The origin of a flora is one of the most interesting and challenging topics in bryology. However, it is a very difficult problem, and considerable effort is needed to accumulate more detailed information on the distribution and taxonomy of bryophytes than is available today

Iwatsuki, Zennoske

English

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139Tropical Bryology 2: 139-148, 1990Origin of the New Caledonian Bryophytes“Studies on Oceanic bryophytes 18”Zennoske IwatsukiBotanical Institute, Faculty of Science, Hiroshima University, Higashi-senda-machi,Hiroshima 730andThe Hattori Botanical Laboratory, Obi, Nichinan-shi, Miyazaki-ken 889-25, JapanThe origin of a flora is one of the mostinteresting and challenging topics inbryology.  However, it is a very difficultproblem, and considerable effort is neededto accumulate more detailed informationon the distribution and taxonomy ofbryophytes than is available today.In 1982 there was a short bryologicalexpedition to New Caledonia and Fiji(Iwatsuki & Kitagawa, 1985).  Membersof the party consisted of Dr. N. Kitagawaand myself.  An additional member, Mr.T. Suzuki, a specialist of Fissidens, joinedus for about ten days.  Dr. GordonMcPherson of the Missouri BotanicalGarden, who was in Nouméa at that time,helped us in many ways.  During thissurvey, 4761 specimens of bryophyteswere collected.  We are still studying thespecimens, but so far we have published16 papers on the New Caledonianbryophytes.One of the reasons we selected NewCaledonia for our study area was that somany endemic taxa of bryophytes hadbeen described from that region, as pointedout by Herzog (1926).  The NewCaledonian bryoflora includes manyendemic or remarkable taxa, e.g., the mo-notypic genus Perssoniella, representingPerssoniellaceae, a suborder ofJungermanniales.The small island (20,000 km2) lies aboutthe same distance, nearly 1600 km, fromNew Guinea, Australia, and New Zealand.According to Good (1974) the vascularflora of New Caledonia is oceanic inisolation and continental in structure.  It isof the greatest interest not only for itsmarked endemism but also because of itsfloral relationships.  The vascular floraconsists of 2600 species and 600 genera,which include more than 130 endemicgenera in nearly 30 families.  About halfthe genera are monotypic.  Moreover, thequality of endemism is high; several of thegenera form separate families (5 endemicfamilies).  These are Amboleaceae,Oceanopapaveraceae, Onchothecaceae,Paracryphiaceae, and Strasburgiaceae.The island is comprised partly of igneousrocks, including much serpentine andpartly Permian and younger sedimentaryrocks.  The island was probably separatedfrom Australia 60 - 65 million years ago(end of Cretaceous).There is only one native composite and146Gupta and Udar (1986) reported 21-76um for spores of Indian  species ofFrullania. Origin of Fissidens and Frullania florasof New CaledoniaEvolution of bryophytes is generallythought to be very slow, compared withangiosperms.  However, the rate ofevolution in bryophytes varies amongfamilies and genera.  Schuster (1969)considers that the Hepaticopsida isprobably an old group in which evolutionis generally slow except for a few familiesof mostly epiphytic leafy hepatics.  Hattori(1982) also thinks Frullania evolves ratherfast.As previously explained, I selected twofamilies of bryophytes:  Fissidentaceae(Musci) and Frullaniaceae (Hepaticae) fora study of distribution.  Distributions ofNew Caledonian species of these familiesare remarkably different in number ofendemic species.  Distribution ofFissidentaceae in New Caledonia can beexplained by spore dispersal by wind.Zanten and Pócs (1981) concluded thataerial spore transport over moderately longand long distances within the sameclimatological belt is possible.  This isprobably a common phenomenon forspores smaller than ca. 25 um.  Spores ofmosses are usually smaller than 25 um.  Istudied the moss flora of the Bonin andVolcano Islands (Iwatsuki 1985), andfound that the moss flora of these islandsis very similar to that of the RyukyuArchipelago, which is about 1600 kmwest of the Bonin Islands.  The moss floraof the Bonin and Volcano Islands is notrelated to that of Guam, which is about1450 km southward.  This suggests thatspores of mosses were carried by westerlywinds from the Ryukyu Archipelago tothe Bonin and Volcano Islands.As mentioned before, many species ofFissidens recorded from New Caledoniaare monoicous (autoicous or synoicous)and many specimens are fertile.  Theirspores are small, less than 20 um indiameter.  Many of them occur on newlyeroded soil, trail side banks, etc.  Theirdistribution can be explained by sporedispersal.In 1972, Iwatsuki discussed spore dispersalof mosses, and wrote ‘According to thepresent distribution of bryophytes wecannot assume that wind is a very effectiveagent for long-range dispersal, except fora few weedy species.’  From distributionaldata on New Caledonian Fissidens, manyof them are ‘weedy’ species.As shown in Table 3, many species ofFissidentaceae, especially those ofFissidens, are common to Asia.  Somespecies have pantropical distribution,including Asia, Oceania, the Americas,and Africa.  Fissidens species withpantropical distribution will increase whenwe study more specimens from tropicalAmericas and Africa.  For example,recently Iwatsuki and Suzuki (1982)pointed out that Fissidens mangarevensis(=F. oblongifolius)  might be found intropical America.At the moment, we have only sevenspecies of New Caledonian Fissidentaceaecommon to Australia, but I am sure thisnumber will increase when AustralianFissidens, especially in northern tropicalareas, have been thoroughly investigated.Nanobryum thorsbornei Stone, a veryinteresting tiny species, is recorded only innorthern Queensland and New Caledonia.This species may be a reduced form ofFissidens, and might be better classified inthe genus Fissidens.  The genusNanobryum has been recorded in Africa,Australia, and New Caledonia.As a whole, the New Caledonian Fissidensflora is very closely related to that oftropical Asia.  Long range dispersal by148Species of Frullaniaceae in NewCaledonia are probably Gondwanalandicin derivation.  However, they seem tohave become highly differentiated on thissmall island.  They include Schusterella, asmall genus of the Frullaniaceae whosecenter of differentiation is New Caledo-nia; all three species of Schusterella areendemic to New Caledonia.The subgenus Diastaloba of Frullania isstrongly differentiated, and nine speciesare endemic to the island.  Furthermore,the monotypic subgenus Huerlimannia isknown only from this island (Table 6).The Frullania flora of New Caledoniaincludes only four species that are commonto Australia and New Zealand.  However,there are eleven species common to NewGuinea, nine species to the MalayArchipelago, and nine species to thePacific Islands.  The lesser number ofspecies common to Australia and NewZealand is difficult to explain, but after adetailed study of more specimens fromnorthern Australia, this number might bemuch greater.AcknowledgmentsI wish to acknowledge Dr. G. McPherson and Dr. Bruce Allenof the Missouri Botanical Garden for providing me informationon New Caledonian plants.  I thank Dr. R. A. Pursell ofPennsylvania State University and Dr. J. M. Glime of MichiganTechnological University for help with the English text.Literature CitedAllen, B.  (1989)  Dicnemonaceae in New Caledonia.  Hikobia10:   245-267.Fleischer, M.  (1904)  Die Musci der Flora von Buitenzorg.Bd.   1: 643 pp.  Leiden.Froehlich (1953)  Die von Prof. Dr. Viktor Schiffner in denJahren 1893/94 in Ceylon, Penang, Singapore, Sumatra undJava   gesammelten Laub- und Torfmoose.  Ann. Naturhist.Muse. Wien   59: 66-116.Gangulee, H. C.  (1971)  Mosses of eastern India andadjacent   regions.  Fasc. 2:  171-566.  Calcutta.Good, R.  (1974)  The geography of flowering plants.  4th ed.,557 pp.  Longman Group Ltd., London.Guillaumin.  (1948)  In Good, R.  (1974)  The geography offlowering plants.  4th ed., 557 pp.  Longman Group Ltd.,   London.Gupta, A. & Udar, R. (1986)  Palyno-taxonomy of selectedIndian   liverworts.  Bryoph. Bibliotheca 29: 1-141, pl. I-LIX.Hattori, S.  (1976)  Notes on some species of the NewCaledonian   Frullaniaceae.  Bull. Nat. Sc. Mus. Tokyo, ser. b,2(3):76-86._______.  (1977)  Dr. Hürlimann’s collection of NewCaledonian   Frullaniaceae.  J. Hattori Bot. Lab. 43: 409-438. _______.  (1982)  A synopsis of New Guinean Frullania,Hepaticae.    J. Hattori Bot. Lab. 51: 203-271._______.  (1986)  A synopsis of New Caledonian Frullania-ceae.  J.   Hattori Bot. Lab. 60: 203-237.Hattori,S. & Piippo, S.  (1986)  Bryophyte flora of the HuonPeninsula, Papua New Guinea.  XV.  Frullania (Frullaniaceae,Hepaticae).  Acta Bot. Fennica 133: 25-58.Herzog, T.  (1926)  Geographie der Moose.  439 pp.m Taf. 1-8.   Jena.Iwatsuki, Z.  (1972)  Geographic isolation and speciation ofbryophytes in some islands of eastern Asia.  J. Hattori Bot.   Lab.35: 126-141._______.  (1982)  Speciation of the moss genus Fissidens inNew   Caledonia (preliminary report).  J. Hattori Bot. Lab. 52:113-   126._______.  (1985)  Bryophytes of the Bonin Islands and theVolcano   Islands (5).  Bull. Nat. Sci. Mus. Tokyo ser. b, 11: 57-68.Iwatsuki, Z. & Kitagawa, N.  (1985)  Bryological expeditionto   New Caledonia and Fiji, 1982.  J. Hattori Bot. Lab. 58: 83-86.Iwatsuki, Z. & Mohamed, H.  (1987)  The genus Fissidensin   peninsular Malaysia and Singapore (a preliminary study).  J.Hattori Bot. Lab. 62: 339-360.Iwatsuki, Z. & Suzuki, T.  (1982)  A taxonomical revisionof the   Japanese species of Fissidens (Musci).  J. Hattori Bot. Lab.51: 329-508.Iwatsuki, Z. & Suzuki, T.  (1989)  Fissidens in NewCaledonia.    J. Hattori Bot. Lab. 67: (in press).Iwatsuki, Z. & Tan, B. C.  (1979)  Checklist of Philippinemosses.  Kalikasan, Philipp. J. Biol. 8(2): 179-210.Kitagawa, N.  (1983)  Hepaticae and Anthocerotae of NewCaledonia.  Proc. Bryol. Soc. Japan 3: 119-122 (in Japanese).Kuo & Chiang  (1987) Index of Taiwan mosses. Taiwania 32:119-207.Li, Z. H.  (1985).  A revision of the Chinese species of   Fissidens(Musci, Fissidentaceae).  Acta Botanica Fennica 129,   65 pp.Miyoshi, N.  (1966)  Spore morphology of Hepaticae inJapan.    Bull. Okayama Coll. Sci. 2: 1-46.Norris, D. H. & Koponen, T.  (1987)  Bryophyte flora ofthe Huon   Peninsula, Papua New Guinea.  XX.  Fissidentaceae,Mitteniaceae, Phyllodrepaniaceae, Phyllogoniaceae andSorapillaceae (Musci).  Ann. Bot. Fennici 24: 177-219.Pursell, R. A. & Hoe, W. J.  (1977)  Fissidens in Hawaii.J.   Hattori Bot. Lab. 43: 81-106.Pursell, R. A. & Reese, W. D.  (1982)  The mossesreported from   New Caledonia.  J. Hattori Bot. Lab. 53: 449-482.Sainsbury, G. O. K.  (1955)  A handbook of the NewZealand   mosses.  490 pp.  Dunedin.Schuster, R. M.  (1969)  Problems of antipodal distributionin   lower land plants.  Taxon 18: 46-91.Touw, A.  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Origin of the New Caledonian bryophytes

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