Howard Crum

Mosses of Eastern North America

IntroductionVOLUME 1Class SphagnopsidaClass AndreaeopsidaBryopsidaSubclass ArchidiidaeSubclass BryidaeOrder FissidentalesOrder ByroxiphialesOrder SchistostegalesOrder DicranalesOrder PottialesOrder GrimmialesOrder FunarialesOrder BryalesVOLUME 2Order IsobryalesOrder HookerialesOrder HypnobryalesSubclass BuxbaumiidaeSubclass TetraphididaeSubclass PolytrichidaeConspectus of TaxaGlossaryBibliographyIndex

Disjunctions in Bryophytes

In spite of the more general distribution of many bryophytes, dramatic disjunctions exist, many of them similar to those shown by vascular plants. Various explanations have been offered to explain these disjunctions including continental drift, long-distance dispersal, and the fragmentation of a once more continuous distribution. No single hypothesis is sufficient to accommodate all species within any disjunctive pattern. The most serious difficulty is the inadequacy of exploration of considerable areas of the globe. Various bryophyte disjunctions are mapped in 51 distribution maps, and details of sexual patterns and dispersal mechanisms are assessed. Most bryophytes are widely distributed. In the Northern Hemisphere more than 60% of the flora of arctic and boreal regions is made up of the same species. Within this wide range, however, each species has highly specific requirements and some are exceedingly local. Because bryophytes have air-borne diaspores their means of dissemination would appear to guarantee a wide distribution of all species. That disjunctions exist at all would seem somewhat anomalous, yet such disjunctions do exist, some of them very dramatic. The explanation of these disjunctions has led to numerous intriguing hypotheses, many of which have been derived from similar studies of flowering plant disjunctions. In North America the disjunctions that have received the greatest attention are eastern American-East Asian disjuncts (Iwatsuki, 1958a, 1958b, 1958c; Iwatsuki & Sharp, 1967, 1968; Sharp & Iwatsuki, 1965. See Maps 1-7); tropical and subtropical taxa in the Southern Appalachians (Anderson, 1951; Andrews, 1920; Billings & Anderson, 1966; Crum, 1966; Sharp, 1936, 1938, 1939, 1941. See Maps 8-9); amphi-Pacific taxa (Ando, 19166; Ando, Persson & Sherrard, 1957; Crum, 1965; Hattori, 1952, 1963, 1966b; Hattori & Sharp, 1968; Hattori et al., 1968; Horikawa & Ando, 1957; Lazarenko, 1957; Noguchi & Saito, 1970; Persson, 1946a, 1946b, 1947, 1949, 1952, 1958, 1962, 1968; Persson & Gjaervoll, 1957; Schofield, 1962, 1965, 1966a, 1966b, 1966c, 1968a, 1968b, 1969; Sharp & Hattori, 1967; Steere, 1969; Steere & Schofield, 1956; Steere & Schuster, 1960. See Maps 10-20); bipolar disjuncts (Martin, 1946, 1949, 1952a, 1952b; Du Rietz, 1940; Sainsbury, 1942; Schuster, 1969. See Maps 21-22); taxa disjunctive between Europe and western North America (Abramova & Dildarin, 1969; Evans, 1914; Harvill, 1950; Haynes, 1915; Koch, 1956; Paton, 1966; Schofield, 1968a, 1968b, 1969; Whitehouse, 1961, 1963. See Maps 23-33); amphi-Atlantic taxa (Andrews, 1938, 1961; Maass, 1965, 1966a, 1966b; Schuster, 1962; Sharp, 1941. See Maps 34-37); and arctic-alpine taxa, that have received surprisingly little attention (Gams, 1955; Schuster, 1958a, 1958b; Steere, 1937, 1938, 1953, 1965. See Maps 38-40). 1 Department of Botany, University of British Columbia, Vancouver 8, British Columbia, Canada. 2 The Herbarium, University of Michigan, Ann Arbor, Michigan 48104. ANN. MISSOURI BOT. GARD. 59: 174-202. 1972. This content downloaded from 157.55.39.183 on Thu, 26 May 2016 04:31:58 UTC All use subject to http://about.jstor.org/terms 1972] SCHOFIELD & CRUM-DISJUNCTIONS IN BRYOPHYTES 175 Irmscher (1929) studied moss disjunctions particularly in their contribution to an insight reflecting plant distributions as influenced by continental drift. Herzog (1926) in his monumental Geographie der Moose treats bryophyte disjunctions briefly. Details from other areas are treated by Abramova & Abramov (1969), Bowers (1970), Croizat (1962, 1966), Crum (1966), Crum & Anderson (1964), Fulford (1951), Greig-Smith (1950), Horikawa (1955), Koch (1954), Lazarenko (1957), Muller (1916, 1954), Schelpe (1969), Schofield (1969), and Schuster (1967,1968,1969). Hypotheses explaining these disjunctions have been based mainly on historical factors. In most cases the opinion is that these various disjunct populations represent remnants of a more continuous distribution of the past, always placed preceding the Pleistocene glaciations and usually suggested to be as early as the Tertiary. Information from fossil bryophyte material is generally sparse; thus the assumption is made that bryophytes were associated with a vascular vegetation which exhibited a more continuous distribution during the Tertiary. One bryophyte showing disjunctive European-western North American distribution is Claopodium wchippleanum (Noguchi, 1952. See Map 33). The species is also present in the Hawaiian Islands. Abramova and Abramov (1969) indicate that they discovered it as fossil material from Middle Pliocene deposits in the Caucasus region. In Europe the species is presently restricted to a number of localities in Portugal and Spain. Another species of similar distribution, but with the Caucasus showing the only extant European population, is Crumia latifolia (see Map 26). This is discussed by Abramova and Dildarin (1969). These distributions strongly suggest a more continuous distribution of the past. Remarkable features of a high proportion of disjunctive species is that they are dioicous and in many cases produce no specialized propagula that would make them readily disseminated. This complete reliance on asexual reproduction by simple fragmentation leads to a paucity of biotypes. These combined features mean that the disjuncts tend to be, within their disjunctive areas, highly localized as relatively small populations with very specific environmental requirements. In most cases they are very poor competitors with the more generally distributed flora and are often confined to sites that are continually undergoing ecesis (Lye, 1967). Areas particularly rich in disjunctive taxa often have hyperoceanic climates. This subject has been treated thoroughly by Ratcliffe (1968) and Stormer (1969) in particular, but also discussed by Amann (1 929), Billings and Anderson (1966), Courtejaire (1962), Gaume (1952-1954), Iwatsuki (1958a), Lye (1967), Nicholson (1930), Persson (1949), Schuster (1962), and Touffet (1964). The areas richest in disjunctive species confined largely to hyperoceanic climates are: Britain and Norway, the Faeroes, the Alps, the Himalayas, high mountains of Japan and Taiwan, North Pacific North America, and to a certain degree, high mountain slopes in the Hawaiian Islands. In some cases identical disjunctive species are found in many of these widely separated areas, for example: Geheebia gigantea (Map 41), Mastigophora twoodsii (Map 42), Scapania ornithopodioides (Map 43), Anastrepta orcadensis (Map 44), Anastrophyllum doonianum (Map 45), Pleurozia purpurea (Map 46), Bazzania pearsonii, Cephaloziella pearThis content downloaded from 157.55.39.183 on Thu, 26 May 2016 04:31:58 UTC All use subject to http://about.jstor.org/terms 176 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. 59 sonii, Campylopus atrovirens, and others. Species of the East Asian-eastern American disjunction, the European-western American disjunction and amphiPacific disjunction are also largely of oceanic climates. In the Southern Hemisphere disjunctions are also richly represented in hyperoceanic climates: the southern Australasian-southern South American disjunction is the most pertinent example, shown in Maps 47-48 (see especially

Miscellaneous Notes on the Genus Sphagnum. 1-2

An account is given of 10 previously described South American Sphagna, including one that is elevated from varietal to specific rank and named S. cribriforme Crum (S. perforatum var. rotundifolium Warnst.). Through the kindness of Allan Fife, I was able to study a number of South American specimens from the Botany Institute of the Department of-Scientific and Industrial Research at Christchurch, New Zea- land (CHR). Among them were a number of isotypes, as well as other authentically named specimens cit- ed by Warnstorf in his Sphagnologia Universalis (1911), particularly specimens from Brazil collected by Ernst Ule. Because many of Warnstorfs types are no longer in existence or are represented only as duplicates in herbaria around the world, an ac- count of these specimens seems warranted. All taxa are Brazilian, except as noted.

Notes on tropical American mosses

Nomenclatural novelties include 12 new combinations in Schizomitrium, as well as Tristichium apodum (Herz. as T. lorentzii var.) n. stat.; Dicranella calcarea (Bartr., as Dicrano- weisia) n. comb. (Guatemala); Cynodontium guatemalense (Bartr., as Oncophorus) n. comb. (Gua- temala); Thudium delicatulum var. peruvianum (Mitt., as sp.) n. stat.; Barbula jacksharpii n. sp. (Guatemala); Lepyrodon duellii n. sp. (Mexico); Curviramea n. gen. & C. mexicana (ThOr., as Pilotrichum) n. comb. (Mexico); Callicostaceae nom. nov. for Pilotrichaceae; Rigodiaceae n. fam.; Bryohaplocladium praelongum (Schimp. ex Besch., as Pseudoleskea) n. comb. (Arizona to Bolivia); Thuidium sharpii n. sp. (Mexico and Guatemala); Pylaisiadelpha brasiliensis n. sp. (Brazil). Di- plostichum Mont. should replace Eustichia (Brid.) C. M. Meiothecium commutatum and Pty- chomitrium chimborazense are recorded as new to Mexico, and Acritodon nephophilus is found to have peristome segments alternating with the teeth. In the course of many years of interrupted study on tropical American mosses, I have accumulated a miscellany of nomenclatural and taxonomic nov- elties, together with some observations worthy of note. These are presented here partly as a contri- bution prefatory to the publication of a Mexican moss flora long in preparation by A. J. Sharp and numerous collaborators, including myself.

A New Species of Cryptothallus from Costa Rica

Cryptothallus hirsutus is described as a new species from Costa Rica differing from C. mirabilis of oceanic northwestern Europe in its indehiscent capsules with no elaterophore and spores not coherent as tetrads. The faintly green gametophytes are particularly hairy and rhizoidal on both surfaces. The wide disjunction of these two underground species, surely disadvantaged in spore dispersal, supports recognition at a specific level. Curiously non-green, much branched thalli found underground on the Cerro de la Muerte in Costa Rica were presumed to be gametophytes of a Ly- copodium, sporophytes of which were associated with them. Joseph Beitel, who was present at the time of discovery, recognized them as a Crypto- thallus, and on closer examination, they indeed proved to be a thallose liverwort of the Metzgeri- ales similar to Aneura in having fleshy thalli and shoot calyptrae that enlarge and protect sporophytes throughout their growth and development. In sub- terranean habitat, pale, coralloid aspect, and struc- ture, the plants (Figs. 1, 3-4) greatly resemble

Revision of the Genus Rozea (Musci)

Rozea is a small genus of pleurocarpous mosses which occurs in the highlands of Mexico, Central America, northern South America and Himalayan Asia. The species are, at best, ill-defined, and the genus has been overclassified. We recognize R. chrysea Besch. (syn. R. stricta Besch.), R. chrysea fo. andrieuxii (C. Muell.) stat nov., R. chrysea var. bourgaeana (Besch.) stat. nov. (syn. R. schimperi Besch. and R. viridis Besch.) and R. subjulacea Besch. in the Americas. The Asiatic species are R. pterogonioides (Harv.) Jaeg. (syn. R. myura Herz.), R. microcarpa Broth., R. microcarpa fo. elongata fo. nov. and R. kenoyeri n. sp. (from Uttar Pradesh, India). Rozea diversifolia Broth. appears to be a Homalothecium. Rozea roseorum Williams is a Juratzkaea species. Micrographs of peristomes provide evidence that Rozea may have some relation- ship to the hypnoid group of families rather than to the Entodontaceae. Rozea is a small moss genus which occurs disjunctively in the highlands of tropical America, from Mexico to northern South America, and in Himalayan Asia. The species are difficult to define, especially in sterile condition, but the genus is easily recognizable. The plants, in aspect similar to Brachythecium or Orthothecium, are relatively slender and shiny, golden to red-bronze or sometimes (apparently in shade forms) green. Creeping stems are freely branched, and the branches are subjulaceous, ascending and often curved. Leaves characteristically have two well-marked plicae or a single median fold. Leaf margins are revolute, nearly throughout, and serrulate near the apex. The costa is single but sometimes forked. Leaf cells are smooth and linear-rhomboidal in the upper regions but short and broad in two to six rows across the insertion. Alar regions extend downward as short decurrencies which merge with short, broad, hyaline cells of the stem epidermis which strips off with the leaves on dissection. Capsules are erect and sym- metric, with a somewhat reduced peristome of hypnoid structure (Fig. 5-6). The exostome teeth are cross-striolate below. Segments of the endostome which are rather broad and keeled, arise from a fairly well-developed basal membrane. Cilia are lacking or rudimentary. Because of the hypnoid nature of the peristome, a relationship to Entodon seems questionable. Even though Fleischer (1915), Brotherus (1925) and more recent authors, e.g. Mizushima, 1960, included Rozea in the Entodontaceae, it may have closer ties with some of the Hypnaceae with erect capsules and variously reduced peristomes, such as Ctenidiadelphus cylindricarpus (Card.) Bartr. (Fig. 3-4). In its best development the hypnoid peristome has the following features: exostome teeth abruptly tapered, bordered

Taxonomic and Nomenclatural Notes on Middle American Mosses

The family Rigodiaceae is described as new to accommodate the genus Rigodium, formerly in the Lembophyllaceae. Trichostomum stenocarpum (Ther.) n. comb. is transferred from Weisiopsis. Rhizogonium sublimbatum is described as a new species from Costa Rica. Study leading toward completion of the Mexican moss flora, under the direction of A. J. Sharp, has brought forth a few taxonomic and nomenclatural matters that are given consideration below. These include the recognition of a new family, the Rigodiaceae, which is removed from the poorly defined Lembophyllaceae and shifted to the Hypnales (= Hypnobryales) because of peristomial differences from the Isobryales. A new species of Rhizogonium having thickened leaf margins is described from a Costa Rican type, and the Mexican Weisiopsis stenocarpa Th6r. is transferred to the genus Trichostomum be- cause of bifid peristome teeth.

Sphagnum inretortum, a new species in a new section from Bolivia

A new species from Bolivia, Sphagnum inretortum, is assigned to a new section of the genus-sect. Inretorta--because of branch leaves neither truncate nor toothed at the apex but margined by a resorption furrow and epidermal cells of branches uniform in shape and size and lacking

Three New Moss Species from the Hawaiian Islands

Fissidens kilaueae from the island of Hawaii, and Anoectangium rubrigemmium and Ulota cervina from Maui, are described as new. This Ulota represents the first record of the genus in Hawaii. Fissidens kilaueae Hoe & Crum, sp. nov. (Fig. 1-5) Plantae dispersae vel gregariae, perpusillae, albidae. Folia 3-5-juga, usque ad 0.8-0.9 mm long, sicca non contorta, oblongo-lanceolata, acuta vel acuminata, toto ambitu grosse dentata, non limbata; costa pellucida, 2-4 cellulas sub apicem evanida; cellulae superiores 6-9t, hexagonae, valde unipapillosae. Autoicus; plantae masculae minutae, plerumque prope basin plantarum foeminarum. Calyptra parva, mitriformis, ubique scabrida. Very small, whitish plants about 1-1.5 mm high (including leaves), scattered. Leaves in 3-5 pairs, small below, progressively larger toward the stem tip, the upper-

Bestia, Tripterocladium, and Isothecium: an explication of relationships

Bestia is divided into two components, the lectotype species, B. longipes, going to the Brachytheciaceae and the other, B. vancouveriensis, to the Thamnobryaceae under a new combi- nation as Porotrichum. Tripterocladium is positioned in the Hypnaceae and Isothecium in the Brachytheciaceae. Bestia, Tripterocladium, and Isothecium, from their inception have had a history of confusion one with another. They have now been purged of extra- neous elements, except in the case of Bestia, so that the genera have gained in definition and meaning. However, the familial linkages need some elucida- tion.