Plant species occupy different habitats on the fortress walls in Elbasan, Albania

  • Ermelinda Gjeta Biology Dept., Universiteti 'Aleksander Xhuvani', Elbasan, Albania
  • Jonathan Titus Biology Dept., SUNY-Fredonia, Fredonia, NY 14063 USA
  • Priscilla Titus Western New York Land Conservancy, East Aurora, NY USA
Keywords: colonization, urban ecology, wall flora


Vegetated walls are an important habitat for urban biodiversity. We conducted an analysis of the plant species that grow on the Elbasan, Albania fortress walls. Walls vary in age from 4th to 21st century, and in composition. On 71 walls we assessed 2,787 plants of 35 species and recorded plant size, presence of flowers or fruits, height from the ground, crevice depth, wall aspect, wall age and composition, and distance to nearest opposing wall. Eleven species, two of which were ferns, composed 93.8% of the plants. The vast majority of plants flowered and fruited on the walls. Plant density ranged from 0.1-70 plants/m2. Species distributions varied significantly based on height on the wall, crevice depth, aspect and distance to the opposing wall. These differences may be influenced by dispersal mechanisms, moisture, substrate composition, and other important environmental factors. For example, Antirrhinum majus was generally found high on south facing walls in deep cracks whereas Umbilicus rupestris was found lower on north facing walls in shallower cracks. It is important that older walls colonized by plants be maintained such that a native flora can persist where natural rocky features are lacking.


Download data is not yet available.


Angold, P. G., Sadler, J. P., Hill, M.O., Pullin, A., Rushton, S., Austin, K., Small, E., Wood, B., Wadsworth, R., Sanderson, R. & Thompson, K. 2006: Biodiversity in urban habitat patches. Science of the Total Environment 360: 196–204. doi: 10.1016/j.scitotenv.2005.08.035

Baldock, K., Goddard, M. A., Hicks, D. M., Kunin, W. E., Mitschunas, N., Osgathorpe, L. M., Potts, S. G., Robertson, K. M., Scott, A. V., Stone, G. N., Vaughan, I. P. & Memmott, J. 2015: Where is the UK’s pollinator biodiversity? The importance of urban areas for flower-visiting insects. Proceedings of the Royal Society B 282: 20142849.

Bates, R. G., Bower, V. E. & Smith, E. R. 1956: Calcium Hydroxide as a Highly Alkaline pH Standard. Research Paper 2680. Journal of Research of the National Bureau of Standards 56: 305-311.

Brandes, D. 2010: Contributions to the wall flora of Albania.

Celesti-Grapow, L., Pysek, P., Jarosık, V. & Blasi, C. 2006: Determinants of native and alien species richness in the urban flora of Rome. Diversity and Distributions 12: 490–501.

Chocholouskova, Z. & Pysek, P. 2003: Changes in composition and structure of urban flora over 120 years: a case study of the city of Plzen. Flora 198: 366– 376.

Darlington, A. 1981: Ecology of Walls. Heinemann Educational, London, 64pp.

Duchoslav, M. 2002: Flora and vegetation of stony walls in East Bohemia (Czech Republic). Preslia 74: 1–25.

Francis, R. A., 2011: Wall ecology: A frontier for urban biodiversity and ecological engineering. Prog. Phys. Geograph. 35: 43–63.

Gaston, K.J. ed. 2010: Urban Ecology. Cambridge University Press, Cambridge, UK.

Geslin B., Gauzens, B., Thebault, E. & Dajoz, I. 2013: Plant-pollinator networks along a gradient of urbanisation. PLoS ONE 8, e63421. (doi: 10.1371/journal.pone.0063421)

Godefroid, S. & Koedam, N. 2007: Urban plant species patterns are highly driven by density and function of built-up areas. Landscape Ecology 22: 1227–1239.

Grime, J. P. 2001: Plant strategies, vegetation processes, and ecosystem properties. John Wiley & Sons, Chichester, UK.

Groves, R. H. & di Castri. F., eds. 1991: Biogeography of Mediterranean invasions. Cambridge University Press, Cambridge.

Holland, P. G. 1972: The pattern of species density of old stone walls in western Ireland. Journal of Ecology 3: 799–805.

Jim, C. Y. 1998: Old stone walls as an ecological habitat for urban trees in Hong Kong. Landscape and Urban Planning 42: 29-43.

Jim, C. Y. & Chen, W. Y. 2010: Habitat effect on vegetation ecology and occurrence on urban masonry walls. Urban Forestry & Urban Greening 9: 169-178.

Kolbek, J. 1997: Plant communities on walls in the Czech Republic – preliminary notes. Zpráv České Botanické Společnosti 32 Mater 15: 61–67.

Kolbek, J, Valachovic, M. & Misikova, K. 2015: Wall vegetation in old royal mining towns in central Slovakia. Hacquetia 14: 249-263.

Kuhn, I., Brandl, R., May, R. & Klotz, S. 2004: The flora of German cities is naturally species rich. Evolutionary Ecology Research 6: 749–764.

Láníková, D. & Lososová, Z. 2009: Rocks and walls: natural versus secondary habitats. Folia Geobotanica 44: 263–280.

Lisci, M. & Pacini, E. 1993: Plants growing on the walls of Italian towns. 1. Sites and distribution. Phyton (Horn, Austria) 33: 15-26.

Lososová, Z & Láníková, D. 2010: Differences in trait compositions between rocky natural and artificial habitats. Journal of Vegetation Science 21: 520-530.

Lundholm, J. T. & Richardson, P. J. 2010: Mini-review: Habitat analogues for reconciliation ecology in urban and industrial environments. Journal of Applied Ecology 47: 966–975.

Madrea, F., Clergeau, P., Machona, N & Vergnes, A. 2015: Building biodiversity: Vegetated façades as habitats for spider and beetle assemblages. Global Ecology and Conservation 3: 222–233.

McCune, B. & Mefford, M. J. 2006: PC-ORD, multivariate analysis of ecological data, version 5.0. MjM Software Design, Gleneden Beach, OR, USA.

McIntyre, N. E., Rango, J., Fagan, W. F. & Faeth, S. H. 2001: Ground arthropod community structure in a heterogeneous urban environment. Landscape and Urban Planning 52: 257–274. (doi: 10.1016/S0169-2046(00)00122-5)

Naveh, Z. & Vernet, J.-L. 1991: The palaeohistory of the Mediterranean biota. In: Groves, R. H. & di Castri, F. (eds.): Biogeography of Mediterranean invasions. Cambridge University Press, Cambridge, pp. 19–32.

Ricotta, C., Celesti-Grapow, L., Avena, G. & Blasi, C. 2001: Topological analysis of the spatial distribution of plant species richness across the city of Rome (Italy) with the echelon approach. Landscape and Urban Planning 57: 69–76.

Simonová, D. 2008: Alien flora on walls in southern and western Moravia (Czech Republic). In: TokarskaGuzik, B., Brock, J. H., Brundu, G., Child, L., Daehler, C. C. & Pysek, P. (eds.): Plant invasions: human perception, ecological impacts and management. Backhuys Publishers, Leiden, pp. 317–332.

SPSS. 2017: IBM SPSS Statistics for Windows, Version 25.0. Armonk, NY: IBM Corp.

Sternberg, M. & Shoshany, M. 2001: Influence of slope aspect on Mediterranean woody formations: comparison of a semiarid and an arid site in Israel. Ecological Research 16: 335–345.

Świerkosz, K. 2004: Notes on the syntaxonomy of the Asplentia trichomanis class in Poland. Polish Botanical Journal 49: 203-213.

Świerkosz, K. 2012: Notes on the wall vegetation of the Lake Garda surroundings (North Italy) and its consequences to the syntaxonomy of Tortulo-Cymbalarietalia Segal 1969. Časopsis Slezskeho Muzea, serie A 61: 11-22.

ter Braak, C. J. F. & Smilauer, P. 2002: CANOCO reference manual and CanoDraw for Windows users guide. Software for Canonical Community Ordination (version 4.5). Biometris, Wageningen, NL.

Thompson, K. & McCarthy, M. A. 2008: Traits of British alien and native urban plants. Journal of Ecology 96: 853–859.

Williams, N. S. G., Schwartz, M. W., Vesk, P. A., McCarthy, M. A., Hahs, A. K., Clemants, S. E., Corlett, R. T., Duncan, R. P., Norton, B. A. & Thompson, K. 2009: A conceptual framework for predicting the effects of urban environments on floras. Journal of Ecology 97, 4–9.

Wittig, R. 2002: Siedlungsvegetation. Ulmer, Stuttgart.

Woodell, S. & Rossiter, J. 1959: The flora of Durham walls. Proceedings of the Botanical Society of the British Isles 2: 257–273.

Yalcinalp, E. & Meral, A. 2017: Wall vegetation characteristics of urban and sub-urban areas. Sustainability 9: 1691.

How to Cite
GjetaE., TitusJ. and TitusP. (2020) “Plant species occupy different habitats on the fortress walls in Elbasan, Albania”, Hacquetia, 20(1), pp. 81–90. Available at: (Accessed: 8May2021).