Abstract
Habitat fragmentation is one of the main causes of the global loss of plant biodiversity. It is also one of the major challenges in Ethiopia, where fragmented forest patches of different sizes have been observed. These patches of forest, especially in the highlands of northern Ethiopia, are mainly confined to churches. These remnant forest patches have long been said to have negative impacts (habitat amount hypothesis). Recently, however, there is evidence that these small patches of remnant forest may harbour more species than relatively large patches of the same area. We tested this hypothesis in the remnant church forests of the Ethiopian highlands using different plant growth forms. Ten church forests of different sizes were selected in which transects were established from the forest edges to the interior. A total of 56 20 × 20 m plots were used for plant sampling. All trees, shrubs and herbs within the plots were recorded, collected and identified. We found that the effects of forest fragmentation on plant diversity, abundance and composition varied with plant growth forms. In contrast to the habitat amount hypothesis, small forest fragments were found to support more tree species than relatively large forest fragments, suggesting the role of small forest fragments in maintaining species diversity. We found that soil moisture changes with fragment size but has no significant effect on plant abundance and diversity of plant growth forms. Our results indicate that a shift in conservation priorities may be needed to recognize the value of small fragmented patches of forest for biodiversity conservation, as a lack of protection of small patches of forest can lead to high cumulative impacts on biodiversity loss.
This is a preview of subscription content, log in via an institution to check access.
Similar content being viewed by others
Data availability
The dataset will be available from the corresponding author upon request.
References
Arroyo-Rodríguez V, Rös M, Escobar F et al (2013) Plant β-diversity in fragmented rain forests: testing floristic homogenization and differentiation hypotheses. J Ecol 101:1449–1458. https://doi.org/10.1111/1365-2745.12153
Asefa M, Cao M, He Y et al (2020) Ethiopian vegetation types, climate and topography. Plant Divers 42:302–311. https://doi.org/10.1016/j.pld.2020.04.004
Barbier S, Gosselin F, Balandier P (2008) Influence of tree species on understory vegetation diversity and mechanisms involved-A critical review for temperate and boreal forests. Ecol Manage 254:1–15. https://doi.org/10.1016/j.foreco.2007.09.038
Baruch Z (1984) Ordination and classification of vegetation along an altitudinal gradient in the Venezuelan páramos. Vegetation 55:115–126
Bates D, Mächler M, Bolker B, Walker S (2015) Fitting Linear mixed-effects models using lme4. J Stat Softw 67(1):1–48
Belay T, Mengistu D (2019) Land use and land cover dynamics and drivers in the Muga watershed, Upper Blue Nile basin, Ethiopia. RSASE 5:100249
Bennett A, Saunders D (2010) Habitat fragmentation and landscape change. Conserv Biol 93:1544–1550
Bunyan M, Jose S, Fletcher R (2012) Edge effects in Small Forest fragments: why more is better? Am J Plant Sci 03:869–878. https://doi.org/10.4236/ajps.2012.37104
Cardelús CL, Mekonnen AB, Jensen KH et al (2020) Edge effects and human disturbance influence soil physical and chemical properties in Sacred Church forests in Ethiopia. Plant Soil 453:329–342. https://doi.org/10.1007/s11104-020-04595-0
CDOA (2020) Chilga District Agricultural Office Annual Report, Aykel, Ethiopia
Chapin F (1991) Integrated response of plants to strees. Bioscience 41:29e36
Chase JM, Blowes SA, Knight TM et al (2020) Ecosystem decay exacerbates biodiversity loss with habitat loss. Nature 584:238–243
Chen J, Saunders SC, Crow TR et al (1999) Microclimate in forest ecosystem and landscape ecology: variations in local climate can be used to monitor and compare the effects of different management regimes. Bioscience 49:288–297. https://doi.org/10.2307/1313612
Daye DD, Healey JR (2015) Impacts of land-use change on sacred forests at the landscape scale. Glob Ecol Conserv 3:349–358. https://doi.org/10.1016/j.gecco.2014.12.009
Deane D, He F (2018) Loss of only the smallest patches will reduce species diversity in most discrete habitat networks. Glob Chang Biol 24:5802–5814
Diamond JM (1975) The island dilemma: lessons of modern biogeographic studies for the design of natural reserves. Biol Conserv 7:129–146. https://doi.org/10.1016/0006-3207(75)90052-X
Dormann CF, Bagnara M, Boch S et al (2020) Plant species richness increases with light availability, but not variability, in temperate forests understorey. BMC Ecol 20:1–9. https://doi.org/10.1186/s12898-020-00311-9
Ewers RM, Didham RK (2006) Confounding factors in the detection of species responses to habitat fragmentation. Biol Rev Camb Philos Soc 81:117–142. https://doi.org/10.1017/S1464793105006949
Fahrig L (2003) Effects of Habitat Fragmentation on Biodiversity. Annu Rev Ecol Evol Syst 34:487–515. https://doi.org/10.1146/annurev.ecolsys.34.011802.132419
Fahrig L (2017) Ecological responses to Habitat Fragmentation per Se. Annu Rev Ecol Evol Syst 48:1–23. https://doi.org/10.1146/annurev-ecolsys-110316-022612
Fahrig L (2020) Why do several small patches hold more species than few large patches? Glob Ecol Biogeogr 29:615–628. https://doi.org/10.1111/geb.13059
Fox JW, Vasseur D, Cotroneo M et al (2017) Population extinctions can increase metapopulation persistence. Nat Ecol Evol 1:1271–1278. https://doi.org/10.1038/s41559-017-0271-y
Friis I, Sebsebe D, Breugel P (2010) Atlas of the potential vegetation. of Ethiopia Atlas of the potential vegetation of Ethiopia
Gebrehiwot K, Demissew S, Woldu Z et al (2019) Elevational changes in vascular plants richness, diversity, and distribution pattern in Abune Yosef mountain range, Northern Ethiopia. Plant Divers 41:220–228. https://doi.org/10.1016/j.pld.2019.06.005
Haddad NM, Brudvig LA, Clobert J et al (2015) Habitat fragmentation and its lasting impact on Earth’s ecosystems. Sci Adv 1:1–10. https://doi.org/10.1126/sciadv.1500052
Hammill E, Clements CF (2020) Imperfect detection alters the outcome of management strategies for protected areas. Ecol Lett 23:682–691. https://doi.org/10.1111/ele.13475
Hanski I (2015) Habitat fragmentation and species richness. J Biogeogr 42:989–993. https://doi.org/10.1111/jbi.12478
Honnay O, Verheyen K, Hermy M (2002) Permeability of ancient forest edges for weedy plant species invasion. Ecol Manage 161:109–122. https://doi.org/10.1016/S0378-1127(01)00490-X
Jacobs BF (2004) Palaeobotanical studies from tropical Africa: relevance to the evolution of forest, woodland and savannah biomes. Philos Trans R Soc B Biol Sci 359:1573–1583. https://doi.org/10.1098/rstb.2004.1533
King SA, Buckney RT (2001) Exotic plants in the soil-stored seed bank of urban bushland. Aust J Bot 49(6):717–720
Laurance W (2002) Hyperdynamism in fragmented habitats. J Veg Sci 13:595–602
Lin L, Cao M (2009) Edge effects on soil seed banks and understory vegetation in subtropical and tropical forests in Yunnan, SW China. Ecol Manage 257:1344–1352. https://doi.org/10.1016/j.foreco.2008.12.004
Lovett J (2010) Tanzanian Forest Tree Plot Diversity and Elevation Author (s): Jon C. Lovett Published by : Cambridge University Press Stable URL : http://www.jstor.org/stable/2560212. Diversity 15:689–694
McCann JC (1997) The plow and the forest: narratives of deforestation in Ethiopia, 1840–1992. Environ Hist Durh N C 2:138–159. https://doi.org/10.2307/3985505
Mcdonald RI, Urban DL (2006) Edge effects on species composition and exotic species abundance in the North Carolina Piedmont. 1049–1060. https://doi.org/10.1007/s10530-005-5227-5
Mendes CB, Prevedello JA (2020) Does habitat fragmentation affect landscape-level temperatures ? A global analysis. Landsc Ecol 35:1743–1756. https://doi.org/10.1007/s10980-020-01041-5
Mittermeier RA, Gil PR, Hoffman M et al (2004) Hotspots revisited: Earth’s biologically richest and most endangered terrestrial ecoregions. Conservation International, Washington
Negassa MD, Mallie DT, Gemeda DO (2020) Forest cover change detection using Geographic Information Systems and remote sensing techniques: a spatio-temporal study on Komto protected forest priority area, East Wollega Zone, Ethiopia. Environ Syst Res 9:1–14. https://doi.org/10.1186/s40068-020-0163-z
Normann C, Tscharntke T, Scherber C (2016) How forest edge-center transitions in the herb layer interact with beech dominance versus tree diversity. J Plant Ecol 9:498–507. https://doi.org/10.1093/jpe/rtw004
Oksanen J, Simpson G, Blanchet F et al (2022) vegan: Community Ecology Package_. R package version 2.6-4, https://CRAN.R-project.org/package=vegan
Parra-Sanchez E, Banks-Leite C (2020) The magnitude and extent of edge effects on vascular epiphytes across the Brazilian Atlantic Forest. Sci Rep 10:1–11. https://doi.org/10.1038/s41598-020-75970-1
Pasion BO, Roeder M, Liu J et al (2018) Trees represent community composition of other plant life-forms, but not their diversity, abundance or responses to fragmentation. Sci Rep 8:2–14. https://doi.org/10.1038/s41598-018-29635-9
Phillips OL, Rose S, Mendoza AM, Vargas PN (2006) Resilience of southwestern Amazon forests to anthropogenic edge effects. Conserv Biol 20:1698–1710. https://doi.org/10.1111/j.1523-1739.2006.00523.x
Rahbek C (2005) The role of spatial scale and the perception of large-scale species-richness patterns. Ecol Lett 8:224–239. https://doi.org/10.1111/j.1461-0248.2004.00701.x
Rantalainen ML, Haimi J, Fritze H et al (2008) Soil decomposer community as a model system in studying the effects of habitat fragmentation and habitat corridors. Soil Biol Biochem 40:853–863. https://doi.org/10.1016/j.soilbio.2007.11.008
Riva F, Fahrig L (2022) The disproportionately high value of small patches for biodiversity conservation. Conserv Lett 15:1–7. https://doi.org/10.1111/conl.12881
Riva F, Fahrig L (2023) Landscape-scale habitat fragmentation is positively related to biodiversity, despite patch-scale ecosystem decay Landscape-scale habitat fragmentation is positively related to biodiversity, despite patch-scale ecosystem decay. Ecol Lett 26(2):268–277. https://doi.org/10.1111/ELE.14145
Rodríguez-Loinaz G, Amezaga I, Onaindia M (2012) Does forest fragmentation affect the same way all growth-forms? J Environ Manage 94:125–131. https://doi.org/10.1016/j.jenvman.2011.06.024
Ruokolainen K, Tuomisto H, Macía MJ et al (2018) Are Floristic and Edaphic Patterns in Amazonian Rain Forests Congruent for Trees, Pteridophytes and Melastomataceae ? Stable URL : https://www.jstor.org/stable/4499065 REFERENCES Linked references are available on JSTOR for this article : Are floristic a. 23:13–25
Ruwanza S (2019) The edge effect on plant diversity and soil properties in abandoned fields targeted for ecological restoration. Sustain 11. https://doi.org/10.3390/su11010140
Rybicki J, Abrego N, Ovaskainen O (2020) Habitat fragmentation and species diversity in competitive communities. Ecol Lett 23:506–517
Santana LD, Prado-Junior JA, José JH et al (2021) Edge effects in forest patches surrounded by native grassland are also dependent on patch size and shape. Ecol Manage 482. https://doi.org/10.1016/j.foreco.2020.118842
Sharma CM, Suyal S, Gairola S, Ghildiyal SK (2009) Species richness and diversity along an altitudinal gradient in moist temperate forest of Garhwal Himalaya. J Am Sci 5:119–128
Simberloff D, Abele LG (1982) Refuge design and island biogeographic theory: effects of fragmentation. Am Nat 120:41–50. https://doi.org/10.1086/283968
Slik JWF, Paoli G, Mcguire K et al (2013) Large trees drive forest aboveground biomass variation in moist lowland forests across the tropics. Glob Ecol Biogeogr 22:1261–1271. https://doi.org/10.1111/geb.12092
Trigas P, Panitsa M, Tsiftsis S (2013) Elevational Gradient of Vascular Plant Species Richness and Endemism in Crete - the Effect of Post-isolation Mountain Uplift on a Continental Island System. PLoS ONE 8. https://doi.org/10.1371/journal.pone.0059425
Tuff KT, Tuff T, Davies KF (2016) A framework for integrating thermal biology into fragmentation research. Ecol Lett 19:361–374. https://doi.org/10.1111/ele.12579
Tulloch AIT, Barnes MD, Ringma J et al (2016) Understanding the importance of small patches of habitat for conservation. J Appl Ecol 53:418–429. https://doi.org/10.1111/1365-2664.12547
Tullus T, Lutter R, Randlane T et al (2022) The effect of stand age on biodiversity in a 130-year chronosequence of Populus tremula stands. Ecol Manage 504:119833
Valadi G, Rad JE, Khodakarami Y et al (2022) Plant Ecology Edge influence on herbaceous plant species, diversity and soil properties in sparse oak forest fragments in Iran. J Plant Ecol 15:413–424
Villard MA, Metzger JP (2014) Beyond the fragmentation debate: a conceptual model to predict when habitat configuration really matters. J Appl Ecol 51:309–318. https://doi.org/10.1111/1365-2664.12190
Wang X, Blanchet FG, Koper N (2014) Measuring habitat fragmentation: an evaluation of landscape pattern metrics. Methods Ecol Evol 5:634–646. https://doi.org/10.1111/2041-210X.12198
Wassie A, Sterck FJ, Bongers F (2010) Species and structural diversity of church forests in a fragmented Ethiopian Highland landscape. J Veg Sci 21:938–948
Weiher E, Keddy P (1995) Assembly rules, null models, and trait dispersion – new questions front old patterns. Oikos 74:159–164
Wickham H (2016) ggplot2: elegant graphics for data analysis, 2nd edn. Springer International Publishing
Wintle BA, Kujala H, Whitehead A et al (2019) Global synthesis of conservation studies reveals the importance of small habitat patches for biodiversity. Proc Natl Acad Sci U S A 116:909–914. https://doi.org/10.1073/pnas.1813051115
Acknowledgements
We would like to thank the University of Gondar for its financial support. We would also like to thank the Ethiopian Orthodox Tewahido Christian churches for the permission for this study to be conducted in their owned church forests.
Author information
Authors and Affiliations
Contributions
AA and MA designed the study; AA collected the data; MA analyzed the data; AA, MA and EK wrote the manuscript. All authors greatly contributed to the manuscript and provided final approval for the publication.
Corresponding author
Ethics declarations
Competing interests
The authors declare no competing interests.
Additional information
Communicated by Arshad Ali.
Publisher’s Note
Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Electronic supplementary material
Below is the link to the electronic supplementary material.
Rights and permissions
Springer Nature or its licensor (e.g. a society or other partner) holds exclusive rights to this article under a publishing agreement with the author(s) or other rightsholder(s); author self-archiving of the accepted manuscript version of this article is solely governed by the terms of such publishing agreement and applicable law.
About this article
Cite this article
Abuhay, A., Kassaw, E., Masresha, G. et al. Small forest patches in Ethiopian highlands uniquely support high plant biodiversity. Biodivers Conserv 33, 1711–1727 (2024). https://doi.org/10.1007/s10531-024-02827-8
Received:
Revised:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1007/s10531-024-02827-8