Low floristic richness of afro-alpine vegetation of Mount Kenya is related to its small area

  • Vladimir G. Onipchenko
  • Natalia A. Kopylova
  • Alii M. Kipkeev
  • Tatiana G. Elumeeva
  • Andrei Azovsky
  • Sergei V. Dudov
  • Justine M. NyagaEmail author
Original Article


Diversity patterns of tropical alpine vegetation is poorly studied. We estimated vascular plant floristic richness for two typical afro-alpine communities in Mount Kenya using a series of nested plots sized from 25 to 100 m2. The α-diversity was low for all plot sizes (4.8–7.8 and 12.8 species per one and 100 m2, respectively). Comparative analysis of α-, β- and γ-diversity across 22 plant communities from five Mountain regions (Mount Kenya, European Alps, Caucasus, Tibet, New Zealand Alps) revealed that area of mountain system was the only significant variable shaping the local richness; this effect became stronger with increasing spatial scale. Beta-diversity, by contrast, showed neither latitudinal trends nor significant correlation with other geographical or climatic variables. We conclude that the total area of mountain system is one of the main factors determining the regional species pool and, ultimately, the local diversity of alpine plant communities (the “echo-effect”). Small area and isolation of Mt. Kenya are considered as the main reasons for low local richness of its afro-alpine vegetation.


Species–area relationship Floristic richness Alpha-diversity Beta-diversity Mt. Kenya Afro-alpine Tropical alpine Plant communities 



We thank M. Pärtel for useful notes for the paper. The work was supported by the governmental contract of MSU AAAA-A16-116021660037-7.

Author contributions

VGO, NAK, AMK and JMN designed the research, VGO, AMK and JMN conducted the research and VGO, NAK, AMK, TGE, AA, SVD and JMN analyzed the data and wrote the article.

Compliance with ethical standards

Conflict of interest

The authors declare that they have no conflict of interest.

Ethical approval

This article does not contain any data collected from human participants or animals and, therefore, did not require any ethical approval.


  1. Arrhenius O (1921) Species and area. J Ecol 9:95–99CrossRefGoogle Scholar
  2. Azovsky AI (2011) Species–area and species–sampling effort relationships: disentangling the effects. Echography 34:18–30CrossRefGoogle Scholar
  3. Belyea LR, Lancaster J (1999) Assembly rules within a contingent ecology. Oikos 86:402–416CrossRefGoogle Scholar
  4. Bengtsson J, Fagerstrom T, Rydin H (1994) Competition and coexistence in plant communities. Trends Ecol Evol 9:246–250PubMedCrossRefGoogle Scholar
  5. Cantero JJ, Pärtel M, Zobel M (1999) Is species richness dependent on the neighbouring stands? An analysis of the community patterns in mountain grasslands of central Argentina. Oikos 87:346–354CrossRefGoogle Scholar
  6. Chao A (1987) Estimating the population size for capture-recapture data with unequal catchability. Biometrics 43:783–791PubMedCrossRefGoogle Scholar
  7. Chytrý M, Ermakov N, Danihelka J, Hajek M, Hajkova P, Horsak M, Koci M, Kubesova S, Lustyk P, Otypkova Z, Pelankova B, Valachovich M, Zeleny D (2012) High species richness in semiboreal forests of the northern Russian Altai, southern Siberia. J Veg Sci 23:605–616. CrossRefGoogle Scholar
  8. Coe MJ (1967) The ecology of the alpine zone of Mount Kenya. Dr W Junk, The HagueCrossRefGoogle Scholar
  9. Cowling RM, Potts AJ, Bradshow PL, Colville J, Arianoutsou M, Ferrier S, Forest F, Fellas NM, Hopper SD, Ojeda F, Proches S, Smith RJ, Rundel PW, Vassilakis E, Zutta BR (2015) Variation in plant diversity in mediterranean-climate ecosystems: the role of climatic and topographical stability. J Biogeogr 42:552–564. CrossRefGoogle Scholar
  10. Cuesta F, Muriel P, Llambi LD, Halloy S, Aguirre N, Beck S, Carilla J, Meneses RI, Cuello S, Grau A et al (2017) Latitudinal and altitudinal patterns of plant community diversity on mountain summits across the tropical Andes. Ecography 40:1381–1394. CrossRefGoogle Scholar
  11. Dolnik C, Breuer M (2008) Scale dependency in the species–area relationship of plant communities. Folia Geobotanica 43:305–318. CrossRefGoogle Scholar
  12. Fick SE, Hijmans RJ (2017) WorldClim 2: new 1-km spatial resolution climate surfaces for global land areas. Int J Climatol 37:4302–4315. CrossRefGoogle Scholar
  13. Gehrke B, Linder HP (2014) Species richness, endemism and species composition in the tropical Afroalpine flora. Alp Bot 124:165–177. CrossRefGoogle Scholar
  14. Ghalambor CK, Huey RB, Martin PR, Tewksbury JJ, Wang G (2006) Are mountain passes higher in the tropics? Janzen’s hypothesis revisited. Integr Comp Biol 46(1):5–17PubMedCrossRefGoogle Scholar
  15. Gleason HA (1922) On the relation between species and area. Ecology 3:158–162CrossRefGoogle Scholar
  16. Grubb PJ (1986) Problems posed by sparse and patchily distributed species in species-rich plant communities. In: Diamond J, Case T (eds) Community ecology. Harper and Row, New York, pp 207–226Google Scholar
  17. Gurung H (1999) Mountains of Asia—a regional inventory. International Centre for Integrated Mountain Development, KathmanduGoogle Scholar
  18. Hedberg O (1964) Features of Afro-Alpine plant ecology. Acta Phytogeographica Svecica 49:1–44Google Scholar
  19. Jakob M, Annys S, Frankl A, De Ridder M, Beeckman H, Guyassa E, Nyssen J (2015) Tree line dynamics in the tropical African highlands—identifying drivers and dynamics. J Veg Sci 26:9–20CrossRefGoogle Scholar
  20. Kammer PM, Mohl A (2002) Factors controlling species richness in alpine plant communities: an assessment of the importance of stress and disturbance. Arct Antarct Alp Res 34:398–407CrossRefGoogle Scholar
  21. Körner C (2003) Alpine plant life, 2nd edn. Springer, Berlin eaCrossRefGoogle Scholar
  22. Körner C, Jetz W, Paulsen J, Payne D, Rudmann-Maurer K, Spehn EM (2017) A global inventory of mountains for bio-geographical applications. Alp Bot 127:1–15CrossRefGoogle Scholar
  23. Kraft NJB, Comita LS, Chase JM, Sanders NJ, Swenson NG, Crist TO et al (2011) Disentangling the drivers of beta diversity along latitudinal and elevational gradients. Science 333:1755–1758PubMedCrossRefGoogle Scholar
  24. Levesque E (1996) Minimum area and cover-abundance scales as applied to polar desert vegetation. Arct Alp Res 28:156–162CrossRefGoogle Scholar
  25. MacArthur RH, Wilson EO (1967) The theory of island biogeography. Princeton University Press, PrincetonGoogle Scholar
  26. Matthews TJ, Guihaumon F, Triantis KA, Borregaard MK, Whittaker RJ (2016) On the form of species–area relationships in habitat islands and true islands. Glob Ecol Biogeogr 25:847–858. CrossRefGoogle Scholar
  27. McFadden IR, Sandel B, Tsirogiannis C, Morueta-Holme N, Svenning JC, Enquist BJ, Kraft NJ (2019) Temperature shapes opposing latitudinal gradients of plant taxonomic and phylogenetic β diversity. Ecol Lett. CrossRefPubMedGoogle Scholar
  28. Minchin PR (1989) Montane vegetation of the Mt. Field massif, Tasmania: a test of some hypothesizes about properties of community patterns. Vegetatio 83:97–110CrossRefGoogle Scholar
  29. Mizuno K, Fujita T (2014) Vegetation succession on Mt. Kenya in relation to glacial fluctuation and global warming. J Veg Sci 25:559–570. CrossRefGoogle Scholar
  30. Moravec J (1973) The determination of the minimal area of phytocenoses. Folia Geobot et Phytotax 8:23–47CrossRefGoogle Scholar
  31. Myers JA, Chase JM, Jimenez I, Jorgensen PM, Araujo Murakami A, Paniagua-Zambrana N et al (2013) Beta-diversity in temperate and tropical forests reflects dissimilar mechanisms of community assembly. Ecol Lett 16:151–157PubMedCrossRefGoogle Scholar
  32. Onipchenko VG, Pavlov VN (2009) Local plant species richness depends on total area of alpine communities. Dokl Akad Nauk SSSR 427:710–712Google Scholar
  33. Onipchenko VG, Semenova GV (1995) Comparative analysis of the floristic richness of alpine communities in the Caucasus and the Central Alps. J Veg Sci 6:299–304CrossRefGoogle Scholar
  34. Onipchenko VG, Mark A, Wells G (2005) Floristic richness of three perhumid New Zealand alpine plant communities in comparison with other regions. Austral Ecol 30:518–525. CrossRefGoogle Scholar
  35. Onipchenko VG, Shulakov AA, Zernov AS, Elumeeva TG, Wu Y, Wang Q, Werger M (2014) Contrasting floristic richness of alpine plant communities on the eastern Qinghai-Tibetan Plateau. Botanica Pacifica 3:33–37CrossRefGoogle Scholar
  36. Partel M, Zobel M, Zobel K, van der Maarel E (1996) The species pool and its relation to species richness: evidence from Estonian plant communities. Oikos 75:111–117CrossRefGoogle Scholar
  37. Qian H, Ricklefs RE (2007) A latitudinal gradient in large-scale beta diversity for vascular plants in North America. Ecol Lett 10:737–744PubMedCrossRefGoogle Scholar
  38. Rehder H, Beck E, Kokwaro JO (1988) The afroalpine plant communities of Mt. Kenya (Kenya). Phytocoenologia 16:433–463CrossRefGoogle Scholar
  39. Rosenzweig ML (1995) Species diversity in space and time. Cambridge University Press, CambridgeCrossRefGoogle Scholar
  40. Rosenzweig ML, Ziv Y (1999) The echo pattern of species diversity: pattern and process. Ecography 22:614–628CrossRefGoogle Scholar
  41. Rosindell J, Cornell SJ (2009) Species–area curves, neutral models, and long-distance dispersal. Ecology 90:1743–1750PubMedCrossRefGoogle Scholar
  42. Scheiner SM (2003) Six types of species–area curves. Glob Ecol Biogeogr 12:441–447CrossRefGoogle Scholar
  43. Scheiner SM, Chiarucci A, Fox GA, Helmus MR, McGlinn DJ, Willig MR (2011) The underpinnings of the relationship of species richness with space and time. Ecol Monogr 81:195–213CrossRefGoogle Scholar
  44. Sedel’nikov VP (1979) Flora i rastitel’nost’ vysokogorii Kuznetskogo Alatau [Flora and Vegetation of High Mountain Area of the Kuznetskii Alatau]. Nauka, Novosibirsk (in Russian) Google Scholar
  45. Shlothauer SD (1990) Vegetation of the Suboceanic High Mountains. Nauka, Moscow (in Russian) Google Scholar
  46. Sklenar P, Hedberg I, Cleef AM (2014) Island biogeography of tropical alpine floras. J Biogeogr 41:287–297. CrossRefGoogle Scholar
  47. Storch D (2016) The theory of the nested species–area relationship: geometric foundations of biodiversity scaling. J Veg Sci 27:880–891. CrossRefGoogle Scholar
  48. Tanfil’ev VG, Dzybov DS, Petrov LN, Pishchugina NS (1979) Geobotanical description of Rocky Mountains summits and Bechasyn Plato in the Northern Caucasus. Problemy Botaniki 14:121–125 (in Russian) Google Scholar
  49. Triantis KA, Nogues-Bravo D, Hortal J, Borges PAV, Adsersen H, Fernandez-Palacios JM, Araujo MB, Whittaker RJ (2008) Measurements of area and the (island) species–area relationship: new directions for an old pattern. Oikos 117:1555–1559. CrossRefGoogle Scholar
  50. Tsuyuzaki S, Urano S, Tsujii T (1990) Vegetation of alpine marshland and its neighboring areas, northern part of Sichuan Province, China. Vegetatio 88:79–86CrossRefGoogle Scholar
  51. Vonlanthen CM, Kammer PM, Eugster W, Buhler A, Veit H (2006) Alpine vascular plant species richness: the importance of daily maximum temperature and pH. Plant Ecol 184:13–25. CrossRefGoogle Scholar
  52. Whitmore TC, Peralta R, Brown K (1985) Total species count in a Costa Rican tropical rain forest. J Trop Ecol 1:375–378CrossRefGoogle Scholar
  53. Whittaker RJ, Matthews TJ (2014) The varied form of species–area relationship. J Biogeogr 41:209–210. CrossRefGoogle Scholar
  54. Wilson JB, Sykes MT (1988) Some tests for niche limitation by examination of species diversity in the Dunedin area, New Zealand. N Z J Bot 26:237–244CrossRefGoogle Scholar
  55. Wilson JB, Peet RK, Dengler J, Partel M (2012) Plant species richness: the world records. J Veg Sci 23:796–802. CrossRefGoogle Scholar
  56. Young TP, Peacock MM (1992) Giant senecios and alpine vegetation of Mount Kenya. J Ecol 80:141–148CrossRefGoogle Scholar
  57. Zhou Y, Chen S, Hu G, Mwachala G, Yan X, Wang Q (2018) Species richness and phylogenetic diversity of seed plants across vegetation zones of Mount Kenya, East Africa. Ecol Evol 8:8930–8939. CrossRefPubMedPubMedCentralGoogle Scholar
  58. Zobel M, Otsus M, Liira J, Moora M, Mols T (2000) Is small-scale species richness limited by seed availability or microsite availability. Ecology 81:3274–3282CrossRefGoogle Scholar
  59. Zobel M, Otto R, Laanisto L, Naranjo-Cigala A, Partel M, Fernandez-Palacios JM (2011) The formation of species pools: historical habitat abundance affects current local diversity. Glob Ecol Biogeogr 20:251–259. CrossRefGoogle Scholar

Copyright information

© Swiss Botanical Society 2019

Authors and Affiliations

  1. 1.Biological FacultyMoscow State Lomonosov UniversityMoscowRussia
  2. 2.Department of Biological SciencesUniversity of EmbuEmbuKenya

Personalised recommendations