Abstract
Deforestation generates forest fragments within modified matrices with frequently detrimental outcomes for species persistence. In historically fragmented forests, however, species have evolved under fragmented conditions and might be more resistant to matrix modification. We examined the effect of contemporary matrix modification on forest regeneration potential in historically fragmented scarp-forest fragments in KwaZulu-Natal, South Africa. We compared tree, seedling and sapling richness, abundance and composition in four forest-fragment types varying in size and matrix modification: large and small forest fragments surrounded by indigenous grassland and small forest fragments enclosed by eucalypt and sugarcane plantations. Additionally, seedlings and saplings were classified according to their origin as local/immigrant to assess matrix permeability for seed dispersers. Trees and saplings were slightly, and seedlings were strongly affected by matrix modification: seedling richness and abundance were significantly lower in fragments with sugarcane compared to fragments with grassland matrix. Furthermore, all stage classes revealed compositional differences with more light-tolerant species occurring in fragments with modified matrices. We suggest that human activities associated with matrix modification have altered abiotic factors favouring light-tolerant species. Changes in biotic interactions seemed to be less important as immigrant recruitment occurred in all fragments implying matrix permeability. Our findings demonstrate that contemporary matrix modification might threaten forest regeneration potential in historical scarp-forest fragments. Reduced seedling recruitment might indicate a time lag in local species extinction, as trees regenerate slowly. We recommend using both recruitment stages as early warning indicators. Further, including small scarp-forest fragments into protected area networks will minimise human impact associated with matrix modification.
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Acknowledgments
We thank the Robert Bosch foundation for funding this project. We are grateful to Ezemvelo KZN Wildlife for granting us access to their nature reserves. We thank David Johnson, Tony Abbott and Christina Potgieter for their help with plant identification and categorising tree species according to their successional status. We are thankful to Megan E. Griffiths for providing guidance in categorising species according to the dispersal mode. We gratefully acknowledge the valuable feedback on this manuscript offered by Eike Lena Neuschulz, Dana G. Berens and Jörg Albrecht as well as two anonymous reviewers. We also thank Friederike A. Voigt and Lackson Chama for scientific advice and Mike and Heidi Neethling, George Archibald, Pravin Pillay and Sandy-Lynn Steenhuisen for manifold support.
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Communicated by Shayne Martin Jacobs.
Appendix
Appendix
Light tolerance (lt: E = edge I = interior), dispersal mode (dm: G = gravity, W = wind, Z = zoochorous) and presence (+) or absence (−) of tree (tr), seedling (se) and sapling (sa) species in large natural forest fragments (ForFra), small natural forest fragments surrounded by grassland (GraFra) and in small modified forest fragments surrounded by plantations (PlaFra) and agriculture (AgrFra) in KwaZulu-Natal, South Africa (modified from Botzat et al. 2013).
Forest-fragment type | lt | dm | ForFra | GraFra | PlaFra | AgrFra | ||||||||
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Species (family) | tr | se | sa | tr | Se | sa | tr | se | Sa | Tr | se | sa | ||
Acacia caffra (Fabaceae) | E | Z | − | − | − | + | − | − | − | − | − | − | − | − |
Acacia mearnsii (Fabaceae) | E | Z | − | − | − | − | − | − | − | + | + | − | − | − |
Acalypha glabrata var. glabrata (Euphorbiaceae) | E | Z | + | + | + | − | − | + | − | − | − | + | − | − |
Acokanthera oppositifolia (Apocynaceae) | I | Z | − | − | + | − | − | − | − | − | − | + | − | + |
Acridocarpus natalitius var. natalitius (Malpighiaceae) | I | W | − | − | + | − | − | − | − | − | − | − | − | − |
Albizia adianthifolia var. adianthifolia (Fabaceae) | E | Z | + | − | − | − | − | − | − | − | − | + | − | + |
Allophylus africanus var. africanus (Sapindaceae) | E | Z | + | − | − | − | + | − | − | + | − | + | − | + |
Allophylus dregeanus (Sapindaceae) | E | Z | − | − | − | + | + | − | + | + | − | + | − | − |
Antidesma venosum (Euphorbiaceae) | E | Z | + | − | − | − | − | − | − | − | − | + | − | − |
Apodytes dimidiata subsp. dimidiata (Icacinaceae) | E | Z | − | − | − | + | + | − | + | − | − | + | + | + |
Bachmannia woodii (Capparaceae) | I | Z | − | − | + | − | − | − | − | − | − | − | − | − |
Baphia racemosa (Fabaceae) | I | G | + | + | + | − | − | − | − | − | − | − | − | − |
Bersama swinnyi (Melianthaceae) | I | Z | − | − | − | − | − | − | − | − | − | + | − | − |
Bersama tysoniana (Melianthaceae) | I | Z | + | − | − | + | − | + | − | − | − | + | + | − |
Brachylaena discolour (Asteraceae) | E | W | − | − | − | + | − | − | − | − | − | + | − | − |
Brachylaena elliptica (Asteraceae) | E | W | + | − | − | − | − | − | − | − | − | − | − | − |
Brachylaena uniflora (Asteraceae) | E | W | + | − | − | − | − | − | − | − | − | − | − | − |
Bridelia micrantha (Euphorbiaceae) | I | Z | + | − | − | + | − | − | − | − | − | + | − | − |
Burchellia bubalina (Rubiaceae) | E | Z | − | − | − | − | − | − | − | − | − | − | − | + |
Calodendrum capense (Rutaceae) | I | Z | − | − | − | − | − | − | − | − | − | + | − | − |
Calpurnia aurea subsp. aurea (Fabaceae) | E | Z | − | − | + | − | + | − | − | − | − | − | − | − |
Canthium ciliatum (Rubiaceae) | E | Z | − | − | − | + | + | + | − | + | + | + | + | + |
Canthium inerme (Rubiaceae) | E | Z | + | − | − | − | + | − | − | − | − | + | − | + |
Canthium spinosum (Rubiaceae) | E | Z | -− | − | − | + | − | + | + | − | + | + | + | + |
Canthium suberosum (Rubiaceae) | I | Z | + | − | − | − | − | − | − | − | − | − | − | − |
Carissa macrocarpa (Apocynaceae) | E | Z | − | − | − | − | − | − | − | − | − | + | − | − |
Cassine peragua subsp. peragua (Celastraceae) | E | Z | − | − | − | − | − | − | − | − | − | + | − | − |
Cassipourea gummiflua var. verticillata (Rhizophoraceae) | I | Z | + | + | − | + | + | + | − | − | − | − | − | − |
Cassipourea malosana (Rhizophoraceae) | I | Z | + | − | + | + | + | + | − | − | − | + | − | − |
Caturanegam obovata (Rubiaceae) | E | Z | − | − | − | + | + | + | − | − | − | − | − | − |
Celtis africana (Celtidaceae) | E | Z | + | + | − | + | + | + | + | + | + | + | − | − |
Cestrum laevigatum (Solanaceae) | E | Z | − | − | − | − | − | − | + | + | + | − | − | − |
Chaetachme aristata (Celtidaceae) | I | Z | + | − | − | + | + | − | − | − | − | − | − | − |
Clausena anisata (Rutaceae) | E | Z | + | + | − | − | − | − | − | − | − | + | − | + |
Clerodendrum glabrum var. glabrum (Lamiaceae) | E | Z | + | − | − | + | − | − | + | − | + | + | − | − |
Cnestis polyphylla (Connaraceae) | I | Z | − | − | − | − | − | − | − | + | + | − | − | + |
Combretum erythrophyllum (Combretaceae) | E | Z | + | − | − | + | + | − | + | + | + | + | + | + |
Combretum kraussii (Combretaceae) | E | W | + | + | + | + | + | + | + | + | + | + | + | + |
Commiphora harveyi (Burseraceae) | I | Z | − | − | − | − | − | − | + | − | − | − | − | − |
Commiphora woodii (Burseraceae) | I | Z | − | − | − | − | + | − | − | − | − | − | − | − |
Croton sylvaticus (Euphorbiaceae) | E | Z | + | − | − | + | − | − | + | − | − | + | − | − |
Cryptocaria myrtifolia (Lauraceae) | I | Z | − | − | − | − | − | + | − | − | − | − | − | − |
Cryptocaria woodii (Lauraceae) | I | Z | − | − | − | − | − | − | − | − | − | + | − | + |
Cryptocaria wyliei (Lauraceae) | E | Z | − | − | − | − | − | − | − | − | − | + | − | + |
Cunonia capensis (Cunoniaceae) | E | W/Z | − | − | − | + | − | − | − | − | − | + | − | − |
Cussonia sphaerocephala (Araliaceae) | I | Z | + | − | − | + | − | − | − | − | − | + | − | − |
Cussonia spicata (Araliaceae) | E | Z | + | − | − | − | − | − | − | − | − | + | − | − |
Deinbollia oblongifolia (Sapindaceae) | E | Z | + | − | − | − | − | − | − | − | − | − | − | − |
Diospyros lycioides subsp. sericea (Ebenaceae) | E | Z | + | + | + | − | − | + | + | + | + | + | − | − |
Diospyros villosa var. villosa (Ebenaceae) | E | Z | − | − | − | − | − | − | + | − | − | − | − | − |
Dombeya burgessiae (Sterculiaceae) | E | G/W | + | − | − | − | − | − | − | − | − | + | − | − |
Dombeya tiliaceae (Sterculiaceae) | E | G/W | − | − | − | − | − | + | − | − | + | − | − | − |
Dovyalis lucida (Flacourtiaceae) | E | Z | − | − | − | + | − | − | − | − | − | − | − | − |
Dovyalis rhamnoides (Flacourtiaceae) | E | Z | − | − | − | − | − | + | + | + | + | + | − | − |
Dracaena aletriformis (Dracaenaceae) | E | Z | + | − | − | + | − | − | + | + | + | + | − | + |
Drypetes arguta (Euphorbiaceae) | I | Z | + | + | + | − | − | + | − | − | − | − | − | + |
Drypetes gerrardii var. gerrardii (Euphorbiaceae) | I | Z | + | + | − | + | − | + | + | − | − | + | + | + |
Ekebergia capensis (Meliaceae) | E | Z | − | − | − | − | − | − | − | − | − | + | − | − |
Elaeodendron croceum (Celastraceae) | E | Z | + | + | + | − | − | − | − | − | − | + | − | + |
Englerophytum natalense (Sapotaceae) | I | Z | + | + | + | + | + | + | − | − | − | + | − | − |
Erythrina caffra (Fabaceae) | E | Z | − | − | − | + | − | − | − | − | − | − | − | − |
Erythrina lysistemon (Fabaceae) | E | Z | − | − | − | − | − | − | − | − | + | − | − | − |
Erythrococca sp. nov. (Euphorbiaceae) | I | Z | − | − | − | − | − | − | − | − | − | + | − | − |
Eucalyptus grandis (Myrtaceae) | E | W | − | − | − | − | − | − | + | − | − | − | − | − |
Euclea natalensis subsp. natalensis (Ebenaceae) | I | Z | − | − | − | − | − | − | − | − | − | + | − | + |
Eugenia natalitia (Myrtaceae) | I | Z | − | − | − | + | + | + | + | − | + | + | − | + |
Eugenia umtamvunensis (Myrtaceae) | I | Z | + | + | + | − | − | − | − | + | − | − | − | − |
Eugenia woodii (Myrtaceae) | E | Z | + | − | − | − | − | − | − | − | − | − | − | − |
Faurea saligna (Proteaceae) | E | W | − | − | − | − | − | − | − | − | + | − | − | − |
Ficus burkei (Moraceae) | E | Z | + | − | − | + | − | − | − | − | − | − | − | − |
Ficus burtt-davyi (Moraceae) | E | Z | + | − | − | − | − | − | − | − | − | − | − | − |
Ficus craterostoma (Moraceae) | I | Z | − | − | − | + | − | − | + | − | − | + | − | − |
Ficus natalensis subsp. natalensis (Moraceae) | E | Z | + | − | − | − | − | − | − | − | − | + | − | − |
Ficus sur (Moraceae) | E | Z | − | − | − | + | − | − | + | − | − | + | − | − |
Gardenia thunbergia (Rubiaceae) | I | Z | + | − | − | − | − | − | − | − | − | − | − | − |
Grewia lasiocarpa (Tiliaceae) | E | Z | − | − | − | + | + | + | + | + | + | + | − | + |
Grewia occidentalis var. occidentalis (Tiliaceae) | E | Z | − | − | − | + | − | − | + | − | − | + | − | − |
Gymnosporia buxifolia (Celastraceae) | E | Z | − | − | − | − | − | + | − | − | − | + | − | − |
Gymnosporia harveyana (Celastraceae) | I | Z | + | − | − | − | + | + | − | − | − | + | + | + |
Gymnosporia nemorosa (Celastraceae) | E | Z | + | − | − | − | − | − | − | − | − | − | − | − |
Halleria lucida (Scrophulariaceae) | E | Z | − | − | − | − | − | − | + | − | + | + | − | + |
Harpephyllum caffrum (Anacardiaceae) | I | Z | − | − | − | − | − | − | − | − | − | + | − | − |
Heywoodia lucens (Euphorbiaceae) | E | G | + | − | − | − | − | − | − | − | − | − | − | − |
Hippobromus pauciflorus (Sapindaceae) | E | Z | + | − | − | − | + | − | − | − | − | − | − | − |
Hyperacanthus amoenus (Rubiaceae) | I | Z | + | − | − | − | − | − | − | − | − | − | − | − |
Kiggelaria africana (Flacourtiaceae) | E | Z | − | − | − | − | − | − | + | − | + | − | − | − |
Macaranga capensis (Euphorbiaceae) | E | Z | − | − | − | + | − | − | − | − | − | − | − | − |
Maerua cafra (Capparaceae) | E | Z | − | − | − | − | − | − | − | − | − | + | − | + |
Maesa lanceolata (Maesaceae) | E | Z | + | − | − | − | − | + | + | + | + | + | − | + |
Margaritaria discoidea var. fagifolia (Euphorbiaceae) | I | Z | + | − | − | − | − | − | − | − | − | − | − | − |
Maytenus abbottii (Celastraceae) | E | Z | − | − | − | − | − | − | − | − | − | + | − | + |
Maytenus acuminata (Celastraceae) | E | Z | − | − | − | − | − | − | − | + | − | − | − | − |
Maytenus peduncularis (Celastraceae) | I | Z | − | − | − | − | − | − | − | − | − | + | − | − |
Maytenus undata (Celastraceae) | E | Z | − | − | − | − | − | − | + | − | − | − | − | − |
Memecylon natalense (Melastomataceae) | I | Z | − | + | − | + | − | − | − | − | − | − | − | − |
Millettia grandis (Fabaceae) | E | G | + | + | + | − | − | − | − | − | − | − | − | − |
Mimusops obovata (Sapotaceae) | E | Z | + | − | − | − | − | − | − | − | − | + | − | − |
Nectaropetalum capense (Erythroxylaceae) | I | Z | + | − | + | − | − | − | − | − | − | − | − | − |
Obetia tenax (Urticaceae) | E | G | + | − | + | − | − | − | − | − | − | + | − | − |
Ochna arborea var. arborea (Ochnaceae) | I | Z | + | + | + | − | + | − | − | − | − | − | − | − |
Ochna serrulata (Ochnaceae) | E | Z | − | − | + | − | + | + | − | + | − | − | + | − |
Olea capensis subsp. macrocarpa (Oleaceae) | I | Z | − | − | + | − | − | − | − | − | − | − | − | − |
Oricia bachmannii (Rutaceae) | I | Z | + | − | + | − | − | − | − | − | − | − | − | − |
Pavetta bowkeri (Rubiaceae) | I | Z | − | − | − | − | − | − | − | + | + | − | − | + |
Pavetta lanceolata (Rubiaceae) | E | Z | − | − | − | − | − | − | + | + | + | + | − | − |
Peddiea africana (Thymelaeaceae) | I | Z | + | + | − | + | + | + | + | + | + | − | + | + |
Phoenix reclinata (Arecaceae) | E | Z | + | + | + | + | + | + | + | + | + | − | − | − |
Pittosporum viridiflorum (Pittosporaceae) | E | Z | + | − | − | + | − | − | − | − | − | + | − | + |
Pleurostylia capensis (Celastraceae) | I | Z | + | − | + | + | + | + | + | − | − | + | − | + |
Podocarpus latifolius (Anacardiaceae) | I | Z | − | − | − | − | − | + | − | + | + | − | − | − |
Protorhus longifolia (Celastraceae) | E | Z | + | + | − | + | + | + | + | + | + | + | + | + |
Pseudoscolopia polyantha (Flacourtiaceae) | E | Z | − | − | − | − | − | − | − | − | + | − | − | − |
Psychotria capensis subsp. capensis var. capensis (Rubiaceae) | E | Z | + | − | + | + | + | + | + | + | + | + | + | + |
Rapanea melanophloeos (Myrsinaceae) | E | Z | + | − | − | + | + | + | − | − | − | + | + | + |
Rauvolfia caffra (Apocynaceae) | E | Z | + | + | − | − | + | − | − | − | − | − | − | − |
Rawsonia lucida (Flacourtiaceae) | I | G | − | − | − | + | + | + | − | − | − | − | − | − |
Rothmannia globosa (Rubiaceae) | I | Z | + | − | − | − | + | − | − | − | − | + | + | + |
Schrebera alata (Oleaceae) | E | W | + | − | − | − | − | − | − | − | − | − | − | − |
Scolopia zeyheri (Flacourtiaceae) | E | Z | − | − | − | + | − | − | − | − | − | − | − | − |
Searsia chirindensis (Anacardiaceae) | E | Z | + | − | − | + | − | + | + | − | − | + | − | + |
Searsia dentata (Anacardiaceae) | E | Z | − | − | − | − | − | − | − | − | − | + | − | + |
Strelitzia nicolai (Strelitziaceae) | E | Z | + | − | + | + | + | + | + | − | + | − | − | − |
Strychnos decussata (Strychnaceae) | I | Z | + | − | − | − | − | − | − | − | − | − | − | − |
Strychnos henningsii (Strychnaceae) | I | Z | + | − | − | − | − | − | − | − | − | − | − | − |
Strychnos usambarensis (Strychnaceae) | I | Z | + | + | + | − | − | − | − | − | − | − | − | − |
Syzygium cordatum (Myrtaceae) | E | Z | + | − | − | + | − | − | + | − | + | + | − | − |
Tarenna pavettoides subsp. pavettoides (Rubiaceae) | E | Z | + | + | − | − | − | − | − | − | − | − | − | − |
Teclea gerrardii (Rutaceae) | I | Z | + | − | − | − | − | − | − | − | − | − | − | − |
Teclea natalensis (Rutaceae) | I | Z | + | + | − | + | − | − | − | − | − | − | − | − |
Trema orientalis (Celtidaceae) | E | Z | − | − | − | + | − | − | + | + | − | + | − | − |
Tricalysia capensis var. capensis (Rubiaceae) | E | Z | − | − | − | − | − | − | − | − | − | + | − | + |
Tricalysia sonderiana (Rubiaceae) | E | Z | + | − | − | − | − | − | − | − | − | − | − | − |
Trichilia dregeana (Meliaceae) | E | Z | − | − | − | + | − | − | − | − | − | − | − | − |
Vangueria infausta (Rubiaceae) | E | Z | − | − | − | − | + | − | − | − | + | + | − | − |
Vangueria parviflora (Rubiaceae) | E | Z | − | − | − | − | − | − | − | − | + | − | − | − |
Vepris lanceolata (Rutaceae) | E | Z | − | − | − | + | − | + | + | − | − | + | + | + |
Xylotheca kraussiana (Flacourtiaceae) | E | Z | + | − | − | − | − | − | − | − | − | − | − | − |
Xymalos monospora (Monimiaceae) | I | Z | + | − | − | − | + | − | − | − | − | − | − | − |
Zanthoxylum capense (Rutaceae) | E | Z | − | − | − | + | + | − | + | − | + | + | − | − |
Zanthoxylum davyi (Rutaceae) | I | Z | − | − | − | + | − | − | − | − | − | − | − | + |
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Botzat, A., Fischer, L. & Farwig, N. Regeneration potential in South African forest fragments: extinction debt paid off or hampered by contemporary matrix modification?. Plant Ecol 216, 535–551 (2015). https://doi.org/10.1007/s11258-015-0457-9
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DOI: https://doi.org/10.1007/s11258-015-0457-9