Abstract
This review deals with the forest vegetation of the Himalaya with emphasis on: paleoecological, phytogeographical, and phytosociological aspects of vegetation; structural and functional features of forest ecosystem; and relationship between man and forests.
The Himalayan mountains are the youngest, and among the most unstable. The rainfall pattern is determined by the summer monsoon which deposits a considerable amount of rain (often above 2500 mm annually) on the outer ranges. The amount of annual rainfall decreases from east to west, but the contribution of the winter season to the total precipitation increases. Mountains of these dimensions separate the monsoon climate of south Asia from the cold and dry climate of central Asia. In general, a rise of 270 m in elevation corresponds to a fall of 1°C in the mean annual temperature up to 1500 m, above which the fall is relatively rapid.
Large scale surface removals and cyclic climatic changes influenced the course of vegetational changes through geological time. The Himalayan ranges, which started developing in the beginning of the Cenozoic, earlier supported tropical wet evergreen forests throughout the entire area (presently confined to the eastern part). The Miocene orogeny caused drastic changes in the vegetation, so much so that the existing flora was almost entirely replaced by the modern flora. Almost all the dominant forest species of the Pleistocene continue to maintain their dominant status to the present. Presently the Himalayan ranges encompass Austro-Polynesian, Malayo-Burman, Sino-Tibetan, Euro-Mediterranean, and African elements. While the Euro-Mediterranean affinities are well represented in the western Himalayan region (west of 77°E long.), the Chinese and Malesian affinities are evident in the eastern region (east of 84°E long.). However, the proportion of endemic taxa is substantial in the entire region.
A representation of formation types in relation to climatic factors, viz., rainfall and temperature, indicates that boundaries between the types are not sharp. Formation types often integrate continuously, showing broad overlaps. Climate does not entirely determine the formation type, and the influence of soil, fire, etc., is also substantial. The ombrophilous broad leaf forests located in the submontane belt (< 1000 m) of the eastern region are comparable to the typical tropical rain forests. On the other extreme, communities above 3000 m elevation are similar to sub-alpine and alpine types. From favorable to less favorable environments, as observed with decreasing moisture from east to west, or with decreasing temperature from low to high elevations, the forests become increasingly open, shortstatured and simpler, with little vertical stratification. Ordination of forest stands distributed within 300–2500 m elevations of the central Himalaya, by and large indicates a continuity of communities, with scattered centers of species importance values in the ordination field. Within the above elevational transect, sal (Shorea robusta) and oak (Quercus spp.) forests may be designated as the climax communities, respectively, of warmer and cooler climates. The flora of a part of the central Himalayan region is categorized as therohemigeophytic and that of a part of the western Himalayan region as geochamaephytic.
An analysis of population structure over large areas in the central Himalaya, based on density-diameter distribution of trees, suggests that oldgrowth forests are being replaced by even-aged successional forests, dominated by a few species, such asPinus roxburghii. Paucity of seedlings of climax species, namelyShorea robusta andQuercus spp. over large areas is evident.
The Himalayan catchments are subsurface-flow systems and, therefore, are particularly susceptible to landslips and landslides. Loss of water and soil in terms of overflow is insignificant.
Studies on recovery processes of forest ecosystems damaged due to shifting cultivation or landslides indicate that the ecosystems can recover quite rapidly, at least in elevations below 2500 m. For example, on a damaged forest site, seedlings of climax species (Quercus leucotrichophora) appeared only 21 years after the landslide.
In the central Himalaya, the biomass of a majority of forests (163-787 t ha−1) falls within the range (200-600 t ha−1) given for many mature forests of the world, and the net primary productivity (found in the range of 11.0–27.4 t ha−1 yr−1) is comparable with the range of 20–30 t ha−1 yr−1 given for highly productive communities of favorable environments. In most of the forests of this region, the litter fall values (2.1-3.8 t C ha−1 yr−1) are higher than the mean reported for warm temperate forests (2.7 t C ha−1 yr−1). Of the total litter, the tree leaves account for 54–82% in the Himalayan forests.
The rate of decomposition of leaves in some broadleaf species of submontane belt (0.253-0.274% day−1) are comparable with those reported for some tropical rain forest species. Because of the paucity of microorganisms and microarthropods in the forest litter and soil, high initial C:N ratio and high initial lignin content in leaves, the rate of leaf litter decomposition inPinus roxburghii is markedly slower than in other species of the central Himalaya. The fungal species composition of the leaf litterof Pinus roxburghii is also distinct from those of other species.
A greater proportion of nutrients is accumulated in the biomass component of the Himalayan forests than in the temperate forests. Although litter fall is the major route through which nutrients return from biomass to the soil pool, a substantial proportion of the total return is in the form of throughfall and stemflow. Among the dominant species of the central Himalaya, retranslocation of nutrients from the senescing leaves was markedly greater inPinus roxburghii than inQuercus spp. andShorea robusta. Consequently, the C:N ratio of leaf litter is markedly higher inPinus roxburghii than in the other species. Immobilization of nutrients by the decomposers of the litter with high C:N ratio is one of the principal strategies through whichPinus roxburghii invades other forests and holds the site against possible reinvasion by oaks.
Observations on the seasonality of various ecosystem functions suggest that Himalayan ecosystems are geared to take maximum advantages of the monsoon period (rainy season).
Most of the human population depends on shifting-agriculture in the eastern region and on settled agriculture in the central and western regions. Either of these is essentially a forest-dependent cultivation. Each unit of agronomic energy produced in the settled agriculture entails about seven units of energy from forests. Consequently, forests with reasonable crown cover account for insignificant percentage of the land. Tea plantations and felling of trees for timber, paper pulp, etc., are some of the major commercial activities which adversely affected the Himalayan forests.
Résumé
Cette revue concerne la végétation forestière de l’Himalaya. Elle précise l’information concernant la paléoécologie, la phytogéographie, la phytosociologie, le structure et le fonctionnement des écosystèmes et le rapport entre l’homme et la forêt.
Les montagnes de l’Himalaya sont les plus jeunes et parmi les plus instables. La pluviométrie dépend surtout de la mousson d’été et les chaînes extérieures sont bien arrosées (>2500 mm par an). Les précipitations annuelles décroissent de l’Est vers l’Ouest tandis que la composante hivernale augmente. Ces montagnes séparent les climats de mousson de l’Asie du Sud des climats froids et secs de l’Asie Centrale.
L’érosion du sol sur une grand étendue et des changemenets cycliques du climat ont déterminé des changements dans le couvert végétal tout au long des temps géologiques. Les chaînes Himalayennes qui ont commencés leur soulèvement au commencement du coenozoïque étaient entièrement couvertes d’une forêt ombrophile tropicale. (Ce type se trouve encore de nos jours dans la partie orientale de l’Himalaya.) L’orogénie miocène provoqua de tels changements dans la végétation que la flore de cette époque a été entièrement remplacée par la flore moderne. Les espèces forestières dominantes du pleistocène gardent leur importance dans les forêts actuelles.
Des éléments floraux Austro-Polynésiens, Malais-Birmans, Sino-Tibetains, Euro-Méditerranéens et Africains sont actuellement présents sur les montagnes himalayennes. Tandis que les affinités Euro-Mediterranéennes sont bien représentées dans l’Himalaya occidental (à l’Ouest du 77° Est), les affinités Chinoises et Malaises sont évidentes dans la partie orientale (à l’Est de 84°E). Cependant la proportion des éléments endémiques est importante dans toute la région.
La relation entre les types de formations et les facteurs climatiques (pluviosité, température) indique que les limites entre les types sont approximatives. D’ailleurs, le climat lui même ne détermine pas exclusivement les types et les effets du sol, du feu, etc., peuvent être importantes. Les forêts feuillues ombrophiles localisées dans l’étage sous-montagnard (< 1000 m) de la région orientale sont comparables aux forêts ombrophiles tropicales typiques. A l’opposé les communautés qui se trouvent au-dessus de 3000 m d’altitude sont comparables aux types subalpins et alpins. En allant des conditions favorables vers le moins favorables soit par exemple d’Est en Ouest le long de l’axe de diminution des précipitations soit en suivant les gradient altitudinal de baisse des températures les forêts deviennent de plus en plus ouvertes, basses et structurellement simples avec peu de stratification verticale. L’ordination des peuplements forestières situés entre 300–2500 m dans l’Himalaya central indique une continuité des communautés avec des centres de valeurs d’importance des espèces dispersés dans le champ d’ordination. Dans ce transect altitudinal, les forêts à sal (Shorea robusta) et à chêne (Quercus spp.) peuvent être désignés comme des communautés climax pour les climats chaud et froid respectivement.
Le spectre biologique basé sur les formes biologiques de Raunkiaer est du type Théro-Hémi-Géophytique dans l’Himalaya central tandis que celui de l’Himalaya occidental est du type Géo-Chamaephytique.
L’analyse de la structure du peuplement couvrant une superficie assez importante dans l’Himalaya central basé sur la répartition des arbres par densité-diamètre suggère que les anciennes forêts sont en train d’être remplacées par des forêts équiennes de succession, dominées par un petit nombre d’espèces, tel quePinus roxburghii. La mauvaise régénération des espèces climax, à savoir le sal (Shorea robusta) et le chêne (Quercus) sur une aire assez vaste est un fait bien établi.
Les bassins versants de l’Himalaya sont du type à l’écoulement hypodermique et sont donc sensibles aux glissements de terrain. La perte d’eau et de sol en terme d’épanchement est peu important.
L’étude de la reconstitution des écosystèmes forestiers dégradés par les cultures itinérantes ou par les glissements de terrain montre que les écosystèmes endommagés peuvent se reconstituer assez rapidement au moins en dessous de 2500 m.
Par exemple, sur un site forestière dégradée, les plantules de l’espèce climax (Quercus leucotrichophora) sont réapparues seulement 21 ans après le glissement du terrain.
Dans l’Himalaya central, la biomasse de la majorité des forêts (163783 t ha−1) tombe dans la classe de la plupart des forêts du monde (200600 t ha−1) et la productivité nette primaire (11.0-27.4 t ha−1 an−1) est comparable à celle des meilleures forêts (20-30 t ha−1 an−1) soumises à des conditions favorables. Les valeurs de la chute de litière des forêts de cette région (2.1-3.8 t C ha−1 an−1) sont plus élevées que la moyenne de celles des forêts tempérées chaudes (2.7 t C ha−1 an−1). La contribution des feuilles d’arbre à la litière totale est entre 54 et 82 pourcent dans les forêts Himalayennes.
Le taux de décomposition des feuilles chez certains feuillus de l’étage sous-montagnard (0.253–0.274% par jour) est comparable à celui de certaines espèces de la forêt ombrophile. C’est à cause d’une pauvreté de la litière forestière et du sol en microorganismes et des microarthropodes, du rapport initial élevé C:N et du pourcentage initial élevé en lignine des feuilles, que le taux de décomposition des aiguilles dePinus roxburghii est significativement plus lent que chez les autres espèces de l’Himalaya central. D’ailleurs la composition de la flore fongique de la litière des aiguilles dePinus roxburghii est bien différente de celles des autres espèces.
Une plus grande proportion d’éléments biogéochimiques est accumulée dans la composante biomasse des forêts Himalayennes par rapport aux forêts tempérés.
Bien que la chute du litière constitue la voie principale par laquelle les éléments de la biomass retournent au sol, une fraction assez importante est restituée sous formes de pluviolessivats et d’écoulements sur le tronc.
Parmi les espèces dominantes d l’Himalaya central, la redistribution des éléments des feuilles senescentes est plus important chezPinus roxburghii que chezShorea ouQuercus. Par conséquent, le rapport C:N de la litière de feuille est plus élevé chez le pin que chez les autres espèces. L’immobilisation des éléments par les décomposeurs de la litière à rapport C:N élevé est une des stratégies principales par laquellePinus roxburghii envahit les autres forêts et évite la reconquête par les autres espèces.
L’étude saisonnière des divers fonctionnements de l’écosystème met en évidence des liens étroits avec la régime des pluies de mousson.
La majorité de la population humaine pratique la culture itinérante dans la région orientale et l’agriculture sédentaire dans les parties central et occidentale. Ces deux types d’agriculture sont très liés à la forêt. Chaque unité d’énergie agronomique produite en agriculture sédentaire demande sept unités d’énergie des forêts. Par conséquent, les forêts peu ouvertes à canopées assez fermées n’occupent plus qu’un pourcentage négligeable de ces régions. Les plantations de thé et l’exploitation forestière (bois d’oeuvre, pâte à papier, etc.) sont parmi les activités qui ont contribué à dégrader les forêts de l’Himalaya.
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Singh, J.S., Singh, S.P. Forest vegetation of the Himalaya. Bot. Rev 53, 80–192 (1987). https://doi.org/10.1007/BF02858183
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DOI: https://doi.org/10.1007/BF02858183