and more or less laboratory simulations, that the well-defined regularities between the combination of high temperature and low geographical distribution patterns of bryo- light intensity is responsible for the phytes and their altitudinal ranges on the phenomenon, since it does not allow Huon Peninsula are considered. sufficient net assimilation for bryophyte growth. With increasing altitude the mean Materials and Methods temperature falls by about 0.4-0.7°C/100 m (Richards 1984) - lapse rates between This study is based on the bryophytes 0.53-0.60°C have been given for the New (ca. 17500 specimens) collected by Guinea mountains (Barry 1980) -, the tree Prof. Timo Koponen (Helsinki) and Prof. canopy becomes lower and more open, Daniel H. Norris (Arcata) on the Huon allowing more light to penetrate to the Peninsula, Papua New Guinea, in 1981. forest interior, and the occurrence of clouds The background and methods of the and mist become more frequent or even expedition, and introduction to the study continuous. These factors together allow area and its bryological exploration were a more luxuriant bryophyte vegetation to presented by Koponen & Norris (1983a). develop. At present, 31 papers under the title Bryophyte flora of the Huon Peninsula, Although numerous workers, in revisions Papua New Guinea have been published of tropical taxa and in local or regional in Acta Botanica Fennica and Annales bryophyte floras, have given more or less Botanici Fennici. Despite the title of these detailed information regarding the papers, the bryophyte floras of Western altitudinal ranges of bryophytes, Melanesia (i.e. West Irian, Papua New comprehensive and thorough studies on Guinea, and the Solomon Islands) are also the altitudinal zonation of bryophytes in treated. the paleotropical region are lacking. However, an elaborate study on the subject On the Huon Peninsula 55 moss families was carried out by Reenen & Gradstein (Koponen pers. comm.) and 39 hepatic (1983, 1984) on the Sierra Nevada de families (Grolle & Piippo 1984b) are

66 bryologically poorer, viz. in lowland bryophyte taxa along the altitudinal forests on the one hand and above ca. gradient. In general terms, the 3000 meters on the other (cf. Fig. 1). diversity increases from sea level and is highest at 2200-2300 m in The primary working hypothesis used in the mossy forests. This implies that this study is that if there is an altitudinal below approximately 2200 m the zonation (instead of a complete continuum) lower limits of taxa are more of bryophytes on the Huon Peninsula, the numerous, and apparently also limits of the zones will be indicated by more clearly concentrated at floristic discontinuities (cf. Gradstein & particular altitudes, than the upper Frahm 1987). Accordingly, the altitudinal ones, whereas above that altitude limits of the included taxa are determined. the gradual impoverishment of the The numbers of these limits along the bryoflora implies a higher amount altitudinal gradient are graphed separately of upper limits (Figs. 2- 4). It for hepatics and mosses, and for the groups logically follows from this, that at together. In these graphs, the altitudinal lower altitudes the boundaries of boundaries of the taxa have been rounded the zones must mainly be based on off to the nearest even hundred; if the the lower limits of individual taxa, lower limit of a taxon was at, say, 650 m, but higher up - above 2200 m, that it has been rounded off to 600 m, and if the is - mainly upper limits should be upper limit of a taxon was at, say, 3250 m, used in the definition of the zones. it has been rounded off to 3300 m. The total number of included taxa along the The altitudinal limits of individual altitudinal gradient are calculated and also hepatic taxa are most strongly graphically displayed. The phytogeogra- concentrated at 300, 800, 1200, phical concepts, employed in the compa- 1500, 1800, 2200 (- 2300), 2500, rison of bryophytes’ altitudinal ranges on 2800, and 3400 m (Fig. 2), and the Huon Peninsula with their total those of individual moss taxa at geographical distributions, follow 300, 700, 1200, 1500, 1800, 2100 Hyvönen (1989a). (-2200), 2500, and 2900 m (Fig. 3). From Fig. 4 we observe that, when the data for both groups are Results and Discussion combined, the highest numbers of altitudinal limits occur at 300, (800), A. Altitudinal zones and their 1200, 1500, 1800, (2200-2300), boundaries on the Huon 2500, 2800-2900, and 3400 m. Peninsula Although the upper limits of hepatics are still more numerous at The altitudinal ranges of all taxa 3500 than at 3400 m (Fig. 2), the used in this study are presented in relevance of this is doubtful (see Table 10. The numbers of lower Materials and Methods section and upper altitudinal limits of above). hepatic and moss taxa at different altitudes on the Huon Peninsula It is obvious that some of the peaks are graphed in Figs. 2 and 3, in Figs. 2-4 indicating high respectively. The numbers of these concentrations of altitudinal limits limits for the combined groups are are artificial, caused by higher graphed in Fig. 4. intensity of collecting at some altitudes than at others. For Fig. 5 presents all of the included example, at 1700-1800 and 2400- 67 2500 m collecting was pursued at plains and fans, and typical tropical 16 sites, at 1500-1600 and 2100- rain forests (with several canopy 2200 m at 14 sites. On the other layers and buttressed trees, hand, at 1600-1700 and 1900- abundant lianas and vascular 2000 m collecting was less epiphytes). On the Huon Peninsula intensive, and both of these altitudes extensive alluvial plains and fans show low numbers limits. It is not are restricted to a narrow belt in the easy to determine the effect on the western part of the southern coast present study of varying collecting (around Lae), the remaining coast intensities. However, it seems rea- being mostly formed of foothills, sonable to hypothesize that it has a low mountains and a belt of grass- more pronounced effect in the land along the northern coast environments which have a rich (Paijmans 1975). flora, such as the mossy forests, and a weaker effect in floristically During the Koponen-Norris poorer habitats, such as tropical expedition, bryophyte collecting lowland forests. Based on this was not very intensive below 300 assumption and on the previous m (cf. Fig. 1), but several different literature, I reject as potential zone environments at these altitudes were boundaries the altitudes at which visited (Koponen & Norris 1983a). the the high concentrations of limits As might be expected, the of individual taxa appear to be bryophyte flora is poor, especially most artificial, namely, 1500, 1800, in hepatics (Fig. 5). In the present and 2500 m. material, only Calymperes tenerum, Leucophanes glaucum, and Pin- Because hepatics and mosses in the natella mariei are resticted to below present data show a similar pattern 300 m. Another species which of altitudinal zonation, there is no obviously occurs only in this zone reason to treat them separately. Of is the swamp-dweller Exodictyon course, this does not mean that incrassatum (Ellis 1985, Enroth some zones could not be partly 1990). Mosses which have their characterized by a richer hepatic lower limit at or near sea level flora or the abundance of epiphytic include, some Calymperaceae mosses, but simply that it would be (Calymperes dozyanum, C. pointless to distinguish separate, taitense, Mitthyridium obtusifo- independent zones for hepatics and lium), Leucobryaceae (Exostratum mosses. blumei, Octoblepharum albidum), Neckeraceae (Neckeropsis ZONE I, 0 - 300 m. gracilenta, Pinnatella kuehliana, P. mucronata, P. nana), Pottiaceae This zone is equivalent to Lane- (Barbula arcuata, B. indica, B. Poole’s (1925) 'Lowland forests' javanica, B. subcomosa, Hyophila (see also Richards 1957: 349), and involuta, Oxystegus crassicostatus) it also has exactly the same upper and Thuidiaceae (Pelekium boundary. This lowland area bifarium, P. velatum, Thuidium embraces several different plumulosum). The most strongly environments. These include represented hepatic family is the mangroves, beach ridges and flats, Radulaceae, with seven species swamps (saline and brackish as (Radula anceps, R. apiculata, R. well as fresh water ones), alluvial javanica, R. koponenii, R. nymanii,

70 R. tjibodensis, R. vrieseana). In than in zone I, and becomes addition, there occur few gradually more so with increased Frullaniaceae (Frullania apiculata, altitude. At the lower boundary of F. ericoides, F. multilacera) the zone, the moss flora, suddenly Geocalycaceae (Heteroscyphus becomes more diverse: 17 species argutus, H. coalitus), and have their lower limits at 300 m. As Dumortiera hirsuta of the examples may be mentioned some Wiesnerellaceae. Neckeraceae (Himantocladium plumula, Homaliodendron flabel- ZONE II, 300 - 1200 m. latum, Neckeropsis lepineana) and Pterobryaceae (Calyptothecium I have accepted 1200 m as the recurvulum, C. urvilleanum, demarcation altitude between zones Garovaglia angustifolia, G. II and III, because its high number elegans, G. powellii). Mosses also of lower limits of individual bryo- remain more numerous than hepa- phyte taxa is not caused by high tics throughout the zone. A distinct collecting intensity. Instead, feature of this zone is the strong collecting at 1500 m was rather representation of the intensive, and the peaks in Figs. 2- Calymperaceae. Species such as 4 at that altitude are probably Calymperes afzelii, C. strictifolium, artifacts. However, it should be Syrrhopodon muelleri, and noted that Lane-Poole (1925) Mitthyridium fasciculatum are placed the upper limit of his 'Foothill exclusively, or almost so, restricted forests' - which I consider roughly zone II. Other taxa not found equivalent to my zone II - at 4500- outside zone II are Plagiochila lon- 5500 ft (approximately 1350-1650 gispica, Radula multiflora var. m). reflexilobula, Fissidens obscurirete, Aerobryidium crispifolium, Lane-Poole’s (1925) account of Aerobryopsis leptostigmata, and the 'Foothill forests' gives the Himantocladium plumula. following general characters: The forests are not as tall as, and are less ZONE III, 1200 - 2200(-2300) hygrophytic than, the lowland rain m. forests, which may be due to the lack of retention of moisture on the The lower limit of zone III is marked steeper slopes; buttressed trees are by a relatively strong increase of fewer; lianas and vascular bryophyte taxa, notably hepatics. epiphytes, notably ferns in the Several Frullaniaceae (Frullania crowns of trees, are clearly less carrii, F. curvistipula, F. epiphylla, abundant; the ground-cover is F. junghuhniana, F. prominula, F. mainly composed of ferns and subnigricaulis var. subtruncata) Elatostema spp. (Urticaceae); the and Plagiochilaceae (Plagiochila characteristic tree is Quercus cymata, P. loriana, P. odorata, P. junghuhnii; conifers (Podocarpus, tamiensis, P. trapezoidea, P. velata) Araucaria) are scarce, but become do not occur below this altitude. In somewhat more frequent towards addition to these, 12 species, be- the upper boundary. longing to the families Geocalycaceae, Jungermanniaceae, As can be seen from Fig. 5, the Lepidoziaceae, Lophoziaceae, bryophyte flora in zone II is richer Pleuroziaceae, Schistochilaceae, 71 and Trichocoleaceae have their altitudinal zone. The topography, lower boundaries here. Of the exposure of the slopes, prevailing mosses which appear in the flora at wind direction, among other things 1200 m (13 taxa) may be mentioned determine the terms for their Campylopodium medium, presence. Papillaria fuscescens, Pinnatella anacamptolepis, Pogonatum According to Lane-Poole (1925), neesii, Elmeriobryum the “Midmountain forests” are philippinensis, and Thuidium dominated by species of Quercus contortulum. Hepatics more or less and several conifers, such as restricted to zone III include Araucaria cunninghamii, Tylimanthus saccatus, several Podocarpus spp. and Phyllocla- Frullaniaceae, Heteroscyphus dus spp. As a whole, there are fewer wettsteinii, Jungermannia tree species than at the lower hasskarliana, Notoscyphus lutes- altitudes, and buttressed trees are cens, Psiloclada clandestina, entirely absent; additionally, “the Telaranea kaindina, T. trisetosa, ground is decidedly mossy, and Porella grollei, and P. viridissima. the trunks and branches carry moss Mosses not encounteterd outside and lichen in crannies and nooks.” this zone are Syrrhopodon (Lane- Poole 1925:221). japonicus, S. tristichus, Brotherobryum undulatifolium, In general bryological terms zone Campylopodium medium, III may be characterized by a gain Microdus miquelianus, Fabronia in hepatic diversity and abundance curvirostris, Himantocladium (Frullaniaceae, Plagiochilaceae, submontanum, Pinnatella Lepidoziaceae, Schistochilaceae), anacamptolepis, Pogonatum and, near the upper limit of the neesii, and Pyrrhobryum zone, hepatics outnumber mosses. latifolium. A few large moss families, such as the Bryaceae, Dicranaceae, The upper limit of zone III coincides Fissidentaceae, and Meteoriaceae, closely with that of Lane- Poole’s also become more frequent and (1925) “Midmountain forests” abundant. It is noteworthy that the (7500 ft, approximately 2300 m). Calymperaceae are very scarce in At 2200 m both hepatics and mosses the upper parts of the zone. Of the reach their peak, in the sense that species included in this study, only the number of taxa is higher there Syrrhopodon prolifer was found than at any other altitude. Accor- as high as 2400 m. The more ding to Koponen & Norris (1983a), favorable temperature/light inten- mossy forests occur between 1700- sity ratio, and the locally continuous 3300 m on the Huon Peninsula, dampness and occurrence of mist and it is not suprising that the enhance the growth of a luxuriant culmination point of the bryophyte and rich bryophyte vegetation. flora and vegetation lies within these forests. It should be remem- bered, however, that since the de- ZONE IV, 2200(-2300) - 2800(- velopment of mossy forests is 2900) m. dependent on the more or less continuous occurrence of mist they As Figs. 4 and 5 demonstrate, 2200- do not form a distinct and uniform 2300 m is the altitude where the 72 number of bryophyte taxa begins Meteoriaceae are very scarce. The to decline; at 2300 m the number of Thuidiaceae become scarce towards the upper limits of individual taxa, the upper boundary. Most notably hepatics (Fig. 2), for the remarkably, Fissidens spp. do not first time clearly exceeds the number occur at all above the upper of the lower limits. It may be boundary of zone IV, and only supposed that at this altitude some Fissidens plagiochiloides, F. physical factor, or a combination strictulus and F. taxifolius reach of physical factors, becomes 2900 m. Mosses restricted to zone unfavorable for many bryophytes IV include Leptobryum pyriforme, which thrive lower down on the Braunfelsia enervis, mountains. The most obvious Dicranodontium fleischerianum, physical factor to restrict the growth Fissidens plagiochiloides, of bryophytes at these moderately Dawsonia beccarii, and Pogona- high altitudes is the lower average tum subtortile. It must also be temperature. mentioned that the monotypic Orthothuidium (O. curtisetum) As mentioned previously, Koponen occurs between 2100-2900 m, and & Norris (1983a) give an altitudinal is therefore nearly restricted to this range of 1700-3300 m for the zone. mossy forests on the Huon Peninsula, and zone IV is As regards hepatics, the completely within this range. Frullaniaceae, Geocalycaceae, However, the environments of the Lepidoziaceae, and Plagiochilaceae collecting sites of the Koponen- are well represented. The Norris expedition between 2200- Radulaceae become scanty, 2900 m vary from open grasslands especially towards the upper and treefern savannahs to well boundary, and only 6 species out developed mossy forests; in fact, of the 21 included in the study only 15 of the numerous sites were occur above 2900 m. Hepatics described as mossy forests or elfin which have their lowermost forests/montane rainforests occurrences at 2200-2300 m are approaching mossy forests. Isotachis armata, eight Frullania species, Jungermannia A number of features characterize diversiclavellata, J. stephanii, J. zone IV. The Dicranaceae and the herzogiana, Lepidozia bursifera, Bryaceae are well represented. L. borneensis, L. holorhiza, Members of the Polytrichaceae Zoopsidella caledonica, Chando- (Dawsonia spp., for example) nanthus pilifer, Jensenia decipiens, become more abundant Plagiochila tecta, Plagiochilion (Polytrichum juniperinum occurs braunianum, Pseudolepicolea lowermost at 2450 m). The grolleana, Radula Calymperaceae is virtually absent. madagascariensis, R. ovalilobu- The Neckeraceae is represented only la, and Schistochila schulzei. A list by Homaliodendron flabellatum. of the hepatics not found outside The Leucobryaceae is represented zone IV includes Frullania only by two species, Leucobryum armatifolia, F. irregularis, F. sanctum and L. javense (up to 2550 papillata, F. pauciramea var. and 2900 m, respectively). The pauciramella, F. seriatifolia, F. Pterobryaceae and the subcaduca, F. subdentata, F. van-

74 The upper boundary of zone V, at above it; the Radulaceae practically 3400 m, should perhaps be vanish approximately at 3100- recognized with some reservations 3250 m, just one (Radula (see Materials and Methods), and it tjibodensis) being encountered is possible that the zone is wider above 3400 m. Of the mosses, the than is defined here. The occurrence Bryaceae, Dicranaceae, of a zone boundary at 3400 m is Ditrichaceae, Polytrichaceae, supported by the fact that a few Pottiaceae, and perhaps also the species were collected lowermost Rhizogoniaceae are well at that altitude (Marsupella revoluta; represented. The Meteoriaceae is Holomitrium obliquum, Ditrichum scarce - Floribundaria floribunda, sericeum, Rhacocarpus alpinus, F. sparsa, and Meteorium R. purpurascens) and the buchananii are present but not lowermost collecting sites defined common - and vanish completely as 'alpine grasslands' by Koponen at 3250 m; the highest occurring & Norris (1983a) were at 3400 m. Neckeraceae, Homaliodendron The species not common above flabellatum, vanishes at 3350 m. 3400 m are rather numerous: the Only 3 Pterobryaceae are present: hepatics Southbya grollei, Garovaglia plumosa is not found Reboulia hemisphaerica, above 3100 m, and Frullania reflexistipula, Herbertus Calyptothecium urvilleanum and armitanus, Triandrophyllum Trachyloma indicum disappear heterophyllum, Jungermannia from the flora at the upper boundary appressifolia, Lepidozia hasskar- of the zone. liana, L. holorhiza, Plagiochila tecta, Schistochila nymanii, S. Zone V apparently coincides in sciurea, Trichocolea tomentella, part with Lane-Poole’s (1925) and Dumortiera hirsuta; and the 'High mountain forests', but it also mosses Anomobryum bulbiferum, embraces the highermost mossy A. hyalinum, Bryum wabagense, forests. Lane-Poole placed the lower B. argenteum, Buxbaumia limit of the 'High mountain forests' javanica, Dicranum armittii, D. at 3050-3350 m (10000-11000 blumii, Homaliodendron ft). The forest is taller than the flabellatum, Dawsonia longifolia, mossy forest and dominated by Pogonatum cirratum, conifers (Dacrydium, Libocedrus, Leptodontium aggregatum, Phyllocladus, Podocarpus). The Timmiella anomala, climate is drier than in the lower Calyptothecium urvilleanum, Tra- 'cloud belt', which, along with the chyloma indicum, Rhizogonium still lower mean temperatures, hattorii, and R. lamii. causes an impoverishment of the bryoflora. In the upper parts of The impoverishment of the hepatic zone V and above it forests are flora in zone V is clearly stronger patchy and give way to alpine than that of the moss flora (Fig. 5). grasslands and heaths. This change For example, of the 46 taxa of in the vegetation is accompanied Frullania included in this study, by an increase in the number of 32 occur only below this zone, and terrestrial and saxicolous only 6 reach above 3400 m; 14 bryophytes, such as numerous Plagiochila species out of 26 are Bryaceae, Ditrichaceae and represented in this zone, but only 5 Polytrichaceae, and a strong 75 decrease in epiphytic bryophytes. Gradstein & Frahm record a considerable increase in the number Table 9 presents a comparison of of species at 1300 m, and include different authors’ views of the Syrrhopodon prolifer and Jubula altitudinal zonation of vegetation hutchinsiae in the list of the species on tropical mountains. Zonation characterizing the 'Montane Stufe' schemes under the numbers 1, 2 between 1300-1900 m. On the and 7 are based on bryophytes, 3- Huon Peninsula a remarkable 6 mainly on phanerogams. increase in the diversity of the bryoflora takes place at 1200 m, As can be seen from Table 9, Reenen which is also the lower limit of & Gradstein’s (1983) and Gradstein Syrrhopodon prolifer; Jubula & Frahm’s (1987) results from hutchinsiae subsp. javanica occurs Colombia and Peru, respectively, between 1500-2800 m. are very similar despite the different methods used. It should be noted, Above these zones, however, the however, that some of the subzones boundaries in Colombia and Peru of the latter authors are considered do not very well match those main zones by the former authors. observed from the Huon Peninsula. This can not be regarded as very This might be expected, because important, since the boundaries of the Andes are very different in the altitudinal zones (or subzones) character from the mountain ran- agree very well. Gradstein & Frahm ges on the Huon Penisula. The (1987: 109) conclude: 'Die upper boundary of my zone V, at Übereinstimmung dieser 3400 m, probably indicates the Ergebnisse, trotz der sehr lower limit of alpine vegetation, unterschiedlichen benutzten since some of the collecting sites at Methoden, lässt eine Verallgemei- or above that altitude (along transect nerung zu. Es scheint, dass sich die 6) were described as 'alpine hier beschriebene Zonierung auf grasslands' by Koponen & Norris orographische und (1983a). The 3400 m boundary makroklimatische Bedingungen coincides with the lower limit (3300 zurückführen lässt, die auch m) of the 'open páramo' on the anderswo in den Anden auftreten Sierra Nevada de Santa Marta in und die nicht auf zufällige lokale Colombia (Reenen & Gradstein Bedingungen zurückzuführen 1983), and the lower limit (3200 sind.' m) of the 'Tropisch alpine Stufe' in Peru (Gradstein & Frahm 1987). The agreement of the above Consequently, my zone V zonations from South America with apparently corresponds to the latter those achieved in the present study authors’ 'Subalpine Stufe' between is in part problematic. Obviously, 2800-3200 m, and to the former my zone I (0-300 m) corresponds authors’ zone IV. to Gradstein & Frahm’s (1987) 'Tropische Tieflandstufe' (0-500 m), It should be noted that the studies and my zone II (300- 1200 m) to in Colombia and Peru are based on their 'Submontane Stufe' (500- a single transect along a mountain 1300 m) and to Reenen & slope, whereas the Huon Peninsula Gradstein’s (1983) zone I (500- material was collected at 16 separate 1400 m). It is noteworthy that transects on a number of mountain 76 ranges (fig. 4 in Koponen & Norris represented by Dicranoloma assi- 1983a). Hence, the present study mile, D. braunii and Symblepharis displays an average zonation in the reinwardtii, and the study area, rather than a zonation Rhizogoniaceae by Rhizogonium on any particular mountain. spiniforme (= Pyrrhobryum spini- Although Seifriz’s (1924) study forme), etc. Seifriz (1924) concludes on the altitudinal distribution of that 'no species is common to both lichens and mosses on Mt Gedeh, zones'. However, Seifriz did not Java, concerned only five collect bryophytes very intensively 'subzones' between 1400-2850 m in either zone, since he reports only (4600-9400 ft), the bryological 17 mosses and 3 hepatics from the aspect is briefly reviewed here. The 'Podocarpus subzone ', and 14 'Rasamala subzone' (1400- 1700 mosses and 6 hepatics from the m, 4600-5500 ft) and the 'Vaccinium subzone'. In fact, many 'Herbaceous subzone' (2150-2450 of the species which he reports as m, 7000-8000 ft) are poor in bryo- occurring only in one of his phytes, whereas the 'Podocarpus subzones have wide altitudinal subzone' (1700-2150 m, 5500- distributions on the Huon 7000 ft) and the 'Vaccinium subzo- Peninsula: Aerobryopsis ne' (2450-2750 m) are longissima 400-1500 m, Floribun- bryologically rich. According to daria floribunda 100-3250 m, Pa- Seifriz (1924), there is a striking pillaria fuscescens 1200-2400 m, difference in the life-forms of Dicranoloma assimile 900-3500 mosses between the latter two m, and Pyrrhobryum spiniforme subzones: in the Podocarpus 500- 2800 m. Although the floristic subzone 'Every tree is festooned differences between Seifriz’s with pendent mosses', whereas in (1924) 'moss zones' are obviously the Vaccinium subzone '...the mos- not quite as clear-cut as he sugge- ses are compact tufted forms sted, it is evident that they really clinging close to the tree trunks, represent separate zones. In New and forming thick reeking wet Guinea, Podocarpus spp. are wads'. The bryofloristic differences especially common at moderately between these two subzones are high altitudes, in the midmountain very clear. Papillaria fuscescens is forests of Lane- Poole (1925), or in 'perhaps, the most abundant of the the mid-montane forests of Johns second subzone epiphytic mosses'; (1982). It is likely that the common are, e.g., some other bryologically poorer 'Herbaceous Meteoriaceae (Floribundaria subzone' between 2150-2450 m floribunda, F. aurea, Aerobryopsis on Mt Gedeh is but a local pheno- longissima), Hookeriaceae menon. Indeed, on the Huon (Hypopterygium struthiopteris), Peninsula, the richest bryoflora Hylocomiaceae (Macrothamnium occurs within that range (Fig. 5). macrocarpum), Hypnodendraceae The great abundance of Vaccinium (Hypnodendron reinwardtii, H. spp. at certain altitudes on Mt Gedeh divaricatum), and Sematophylla- may also be a local phenomenon, ceae (Sematophyllum spp.). The but the reported abundance of most abundant moss in the Rhododendron spp. in the 'Vaccinium subzone' is Sematophyl- 'Vaccinium subzone' (Seifriz 1924, lum pinnatum (= Acroporium war- cf. also Lane-Poole 1925: 223) burgii); the Dicranaceae is and Seifriz’s (1924) fig. 1 (plate 77 17) suggest that we are dealing to the one presented in this paper with a 'true' mossy forest, which, (cf. Table 9). Their lowermost zone consequently, occurs between extends from sea level to 660 m. It 2450-2750 m on Mt Gedeh. Thus, comprises lowland rainforests in Seifriz’s (1923, 1924) works are the lower part and hill forests in the quite valuable locally, but their upper, but the authors do not importance as general schemes of indicate any approximate altitude the vertical zonation of vegetation where the shift occurs. The lower in the (paleo)tropics is more montane zone of Gressitt & doubtful. Nadkarni occurs between 660- 1300 m. Therefore, my zone II Lane-Poole’s (1925) zonation embraces that zone and the hill scheme has already been discussed forests below it. Gressitt & above. His results agree well with Nadkarni place the mid montane those reported here (cf. Table 9). zone between 1300-2300 m, and it The single discrepancy between corresponds to the altitudinal range our studies is that he placed the of my zone III. Interestingly, zones upper boundary of his “Foothill IV and V as defined here seem to forests” at an average altitude of subdivide Gressitt & Nadkarni’s 1650 m (5500 ft), whereas my upper montane zone. Finally, they zone II extends only up to 1200 m. establish the lower boundary of the However, he allowed an altitudinal alpine zone at 3300 m, while on the range of 1350-1650 m (4500-5500 Huon Peninsula that boundary ft) for the boundary. apparently occurs at 3400 m. A brief but important conclusion The correspondence of Lane- may be drawn from the close Poole’s (1925) vegetation zones agreement of the above zonation with those defined here is roughly schemes for the New Guinea as follows: vegetation. It is obvious that the altitudinal boundaries of the Lowland forest, 0-300 m —— vegetation zones, based on Zone I, 0-300 m phanerogamic flora and vegetation, are accompanied by distinct Foothill forest, 300-(1350)1650 discontinuities in the bryophyte m —— Zone II, 300-1200 m flora. Consequently, each zone has its characteristic bryophyte species Midmountain forest, 1650-2300 assemblage. This makes sense since m —— Zone III, 1200-2200(2300) changes in the gross structure of m vegetation imply changes in the physical environment within the Mossy forest (1500)2300- forest interior which strongly affects 3050(3350) —— Zone IV, the bryophyte flora and vegetation. (2200)2300- 2800(2900) m B. Correlations of the altitudinal High mountain forest zonation of bryophytes on the (3050)3350-ca. 3600 m —— Zone Huon Peninsula with their V, 2800(2900)-3400m phytogeography Gressitt & Nadkarni’s (1978: fig. Hyvönen (1989a) presented lists of 2) zonation scheme is fairly similar Western Melanesian mosses in each of the 78 phytogeographic units discussed below. hispidissima (III), Anomobryum ochii (III), A. Therefore, this list is not presented here. hyalinum (V), Aerobryidium fuscescens (III), The abbreviations of the Garovaglia subelegans (III), Powellia parvula phytogeographical areas used are (III), Brotherobryum undulatifolium (III), and according to Wijk et. al. (1959). Dicnemon robbinsii (III). It may be noted that none of the endemics occurring at low altitudes 1. Endemics of New Guinea or Western are restricted to any particular zone. Melanesia (Table 1) 2. Malesian endemics (As 4, including species The endemism of the bryoflora in New which in Oceania occur only in the Solomon Guinea was examined by Piippo et al. Islands) (Table 2) (1987). They concluded that most of the According to Hyvönen (1989a), ca. 15% of endemic species in Western Melanesia Western Melanesian mosses represent this occur above 1500 m, with the exception distribution type. of the Frieda River area, where the level of The hepatics in this group show similar endemism is high between 200-1350 m altitudinal distributions with the New (see Norris & Koponen 1985b, Piippo Guinean or Western Melanesian endemics 1986). Based on the study of 22 hepatic discussed above. The percentages are very families and 8 moss families, Piippo et al. high in the zones III and IV. Interestingly, (1987) gave endemism percentages (at the mosses have a somewhat different species level) of 48% for hepatics and altitudinal distribution. The number of 23% for mosses. Hyvönen (1989a) Malesian endemic mosses rises continu- presented endemism percentage of ously from zone I to zone V, and decreases 17% for mosses. The present study only above 3400 m. These results do not includes 25 hepatic families and 31 seem to be in accordance with the reported moss families, and their percentage strong affinities to Malesia of the New of endemism, is 33% (70 out of Guinean lowland phanerogam flora (e.g. 213 taxa) and 24% (54 out of 226 Smith 1980). taxa), respectively. However, the last mentioned percentages include None of the hepatics and only two mosses only taxa which were collected at (Barbula pseudo- ehrenbergii, least twice on the Huon Peninsula. Thamnobryum ellipticum) of this distribution type occur in zone I. Species The present results agree with those of Piippo et restricted to a specific zone include al. (1987). The endemism level of hepatics is Triandrophyllum heterophyllum (zone V), highest in zone IV. This might be expected, since Lepidozia borneensis (IV), Heteroscyphus there are numerous endemic hepatics which occur wettsteinii (III), Frullania armatifolia (IV), only in the mossy forests. The highest number of Aerobryidium crispifolium (II), endemic mosses are encountered in zone III, which Rhizogonium lamii (V), Dawsonia comprises lower montane forests and mossy beccarii (IV), Oligotrichum javanicum forests. (V), Pogonatum subtortile (IV), Buxbau- mia javanica (V), Leptodontium The four endemics occurring in zone I are aggregatum (V), Himantocladium Frullania multilacera, Radula koponenii, Fissi- submontanum (III), Microdus miquelianus dens vanzantenii, and Barbula novoguinensis. (III), Braunfelsia enervis (IV), and The endemic taxa restricted to a single altitudinal Fabronia curvirostris (III). zone include, Porella grollei (zone III), Telara- nea trisetosa (III), T. kaindina (III), Leptoscyphus 3. SE-Asian distribution (As 2 - 4, huonicus (V), Lepicolea norrisii (III), Radula including taxa which in Oceania occur ovalilobula (IV), R. minutilobula (V), Metzgeria only in the Solomon Islands) (Table 3) 79 from SE-Asia and Australasia, and rather About 14% of Western Melanesian mosses extensive bryophyte migration from these have a SE-Asian distribution (Hyvönen regions to Oceania has taken place at low 1989a). altitudes. SE-Asian hepatics and mosses are best represented in zones III and IV. However, Asian-Oceanian species encountered in when compared with the Malesian only one altitudinal zone on the Huon endemics, the SE-Asian species are clearly Peninsula comprise Pleurozia conchifolia more numerous at lower altitudes, notably (III), Jackiella javanica (III), Frullania in zone II. The single hepatic and the four gracilis (III), F. trichodes (III), F. immersa mosses with this distribution pattern in (III), Treubia insignis (III), Radula zone I are Radula vrieseana, Fissidens amentulosa (V), R. acuminata (III), hollianus, F. zippelianus, Barbula javani- Plagiochila longispica (II), Notoscyphus ca, and B. subcomosa. Species occurring lutescens (III), Calymperes strictifolium only in one altitudinal zone include (II), Syrrhopodon japonicus (III), and Southbya grollei (V), Plagiomnium Fissidens obscurirete (II). succulentum (III), Orthodontium infractum (V), Aerobryopsis leptostigma- 5. Asian-Oceanian-Australian ta (II), Pogonatum neesii (III), Dicrano- distribution (Table 5) dontium fleischerianum (IV), and Leuco- bryum neilgherrense (III). Asian-Oceanian-Australian species are relatively few in Western Melanesia, 4. Asian-Oceanian distribution (Table 4) comprising only 11% of the moss flora (Hyvönen 1989a). As might be About 14% of Western Melanesian expected, they show a similar mosses belong to this distribution overall altitudinal distribution type (Hyvönen 1989a). pattern as the Asian-Oceanian species discussed above, but mosses This group is more strongly represented at of this distribution type are not as low altitudes (zones I and II) than any of commonly encountered in the two the preceding distribution groups. In the lowermost zones. Species present three groups dealt with above, hepatics in only one altitudinal zone include had the highest number of taxa in zone IV Jungermannia hasskarliana (III), instead of zone III, where most of the Syrrhopodon muelleri (II), S. Asian- Oceanian distribution hepatics are tristichus (III), Himantocladium concentrated. Significantly, mosses are plumula (II), and Leucophanes even more strongly represented at lower glaucum (I). altitudes, the highest number of species being encountered in zones II and III, but 6. Asian-Australian distribution also the lowermost zone is relatively rich in moss taxa. The moss flora also shows a This distribution group is a small more abrupt impoverishment at high one in Western Melanesia; only 4% altitudes, especially in zone V and above of the mosses belong here (Hyvönen that, than the hepatic flora. The strong 1989a). Of the species included in the representation of Asian-Oceanian bryo- present study, only 3 hepatics and 7 mosses phytes at low altitudes is readily explained are regarded as Asian-Australian, Junger- by the fact that high mountains in Oceania mannia appressifolia (V), Tylimanthus are very few. Many of the Oceanian islands saccatus (III-V), Zoopsidella caledonica are geologically young, and they must (IV), Dawsonia longifolia (III-V), have received their bryophytes mainly Dicranum dicarpum (III-IV), Fissidens 80 nymanii (II-III), Orthomnion elimbatum (II-V), Orthorrhynchium elegans (III-V), Only about 4% of the Western Melanesian Powellia subelimbata (II-III), and mosses are paleotropical in distribution Tridontium tasmanicum (III). (Hyvönen 1989a). Koponen & Norris (1986) regarded The hepatics in this group are Powellia subelimbata as endemic to New Pleurozia gigantea (III-V and Guinea, but according to Zanten & Pócs above), Psiloclada clandestina (1981: 533), it has been collected also in (III), Heteroscyphus splendens (III), Queensland. Frullania apiculata (I-V), Wiesnerella denudata (II-IV), Most of the species named above have Radula madagascariensis (IV), their main distributions in Asia, whence Metzgeria consanquinea (III-V and they probably have dispersed to Australia. above), Gottschelia schizopleura An obvious exception is Powellia (IV-V and above), and the mosses subelimbata, which Zanten & Pócs (1981), Calymperes dozyanum (I-II), following the terminology of Smith (1980), Calymperes taitense (I-II), considered to represent an Australian Mitthyridium fasciculatum (II), element, encountered in New Guinea in Floribundaria floribunda (I-V and eucalyptus woodlands and savannahs. Ko- above), Neckeropsis lepineana (II- ponen & Norris (1986) report that on the III), Pinnatella mariei (I), Huon Peninsula this species also occurs in Rhodobryum aubertii (II-III), R. gardens and disturbed forests. giganteum (III-V), Brachymenium nepalense (II-V and above), The low number of taxa in this phytogeo- Trachypus bicolor (III-V and graphic group prevents reliable above), Leucophanes conclusions as to their altitudinal zonation octoblepharioides (II-III), and L. on the Huon Peninsula. It can be noted angustifolium (II-III). Pinnatella that none of the taxa occur in zone I, and mariei has recently been reported only three are encountered in zone II. from Africa (Mattila & Enroth, They seem to be mainly distributed within 1990). zones III-V. As the relatively scanty material 7. Transpacific distribution suggests, there seem to be slight differences between the altitudinal This type of distribution is represented distributions of paleotropical only by 1 hepatic and 3 mosses: Frullania hepatics and mosses on the Huon gaudichaudii (zones II-III), Barbella Peninsula. The hepatics tend to cubensis (II-III), Campylopus aureus (IV- occur at somewhat higher altitudes V, and up to 3600 m), and Trichostomum than the mosses, the only hepatic crispulum (III-IV). The total range of present in zones I and II being Frullania gaudichaudii is inadequately Frullania apiculata. Most known, since its presence in Australia is paleotropical mosses occur in zones not certain (Hattori 1982). The small II and III on the Huon Peninsula. number of transpacific taxa prevents any reliable conclusions, but it seems that these 9. Pantropical distribution (Table taxa are mainly distributed at moderate 7) altitudes between 1000-3000 m. Pantropical mosses represent 5% 8. Paleotropical distribution (Table 6) of the Western Melanesian flora 81 (Hyvönen 1989a). available suggests that, in general, pantropical mosses are most The species included here are numerous at moderate to relatively Heteroscyphus argutus (I-III), Frullania high altitudes (zones II-IV). This ericoides (I-IV), F. arecae (III-IV), altitudinal distribution is rather Metzgeria epiphylla (IV), M. albinea (II- similar to that of the whole IV), M. australis (II-IV), Bryum bryophyte flora included in this apiculatum (I-III), B. billarderi (III-V and study (cf. Fig. 5). above), B. coronatum (III-V), Ceratodon stenocarpus (IV-V and above), 10. Southern hemisphere Calymperes afzelii (II), C. tenerum (I), distribution Fissidens asplenioides (III-IV), Syrrhopodon prolifer (III-IV), In the present study, hepatics are Octoblepharum albidum (I-III), and not represented in this type of Pyrrhobryum spiniforme (II-IV). The last distribution, and there are only 4 mentioned species was 'with some mosses: Bryum clavatum (zones II- reservations' considered a 'cosmopolitan' V), Catagonium nitens (IV-V), by Hyvönen (1989a). However, Ditrichum difficile (III-V), and judged from the distribution map Rhacocarpus purpurascens (lo- in Iwatsuki (1972: 131), it seems wermost at 3400 m). According to reasonable to regard it as pantropi- Ochi (1985), in the tropics Bryum cal (see also Menzel & Schultze- clavatum occurs only at high Motel 1987). altitudes, but on the Huon Peninsula it has rather a wide altitudinal In addition Hyvönen (1989a) and amplitude between 700-3300 m. Menzel & Schultze-Motel (1987) regard Menzel & Schultze-Motel (1987) Bryum billarderi as a pantropical species. considered Rhacocarpus Ochi (1985) writes: 'Pan-tropical, but purpurascens a pantropical species, extending up or down to temperate regions but since it is apparently restricted in the Northern and Southern hemispheres'. to the southern hemisphere, I prefer Based on information in Norris & Koponen to follow Hyvönen (1989a). Menzel & (1987), Hyvönen (1989a) considered Schultze-Motel (1987) give an altitudinal Fissidens asplenioides a transpaci- distribution of 2100- 4400 m for R. fic species, but the map in Schuster purpurascens in Peru. (1983) shows a pantropical range (see also Menzel & Schultze-Motel Although the number of taxa representing 1987). In this study the species is this distribution type is very small, it may, included in the pantropical element. with some reservations, be concluded that they mostly occur at relatively high It is noteworthy that pantropical altitudes, mainly in zones IV and V. This hepatics do not occur in zone V or seems to have some relevance, since Bryum above. The two mosses encountered clavatum, Catagonium nitens and above 3400 m are Bryum billarderi Rhacocarpus purpurascens apparently and Ceratodon stenocarpus. It belong to the Gondwanaland element (Hy- should be mentioned in this vönen 1989a), which in New connection that Norris & Koponen Guinea mainly occurs on (1988) doubt whether C. mountains and is scarce in the low- stenocarpus can be kept separate lands (Smith 1980). from the 'cosmopolitan' C. purpureus. The meager material 11. Mainly northern hemisphere 82 distribution The three mosses included in this According to Hyvönen (1989a), only distribution are Campylopodium medium, ca. 5% of Western Melanesian mosses Homaliodendron flabellatum and belong to this distribution type. Of the Timmiella anomala. The main distribution bryophytes included in this study, the of Campylopodium medium is Asian- hepatics Jungermannia sphaerocarpa Oceanian- Australian, but it is also known (zone V) and Marsupella revoluta (only from Puerto Rico and Chile. On the Huon above 3400 m) have a mainly northern Peninsula it was collected between 1200- hemisphere distribution. Eight mosses are 2000 m (zone III). Timmiella anomala included here: Pogonatum urnigerum has a wide distribution in North America, (zone V and above), Bryum Central America, Europe and Asia (Japan, microerythrocarpum (III-V and above), India). From the Paleotropics it is only Anomobryum julaceum (IV-V and above), known from Papua New Guinea and the Epipterygium tozeri (II-IV), Pohlia Philippines. The collections from the Huon elongata (V and above), Peninsula come from relatively high Bryoerythrophyllum ferruginascens (III- altitudes (2100-3400 m, zones III-V), V and above), Didymodon rigidicaulis which would suggest that this is a northern (III-V and above), and Fissidens taxifolius hemisphere species (group 11 above). My (III-IV). It is well-known that in the tropics, decision to regard Timmiella anomala as temperate species mostly occur at high a plant of a 'miscellaneous distribution' is altitudes. This is supported by the present based on Hyvönen (1989a) and the data. It should be noted, however, that on opinion of Norris & Koponen the Huon Peninsula Epipterygium tozeri (1989): 'Several species are occurs as low as 750 m (zone II), and included in the world list of this Fissidens taxifolius at 1400 m (zone III). genus but careful review of the described species may well show 12. Miscellaneous distribution them all conspecific with Timmiella anomala.' Two hepatics may be placed here. Chan- donanthus hirtellus has a mainly The extremely variable paleotropical distribution, but it is also Homaliodendron flabellatum known from British Columbia (Canada) (zones II-V) has rather a wide range and New Zealand (Miller et al. 1983: 79). in the tropical regions, where it Plagiochila sciophila also occurs in remains unrecorded only from Arkansas, North America (Piippo 1989). continental Africa and (possibly) Inoue (1984: 127) writes: 'Plagiochila large areas of the Neotropics. In the sciophila is the most common and most latter region (at least on the islands polymorphic taxon...; it is distributed very of the Caribbean) the species has widely in tropical - temperate regions of been known as H. decompositum the Paleotropics and Neotropics, but (see Enroth 1989b). lacking from the African continent.' It is possible that the total range of this variable 13. Global distribution taxon is still inadequately known. Both ('cosmopolitan' (Table 8) Chandonanthus hirtellus and Plagiochila sciophila have wide altitudinal ranges on As might be expected this group the Huon Peninsula. The former occurs comprises mostly 'weedy' plants, between 1200-3500 m (zones III-V and which often occur in exposed, man- above) and the latter between 350-2800 m made habitats, and usually grow (zones II-IV). on soil or cliffs and rocks (cf. Hyvönen 1989a). However, of the four hepatics included here Lophocolea bidentata is exclusively, and Metzgeria leptoneura mainly, epiphytic. The two other hepatics, Dumortiera hirsuta and Reboulia hemisphaerica are epigaeic or epilithic.

The Pottiaceae is represented by eight species, all occurring on soil and/or boulders and rocks: Anoectangium aestivum, Barbula indica, Hymenostylium recurvirostre, Hyophila involuta, Lepto- 83

HEPATICAE (1500-2400 m) F. armatifolia Verd. (2300- Acrobolbaceae (Piippo 1985a) 2800 m) Tylimanthus saccatus (Hook.) F. armitiana Steph. (1500- Mitt. (1700-2400 m) 3500 m) F. attenuata Steph. (900-2800 Adelanthaceae (Piippo 1984b) m) Wettsteinia inversa (Sande Lac.) F. bergmanii Hatt. (1500-2600 Schiffn. (2500-3100 m) m) F. carrii Kamim. & Hatt. Arnelliaceae (Piippo 1985a) (1200-1500 m) Southbya grollei N. Kitag. F. curvistipula Steph. (1200- (3250-3400 m) 2200 m) F. cuspidifolia Steph. (1700- Aytoniaceae (Piippo 1988b) 3300 m) Asterella khasyana (Griff.) F. echinantha Hatt. (2800-3200 Pandé et al. (1300-2900 m) m) Reboulia hemisphaerica (L.) F. epiphylla Hatt. (1250-1500 Raddi (1500-3400 m) m) F. ericoides (Nees) Mont. (0- Balantiopsiaceae (Piippo 2900 m) 1984a) F. eymae Hatt. (600-2200 m) Balantiopsis ciliaris Hatt. F. errans Verd. var. subsp. novoguineensis Hatt. angulistipula Hatt. (2200-2300 (1500-2500 m) m) Isotachis armata (Nees) Gott. F. falsisinuata Hatt. fo. (2200-3100 m) parvistylata (Hatt.) Hatt. & Piippo Cephaloziaceae (Grolle & (3300-3400 m) Piippo 1984a; Piippo 1984b) F. gaudichaudii (Nees & Nowellia borneensis (De Not.) Mont.) Nees & Mont. (550-2000 Schiffn. (2100-2250 m) m) Odontoschisma naviculare F. gracilis (Reinw. et al.) Dum. (Steph.) Grolle (1800-2450 m) (1700-2300 m) F. immersa Steph. (1300-1400 Frullaniaceae (Hattori & Piippo m) 1986) F. irregularis Hatt. & Piippo Frullania apiculata (Reinw. et (2200-2600 m) al.) Dum. (100-3200 m) F. junghuhniana Gott. var. F. appendistipula Hatt. (2300- tenella (Sande Lac.) Grolle & 3500 m) Hatt. F. arecae (Spreng.) Gott. (1200-3350 m) 84 F. leeuwenii Verd. (800-1800 F. van-zantenii Kamim. & Hatt. m) (2300-2500 m) F. macgregorii Steph. (2800- 3600 m) Geocalycaceae (Piippo 1985b) F. multilacera Steph. (200- Heteroscyphus acutangulus 1800 m) (Schiffn.) Schiffn. (1200-3300 F. nepalensis (Spreng.) Lehm. m) & Lindenb. (2000-2600 m) H. argutus (Reinw. et al.) F. nobilis Steph. var. cochleata Schiffn. (200-1600 m) (Steph.) Hatt. & Piippo (2200 H. aselliformis (Reinw. et al.) -2300 m) Schiffn. (1200-3000 m) F. orientalis Sande Lac. (600- H. coalitus (Hook.) Schiffn. 2500 m) (200-3100 m) F. papillata Steph. (2200-2900 H. sarawaketanus Piippo m) (3250-3550 m) F. papuana Verd. (3300-3600 H. splendens (Lehm. & m) Lindenb.) Grolle (1800-2300 m) F. pauciramea Steph. (1300- H. tridentatus (Sande Lac.) 3000 m) Grolle (1850-3600 m) — var. pauciramella Hatt. & H. wettsteinii (Schiffn.) Schiffn. Piippo (2200-2800 m) (1700-2000 m) F. prominula Hatt. & Streim. H. zollingeri (Gott.) Schiffn. (1200-1500 m) (350-1600 m) F. pullei (2200-3500 m) Leptoscyphus huonicus Piippo F. reflexistipula Sande Lac. (2800-3100 m) (1400-3400 m) Lophocolea bidentata (L.) F. reimersii Verd. (2800-3500 Dum. (2850-3300 m) m) L. ciliolata (Nees) Gott. (1800- F. saepis-dentata Hatt. & 3100 m) Streim. (1730-1800 m) L. costata (Nees) Gott. (1500- F. schusteriana Hatt. (1700- 3100 m) 2300 m) L. levieri Schiffn. (1400-3500 F. seriatifolia Steph. (2000- m) 2900 m) Saccogynidium muricellum (De F. serrata Gott. fo. Not.) Grolle (1500-2500 m) crispulodentata Verd. (1500- 2300 m) Gymnomitriaceae (V na & F. serrata var. hamatispina Piippo 1989) (Hatt.) Hatt. (800-3300 m) Marsupella revoluta (Nees) F. serrata subsp. spinistipula Dum. (3400-3570 m) (Hatt.) Hatt. (1000-2500 m) F. subcaduca Hatt. (2400-2800 Haplomitriaceae (Piippo m) 1984a) F. subdentata Steph. (2200- Haplomitrium blumii (Nees) 2900 m) Schust. (1100-3550 m) F. subnigricaulis Hatt. var. subtruncata Hatt. (1200-1700 Herbertaceae (Piippo 1984a) m) Herbertus armitanus (Steph.) F. trichodes Mitt. (1450-2230 H. A. Miller (1700-3400 m) m) H. circinatus (Steph.) H. A. 85 Miller (1950-3500 m) 3500 m) H. longifissus Steph. (1900- Mastigophora diclados (Brid.) 3500 m) Nees (900-3500 m) Triandrophyllum heterophyllum (Steph.) Grolle (3250-3400 m) Lepidoziaceae (Grolle & Piippo 1984a; Piippo 1984b, 1985c, Jackiellaceae (Piippo 1985a) pers. comm.) Jackiella javanica Schiffn. Hygrolembidium boschianum (1500-2000 m) (Sande Lac.) Schust. (2450-3300 m) Jubulaceae (Piippo 1984b) Kurzia pallida Piippo (2400- Jubula hutchinsiae (Hook.) 3550 m) Dum. subsp. javanica (Steph.) Lepidozia bursifera Hatt. & Verd. Grolle (2200-3000 m) (1500-2800 m) L. borneensis Steph. (2300- 2700 m) Jungermanniaceae (Vana & L. cladorhiza (Reinw. et al.) Piippo 1989) Gott. et al. (1900-3100 m) Jungermannia appressifolia L. ferdinandi-muelleri Steph. Mitt. (2800-3350 m) (1200-2400 m) J. comata Nees var. L. gwamii Piippo (2800-3550 novaeguineae Vana (1300-2300 m) m) L. hasskarliana (Lindenb.) J. diversiclavellata Amak. & Steph. (1500-3350 m) Grolle (2300-3300 m) L. holorhiza (Reinw. et al.) J. hasskarliana (Nees) Mitt. Gott. et al. (2200-3350 m) (1250-2000 m) L. stahlii Steph. (2500-3500 m) J. herzogiana Vana (2300-3550 L. trichodes (Reinw. et al.) Nees m) (1000-2500 m) J. hirticalyx Steph. (750-2300 L. wallichiana Gott. (1200- m) 3500 m) J. sphaerocarpa Hook. (3200- Psiloclada clandestina Mitt. 3250 m) (1800-2300 m) J. stephanii (Schiffn.) Amak. Telaranea kaindina Grolle (2280-3550 m) (1800-2250 m) J. stricta (Schiffn.) Steph. T. lawesii (Steph.) Grolle (1200-3550 m) (1200-3100 m) J. tetragona Lindenb. (800- T. trisetosa (Steph. emend. 1400 m) Grolle) Grolle (1350-2300 m) J. truncata Nees (800-2450 m) Zoopsidella caledonica Schust. J. virgata (Mitt.) Steph. (800- (2200-2600 m) 1800 m) Zoopsis liukiuensis Horik. Notoscyphus lutescens (Lehm.) (1000-3000 m) Mitt. (1450-1700 m) Lophoziaceae (Vana & Piippo Lepicoleaceae (Piippo 1984a, 1989b) 1988a) Anastrophyllum revolutum Lepicolea norrisii Piippo Steph. (1800-3600 m) (1900-2300 m) Andrewsianthus cavifolius L. rara (Steph.) Grolle (1800- Grolle & Vana (2500-3600 m) 86 A. bidens (Mitt. ex Steph.) S. similis Grolle (1600-2900 m) Schust. (2450-3600 m) Anomacaulis flaccidus (Steph.) Plagiochilaceae (Piippo 1989) Grolle (2100-2900 m) Plagiochila abietina (Nees) Chandonanthus hirtellus Lindenb. (1500-3300 m) (Web.) Mitt. (1200-3500 m) P. arbuscula (Brid. ex Lehm. & C. pilifer Steph. (2200-3600 m) Lindenb.) Lindenb. (1600-3000 Denotarisia linguifolia (De m) Not.) Grolle (1500-3500 m) P. cymata Inoue & Grolle Gottschelia schizopleura (1200-2100 m) (Spruce) Grolle (2600-3600 m) P. decidua Inoue & Grolle Jamesoniella flexicaulis (Nees) (2500-3000 m) Schiffn. P. dendroides (Nees) Lindenb. Syzygiella securifolia (Nees) (1500-3300 m) Inoue (1500-2500 m) P. gymnoclada Sande Lac. (1500-3600 m) Makinoaceae (Piippo 1988c) P. hampeana Gott. (1900-3500 Makinoa crispata (Steph.) m) Miyake (1500-3000 m) P. javanica (Swartz) Dum. (500-3300 m) Metzgeriaceae (Piippo pers. P. kurokawae Inoue (2800- comm.) 3500 m) Metzgeria consanquinea P. longispica Mitt. (300-600 m) Schiffn. (1750-3550 m) P. loriana Steph. (1200-2800 M. epiphylla Evans (2750- m) 2900 m) P. odorata Inoue & Grolle M. hispidissima Steph. (2000- (1200-2800 m) 2200 m) P. propinqua Sande Lac. (800- M. albinea Spruce (300-2800 2000 m) m) P. pulvinata Steph. (2000-3550 M. australis Steph. (1100-2350 m) m) P. renitens (Nees) Lindenb. M. macveanii Kuwah. (2850- (1800-3300 m) 3500 m) P. schusteri Inoue & Grolle M. hattorii Kuwah. (1500- (2500-3600 m) 3100 m) P. sciophila Nees (350-2800 m) M. leptoneura Spruce (400- P. seemannii Mitt. (300-1800 3550 m) m) P. singularis Schiffn. (1800- Pallaviciniaceae (Grolle & 2800 m) Piippo 1986) P. sumatrana Schiffn. (800- Jensenia decipiens (Mitt.) 1500 m) Grolle (2300-3600 m) P. tamiensis Steph. (1200- Pallavicinia ambigua (Mitt.) 3000 m) Steph. (800-1750 m) P. tecta Inoue & Grolle (2200- Podomitrium malaccense 3400 m) (Steph.) D. H. Campb. (400- P. teysmannii Sande Lac. (400- 2450 m) 1800 m) Symphyogyna rectidens Grolle P. trapezoidea Lindenb. (1200- (1700-2400 m) 3300 m) 87 P. ulata Inoue & Grolle (1500- R. madagascariensis Gott. 1600 m) (2200-2600 m) P. velata Inoue & Piippo R. minutilobula Yamada & (1250-2500 m) Piippo (2850-3100 m) Plagiochilion braunianum R. multiflora Gott. ex Schiffn. (Nees) Hatt. (2300-3500 m) var. reflexilobula Grolle & P. oppositum (Reinw. et al.) Yamada (900-1100 m) Hatt. (1500-3600 m) R. norrisii Yamada & Piippo (2100-3000 m) Pleuroziaceae (Piippo 1984a) R. nymanii Steph. (200-1900 Pleurozia conchifolia (Hook. & m) Arnott) Aust. (1700-2200 m) R. ovalilobula Yamada (2200- P. gigantea (Web.) Lindb. 2800 m) (1200-3500 m) R. parvitexta Steph. (2100- 2900 m) Porellaceae (Piippo 1984a) R. retroflexa Tayl. (350-3250 Porella acutifolia (Lehm. & m) Lindenb.) Trev. (300-1800 m) - - var. fauciloba (Steph.) P. geheebii (Steph.) Hatt. (600- Yamada (350-1900 m) 2800 m) R. sharpii Yamada (750-2850 P. grollei Hatt. (1800-2100 m) m) P. viridissima (Mitt.) Grolle var. R. subpallens Steph. (1450- novoguineensis (Hatt.) Hatt. 2400 m) (2000-2300 m) R. tjibodensis Goebel ex Schiffn. (100-3600 m) Pseudolepicoleaceae (Piippo R. vrieseana Sande Lac. (100- 1984a) 600 m) Pseudolepicolea grolleana (Schust.) Grolle in Piippo (2200 Scapaniaceae (Piippo 1985a) -3550 m) Scapania javanica (1500-3600 Temnoma setigerum (Lindenb.) m) Schust. (1850-2850 m) Schistochilaceae (Piippo 1984a) Radulaceae (Yamada & Piippo Schistochila aligera (Nees & 1989) Blume) Jack & Steph. (1250- Radula acuminata Steph. 3200 m) (1700-1900 m) S. antara Grolle (1500-2900 R. amentulosa Mitt. (2800- m) ca.3100 m) S. blumii (Nees) Trev. (1200- R. anceps Sande Lac. (100- 2800 m) 2200 m) S. nymanii Steph. (800-3400 R. apiculata Sande Lac. ex m) Steph. (100-1900 m) S. philippinensis (Mon.) Jack & R. densifolia Castle (750-3000 Steph. (500-3000 m) m) S. schultzei (Steph. (2200-3200 R. javanica Gott. (40-800 m) m) R. koponenii Yamada & Piippo S. sciurea (Nees) Schiffn. (100-650 m) (1800-3350 m) R. lacerata Steph. (900-2500 m) Targioniaceae (Piippo 1988b) 88 Cyathodium foetidissimum B. microerythrocarpum C. Schiffn. (600-1500 m) Müll. & Kindb. in Macoun (1700-3500 m) Treubiaceae (Piippo 1988c) B. perdecurrens Bartr. (1500- Treubia insignis Goebel (1800- 3500 m) 2100 m) B. pseudoblandum T. Kop. & Norris (2000-3500 m) Trichocoleaceae (Piippo 1984a) B. russulum Broth. & Geh. Trichocolea breviseta Steph. (1400-2200 m) (1250-3100 m) Epipterygium tozeri (Grev.) T. tomentella (Ehrh.) Dum. s. Lindb. (750-2400 m) lat. (800-3350 m) Leptobryum pyriforme (Hedw.) Wils. (2200-2900 m) Wiesnerellaceae (Piippo 1988b) Orthodontium infractum Dozy Dumortiera hirsuta (Sw.) Nees & Molk. (2800-3100 m) (200-3400 m) Pohlia elongata Hedw. (2800- 3600 m) P. flexuosa Hook. (1600-3600 MUSCI m) Rhodobryum aubertii Bryaceae (Koponen & Norris (Schwaegr.) Thér. (400-2300 m) 1984, 1985b) R. giganteum (Schwaegr.) Par. Anomobryum bulbiferum Bartr. (1500-3000 m) (2000-3400 m) A. hyalinum T. Kop. & Norris Buxbaumiaceae (Hyvönen 1986) (2900-3400 m) Buxbaumia javanica C. Müll. (3300- A. julaceum (Gaertn. et al.) 3400 m) Schimp. (2400-3600 m) A. ochii T. Kop. & Norris Calymperaceae (Reese et al. 1986) (1500-1800 m) Calymperes afzelii Sw. (500-600 m) Brachymenium huonii T. Kop. C. dozyanum Mitt. (0-1200 m) & Norris (3250-3600 m) C. strictifolium (Mitt.) G. Roth (500- B. nepalense Hook. in 650 m) Schwaegr. (700-3600 m) C. taitense (Sull.) Mitt. (50-1000 m) - - var. globosum Bartr. (1200- C. tenerum C. Müll. (0-100 m) 1800 m) Syrrhopodon japonicus (Besch.) B. wabagense (Bartr.) T. Kop. Broth. (1800-1900 m) & Norris (1800-3400 m) S. muelleri (Dozy & Molk.) Sande Bryum apiculatum Schwaegr. Lac. (400-1000 m) (0-1500 m) S. prolifer Schwaegr. (1200-2400 m) B. argenteum Hedw. (1750- S. tristichus Nees ex Schwaegr. (1800- 3350 m) 2100 m) B. billarderi Schwaegr. (2000- Mitthyridium fasciculatum (Hook. & 3500 m) Grev.) Robins.(500-1000 m) B. capillare Hedw. (700-2500 M. obtusifolium (Lindb.) Robins. (200- m) 1000 m) B. clavatum (Schimp.) C. Müll. (700-3300 m) Dicnemonaceae (Norris & Koponen B. coronatum Schwaegr. 1990a) (1500-3000 m) Dicnemon novaeguineae (Dix.) B. 89 Allen (2100-3500 m) D. cutlackii Norris & T. Kop. (2800- D. robbinsii (Bartr.) B. Allen (1500- 3500 m) 2000 m) D. dicarpum Nees in Spreng. (1800- 2900 m) Dicranaceae (Frahm et al. 1985; Norris D. geluense (Herz.) Norris & T. Kop. & Koponen 1990a) (1800-3100 m) Atractylocarpus comosus Dix. (2700- D. rugifolium (Bartr.) Norris & T. 3600 m) Kop. (2800-3600 m) A. novoguineensis (Broth. & Geh.) Holomitrium austroalpinum Bartr. Norris & T. Kop. (2450- (2550-3100 m) 3600 m) H. obliquum Salm. (3400-3600 m) Braunfelsia dicranoides (Dozy & H. stenobasis Dix. (2400-3550 m) Molk.) Broth. (1800-3550 m) Microdus miquelianus (Mont.) Besch. B. enervis (Dozy & Molk.) Par. (2450- in Par. (1800-2200 m) 2700 m) Symblepharis reinwardtii (Dozy & Brotherobryum undulatifolium Zanten Molk.) Mitt. (3250-3570 m) (1800-2200 m) Trematodon longicollis Michaux Bryohumbertia walkeri (Mitt.) J.-P. (1750-2450 m) Frahm (1800-2900 m) Campylopodium medium (Duby) Ditrichaceae (Norris & Koponen 1988) Giese (1200-2200 m) Ceratodon stenocarpus Bruch & Campylopus aureus Bosch & Sande Schimp. ex C.Müll. (2430-3600 m) Lac. (2600-3600 m) Distichium capillaceum (Hedw.) C. austrosubulatus Broth. & Geh. B.S.G. (2850-3600 m) (3300-3550 m) Ditrichum colijnii Dix. (3250-3600 m) C. clemensiae Bartr. (1200-3300 m) D. difficile (Duby) Fleisch. (1900-3300 C. comosus (Schwaegr.) Bosch & m) Sande Lac. (1800-3300 m) D. sericeum Bartr. (3400-3570 m) C. crispifolius Bartr. (1500-3300 m) C. exasperatus (Nees & Blume) Brid. Fabroniaceae (Norris & Koponen (1700-3600 m) 1990b) - - var. lorentzii (Fleisch.) J.-P. Frahm Fabronia curvirostris Dozy & Molk. (2500-3400 m) (2070-2200 m) C. involutus (C. Müll.) Jaeg. (750- 2850 m) Fissidentaceae (Norris & Koponen C. macgregorii Broth. & Geh. (3300- 1987) 3550 m) Fissidens asplenioides Hedw. (1480- C. umbellatus (Arnott) Par. (1300- 2500 m) 3600 m) F. braunii (C.Müll.) Dozy & Molk. Dicranella papua-palustris Norris & (300-2600 m) T. Kop. (1900-3570 m) F. flabellulus Thwait. & Mitt. (880- Dicranodontium fleischerianum 2550 m) Schultze-Motel (2330-2500 m) F. flexifolius Bartr. (1100-2400 m) Dicranum arfakianum C. Müll. ex F. geminiflorus Dozy & Molk. (750- Geh. (1800-2500 m) 2200 m) D. armittii C Müll. (1000-3400 m) F. hollianus Dozy & Molk. (0-1600 D. assimile Hampe (900-3500 m) m) D. blumii Nees (1100-3400 m) F. nobilis Griff. (750-2100 m) D. braunii C. Müll. ex Dozy & Molk. F. nymanii Fleisch. (900-1700 m) (1480-2500 m) F. obscurirete Broth. & Par. in Broth. 90 (600-1100 m) Leucophanes angustifolium Ren. & F. plagiochiloides Besch. (2100-2900 Card. (300-1800 m) m) L. glaucum (Schwaegr.) Mitt. (0-20 m) F. rigidulus C. Müll. subsp. L. octoblepharioides Brid. (500-1450 novaguineensis Brugg. m) (1500-2400 m) Octoblepharum albidum Hedw. (0- F. strictulus C. Müll. (1400-2900 m) 1450 m) F. taxifolius Hedw. (1400-2900 m) F. vanzanteni Brugg. (200-2600 m) Meteoriaceae (Norris & Koponen F. wichurae Broth. & Fleisch. (1700- 1985a) 2600 m) Aerobryidium crispifolium (Broth. & F. zippelianus Dozy & Molk. in Zoll. Geh.) Fleisch. (750-1000 m) (100-1300 m) A. fuscescens Bartr. (1400-2200 m) Aerobryopsis longissima (Dozy & Grimmiaceae (Koponen & Norris Molk.) C. Müll. (400-1500 m) 1986) A. leptostigmata (Geh.) Fleisch. (700- Racomitrium javanicum Dozy & 1100 m) Molk. in Zoll. (3250-3550 m) Aerobryum speciosum Dozy & Molk. (1800-2400 m) Hedwigiaceae (Koponen & Norris Barbella cubensis (Mitt.) Broth. (1000- 1986) 1800 m) Rhacocarpus alpinus (Wright) Par. Floribundaria floribunda (Dozy & (3400-3600 m) Molk.) Fleisch. (100-3250 m) R. purpurascens (Brid.) Par. (3400- F. pseudofloribunda Fleisch. (100- 3600 m) 1700 m) F. sparsa (Mitt.) Broth. (1800-3200 m) Hylocomiaceae (Koponen & Norris Meteoriopsis reclinata (C. Müll.) 1985a) Fleisch. (400-2400 m) Leptocladiella flagellaris T. Kop. & M. squarrosa (Hook.) Fleisch. in Norris (1700-3000 m) Broth. (1500-2400 m) Macrothamnium hylocomioides Meteorium buchananii (Brid.) Broth. Fleisch. (1600-3600 m) (400-3200 m) Papillaria flexicaulis (Wils.) Jaeg. Leptostomataceae (Koponen & Norris (1500-2400 m) 1985b) P. fuscescens (Hook.) Jaeg. (1200- Leptostomum intermedium Broth. 2400 m) (2000-3500 m) Pseudobarbella ancistrodes (Ren. & L. perfectum Bartr. (3350-3550 m) Card.) Manuel (500-1900 m)

Leucobryaceae (Enroth 1990a) Mniaceae (Koponen & Norris 1983b) Exostratum blumei (Nees ex Hampe) Orthomnion elimbatum (Nog.) T. L.T. Ellis (100-1750 m) Kop. (350-3300 m) E. sullivantii (Dozy & Molk.) L.T. Plagiomnium cordatum T. Kop. & Ellis (900-1300 m) Norris (2500-3600 m) Leucobryum javense (Brid.) Mitt. P. integrum (Bosch & Sande Lac.) T. (1500-2900 m) Kop. (350-3500 m) L. neilgherrense C. Müll. (1450-1600 - - var. subelimbatum (Dix.) T. Kop. m) & Norris (1250-1900 m) L. sanctum (Brid.) Hampe (1450-2550 P. succulentum (Mitt.) T. Kop. (1550- m) 1900 m)

Neckeraceae (Enroth 1989a) Himantocladium plumula (Nees) Fleisch. (350-650 m) H. submontanum Enroth (1500-2100 m) Homaliodendron flabellatum (Sm.) Fleisch. (300-3350 m) Neckeropsis gracilenta (Bosch & Sande Lac.) Fleisch. (0-800 m) N. lepineana (Mont.) Fleisch. (350- 1900 m) Pinnatella anacamptolepis (C. Müll.)