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Floristics of the South American Páramo Moss Flora

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Floristics of the South American Páramo Moss Flora CORE Metadata, citation and similar papers at core.ac.uk Provided by Hochschulschriftenserver - Universität Frankfurt am Main 127 Tropical Bryology 2: 127-132, 1990 Floristics of the South American Páramo Moss Flora Dana Griffin, III Department of Botany & The Florida Museum of Natural History, University of Florida, Gainesville, Florida 32611 Abstract. The South American paramos appeared in Pliocene times and persist to the present day. The moss flora of this habitat consists of an estimated 400 species that comprise 8 floristic groups. In Venezuela these groups and their percent representation are as follows: neotropical 37%, Andean 26%, cosmopolitan 18%, Andean-African 8%, neotropical-Asiatic 3%, neotropical-Australasian 2%, temperate Southern Hemisphere 2% and northern boreal-temperate 2%. Acrocarpous taxa outnumber pleurocarps by nearly 3:1. The neotropical and Andean floristic stocks likely were present prior to late Pliocene orogenies that elevated the cordillera above climatic timberlines. These species may have existed in open, marshy areas (paramillos) or may have evolved from cloud forest ancestors. Taxa of northern boreal- temperate affinities, including those with Asiatic distributions, probably arrived in the paramos during the Pleistocene, a period which may also have seen the establishment in the Northern Andes of some cosmopolitan elements. Species with temperate Southern Hemisphere and Australasian affinities likely spread first to austral South America thence migrated northward during a cool, moist interval sometime over the past 2.5-3 million years or may have become established in the paramos as a result of long- distance dispersal. Reconstructing a floristic history for the pollen of Andean forest taxa. The páramo mosses of South America requires establishment of this chronology for the addressing a number of questions. When páramo habitat has important implications did the páramo life zone appear and for floristic reconstructions since it sets the supposedly become available to time frame within which the earliest colonizing species? Here we can be rather colonists must have arrived. It does not precise. From the pioneering work of establish; however, where the páramo Thomas van der Hammen and his moss species (or their antecedents) were coworkers (cf. van der Hammen 1974; in the period prior to the development of a van der Hammen & Cleef 1983, 1986), it páramo system (although we can speculate is clear an upper elevational timberline did on this point) nor does it tell us exactly not develop in the Northern Andes before how old a particular páramo species is. late Pliocene time (ca. 2-4 million years B.P.). The development of a páramo What are the broad outlines of the páramo system is revealed by a study of pollen moss flora? Here again, we are in a position deposited in lake sediments and is signa- to give a fairly detailed answer. led by a sharp rise in the frequency of Accelerated field work over the past 25 pollen types assignable to the Poaceae and years in the principal páramo countries of Asteraceae and a concomitant decrease in Ecuador, Colombia and Venezuela has 128 brought into herbaria thousands of fresh Colombia or Ecuador, a distinction that collections a majority of which have been will undoubtedly be reflected in the parti- identified at least to the level of genus. cular percent representation of the Much of this identification work remains following floristic groups in those to be published, but, in general terms, we countries. can observe that the páramo moss flora is distributed among approximately 45 The diverse nature of the páramo flora in families, 120-140 genera and Venezuela can be appreciated from the approximately 400 species. The great ma- fact that while this habitat occupies 4,100 jority of the species belong to acrocarpous square kilometers (Jahn 1931), it represents groups. Acrocarpy appears to be favored less than 1% of the Venezuelan national over pleurocarpy in exposed environments territory yet supports 39% of the total as a check of some selected floras confirms. known moss flora for the country. There For the following floras an acrocarp/ are approximately 700 species of Musci in pleurocarp ratio is given (numbers Venezuela (Griffin 1975, 1977a, 1977b, represent percent of total flora) along with 1979; Griffin, Lopez & Ruiz-Teran 1973; the references consulted: Alaskan tundra Pursell 1973) of which 272 (Griffin, this 75/25 (Steere 1978), alpine zone of Mexico paper) have been found above timberline 93/7 (Delgadillo 1971, 1979, 1982, 1984, in the Venezuelan Andes. 1986, 1987), páramos of Costa Rica 67/ 33 (Bowers 1974), páramos of Venezuela The páramo moss flora of Venezuela can 72/28 (Griffin, this paper), Antarctica 81/ be partitioned into 8 floristic groupings. 19 (Robinson 1972). Vitt (1979) reported These are (with percent representation): a preponderance of acrocarpous taxa in neotropical (37%), Andean (26%), polar floras of both hemispheres and in the cosmopolitan (18%), Andean- African temperate floras of the Southern (8%), neotropical-Asiatic (3%), Hemisphere with a shift toward more neotropical-Australasian (2%), temperate pleurocarpous species in tropical floras Southern Hemisphere (2%) and northern and those of temperate latitudes in the temperate/boreal (2%). Northern Hemisphere. While valid as generalizations, these trends can change Neotropical Element for more localized regions or habitats. I This component of the páramo flora believe the more universal correlative is consists of those taxa of wide distribution degree of exposure (i.e., likelihood of in the neotropics. Few, if any, are restricted periodic drying), regardless of which to habitats above timberline and likely hemisphere is examined. Forested areas were present before the appearance of a tend to support pleurocarp-rich floras, as páramo life zone. These species evidently seen in the case of the Great Lakes area 53/ were preadapted to surviving under the 47 (Crum 1976) or Mont Bokor, more stressful conditions of the páramo Cambodia 41/59 (Tixier 1979), whereas environment. Included in this element are relatively treeless areas appear to favor species found in forests, forest clearings acrocarps. and savannahs. Some representatives of this group are: Sphagnum meridense The high elevation moss flora of Venezuela (Hampe) C. Müll., S. sparsum Hampe, has been studied in some detail, and it can Fissidens repandus Wils., Ditrichum be used as a model for discussing floristic rufescens (Hampe) Broth., Campylopus groups throughout the Andean páramo heterostachys (Hampe) Jaeg., Pilopogon life zone. It should be recognized; however, gracilis (Hook.) Brid., Bryoerythrophyl- that the Venezuelan páramos are lum jamesonii (Tayl.) Crum, Entosthodon comparatively drier than those found in jamesonii (Tayl.) Mitt., Bryum 129 beyrichianum (Hornsch.) C. Mül., Bryum chadelphus ciliatus (Hook. f. & Wils.) grandifolium (Tayl.) C. Müll., Pohlia Mitt. papillosa (C. Müll. ex Jaeg.) Broth., Bartramia potosica Mont., Ptychomitrium Cosmopolitan Element chimborazense (Mitt.) Jaeg., Macromi- trium guatemalense C. Müll., The cosmopolitan element of the páramo Orthotrichum pycnophyllum Schimp., moss flora contains a diverse assortment Zygodon pichinchensis (Tayl.) Mitt., of species which, while sharing Lepyrodon tomentosus (Hook.) Mitt., multicontinental distributions, are Porotrichum longirostre (Hook.) Mitt., otherwise very different. The time of their Daltonia gracilis Mitt., Rigodium toxarion appearance in the páramos and the migra- (Schwaegr.) Jaeg., Brachythecium tory tract by which they arrived are virtually stereopoma (Spr. ex Mitt.) Jaeg., impossible to ascertain. Some were surely Plagiothecium conostegium Herz., present prior to the development of the Catagonium brevicaudatum C. Müll. ex páramos, others may have arrived by long- Broth., Aptychella proligera (Broth.) distance dispersal while still others Herz., Sematophyllum swartzii undoubtedly came in stages utilizing (Schwaegr.) Welch & Crum and Hypnum stepping stone habitats that must have amabile (Mitt.) Hampe. been available during the period of the Pleistocene. Included in this group are Andean Element species which produce sporophytes readily This is a montane component which, like and others for which only gametophytes the neotropical element, most likely was are known. Some representatives are: in place before the appearance of the Sphagnum magellanicum Brid., Andreaea páramo habitat. As such, most of these rupestris Hedw., Fissidens asplenioides species occur in Andean forests as well as Hedw., Ceratodon stenocarpus BSG, páramos. Some typical members of the Distichium capillaceum (Hedw.) BSG, group are: Pleuridium andinum Herz., Campylopus flexuosus (Hedw.) Brid., An- Campylopus argyrocaulon (C.M.) Broth., oectangium aestivum (Hedw.) Mitt., Campylopus cucullatifolius Herz., Kin- Didymodon vinealis (Brid.) Zander, giobryum paramicola H. Robins., Grimmia ovalis (Hedw.) Lindb., Funaria Oreoweisia tunariensis Herz., Aloinella hygrometrica Hedw. var. calvescens venezuelana Griffin, Leptodontium (Schwaegr.) Mont., Anomobryum filifor- erythroneuron Herz., L. stellaticuspis me (Dicks.) Husn., Bryum argenteum Bartr., Racomitrium steerei Griffin, Hedw., Bryum billardieri Schwaegr., Splachnum weberbaueri Reimers, Aulacomnium palustre (Hedw.) Tayloria rubicaule A. Kop., Acidodontium Schwaegr., Philonotis hastata (Dub.) Wijk heteroneuron (Mitt.) Broth., & Marg., Orthotrichum rupestre Schizymenium pseudopohlia Shaw, Schwaegr., Hedwigia ciliata (Hedw.) P.- Bartramia
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