Urban vegetation types and bryophyte diversity: why woodlands matter

Abstract

Various types of vegetative cover, predominantly woodlands, shrublands, and grasslands, provide a range of habitats for urban organisms and are the main contributors to urban ecosystem services. The goal of this study was to assess the potential of different vegetation types in maintaining and enhancing bryophyte diversity in urban areas. Bryophytes, small non-vascular plants, in woodlands, shrublands, and grasslands in Hefei city (eastern China) were investigated, and their species richness, abundance, community composition and relationships with habitat characteristics were analysed. The results show that urban woodlands had the highest species richness and abundance of bryophytes and a distinct community composition compared with the other vegetation types. Relative light intensity had a positive effect on species richness in urban woodlands, whereas litter cover had a negative effect on abundance. Shrub, grass and litter covers, and relative air humidity were the main factors driving the differences in bryophyte species composition between woodlands and other vegetation types. Therefore, owing to their high potential to maintain bryophyte diversity, woodlands should receive increased attention during the construction of urban green spaces. In addition, promoting structural heterogeneity and increasing light availability would be beneficial to bryophyte species diversity.

Graphic abstract

Keywords

Urban green spaces / Vegetation types / Cryptogams / Biodiversity conservation / Habitat heterogeneity

References

[1] Ah-Peng C, Cardoso AW, Flores O, West A, Wilding N, Strasberg D, Hedderson TAJ. The role of epiphytic bryophytes in interception, storage, and the regulated release of atmospheric moisture in a tropical montane cloud forest. J Hydrol, 2017, 548: 665-673

[2] Alatalo JM, Jägerbrand AK, Erfanian MB, Chen SB, Sun SQ, Molau U. Bryophyte cover and richness decline after 18 years of experimental warming in alpine Sweden. AoB Plants, 2020, 12(6): plaa61

[3] Anhui Provincial Bureau of Statistics, Survey Office of the National Bureau of Statistics in Anhui (2025) Anhui statistical yearbook 2024. China Statistics Press, Beijing (in Chinese)

[4] Barbé M, Bouchard M, Fenton NJ. Examining boreal forest resilience to temperature variability using bryophytes: forest type matters. Ecosphere, 2020, 11(8 e03232

[5] Beninde J, Veith M, Hochkirch A. Biodiversity in cities needs space: a meta-analysis of factors determining intra-urban biodiversity variation. Ecol Lett, 2015, 18(6581-592

[6] Boch S, Müller JC, Prati D, Fischer M. Low-intensity management promotes bryophyte diversity in grasslands. Tuexenia, 2018, 38: 311-328

[7] Campos-Silva LA, Piratelli AJ. Vegetation structure drives taxonomic diversity and functional traits of birds in urban private native forest fragments. Urban Ecosyst, 2021, 24(2): 375-390

[8] Caners RT, MacDonald SE, Belland RJ. Bryophyte assemblage structure after partial harvesting in boreal mixedwood forest depends on residual canopy abundance and composition. For Ecol Manag, 2013, 289: 489-500

[9] Chaudhuri S, Roy M. Global ambient air quality monitoring: Can mosses help? A systematic meta-analysis of literature about passive moss biomonitoring. Environ Dev Sustain, 2024, 26(3): 5735-5773

[10] Cui DF. Plant Taxonomy, 2010, Beijing, China Agriculture Press34(in Chinese)

[11] Cusack DF, Halterman SM, Tanner EVJ, Wright SJ, Hockaday W, Dietterich LH, Turner BL. Decadal-scale litter manipulation alters the biochemical and physical character of tropical forest soil carbon. Soil Biol Biochem, 2018, 124: 199-209

[12] de Souza ERF, Silva JB, Pinto AS, de Faria LS. Soil texture and functional traits of trees structure communities of epiphytic mosses in a tropical dry forest. Flora, 2021, 283 151924

[13] Depauw L, Perring MP, Landuyt D, Maes SL, Blondeel H, De Lombaerde E, Brūmelis G, Brunet J, Closset-Kopp D, Decocq G, Den Ouden J, Härdtle W, Hédl R, Heinken T, Heinrichs S, Jaroszewicz B, Kopecký M, Liepiņa I, Macek M, Máliš F, Schmidt W, Smart SM, Ujházy K, Wulf M, Verheyen K. Evaluating structural and compositional canopy characteristics to predict the light-demand signature of the forest understorey in mixed, semi-natural temperate forests. Appl Veg Sci, 2021, 24(1 e12532

[14] Donnelly A, Yu R, Rehberg C, Schwartz MD. Characterizing spring phenology in a temperate deciduous urban woodland fragment: trees and shrubs. Int J Biometeorol, 2024, 68(5): 871-882

[15] Dormann CF, Elith J, Bacher S, Buchmann C, Carl G, Carré G, García Marquéz JR, Gruber B, Lafourcade B, Leitão PJ, Münkemüller T, McClean C, Osborne PE, Reineking B, Schröder B, Skidmore AK, Zurell D, Lautenbach S. Collinearity: a review of methods to deal with it and a simulation study evaluating their performance. Ecography, 2013, 36(1): 27-46

[16] Dormann CF, Bagnara M, Boch S, Hinderling J, Janeiro-Otero A, Schäfer D, Schall P, Hartig F. Plant species richness increases with light availability, but not variability, in temperate forests understorey. BMC Ecol, 2020, 20(1): 43

[17] Doroski DA, Ashton MS, Duguid MC. The future urban forest—a survey of tree planting programs in the Northeastern United States. Urban For Urban Green, 2020, 55 126816

[18] Enkossa T, Nemomissa S, Lemessa D. Effects of land use types and environmental factors on woody species diversity in agricultural landscapes of Jimma Ganati District, Western Oromia. Ethiopia J Nat Conserv, 2024, 77 126531

[19] Fang JY, Tang ZY, Wang ZH, Shen ZH. The protocol for the survey plan for plant species diversity of China’s mountains. Biodivers Sci, 2004, 12(15-9

[20] Ferenc M, Sedláček O, Fuchs R. How to improve urban greenspace for woodland birds: site and local-scale determinants of bird species richness. Urban Ecosyst, 2014, 17(2): 625-640

[21] Fournier B, Frey D, Moretti M. The origin of urban communities: from the regional species pool to community assemblages in city. J Biogeogr, 2020, 47(3): 615-629

[22] Frey D, Vega K, Zellweger F, Ghazoul J, Hansen D, Moretti M. Predation risk shaped by habitat and landscape complexity in urban environments. J Appl Ecol, 2018, 55(5): 2343-2353

[23] Glime JM (2017) The fauna: A place to call home. Chapt. 1. In: Glime JM (ed) Bryophyte Ecology, vol 2. Bryological Interaction. Ebook sponsored by Michigan Technological University and the International Association of Bryologists. http://digitalcommons.mtu.edu/bryophyte-ecology2/

[24] Gosselin M, Fourcin D, Dumas Y, Gosselin F, Korboulewsky N, Toïgo M, Vallet P. Influence of forest tree species composition on bryophytic diversity in mixed and pure pine (Pinus sylvestris L.) and oak (Quercus petraea (Matt.) Liebl.) stands. For Ecol Manag, 2017, 406: 318-329

[25] Guida G, Nicosia A, Settanni L, Ferro V. A review on effects of biological soil crusts on hydrological processes. Earth Sci Rev, 2023, 243 104516

[26] Han D, Wang C, Sun ZK, She JY, Yin LQ, Bian Q, Han WJ. Microhabitat preferences of butterflies in urban parks: Both vegetation structure and resources are decisive. Urban For Urban Green, 2022, 71 127552

[27] Hansen R, Mattes A, Meier M, Kurths A. Reorienting urban green infrastructure planning towards biodiversity–Perspectives and ongoing debates from Germany. Urban For Urban Green, 2023, 90 128155

[28] Hedwall PO, Holmström E, Lindbladh M, Felton A. Concealed by darkness: how stand density can override the biodiversity benefits of mixed forests. Ecosphere, 2019, 10(8 e02835

[29] Helbach J, Frey J, Messier C, Mörsdorf M, Scherer-Lorenzen M. Light heterogeneity affects understory plant species richness in temperate forests supporting the heterogeneity-diversity hypothesis. Ecol Evol, 2022, 12(2 e8534

[30] Hespanhol H, Cezón K, Muñoz J, Mateo RG, Gonçalves J. How vulnerable are bryophytes to climate change? Developing new species and community vulnerability indices. Ecol Indic, 2022, 136 108643

[31] Hutsemékers V, Mouton L, Westenbohm H, Collart F, Vanderpoorten A. Disentangling climate change from air pollution effects on epiphytic bryophytes. Glob Chang Biol, 2023, 29(14): 3990-4000

[32] Ilić M, Igić R, Ćuk M, Veljić M, Radulović S, Orlović S, Vukov D. Environmental drivers of ground-floor bryophytes diversity in temperate forests. Oecologia, 2023, 202(2): 275-285

[33] Jean M, Melvin AM, Mack MC, Johnstone JF. Broadleaf litter controls feather moss growth in black spruce and birch forests of interior Alaska. Ecosystems, 2020, 23(1): 18-33

[34] Jia Y, He S. Species catalogue of China: plants-bryophytes, 2013, Beijing, Science Press(in Chinese)1

[35] Jourgholami M, Feghhi J, Picchio R, Tavankar F, Venanzi R. Efficiency of leaf litter mulch in the restoration of soil physiochemical properties and enzyme activities in temporary skid roads in mixed high forests. CATENA, 2021, 198 105012

[36] Konijnendijk CC. Evidence-based guidelines for greener, healthier, more resilient neighbourhoods: introducing the 3–30-300 rule. J For Res, 2023, 34(3): 821-830

[37] Kovács B, Tinya F, Ódor P. Stand structural drivers of microclimate in mature temperate mixed forests. Agric For Meteor, 2017, 234: 11-21

[38] Kutnar L, Kermavnar J, Sabovljević MS. Bryophyte diversity, composition and functional traits in relation to bedrock and tree species composition in close-to-nature managed forests. Eur J For Res, 2023, 142(4): 865-882

[39] Lintott PR, Bunnefeld N, Fuentes-Montemayor E, Minderman J, Blackmore LM, Goulson D, Park KJ. Moth species richness, abundance and diversity in fragmented urban woodlands: implications for conservation and management strategies. Biodivers Conserv, 2014, 23(11): 2875-2901

[40] Liu GY, Chen LL, Shi XR, Yuan ZY, Yuan LY, Lock TR, Kallenbach RL. Changes in rhizosphere bacterial and fungal community composition with vegetation restoration in planted forests. Land Degrad Dev, 2019, 30(10): 1147-1157

[41] Majasalmi T, Rautiainen M. The impact of tree canopy structure on understory variation in a boreal forest. For Ecol Manag, 2020, 466 118100

[42] Márialigeti S, Németh B, Tinya F, Ódor P. The effects of stand structure on ground-floor bryophyte assemblages in temperate mixed forests. Biodivers Conserv, 2009, 18(8): 2223-2241

[43] Müller J, Boch S, Prati D, Socher SA, Pommer U, Hessenmöller D, Schall P, Schulze ED, Fischer M. Effects of forest management on bryophyte species richness in Central European forests. For Ecol Manag, 2019, 432: 850-859

[44] Natalia S, Lieffers VJ, Landhäusser SM. Effects of leaf litter on the growth of boreal feather mosses: implication for forest floor development. J Veg Sci, 2008, 19(2): 253-260

[45] Neal W, Araya Y, Wheeler PM. Influence of canopy structural complexity on urban woodland butterfly species richness. J Insect Conserv, 2024, 28(5): 1051-1062

[46] Nghiem TPL, Wong KL, Jeevanandam L, Chang CC, Tan LYC, Goh Y, Carrasco LR. Biodiverse urban forests, happy people: experimental evidence linking perceived biodiversity, restoration, and emotional wellbeing. Urban For Urban Green, 2021, 59 127030

[47] Nooten SS, Schultheiss P, Rowe RC, Facey SL, Cook JM. Habitat complexity affects functional traits and diversity of ant assemblages in urban green spaces (Hymenoptera: Formicidae). Myrmecol News, 2019, 29: 67-77

[48] Oishi Y. Influence of urban green spaces on the conservation of bryophyte diversity: the special role of Japanese gardens. Landsc Urban Plan, 2012, 106(1): 6-11

[49] Oishi Y. Urban heat island effects on moss gardens in Kyoto. Japan Landsc Ecol Eng, 2019, 15(2): 177-184

[50] Onoda Y, Saluñga JB, Akutsu K, Aiba SI, Yahara T, Anten NPR. Trade-off between light interception efficiency and light use efficiency: implications for species coexistence in one-sided light competition. J Ecol, 2014, 102(1): 167-175

[51] Petersson L, Nilsson S, Holmström E, Lindbladh M, Felton A. Forest floor bryophyte and lichen diversity in Scots pine and Norway spruce production forests. For Ecol Manag, 2021, 493 119210

[52] Pharo EJ, Lindenmayer DB, Taws N. The effects of large-scale fragmentation on bryophytes in temperate forests. J Appl Ecol, 2004, 41(5): 910-921

[53] Qiu DX, Xiao B, Algora C, Delgado-Baquerizo M. Litter covering decreases moss activity, mediating the relationships between moss biocrusts and shrub patches in semiarid dryland ecosystems. Plant Soil, 2024

[54] R Development Core Team (2024) R: a language and environment for statistical computing, version 4.4.1. Foundation for Statistical Computing, Vienna, Austria

[55] Rajaniemi TK, Goldberg DE, Turkington R, Dyer AR. Local filters limit species diversity, but species pools determine composition. Perspect Plant Ecol Evol Syst, 2012, 14(6): 373-380

[56] Riffo-Donoso V, Osorio F, Fontúrbel FE. Habitat disturbance alters species richness, composition, and turnover of the bryophyte community in a temperate rainforest. For Ecol Manag, 2021, 496 119467

[57] Rola K, Plášek V, Rożek K, Zubek S. Effect of tree species identity and related habitat parameters on understorey bryophytes–interrelationships between bryophyte, soil and tree factors in a 50-year-old experimental forest. Plant Soil, 2021, 466(1): 613-630

[58] Salminen H, Tukiainen H, Alahuhta J, Hjort J, Huusko K, Grytnes JA, Pacheco-Riaño LC, Kapfer J, Virtanen R, Maliniemi T. Assessing the relation between geodiversity and species richness in mountain heaths and tundra landscapes. Landsc Ecol, 2023, 38(9): 2227-2240

[59] Schüle M, Heinken T. Allometric estimation models for aboveground and belowground biomass of pre-fire and post-fire vegetation in Scots pine forests. Forestry, 2025, 98(1): 84-97

[60] Sebek P, Vodka S, Bogusch P, Pech P, Tropek R, Weiss M, Zimova K, Cizek L. Open-grown trees as key habitats for arthropods in temperate woodlands: the diversity, composition, and conservation value of associated communities. For Ecol Manag, 2016, 380: 172-181

[61] Simkin RD, Seto KC, McDonald RI, Jetz W. Biodiversity impacts and conservation implications of urban land expansion projected to 2050. Proc Natl Acad Sci USA, 2022, 119(12 e2117297119

[62] Slate ML, Antoninka A, Bailey L, Berdugo MB, Callaghan DA, Cárdenas M, Chmielewski MW, Fenton NJ, Holland-Moritz H, Hopkins S, Jean M, Kraichak BE, Lindo Z, Merced A, Oke T, Stanton D, Stuart J, Tucker D, Coe KK. Impact of changing climate on bryophyte contributions to terrestrial water, carbon, and nitrogen cycles. New Phytol, 2024, 242(6): 2411-2429

[63] Standing Committee of the Hefei Municipal People’s Congress (2018) Regulations on urban greening management of Hefei City. http://www.ahrd.gov.cn/article.jsp?strId=1529651373681336&strColId=f2a446e22ed74b6da8f79cea33b4d4b5/

[64] Su P, Wang XX, Lin QD, Peng JL, Song JX, Fu JX, Wang SQ, Cheng DD, Bai HF, Li Q. Variability in macroinvertebrate community structure and its response to ecological factors of the Weihe River Basin. China Ecol Eng, 2019, 140 105595

[65] Threlfall CG, Mata L, Mackie JA, Hahs AK, Stork NE, Williams NSG, Livesley SJ. Increasing biodiversity in urban green spaces through simple vegetation interventions. J Appl Ecol, 2017, 54(6): 1874-1883

[66] Tinya F, Ódor P. Congruence of the spatial pattern of light and understory vegetation in an old-growth, temperate mixed forest. For Ecol Manag, 2016, 381: 84-92

[67] Tinya F, Kovács B, Bidló A, Dima B, Király I, Kutszegi G, Lakatos F, Mag Z, Márialigeti S, Nascimbene J, Samu F, Siller I, Szél G, Ódor P. Environmental drivers of forest biodiversity in temperate mixed forests—a multi-taxon approach. Sci Total Environ, 2021, 795 148720

[68] Trujillo-González JM, Zapata-Muñoz YL, Torres-Mora MA, García-Navarro FJ, Jiménez-Ballesta R. Assessment of urban environmental quality through the measurement of lead in bryophytes: case study in a medium-sized city. Environ Geochem Health, 2020, 42(10): 3131-3139

[69] Vega KA, Küffer C. Promoting wildflower biodiversity in dense and green cities: the important role of small vegetation patches. Urban For Urban Green, 2021, 62 127165

[70] Wang XL, Rahman MA, Mokroš M, Rötzer T, Pattnaik N, Pang YL, Zhang YR, Da LJ, Song K. The influence of vertical canopy structure on the cooling and humidifying urban microclimate during hot summer days. Landsc Urban Plan, 2023, 238 104841

[71] Williams SE, Marsh H, Winter J. Spatial scale, species diversity, and habitat structure: small mammals in Australian tropical rain forest. Ecology, 2002, 83(5): 1317-1329

[72] Wu JY, Yang MX, Xiong LY, Wang CF, Ta N. Health-oriented vegetation community design: innovation in urban green space to support respiratory health. Landsc Urban Plan, 2021, 205 103973

[73] Wu XZ, Lei FM, Shan YY, Yu J. Distribution pattern of bryophyte diversity and environmental impact factors in urban parks of Shanghai. Biodivers Sci, 2024, 32(2): 62-72(in Chinese)

[74] Yin S, Chen WY, Liu CJ. Urban forests as a strategy f1or transforming towards healthy cities. Urban For Urban Green, 2023, 81 127871

[75] Zar JH. Biostatistical analysis, 1999, New Jersey, Prentice-Hall Inc

[76] Zhang Y, Meng WQ, Yun HF, Xu WB, Hu BB, He MX, Mo XQ, Zhang L. Is urban green space a carbon sink or source? A case study of China based on LCA method. Environ Impact Assess Rev, 2022, 94 106766

[77] Zhang YL, He NP, Liu Y. Temperature factors are a primary driver of the forest bryophyte diversity and distribution in the southeast Qinghai-Tibet Plateau. For Ecol Manag, 2023, 527 120610

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