Distribution Pattern of Tree Species and Richness along an Altitude Gradient in the Sub-Alpine Temperate Zone of Hindu Kush Mountainous Forests, Pakistan

Tree species richness along elevation gradient


  • Matiullah Department of Botany, PMAS Arid Agriculture University, Rawalpindi, Pakistan
  • Amjad ur Rahman Department of Botany, University of Swabi, Pakistan
  • Zahid Ullah Center for Plant Sciences and Biodiversity, University of Swat, Pakistan
  • Aamir Saleem Department of Forestry and Range Management, PMAS Arid Agriculture University, Rawalpindi, Pakistan
  • Rahmatullah Qureshi Department of Botany, PMAS Arid Agriculture University, Rawalpindi, Pakistan
  • David F.R.P. Burslem Department of Plant and Soil Science, University of Aberdeen, Cruickshank Building, St Machar Drive, Aberdeen AB24 3UU, United Kingdom
  • Zia-ur-Rehman Mashwani Department of Botany, PMAS Arid Agriculture University, Rawalpindi, Pakistan




Lalkoo Forest, Hindukush Mountains, Pakistan, Regeneration, Elevation gradient, Species richness, Diversity, domain effect


Lal-Koo Mountains Forest (LMF) is the most extensive vegetation type in the largest Hindukush Mountainous ranges of Pakistan, however highly overlooked compared to the Himalayan and Karakorum ranges. Here, we studied the conifer tree species regeneration, diversity, basal area, density, and species richness of the LalKoo Mountains Forest (LMF) along the altitude gradient. We used the quadrate (10 m × 10 m) sampling method for vegetation analysis at 54 different locations between 1970-3120 m elevations. We found a total of 115 species belonging to 58 families. We find the maximum value of Shannon’s -Winner index 3.603 at 2115 m and Simpson’s Diversity Index at 0.91 at 2290 m along an altitude gradient in lower elevation ranges. The current finding revealed that observed tree species richness shows a unimodal pattern with a peak at 2400 m in the mid–elevational range followed by a basal area peaked at 2300 m across the elevation gradient. We concluded that the high growth ratio of regenerates is due to open areas (free canopy space) likely available due to severe deforestation at low altitudes. In Lalkoo, tree density did not follow a regular trend, although the highest values were obtained around 2400 m and 2600 m along altitude. Our results also indicate that there is a narrow elevational range at high altitudes (near the timberline) measured from 2750 m – 3120 m, of the gradient. Furthermore, we discovered broader altitude ranges in the midst (relating well with the theory behind the mid-domain effect) in the range of 2345 m – 2750 m, but the lower altitude range assessed from 1970 - 2345 m does not reveal precise data for the reported species richness, which is a deviation from Stevens elevational Rapaport rule. 


K. Manish, M. K. Pandit, Y. Telwala, D. C. Nautiyal, L. P. Koh, and S. Tiwari. Elevational plant species richness patterns and their drivers across nonendemics, endemics and growth forms in the Eastern Himalaya. Journal of Plant Research 130(5): 829-844 (2017).

A. U. Rahman. S. M. Khan, Z. Saqib, Z. Ullah, Z. Ahmad, S. Ekercin, and H. Ahmad. Diversity and abundance of climbers in relation to their hosts and elevation in the monsoon forests of Murree in the Himalayas. Pakistan. Journal of Botany 52(2): 601-612 (2020).

M. Ilyas, R. Qureshi, N. Akhtar, M.K.A. Ziaul-Haq and A. M. Khan. Floristic diversity and vegetation structure of the remnant subtropical broad-leaved forests from Kabal valley, Swat, Pakistan. Pakistan Journal of Botany 50(1): 217-230 (2018).

C.M. Oswalt, W.K. Clatterbuck, and E.A. Houston. Impacts of deer herbivory and visual grading on the early performance of high-quality oak planting stock in Tennessee, USA. Forest Ecology and Management 229(1-3): 128-135 (2006).

B. Rawat, S. Gairola, L.M. Tewari, and R.S. Rawal. Long-term forest vegetation dynamics in Nanda Devi Biosphere Reserve, Indian west Himalaya: evidence from repeat studies on compositional patterns. Environmental Monitoring and Assessment 193(8): 1-17 (2021).

M. Begon, C.R. Townsend, and J. L. Harper. Ecology: From Individuals to Ecosystems. Blackwell Scientific Publications,UK (2006).

X.M. Mou, Y.W. Yu, X.G. Li, and A.A. Degen. Presence frequency of plant species can predict spatial patterns of the species in small patches on the Qinghai-Tibetan Plateau. Global Ecology and Conservation 21: e00888 (2020).

R.K. Colwell, and D.C. Lees. The mid-domain effect: geometric constraints on the geography of species richness. Trends in Ecology & Evolution 15(2): 70-76 (2000).

J.G. Pausas, and M. P. Austin. Patterns of plant species richness in relation to different environments: an appraisal. Journal of Vegetation Science 12(2): 153-166 (2001).

P. Bromley. The Effects of Elevation Gain on Soil. (1995) (Accessed April 7, 2016)

D. Conway, K. Bhattarai, and N.R. Shrestha. Population–environment relations at the forested frontier of Nepal: Tharu and Pahari survival strategies in Bardiya. Applied Geography 20(3): 221-242 (2000).

B.A. Hawkins, J.A.F. Diniz-Filho, C.C.A. Jaramillo, and S.A. Soeller. Climate, niche conservatism, and the global bird diversity gradient. The American Naturalist 170(S2): S16-S27 (2007).

K. Gebrehiwot, S. Demissew, Z. Woldu, M. Fekadu, T. Desalegn, and E. Teferi. Elevational changes in vascular plants richness, diversity, and distribution pattern in Abune Yosef mountain range, Northern

M. Ilyas, Z.K. Shinwari, and R. Qureshi. Vegetation composition and threats to the montane temperate forest ecosystem of Qalagai hills, Swat, Khyber Pakhtunkhwa, Pakistan. Pakistan Journal of Botany 44: 113-122 (2012).

S. Saima, A. Altaf, M.H. Faiz, F. Shahnaz, and G. Wu. Vegetation patterns and composition of mixed coniferous forests along an altitudinal gradient in the Western Himalayas of Pakistan. Austrian Journal of Forest Science 135(2): 159-180 (2018).

S. Kamran, S.M. Khan, Z. Ahmad, A.U. Rahman, M. Iqbal, F. Manan, and S. Ullah. The role of graveyards in species conservation and beta diversity: a vegetation appraisal of sacred habitats from Bannu, Pakistan. Journal of Forestry Research 31(4): 1147-1158 (2020).

R.A. Mittermeier, P. Robles-Gil, M. Hoffmann, J.D. Pilgrim, T.M. Brooks, C.G. Mittermeier, and G. Fonseca. Hotspots Revisited: Earth’s Biologically Richest and Most Endangered Ecoregions (Cemex, Mexico City) (2005).

C.L. Cardelús, R.K. Colwell, and J.E. Jr Watkins. Vascular epiphyte distribution patterns: explaining the mid-elevation richness peak. Journal of Ecology 144-156 (2006).

M. Chhetri, K.L. Maskey, N.R. Chapagain, and B.D. Sharma. Mustang-the Land of Fascination. King Mahendra Trust for Nature Conservation (KMTNC), Nepal (2004).

N.J. Sanders, J.P. Lessard, M.C. Fitzpatrick, and R.R. Dunn. Temperature, but not productivity or geometry, predicts elevational diversity gradients in ants across spatial grains. Global Ecology and Biogeography 16(5): 640-649 (2007).

C. Shen, and A.S. Nelson. Natural conifer regeneration patterns in temperate forests across the Inland Northwest, USA. Annals of Forest Science 75(2): 1-16 (2018).

IPCC (2007c), Climate Change 2007. Synthesis Report (Geneva: IPCC); at: <http://www.ipcc.ch/90 Matiullah et al

pdf/assessment-report/ar4/syr/ar4_syr. pdf>.

P. Bhattarai, K.P. Bhatta, R. Chhetri, and R.P. Chaudhary. Vascular plant species richness along elevation gradient of the Karnali River valley, Nepal Himalaya. International Journal of Plant, Animal and Environmental Sciences 4(3): 114-126 (2014).

Braun-Blanquet. Journal Plant sociology. New York: Hafner Publishing Company (1965).

P.K.T. Munishi, F. Philipina, R.P.C. Temu, and N.E. Pima. Tree species composition and local use in agricultural landscapes of west Usambaras Tanzania. African Journal of Ecology 46: 66-73 (2008).

C.M. McCain. The mid‐domain effect applied to elevational gradients: species richness of small mammals in Costa Rica. Journal of Biogeography 31(1): 19-31 (2004).

R.K. Colwell, and D.C. Lees. The mid-domain effect: geometric constraints on the geography of species richness. Trends in Ecology & Evolution 15(2): 70-76 (2000).

J. Hortal, P.A. Borges, and C. Gaspar. Evaluating the performance of species richness estimators: sensitivity to sample grain size. Journal of Animal Ecology 75(1): 274-287 (2006).

R.S. Rawal, N.S. Bankoti, S.S. Samant, and Y.P.S. Pangtey. Phenology of tree layer species from the timber line around Kumaun in Central Himalaya, India. Vegetation 93(2): 108-118 (1991).

F. Bokma, J. Bokma, and M. Mönkkönen. Random processes and geographic species richness patterns: why so few species in the north? Ecography 24(1): 43-49 (2001).

P. McCullagh, and J.A. Nelder. Generalized Linear Models. Chapman & Hall, New York (1989).

M. Barbour, J.H. Burk, W.D. Pitts, F.S. Gillians, and M.W. Schwartz. Terrestrial Ecology. Chicago, Illinois: Addson Wesley Longman, Inc (1999).

V.H. Heywood. (Ed.). Global biodiversity assessment. Cambridge: Cambridge University Press (1995).

B.B. Shrestha, B. Ghimire, H.D. Lekhak, and P.K. Jha. Regeneration of treeline birch (Betula utilis D. Don) forest in a trans-Himalayan dry valley in central Nepal. Mountain Research and Development 27(3): 259-267 (2007).

D. Scott, D. Welch, M. Thurlow, and D.A. Elston. Regeneration of Pinus sylvestris in a natural pinewood in NE Scotland following reduction in grazing by Cervus elaphus. Forest Ecology and Management 130(1-3): 199-211 (2000).

F.D. Meyer. Rekonstruktion der KlimaWachstumsbeziehungen und der Waldentwicklung im subalpinen Waldgrenzokoton bei Grindelwald, Schweiz [Dissertation]. Basel, Switzerland: Basel University. 161 p (2000).

B.K. Ghimire, and H.D. Lekhak. Regeneration of Abies spectabilis (D. Don) Mirb. in subalpine forest of upper Manang, north-central Nepal. Local effects of global changes in the Himalayas: Manang, Nepal 139-149 (2007).

N.J. Sanders. Elevational gradients in ant species richness: area, geometry, and Rapoport’s rule. Ecography 25(1): 25-32 (2002).

C. Rahbek. The role of spatial scale and the perception of large‐scale species‐richness patterns. Ecology Letters 8(2): 224-239 (2005).

C. Carpenter. The environmental control of plant species density on a Himalayan elevation gradient. Journal of Biogeography 32(6): 999-1018 (2005).

M.D. Behera, and S.P.S. Kushwaha. An analysis of altitudinal behavior of tree species in Subansiri district, Eastern Himalaya. In: Plant Conservation and Biodiversity (pp. 277-291). Springer, Dordrecht (2006).

J. Li, Q. He, X. Hua, J. Zhou, H. Xu, J. Chen, and C. Fu. Climate and history explain the species richness peak at mid‐elevation for Schizothorax fishes (Cypriniformes: Cyprinidae) distributed in the Tibetan Plateau and its adjacent regions. Global Ecology and Biogeography 18(2): 264-272 (2009).

J.A. Grytnes, and O.R. Vetaas. Species richness and altitude: a comparison between null models and interpolated plant species richness along the Himalayan altitudinal gradient, Nepal. The American Naturalist 159(3): 294-304 (2002).

A. Sánchez‐González, and L. López‐Mata. Plant species richness and diversity along an altitudinal gradient in the Sierra Nevada, Mexico. Diversity and Distributions 11(6): 567-575 (2005).

J.S. Li, Y.L. Song, and Z.G. Zeng. Elevational gradients of small mammal diversity on the northern slopes of Mt. Qilian, China. Global Ecology and Biogeography 12(6): 449-460 (2003).

R.K. Colwell, and D.C. Lees. The mid-domain effect: geometric constraints on the geography of species richness. Trends in Ecology & Evolution 15(2): 70-76 (2000).

R.R. Dunn, C.M. McCain, and N.J. Sanders. When does diversity fit null model predictions? Scale and range size mediate the mid‐domain effect. Global Ecology and Biogeography 16(3): 305-312 (2007).

Tree species richness along elevation gradient 91 48. C. Korner. Why are there global gradients in species richness? Mountains might hold the answer. Trends Ecological Evolution 15: 513-514 (2000).

T.F.L.V.B. Rangel, and J.A.F. Diniz-Filho. Spatial patterns in species richness and the geometric constraint simulation model: a global analysis of mid-domain effect in Falconiformes. Acta Oecologica 24: 203e207 (2003).

T.F. Rangel, J.A.F. Diniz‐Filho, and L.M. Bini. SAM: a comprehensive application for spatial analysis in macroecology. Ecography 33(1): 46-50 (2010).

J.E. Jr. Watkins, C. Cardelús, R.K. Colwell, and R.C. Moran. (2006). Species richness and distribution of ferns along an elevational gradient in Costa Rica. American Journal of Botany 93(1): 73-83.52. Z. Wang, Z. Tang, and J. Fang. Altitudinal patterns of seed plant richness in the Gaoligong Mountains, south‐east Tibet, China. Diversity and Distributions 13(6): 845-854 (2007).

A. Rahman, S.M. Khan, Z. Ahmad, S. Alamri, M. Hashem, M. Ilyas, and S. Ali. Impact of multiple environmental factors on species abundance in various forest layers using an integrative modeling approach. Global Ecology and Conservation 29: e01712 (2021).

M.V. Lomolino. Elevation gradients of species‐density: historical and prospective views. Global Ecology and Bibliography 10(1): 3-13 (2001).

W.F. Laurance, A.K. Albernaz, P.M. Fearnside, H.L. Vasconcelos, and L.V. Ferreira. Deforestation in amazonia. Science 304(5674): 1109-1111 (2004).

P. Koleff, and K.J. Gaston. Latitudinal gradients in diversity: real patterns and random models. Ecography 24(3): 341-351 (2001).




How to Cite

Matiullah, Amjad ur Rahman, Zahid Ullah, Aamir Saleem, Rahmatullah Qureshi, David F.R.P. Burslem, & Mashwani, Z.- ur-R. (2022). Distribution Pattern of Tree Species and Richness along an Altitude Gradient in the Sub-Alpine Temperate Zone of Hindu Kush Mountainous Forests, Pakistan: Tree species richness along elevation gradient. Proceedings of the Pakistan Academy of Sciences: B. Life and Environmental Sciences, 59(4), 81–92. https://doi.org/10.53560/PPASB(59-4)741