The land areas of the Western Hemisphere south of the United States support over 276 species of Fissidens (Wijk et al. 1962, 1969). This number is approximately 30% of the total number of species known. Progress made on a monograph of the family in the neotropics and the adjacent areas is summarized; approximately 50% of the species have been studied. Commonality among the neotropical, African and Asian species of Fissidens is discussed. Changes to be made in the classification of the family are indicated. New characters used in distinguishing species and the classification of the family are enumerated

Pursell, Ronald A.

English

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193Tropical Bryology 2: 193-200, 1990Fissidens in the NeotropicsRonald A. Pursell208 Mueller Laboratory, The Pennsylvania State University, University Park, PA 16802Abstract. The land areas of the Western Hemisphere south of the United States supportover 276 species of Fissidens (Wijk et al. 1962, 1969). This number is approximately30% of the total number of species known. Progress made on a monograph of the familyin the neotropics and the adjacent areas is summarized; approximately 50% of thespecies have been studied. Commonality among the neotropical, African and Asianspecies of Fissidens is discussed. Changes to be made in the classification of the familyare indicated. New characters used in distinguishing species and the classification of thefamily are enumerated.Resumen. En la region del hemisferio Occidental al sur de los Estados Unidos de NorteAmérica, estan representadas mas de 276 especies de Fissidens (Wijk et al. 1962,1969). Esta cifra corresponde aproximadamente al 30% del total de especies conocidasen este genero. En este trabajo, en el cual se han estudiado aproximadamente el 50%de las especies, se resumen los avances de una monografia de la familia Fissidentaceae.También se discuten las semejanzas entre las especies de Fissidens del neotrópico conlas especies de Africa y Asia. Se indican cambios que deben de hacerse en laclasificación de la familia. Además, se enumeran nuevos caracteres usados para laclasificación de la familia y para distinguir entre especies de la familia.This report consists of a review of theoverall classification of the Fissidentaceaeand a synopsis of the progress made towardproducing a monograph of this family forthe neotropics. As defined by theOrganization for Flora Neotropica, theneotropics consist of “the WesternHemisphere continental land mass lyingbetween the Tropics of Cancer andCapricorn, together with adjacent islandsand including the West Indies andGalápagos.” The part of Mexico north ofthe Tropic of Cancer and the portion ofSouth America south of the Tropic ofCapricorn, inclusive of the Falkland (Mal-vina) Islands in the South Atlantic Oceanand the Juan Fernandez Islands in theSouth Pacific Ocean, are also included.The monograph, thus, will embrace all ofthe Western Hemisphere south of theUnited States.The Fissidentaceae are a monogenericfamily represented by Fissidens andconstitute one of the larger families ofmosses. When established by Hedwig(1801), the genus Fissidens consisted of14 species. On the basis of currentinterpretation of the genus, 10 of these 14species are retained in Fissidens. I have noaccurate count of the number of speciescurrently recognized on a worldwide basis.However, a total of 927 accepted species(inclusive of the species in segregategenera) are listed in Index Muscorum(Wijk et al. 1962, 1969). Remarkably,these hundreds of species aredistinguished, for the most part, on195Robinson (1970). Perhaps the best knownand most widely accepted interpretation isthe one originated by Brown (1819) butchampioned by Salmon (1899) and mostrecently by Mishler (1988). In brief,according to this interpretation, thevaginant laminae (the lamina vera)represent the true leaf and the dorsal andventral laminae are an outgrowth fromthis. Robinson, however, elaborating onan interpretation originated by Spruce(1881), considers the whole leaf ofFissidens as the true leaf whose form hasbeen achieved by the “reorientation of themitotic spindle of the apical cell of the leaf. . .” It is not the intent of this report tochoose between these interpretations. Thepoorly developed additional laminae thatare infrequently found on leaves [viz. anunrecorded specimen of F. taxifoliusHedw. Sharp 2405b (PAC), and F.bourgaeanus Besch., see Figure 3] indeedcan support either of these twoexplanations.The need to approach the study of mossesfrom a worldwide perspective has beendemonstrated a number of times, e.g., bythe work of Frahm (1982) on the widelydistributed and taxonomically difficultgenus Campylopus, by the work of Ochi(1972, 1980) on the family Bryaceae, andrecent work on Fissidens subgeneraOctodiceras and Sarawakia (Pursell 1987,Pursell et al., 1988). Knowledge of therelationship between the moss floras ofAfrica and the neotropics and between theneotropics and Asia is increasing (for areview of the relationship between Africaand the neotropics see Reese 1985). Oneexample of a species of Fissidens describedoriginally from the neotropics that ispantropical is F. asplenioides Hedw. Theneotropical species of F. kegelianus C.Müll. is conspecific with the Asiatic F.zollingeri Mont. (Pursell 1979). Moreover,this species is also known from Africa(Bizot & Pócs 1979, Kis 1985, Sappa &Piovano 1947) and, as species from otherareas are studied and names synonymized,probably will be found to be pantropical.A third example is F. garberi Lesq. &James, a species of broad distribution inthe neotropics which extends northwardinto the United States. Iwatsuki and Suzuki(1982) reduced this species to thesnynonymy of F. microcladus Thwait. &Mitt. which occurs throughout much ofAsia (see also Iwatsuki 1982 and Li 1985).Although F. microcladus has not beenrecognized from Africa, Dury’s (1974)review of the African species in sectionSemilimbidium certainly will facilitate theproblem of determining synonymy amongthe many species described.During July 1989, Drs. M. A. Bruggeman-Nannenga and Zennoske Iwatsuki, twoeminent specialists on the systematics ofthe Fissidentaceae in Africa and Asia,respectively, were able to come to theUnited States and, with me, addressproblems about the commonality of speciesin the neotropics, Asia and Africa. Theoverall classification of the familyFissidentaceae was also discussed. Asmentioned below, results of this highlysuccessful period will be published soonin a series of papers.The classification of the Fissidentaceaemost widely used today was published byBrotherus (1901, 1924, see Table I for anoutline of the scheme), based essentiallyon an earlier one by Müller (1901). In thisclassification four genera are recognized,Fissidens, Fissidentella, Moenkemeyeraand Simplicidens. A more recentclassification, based in part onchromosome number correlated withsexuality, was published by Iwatsuki(1985, see Table 2 for an outline of thissystem). According to Iwatsuki, subgeneraAneuron (= Polypodiopsis sensuBrotherus) and Fissidens are characterizedby n = 10 or n = 12 in monoicous specieswhile subgenera Pachyfissidens andSerridium (= section Serridium sensuBrotherus) have n = 12 in dioicous species.198In the restricted sense, Octodicerasconsists of only those aquatic speciescharacterized by rather lax cells in longlinear leaves that are extremely fragilewhen dry.Subgenus Fissidens is the largest and mostdiversified of the subgenera of genusFissidens. The 12 sections recognized byBrotherus are now considered to bedistinguished along unnatural lines. Thestatus of these sections, not discussed byIwatsuki (1985), was recently reviewed(Pursell 1988). Problems relative to someof the sections of Fissidens were resolvedin the discussions in July. At this time, itcan be noted that the resolution of theseproblems involved the use of charactersheretofore not used or seldom used. Forexample, Allen (1980) published a shortpaper describing peristomal variationsobserved in an SEM study of 19 species ofFissidens. Bruggeman-Nannenga &Berendsen (1988) refined and expandedthis work to include 200 speciesrepresenting all subgenera and sections ofthe family.The structure of the costa, heretofore notemphasized to any great extent, appears tobe of considerable diagnostic value.Bruggeman-Nannenga has been studyingthe costa extensively and will soon beprepared to publish on her observations.Moreover, section Amblyothallia,heretofore rather difficult to define, can berecognized readily by the structure of thecosta. A paper dealing with thecircumscription, synonymy, relationship,distribution and illustrations of this sectionwill be published soon.The number of exothecial cells in thecircumference of the theca (Bruggeman-Nannenga & Berendsen 1988) and theaxillary hyaline nodules (Iwatsuki &Pursell 1980), although less important,nevertheless are also useful.Two additional papers will be publishedalso by Iwatsuki, Bruggeman-Nannengaand myself. One will be a world-widerevision of the ‘ecostate’ species ofFissidens, i.e., subgenus Aneuron. Thesecond, based somewhat on Iwatsuki’s(1985) scheme will address the overallclassification of the Fissidentaceae abovethe species level.There are 276 species of Fissidens recordedin Index Muscorum in the area of thewestern hemisphere to be included in themonograph. This is approximately 30%of the total number of 927 species statedpreviously. According to a study by Touw(1974), modern monographic studies havereduced the number of species by 20-45%. If this is the case in Fissidens thefinal number of species recognized in thewestern hemisphere south of the UnitedStates should be between 221 and 152.It has become increasingly evident duringthe course of my studies that subtledifferences, often thought to be highlyvariable, are remarkably consistent andare useful in distinguishing species. Forexample, Fissidens fontanus was thoughtto be a highly variable and widespreadspecies. In this complex (Pursell 1987),not only are there good differences in theperistome but gametophytically such cha-racters as vaginant laminae length/leaflength, the nature of the distal end of theminor lamina (a term introduced byRobinson 1970) and even costa length,are important in distinguishing species.The pluripapillose species in section Se-milimbidium (including species fromsection Pycnothallia) are another example.In the neotropics these species are centerednomenclaturally around F. elegans Brid.Florschütz (1964) recognized only twospecies, but as a recent study (Pursell1984) indicates, differences important indistinguishing species include: 1) thethickness of limbidia; 2) the length oflimbidia; 3) the kinds of stems (sterile,perichaetial and perigonial) on whichlimbate leaves are found; and 4) the199particular leaves of a stem on whichlimbidia are found.As stated earlier, it is becoming increasinglyapparent that large and extensive generaof bryophytes must be studied from aworldwide perspective. From discussionswith Iwatsuki and Bruggeman-Nannengait is apparent that several species ofFissidens of frequent occurrence in theneotropics, in addition to those alreadymentioned, do not differ from rathercommon species found in Asia and Africa.Species already so identified will be enu-merated in a paper soon. As workprogresses this number of species reflectingdegrees of commonality will surelyincrease.I estimate that 50% of the work on themonograph is near completion. Most majorgroups have been studied to some extent.Work on sections Aloma and Crenulariais nearing completion. Much remains tobe done with section Fissidens (includingsection Heterocaulon) and the unipapillosespecies of section Semilimbidium. Within3 years, or at most 4 years, the monographshould be completed.Acknowledgement.The National Science Foundation is gratefully acknowledged forGrant BSR-8905431 to The Pennsylvania State University, whichnot only made this report possible, but also afforded the means tobring Drs. M. A. Bruggeman-Nannenga and Z. Iwatsuki to theUnited States for consultation and cooperative study.LITERATURE CITEDAllen, B. H. 1980. Peristome variations in the genus Fissidens:an SEM study. Bryologist 83:314-327._____. 1981. A reevaluation of the Sorapillaceae. Bryologist84:335-338.Bizot, M. 1963. Nanobryum gladiolum (Mitt.) Biz. comb. nov.Revue Bryologique et Lichnologique 32:82-83._____ & T. Pócs. 1979. East African Bryophytes, III. ActaBotanica Academiae Scientiarum Hungaricae 25:223-261.Brotherus, V. F. 1901. Fissidentaceae. In: A. Engler & K.Prantl, Die natürlichen Pflanzenfamilien 1(3):351-363. Leipzig._____. 1924. Fissidentaceae. In: A. Engler & K. Prantl, Dienatürlichen Pflanzenfamilien, ed. 2, 10:143-155. Leipzig.Brown, R. 1819. Characters and description of Lyellia, a newgenus of mosses, with observations on the section of the order towhich it belongs; and some remarks on Leptostomum andBuxbaumia. 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A new approach to the classification of the Fissidentaceae

A preliminary study of the Fissidens elegans complex in the neotropics.

A reevaluation of the Sorapillaceae.

A revision of African Bryoideae. Musci (first200 part).

A revision of the Chinese species of Fissidens.

A revision of the neotropical Bryoideae. Musci (first part).

A taxonomic revision of Fissidens subgenus Octodiceras (Fissidentaceae).

A taxonomic revision of Fissidens subgenus Sarawakia (Bryopsida: Fissidentaceae).

A taxonomic revision of the Japanese species of Fissidens (Musci).

Axillary hyaline nodules in Fissidens (Fissidentaceae).

Characters and description of Lyellia, a new genus of mosses, with observations on the section of the order to which it belongs; and some remarks on Leptostomum and Buxbaumia.

Classification of the Bryopsida.

Die Musci der Flora von Buitenzorg.

Die natürlichen Pflanzenfamilien 1(3):351-363.

Etude taxonomique des espèces africains de la section Semilimbidium C. Müll. du genre Fissidens Hedw. Thése. Université de Dijon.

Genera Muscorum Frondosorum.

La raccolte briologiche dei

Mosses of south-east tropical Africa. An annotated list with distributional data. 170 p.

Nanobryaceae - ein neuer Name für eine Familie der Laubmoose.

Nanobryum gladiolum (Mitt.)

Nanobryum thornesbornei, a remarkable new moss from Australia.

Observations on the origin of the specialized leaves of Fissidens and Schistostega.

On Fissidens. Nomenclatural and distributional notes on some American species.

On the genus Fissidens. Annals of Botany,

On the peristome types found in the Fissidentaceae and their importance for the classification. In:

Peristome variations in the genus Fissidens: an SEM study.

Relationships between ontogeny and phylogeny, with reference to bryophytes.

Sainsburia, a new genus of Fissidentaceae.

Some notes on taxonomic and floristic research on exotic mosses.

Speciation of the moss genus Fissidens in New Caledonia (preliminary report).

Species Muscorum Frondosorum.

The morphology of the leaf of Fissidens.

The mosses of Suriname. Part I.

The taxonomic status of the Nanobryaceae (Bryopsida).

Tropical lowland mosses disjunct between Africa and the Americas, including Calyptothecium planifrons (Ren. & Par.) Argent, new to the Western Hemisphere. Acta Amazonia, Suplemento

Fissidens in the Neotropics

Fissidens in the Neotropics

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