Register      Login
Australian Journal of Botany Australian Journal of Botany Society
Southern hemisphere botanical ecosystems
RESEARCH ARTICLE

The effect of Helichrysum shrub encroachment on orchids in a tropical, montane grassland ecosystem, Tanzania

Christopher A. Mgimba https://orcid.org/0000-0001-9940-0220 A B * , Issakwisa B. Ngondya A and Anna C. Treydte A C D
+ Author Affiliations
- Author Affiliations

A Department of Sustainable Agriculture, Biodiversity and Ecosystem Management, School of Life Sciences and Bio-engineering, The Nelson Mandela African Institution of Science and Technology, Arusha, Tanzania.

B Department of Earth Sciences, Mbeya University of Science and Technology, Mbeya, Tanzania.

C Department of Physical Geography, Stockholm University, Stockholm 106 91, Sweden.

D Ecology of Tropical Agricultural Systems, Hans Ruthenberg Institute, University of Hohenheim, Stuttgart, Germany.

* Correspondence to: mgimbac@nm-aist.ac.tz

Handling Editor: Noushka Reiter

Australian Journal of Botany 72, BT23107 https://doi.org/10.1071/BT23107
Submitted: 8 December 2023  Accepted: 17 October 2024  Published: 8 November 2024

© 2024 The Author(s) (or their employer(s)). Published by CSIRO Publishing

Abstract

Context

Although shrub encroachment is a common phenomenon in grasslands, which often suppress co-existing herbaceous plants, little is known about how encroaching native shrubs affect endemic and threatened orchid species.

Aims

We assessed the effect of the native dwarf shrub Helichrysum species on orchid species in a protected mountainous grassland system in Tanzania.

Methods

We selected five Helichrysum shrub-dominated blocks and applied four treatments in each, i.e. no or low encroachment (<20% Helichrysum cover; ‘low cover’), high encroachment (>50% Helichrysum cover; ‘high cover’), cutting all stems of Helichrysum shrubs to ground level (‘stem cut’) and removing both stems and roots of all Helichrysum shrubs (‘uprooted’). We then compared orchid species diversity, abundance and functional traits by using a mixed linear model across treatments.

Key results

Orchid species diversity and abundance were significantly lower in ‘high cover’ plots than in other treatments. In ‘high cover’ plots, orchid species such as Disa robusta, Satyrium acutirostrum, and S. sphaeranthum had a significantly lower chlorophyll content than they did in ‘low cover’ plots. The ‘uprooting’ treatment showed significantly higher orchid species diversity in the second field season.

Conclusion

The expansion of Helichrysum shrubs adversely affected orchid abundance, diversity, and individual vigour, which in turn affected the regenerative ability of orchids.

Implications

We suggest that management should focus on shrub removal, because only ‘cutting’ had a beneficial effect on orchids. Shrub removal should be focused on areas of high shrub cover to promote further orchid species growth in this mountainous grassland of Tanzania.

Keywords: Africa, below- and aboveground competition, cutting, Helichrysum splendidum, Kitulo National Park, plant–plant interactions, protected area, uprooting.

References

Acuña-Tarazona M, Mehltreter K, Toledo-Aceves T, Sosa VJ, Flores-Palacios A, Kessler M (2022) Effects of microenvironmental factors on the diversity and composition of fern and orchid assemblages in an Andean paramo in Peru. Flora 293, 152107.
| Crossref | Google Scholar |

Aguiar ACMd, Silva DROd, Basso CJ, Soriani HH, Novello BD, Muraro DS (2019) Interference of volunteer corn in growth and chlorophyll fluorescence of bean. Revista Ceres 66(3), 210-219.
| Crossref | Google Scholar |

Baker NR, Rosenqvist E (2004) Applications of chlorophyll fluorescence can improve crop production strategies: an examination of future possibilities. Journal of Experimental Botany 55(403), 1607-1621.
| Crossref | Google Scholar | PubMed |

Baldelomar M, Atala C, Molina-Montenegro MA (2019) Top-down and bottom-up effects deployed by a nurse shrub allow facilitating an endemic Mediterranean orchid. Frontiers in Ecology and Evolution 7, 466.
| Crossref | Google Scholar |

Ballantyne M, Pickering CM (2015) Shrub facilitation is an important driver of alpine plant community diversity and functional composition. Biodiversity and Conservation 24(8), 1859-1875.
| Crossref | Google Scholar |

Baltieri M, Fantinato E, Del Vecchio S, Buffa G (2020) Intraspecific variability of leaf traits and functional strategy of Himantoglossum adriaticum H.Baumann. Plant Sociology 57(2), 105-112.
| Crossref | Google Scholar |

Bashirzadeh M, Shefferson RP, Farzam M (2022) Plant–plant interactions determine natural restoration of plant biodiversity over time, in a degraded mined land. Ecology and Evolution 12(5), e8878.
| Crossref | Google Scholar |

Bruno JF, Stachowicz JJ, Bertness MD (2003) Inclusion of facilitation into ecological theory. Trends in Ecology & Evolution 18(3), 119-125.
| Crossref | Google Scholar |

Cao X, Liu Y, Cui X, Chen J, Chen X (2019) Mechanisms, monitoring and modeling of shrub encroachment into grassland: a review. International Journal of Digital Earth 12(6), 625-641.
| Crossref | Google Scholar |

Casal JJ (2012) Shade avoidance. The Arabidopsis Book 10, e0157.
| Crossref | Google Scholar | PubMed |

Casper BB, Cahill JF, Hyatt LA (1998) Above-ground competition does not alter biomass allocated to roots in Abutilon theophrasti. New Phytologist 140(2), 231-238.
| Crossref | Google Scholar | PubMed |

Cavieres LA, Badano EI, Sierra-Almeida A, Gómez-González S, Molina-Montenegro MA (2006) Positive interactions between alpine plant species and the nurse cushion plant Laretia acaulis do not increase with elevation in the Andes of central Chile. New Phytologist 169(1), 59-69.
| Crossref | Google Scholar | PubMed |

Challe JFX, Price LL (2009) Endangered edible orchids and vulnerable gatherers in the context of HIV/AIDS in the Southern Highlands of Tanzania. Journal of Ethnobiology and Ethnomedicine 5(1), 41.
| Crossref | Google Scholar |

Chang Y, Xu C, Yang H, Zhou J, Hua W, Zhang S, Zhong Q, Li B (2021) Leaf structural traits vary with plant size in even-aged stands of Sapindus mukorossi. Frontiers in Plant Science 12, 692484.
| Crossref | Google Scholar |

Chen J, Wu S, Dong F, Li J, Zeng L, Tang J, Gu D (2021) Mechanism underlying the shading-induced chlorophyll accumulation in tea leaves. Frontiers in Plant Science 12, 779819.
| Crossref | Google Scholar |

Chengere SA, Steger C, Gebrehiwot K, Wube S, Dullo BW, Nemomissa S (2023) Quantifying shrub encroachment through soil seed bank analysis in the Ethiopian Highlands. PLoS ONE 18(8), e0288804.
| Crossref | Google Scholar | PubMed |

Craine JM, Dybzinski R (2013) Mechanisms of plant competition for nutrients, water and light. Functional Ecology 27(4), 833-840.
| Crossref | Google Scholar |

Cunningham SA (1997) The effect of light environment, leaf area, and stored carbohydrates on inflorescence production by a rain forest understory palm. Oecologia 111(1), 36-44.
| Crossref | Google Scholar | PubMed |

Davenport TRB, Markes S (2018) ‘Touchwood – the beauty and biodiversity of Tanzania’s Southern Highlands.’ (WCS/CEPF: Arusha, Tanzania)

Davenport TRB, Ndangalasi HJ (2003) An escalating trade in orchid tubers across Tanzania’s Southern Highlands: assessment, dynamics and conservation implications. Oryx 37(1), 55-61.
| Crossref | Google Scholar |

Djordjević V, Tsiftsis S (2020) ‘The role of ecological factors in distribution and abundance of terrestrial orchids.’ (Springer Nature: Switzerland)

Djordjević V, Tsiftsis S, Lakušić D, Jovanović S, Stevanović V (2016) Factors affecting the distribution and abundance of orchids in grasslands and herbaceous wetlands. Systematics and Biodiversity 14(4), 355-370.
| Crossref | Google Scholar |

Eldridge DJ, Bowker MA, Maestre FT, Roger E, Reynolds JF, Whitford WG (2011) Impacts of shrub encroachment on ecosystem structure and functioning: towards a global synthesis. Ecology Letters 14(7), 709-722.
| Crossref | Google Scholar | PubMed |

Emeterio-Lara A, García-Franco JG, Hernández-Apolinar M, Toledo-Hernández VH, Valencia-Díaz S, Flores-Palacios A (2021) Is pseudobulb harvest a sustainable management strategy in wild orchid populations? An experiment with Laelia autumnalis. Forest Ecology and Management 491, 119205.
| Crossref | Google Scholar |

Fay MF (2018) Orchid conservation: how can we meet the challenges in the twenty-first century? Botanical Studies 59, 16.
| Crossref | Google Scholar |

Fekete R, Bódis J, Fülöp B, Süveges K, Urgyán R, Malkócs T, Vincze O, Silva L, Molnár V. A (2020) Roadsides provide refuge for orchids: characteristic of the surrounding landscape. Ecology and Evolution 10(23), 13236-13247.
| Crossref | Google Scholar | PubMed |

Forseth IN, Wait DA, Casper BB (2001) Shading by shrubs in a desert system reduces the physiological and demographic performance of an associated herbaceous perennial. Journal of Ecology 89(4), 670-680.
| Crossref | Google Scholar |

Francis B, Gilman RT (2019) Light intensity affects leaf morphology in a wild population of Adenostyles alliariae (Asteraceae). Italian Botanist 8, 35-45.
| Crossref | Google Scholar |

Franklin KA (2008) Shade avoidance. New Phytologist 179(4), 930-944.
| Crossref | Google Scholar | PubMed |

Gosling P, van Der Gast C, Bending GD (2017) Converting highly productive arable cropland in Europe to grassland: a poor candidate for carbon sequestration. Scientific Reports 7(1), 10493.
| Crossref | Google Scholar |

Gottlieb R, Gruntman M (2022) Can plants integrate information on above-ground competition in their directional responses below ground? Annals of Botany 130(5), 763-771.
| Crossref | Google Scholar | PubMed |

Guido A, Salengue E, Dresseno A (2017) Effect of shrub encroachment on vegetation communities in Brazilian forest–grassland mosaics. Perspectives in Ecology and Conservation 15(1), 52-55.
| Crossref | Google Scholar |

Guo X, Liu X-Y, Jiang S-Y, Guo S-X, Wang J-F, Hu Y, Li S-M, Li H-M, Wang T, Sun Y-K, Li M-Y (2023) Allelopathy and arbuscular mycorrhizal fungi interactions shape plant invasion outcomes. NeoBiota 89, 187-207.
| Crossref | Google Scholar |

Hamblin J, Stefanova K, Angessa TT (2014) Variation in chlorophyll content per unit leaf area in spring wheat and implications for selection in segregating material. PLoS ONE 9(3), e92529.
| Crossref | Google Scholar |

He Y, Liu X, Xie Z (2014) Shrub effects on herbaceous vegetation vary with growth stages and herb relative location. Polish Journal of Ecology 62(3), 421-429.
| Crossref | Google Scholar |

Hinsley A, de Boer HJ, Fay MF, Gale SW, Gardiner LM, Gunasekara RS, Kumar P, Masters S, Metusala , D, Roberts DL, Veldman S, Wong S, Phelps J (2018) A review of the trade in orchids and its implications for conservation. Botanical Journal of the Linnean Society 186(4), 435-455.
| Crossref | Google Scholar |

Hodgson JG, Montserrat-Martí G, Charles M, Jones G, Wilson P, Shipley B, Sharafi M, Cerabolini BEL, Cornelissen JHC, Band SR, Bogard A, Castro-Díez P, Guerrero-Campo J, Palmer C, Pérez-Rontomé MC, Carter G, Hynd A, Romo-Díez A, de Torres Espuny L, Royo Pla F (2011) Is leaf dry matter content a better predictor of soil fertility than specific leaf area? Annals of Botany 108(7), 1337-1345.
| Crossref | Google Scholar | PubMed |

Huber M, Nieuwendijk NM, Pantazopoulou CK, Pierik R (2021) Light signalling shapes plant–plant interactions in dense canopies. Plant, Cell & Environment 44(4), 1014-1029.
| Crossref | Google Scholar | PubMed |

Hüppi R, Felber R, Neftel A, Six J, Leifeld J (2015) Effect of biochar and liming on soil nitrous oxide emissions from a temperate maize cropping system. Soil 1(2), 707-717.
| Crossref | Google Scholar |

Iyengar SB, Bagchi S, Barua D, Mishra C, Sankaran M (2017) A dominant dwarf shrub increases diversity of herbaceous plant communities in a Trans-Himalayan rangeland. Plant Ecology 218(7), 843-854.
| Crossref | Google Scholar |

Jacquemyn H, Brys R, Jongejans E (2010) Size-dependent flowering and costs of reproduction affect population dynamics in a tuberous perennial woodland orchid. Journal of Ecology 98(5), 1204-1215.
| Crossref | Google Scholar |

Jiménez-Arias D, García-Machado FJ, Morales-Sierra S, Suárez E, Pérez JA, Luis JC, Garrido-Orduña C, Herrera AJ, Valdés F, Sandalio LM, Borges AA (2019) Menadione sodium bisulphite (MSB): beyond seed-soaking. Root pretreatment with MSB primes salt stress tolerance in tomato plants. Environmental and Experimental Botany 157, 161-170.
| Crossref | Google Scholar |

Kawamura K, Takeda H (2006) Cost and probability of flowering at the shoot level in relation to variability in shoot size within the crown of Vaccinium hirtum (Ericaceae). New Phytologist 171(1), 69-80.
| Crossref | Google Scholar | PubMed |

Koch B, Edwards PJ, Blanckenhorn WU, Walter T, Hofer G (2015) Shrub encroachment affects the diversity of plants, butterflies, and grasshoppers on two swiss subalpine pastures. Arctic, Antarctic, and Alpine Research 47(2), 345-357.
| Crossref | Google Scholar |

Kononchuk O, Pidlisnyuk V, Mamirova A, Khomenchuk V, Herts A, Grycová B, Klemencová K, Leštinský P, Shapoval P (2022) Evaluation of the impact of varied biochars produced from M. × giganteus waste and application rate on the soil properties and physiological parameters of Spinacia Oleracea L. Environmental Technology & Innovation 28, 102898.
| Crossref | Google Scholar |

Kuzmenko M (2021) Dark Green Colour Index (DGCI): a new measurement of chlorophyll. (University of Arkansas: USA)

Landi M, Frignani F, Lazzeri C, Angiolini C (2009) Abundance of orchids on calcareous grasslands in relation to community species, environmental, and vegetational conditions. Russian Journal of Ecology 40(7), 486-494.
| Crossref | Google Scholar |

Lara-Romero C, García C, Morente-López J, Iriondo JM (2016) Direct and indirect effects of shrub encroachment on alpine grasslands mediated by plant–flower visitor interactions. Functional Ecology 30(9), 1521-1530.
| Crossref | Google Scholar |

Laub M, Pataczek L, Feuerbacher A, Zikeli S, Högy P (2022) Contrasting yield responses at varying levels of shade suggest different suitability of crops for dual land-use systems: a meta-analysis. Agronomy for Sustainable Development 42, 51.
| Crossref | Google Scholar |

Legner N, Fleck S, Leuschner C (2014) Within-canopy variation in photosynthetic capacity, SLA and foliar N in temperate broad-leaved trees with contrasting shade tolerance. Trees 28(1), 263-280.
| Crossref | Google Scholar |

Li P, Krüsi B, Li S, Cai X, Yu F (2011) Facilitation associated with three contrasting shrub species in heavily grazed pastures on the eastern Tibetan Plateau. Community Ecology 12(1), 1-8.
| Crossref | Google Scholar |

Li T, Wu S, Yang W, Selosse M-A, Gao J (2021) How mycorrhizal associations influence orchid distribution and population dynamics. Frontiers in Plant Science 12, 647114.
| Crossref | Google Scholar |

Li D, Liu Y, Yang X, Zhang X, Shi Z (2023) Shrub encroachment alters plant trait response to nitrogen addition in a semi-arid grassland. Frontiers in Plant Science 14, 1103371.
| Crossref | Google Scholar |

Liang Y, Urano D, Liao K-L, Hedrick TL, Gao Y, Jones AM (2017) A nondestructive method to estimate the chlorophyll content of Arabidopsis seedlings. Plant Methods 13, 26.
| Crossref | Google Scholar |

Linder HP, Kurzweil H, Johnson SD (2005) The Southern African orchid flora: composition, sources and endemism. Journal of Biogeography 32(1), 29-47.
| Crossref | Google Scholar |

Lovett JC, Prins E (1994) Estimation of land-use changes on Kitulo Plateau, Tanzania, using satellite imagery. Oryx 28(3), 173-182.
| Crossref | Google Scholar |

Mapunda LND (2007) Edible orchids in makete district, the Southern Highlands of Tanzania: distribution, population and status. Masters Thesis, Uppsala University, Sweden.

Marongiu B, Piras A, Porcedda S (2006) Comparative analysis of the oil and supercritical CO2 extract of Artemisia arborescens L. and Helichrysum splendidum (Thunb.) Less. Natural Product Research 20(5), 421-428.
| Crossref | Google Scholar | PubMed |

Martín-Forés I, Bywaters SL, Sparrow B, Guerin GR (2022) Simultaneous effect of habitat remnancy, exotic species, and anthropogenic disturbance on orchid diversity in South Australia. Conservation Science and Practice 4(4), e12652.
| Crossref | Google Scholar |

Matson E, Bart D (2013) Interactions among fire legacies, grazing and topography predict shrub encroachment in post-agricultural Páramo. Landscape Ecology 28(9), 1829-1840.
| Crossref | Google Scholar |

Mgimba CA, Smith SW, Ngondya IB, Treydte AC (2024) The interplay between historical land-use and the distribution of Helichrysum shrubs in an African-protected grassland. African Journal of Ecology 62(2), e13273.
| Crossref | Google Scholar |

Missanjo E, Ndalowa D, Malinga D (2015) Stand age and diameter class effect on seed production of Pinus kesiya Royle ex Gordon grown in Malawi. Scholars Academic Journal of Biosciences 3(2B), 173-177.
| Google Scholar |

Murphy GP, Dudley SA (2007) Above- and below-ground competition cues elicit independent responses. Journal of Ecology 95(2), 261-272.
| Crossref | Google Scholar |

Muzell Trezzi M, Vidal RA, Balbinot Junior AA, von Hertwig Bittencourt H, da Silva Souza Filho AP (2016) Allelopathy: driving mechanisms governing its activity in agriculture. Journal of Plant Interactions 11(1), 53-60.
| Crossref | Google Scholar |

Ndaki PM, Erick F, Moshy V (2021) Challenges posed by climate change and non-climate factors on conservation of edible orchid in southern highlands of Tanzania: the case of Makete district. Journal of Global Ecology and Environment 13(4), 144-167.
| Google Scholar |

Ngondya IB, Treydte AC, Ndakidemi PA, Munishi LK (2017) Invasive plants: ecological effects, status, management challenges in Tanzania and the way forward. Journal of Biodiversity and Environmental Sciences 10(3), 204-217.
| Google Scholar |

Ninčević T, Jug-Dujaković M, Grdiša M, et al. (2021) Population structure and adaptive variation of Helichrysum italicum (Roth) G. Don along eastern Adriatic temperature and precipitation gradient. Scientific Reports 11, 24333.
| Crossref | Google Scholar |

Nyomora AMS (2009) Distribution and abundance of the edible orchids of the Southern Highlands of Tanzania. Tanzania Journal of Science 31(1), 45-54.
| Crossref | Google Scholar |

Olson ME, Soriano D, Rosell JA, Anfodillo T, Donoghue MJ, Edwards EJ, León-Gómez C, Dawson T, Camarero Martínez JJ, Castorena M, Echeverría A, Espinosa CI, Fajardo A, Gazol A, Isnard S, Lima RS, Marcati CR, Méndez-Alonzo R (2018) Plant height and hydraulic vulnerability to drought and cold. Proceedings of the National Academy of Sciences 115(29), 7551-7556.
| Crossref | Google Scholar |

Parajuli R, O’Brien MJ, Timilsina B, Pugnaire FI, Schöb C, Ghimire SK (2021) Facilitation by a dwarf shrub enhances plant diversity of human-valued species at high elevations in the Himalayas of Nepal. Basic and Applied Ecology 54, 23-36.
| Crossref | Google Scholar |

Pérez-Harguindeguy N, Díaz S, Garnier E, Lavorel S, Poorter H, Jaureguiberry P, Bret-Harte MS, Cornwell WK, Craine JM, Gurvich DE, Urcelay C, Veneklaas EJ, Reich PB, Poorter L, Wright IJ, Ray P, Enrico L, Pausas JG, de Vos AC, Buchmann N, Funes G, Quétier F, Hodgson JG, Thompson K, Morgan HD, ter Steege H, van der Heijden MGA, Sack L, Blonder B, Poschlod P, Vaieretti MV, Conti G, Staver AC, Aquino S, Cornelissen JHC (2013) New handbook for standardised measurement of plant functional traits worldwide. Australian Journal of Botany 61(3), 167-234.
| Crossref | Google Scholar |

Poorter H, Niinemets Ü, Ntagkas N, Siebenkäs A, Mäenpää M, Matsubara S, Pons , TL (2019) A meta-analysis of plant responses to light intensity for 70 traits ranging from molecules to whole plant performance. New Phytologist 223(3), 1073-1105.
| Crossref | Google Scholar | PubMed |

Quinet M, Cawoy V, Lefèvre I, Van Miegroet F, Jacquemart A-L, Kinet J-M (2004) Inflorescence structure and control of flowering time and duration by light in buckwheat (Fagopyrum esculentum Moench). Journal of Experimental Botany 55(402), 1509-1517.
| Crossref | Google Scholar | PubMed |

Rani Das K (2016) A brief review of tests for normality. American Journal of Theoretical and Applied Statistics 5(1), 5-12.
| Crossref | Google Scholar |

Rodríguez-Echeverría S, Armas C, Pistón N, Hortal S, Pugnaire FI (2013) A role for below-ground biota in plant–plant facilitation. Journal of Ecology 101(6), 1420-1428.
| Crossref | Google Scholar |

Ruttan A, Filazzola A, Lortie CJ (2016) Shrub-annual facilitation complexes mediate insect community structure in arid environments. Journal of Arid Environments 134, 1-9.
| Crossref | Google Scholar |

Salter RF, Davenport TRB (2011) ‘Orchids and wildflowers of Kitulo Plateau (WILDGuides).’ 1st edn. (Princeton University Press).

Schob C, Callaway RM, Anthelme F, Brooker RW, Cavieres LA, Kikvidze Z, Lortie CJ, Michalet , R, Pugnaire FI, Xiao S, Cranston BH, García M-C, Hupp NR, Llambí LD, Lingua E, Reid AM, Zhao L, Butterfield BJ (2014) The context dependence of beneficiary feedback effects on benefactors in plant facilitation. New Phytologist 204(2), 386-396.
| Crossref | Google Scholar | PubMed |

Schweiger AH, Otieno DO, Kulunge SR, Reineking B, Tenhunen J (2015) The afro-alpine dwarf shrub Helichrysum citrispinum favours understorey plants through microclimate amelioration. Plant Ecology & Diversity 8(3), 293-303.
| Crossref | Google Scholar |

Shefferson RP (2006) Survival costs of adult dormancy and the confounding influence of size in lady’s slipper orchids, genus Cypripedium. Oikos 115(2), 253-262.
| Crossref | Google Scholar |

Shefferson RP, Kull T, Tali K (2006) Demographic response to shading and defoliation in two woodland orchids. Folia Geobotanica 41(1), 95-106.
| Crossref | Google Scholar |

Shefferson RP, Jacquemyn H, Kull T, Hutchings MJ (2020) The demography of terrestrial orchids: life history, population dynamics and conservation. Botanical Journal of the Linnean Society 192(2), 315-332.
| Crossref | Google Scholar |

Siebenkäs A, Schumacher J, Roscher C (2015) Phenotypic plasticity to light and nutrient availability alters functional trait ranking across eight perennial grassland species. AoB PLANTS 7(1), plv029.
| Crossref | Google Scholar |

Singh J, Kumar A, Singh L (2021) Performance of the petiole mobile application on the leaf area estimation as varied with calibration height. The Pharma Innovation Journal 10(4S), 337-341.
| Crossref | Google Scholar |

Slaviero A, Del Vecchio S, Pierce S, Fantinato E, Buffa G (2016) Plant community attributes affect dry grassland orchid establishment. Plant Ecology 217(12), 1533-1543.
| Crossref | Google Scholar |

Sletvold N, Øien D-I, Moen A (2010) Long-term influence of mowing on population dynamics in the rare orchid Dactylorhiza lapponica: the importance of recruitment and seed production. Biological Conservation 143(3), 747-755.
| Crossref | Google Scholar |

Smart SM, Glanville HC, Blanes MdC, Mercado LM, Emmett BA, Jones DL, Cosby BJ, Marrs , RH, Butler A, Marshall MR, Reinsch S, Herrero-Jáuregui C, Hodgson JG (2017) Leaf dry matter content is better at predicting above-ground net primary production than specific leaf area. Functional Ecology 31(6), 1336-1344.
| Crossref | Google Scholar |

Steger C, Boone RB, Dullo BW, Evangelista P, Alemu S, Gebrehiwot K, Klein JA (2022) Collaborative agent-based modeling for managing shrub encroachment in an Afroalpine grassland. Journal of Environmental Management 316, 115040.
| Crossref | Google Scholar |

Swanson EK, Sheley RL, James JJ (2021) Shrubs facilitate perennial bunchgrass recruitment in drylands under experimental precipitation change. Journal of Arid Environments 187, 104432.
| Crossref | Google Scholar |

Swarts ND, Dixon KW (2009) Terrestrial orchid conservation in the age of extinction. Annals of Botany 104(3), 543-556.
| Crossref | Google Scholar | PubMed |

Tripathi S, Bhadouria R, Srivastava P, Devi RS, Chaturvedi R, Raghubanshi AS (2020) Effects of light availability on leaf attributes and seedling growth of four tree species in tropical dry forest. Ecological Processes 9, 2.
| Crossref | Google Scholar |

United Republic of Tanzania (2010) Strengthening the protected area network in southern Tanzania: improving the effectiveness of national parks in addressing threats to biodiversity. United Republic of Tanzania.

Van Auken OW (2009) Causes and consequences of woody plant encroachment into Western North American Grasslands. Journal of Environmental Management 90(10), 2931-2942.
| Crossref | Google Scholar | PubMed |

van der Niet T, Gehrke B (2005) Rare terrestrial orchids on mbeya peak, Southern Tanzania. Journal of East African Natural History 94(2), 279-285.
| Crossref | Google Scholar |

Veldman S (2018) ‘Markets, mixtures and molecular methods.’ (Acta Universitatis Upsaliensis: Uppsala, Sweden)

Venter ZS, Cramer MD, Hawkins H-J (2018) Drivers of woody plant encroachment over Africa. Nature Communications 9(1), 2272.
| Crossref | Google Scholar |

Vogt-Schilb H, Pradel R, Geniez P, Hugot L, Delage A, Richard F, Schatz B (2016) Responses of orchids to habitat change in Corsica over 27 Years. Annals of Botany 118(1), 115-123.
| Crossref | Google Scholar | PubMed |

Weldemariam EC, Jakisa ES, Ahebwe DA (2017) Implication of forest zonation on tree species composition, diversity and structure in Mabira Forest, Uganda. Environment, Earth and Ecology 1(1), 112-122.
| Crossref | Google Scholar |

Whigham DF (1990) The effect of experimental defoliation on the growth and reproduction of a woodland orchid, Tipularia discolor. Canadian Journal of Botany 68(8), 1812-1816.
| Crossref | Google Scholar |

Wilcox BP, Birt A, Fuhlendorf SD, Archer SR (2018) Emerging frameworks for understanding and mitigating woody plant encroachment in grassy biomes. Current Opinion in Environmental Sustainability 32, 46-52.
| Crossref | Google Scholar |

Wilson PJ, Thompson K, Hodgson JG (1999) Specific leaf area and leaf dry matter content as alternative predictors of plant strategies. New Phytologist 143(1), 155-162.
| Crossref | Google Scholar |

Wotavová K, Balounová Z, Kindlmann P (2004) Factors affecting persistence of terrestrial orchids in wet meadows and implications for their conservation in a changing agricultural landscape. Biological Conservation 118(3), 271-279.
| Crossref | Google Scholar |

Wube S, Lemessa D, Dullo WB (2021) Factors driving the expansion of Helichrysum splendidum in Menz-Guassa community conservation area of the Afroalpine ecosystem of Ethiopia. East African Journal of Sciences 15(1), 17.
| Crossref | Google Scholar |

Xu H, Chen P, Tao Y (2021) Understanding the shade tolerance responses through hints from phytochrome A-mediated negative feedback regulation in shade avoiding plants. Frontiers in Plant Science 12, 813092.
| Crossref | Google Scholar |

Yasin M, Rosenqvist E, Andreasen C (2017) The effect of reduced light intensity on grass weeds. Weed Science 65(5), 603-613.
| Crossref | Google Scholar |

Zhang S, Yang Y, Li J, Qin J, Zhang W, Huang W, Hu H (2018) Physiological Diversity of Orchids. Plant Diversity 40(4), 196-208.
| Crossref | Google Scholar | PubMed |

Zimmerman JK (1990) Role of pseudobulbs in growth and flowering of Catasetum viridiflavum (orchidaceae). American Journal of Botany 77(4), 533-542.
| Crossref | Google Scholar | PubMed |

Zimmerman JK, Whigham DF (1992) Eological functions of carbohydrates stored in corms of Tipularia discolor (orchidaceae). Functional Ecology 6(5), 575-581.
| Crossref | Google Scholar |