Tropical Bryology, Volume 2 (1990)
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In many ways, it is presumptuous for me to speak on the mosses of the tropical regions of China. Many consider the knowledge about the taxonomy, ecology, and geography of tropical bryophytes inadequate (Pócs 1982; Schuster 1983; Richards 1984), and this is certainly the case for the bryophytes of the tropical regions of China. The taxonomy of Chinese taxa is generally in a state of disarray. Early workers, both Chinese and others, have tended to describe new species based upon minor or inconsequential morphological characters and without apparent reference to related taxa found outside of China. This is clear from recent monographic studies that compared Chinese taxa with taxa throughout the world.
During the past 5 years intensive bryological explorations were carried out in Tanzania with special emphasize on hitherto undercollected areas (e.g. Nguru mountains, Mafia Island, unknown accesses of Mount Kilimanjaro and Meru) and on special habitats (e.g. rocky semi-desert or heath vegetation and alkaline tolerant epiphytic vegetation along the Rift Valley). These collections (above 8000 numbers) resulted in numerous records, some of them new to the African continent and at least 8 species new to science. The data point to interesting phytogeographical links and help to explain the evolution of the flora of East African volcanoes and crystalline mountains. Hitherto unknown oil bodies of more than 50 liverwort species were investigated. This paper does not give a full account of these studies but only provides examples to illustrate the above points.
The Guianas (French Guiana, Suriname, Guyana) are probably one of the last areas of the world covered largely by virgin lowland rain forest. Species diversity of epiphytic bryophytes was investigated in dry evergreen forest and mixed forest using mountaineering techniques to ascend into the canopy. The results indicate that the lowland rain forest is richer in species than previously believed due to neglect of the canopy flora, which may hold more than 50% of the local species. The mixed forest holds the richest flora and on one single forest tree up to 67 bryophyte species were found (50 on average); 28 trees yielded 154 species. A species/area curve indicates that epiphytic bryophyte species are usually commonly distributed in the forest and a few trees may yield much of the local flora. A recent checklist of the Guianas includes over 600 species of bryophytes: 375 Hepaticae and 234 Musci. As the region lacks in altitude (except on Mt. Roraima) the general character of the bryophyte flora of the Guianas is typically lowland neotropical. Over 80% of the species are rather widespread in tropical America (Amazonian species included), and the remaining are Guayana Highlands, northern Amazonian or Caribbean elements. Endemism is very low: 2.5 %.
The neotropical hepatic flora, predominantly constituted by members of the Jungermanniales and Metzgeriales, includes a disproportionate number of genera which are endemic (over 38) and a number which evidently originated here but have shown slight and in a geological sense, modern dispersal by solitary species. Endemism is confined almost to the Jungermanniales; it is to a large degree of a unique sort: confined to highly apomorphic derivatives, often extremely reduced, sometimes confervoid or thalloid (aside from 'normal' sexual branches). These endemics are derivatives of basically cool-Gondwanalandic suborders, chiefly Lepidoziineae and Cephaloziineae which, in the Antipodes today include a wide range of plesiomorphic taxa. The highest proportion of endemic genera, often stenotypic (1-3 species each) occurs in the upper montane zone: from upper Andean forest to páramo, to the edge of permanent snow and ice; a smaller number occurs at upper elevations of the Guyana Shield, but more occur in the riverine systems that dissect this shield. The taxa found there (i.a., Zoopsidella, Pteropsiella, Schusterolejeunea, Cephalantholejeunea) are among the most apomorphic of all hepatics. The amount of endemism is shown to be higher than in any comparable region of the globe. It is assumed that this is owing to: (a) isolation, exceeding 40 m.y. and probably exceeding 60 m.y.; (b) continuous tectonic activity, preserving the 'raw' and 'pioneer' habitats which are necessary for the survival of 'fugitive', 'shuttle' and other types of pioneer taxa; (c) the antiquity of the Guyana Shield and its riverine system; (d) creation of striking ecological gradients, many biotic islands; (e) fluctuation in extent and degree of isolation of these 'islands', leading to (f) rapid evolution due to genetic drift and perhaps enhanced selection pressures. It is concluded that part of the complexity of the flora is due to preservation of some elements on the old Guyana Shield but most is due to relatively rapid evolution during Tertiary times. A final contributing element has been the fact that movement of the South American plate has been primarily from east to west, so that the relevant land area has not been rafted into regions with very different climatic parameters: the degree of extinction seen in, e.g., India and Australia is not evident here. It is concluded that the amount of endemism seen, and its extreme kinds, 'need' in excess of the 40-60 m.y. time span which seems available. In particular, the large number of high elevation endemics, some (such as Ruizanthus) very isolated, cannot be satisfactorily explained by assuming their evolution in the few million years available since alpine regions were created by the rise of the Andes. It is almost necessary to conclude that limited 'pre-Andes' must have existed and that the ancestors of the isolated taxa seen today in alpine loci in Colombia and Venezuela originated elsewhere. The other side of the outlined scenario is that with the near-total isolation of tropical America until the Andes were elevated, and until the Pliocene connection to North America arose, one would expect to see few and scattered intruders from cool-Gondwanalandic areas and from Laurasia. The modern flora reflects exactly this.
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.
Fissidens in the Neotropics
(1990)
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.
The Frullania taxa on Mount Albert Edward, Papua New Guinea, form many associations that suggest a high degree of niche similarity, but at different altitudes, different associations form. The species diversity of the genus is greatest at the middle altitudes and least in the dry lowlands. This altitudinal separation is apparent in the niche widths of the taxa. The members of the four sibling taxa pairs examined exhibit distinct altitudinal niches, suggesting that the sibling taxa are distinct, with different niche optima.
China is located in eastern Asia, between 18°-55°N and 74°-138°E. The total land area of about 9,600,000 km is divided into 30 administrative areas. The climate varies from subtropic in the south to temperate in the north. The topography is complex. All of the conditions mentioned above cause the bryoflora to be complex. There are estimated to be about 2,000 species of Musci and 1,000 species of Anthocerotae and Hepaticae. The late Professor P.C. Chen considered that China could be divided into seven distributional areas, this view is supported by most Chinese bryologists today.
The growth habit of the Hedwigiaceae has been described variously as acrocarpous, pseudopleurocarpous, or pleurocarpous. Anatomical evidence presented here indicates that Hedwigia ciliata is acrocarpous. The archegonia are terminal on the main shoot, and the branching pattern is sympodial. The main axis of each plant thus consists of a succession of subterminal innovations, rather than a single shoot of indeterminate growth. Since the plants are plagiotropic and are pleurocarpous in appearance, this growth pattern can be also called pseudo-pleurocarpous.