Macrofauna and mesofauna in Hyrcanian Protected Forest: Seasonal patterns of the arthropod diversity and community structure across soil depths

Mehrafrooz Mayvan, Mahmood; Shayanmehr, Masoumeh

doi:10.48311/jibs.12.04.705

Macrofauna and mesofauna in Hyrcanian Protected Forest: Seasonal patterns of the arthropod diversity and community structure across soil depths

Articles in Press

Document Type : Full-length Research Article

Authors

Mahmood Mehrafrooz Mayvan ¹^{, 2}

Masoumeh Shayanmehr ¹

¹ Department of Plant Protection, Faculty of Crop Sciences, Sari University of Agricultural Sciences and Natural Resources (SANRU), Sari, Iran

² Institute of Soil Biology and Biogeochemistry, Biology Centre of the Czech Academy of Sciences, Na Sádkách 7, 370 05 České Budějovice, Czech Republic

10.48311/jibs.12.04.705

Abstract

Hyrcanian forests are considered among the oldest forests in the world, and due to their temperate and humid climate and fertile soil, they have rich fauna and flora. Soil-dwelling arthropods play essential roles in forest ecosystem functioning, contributing to litter decomposition, nutrient cycling, and soil structure maintenance. This study assessed the diversity and vertical distribution of macrofauna and mesofauna in Semeskandeh Forest, a protected Hyrcanian Forest in northern Iran, across different soil depths (leaf litter, 0–3 cm, 3–6 cm) and seasons over one year. Over 10,000 individuals were collected, representing diverse taxonomic groups, including ants, millipedes, springtails, mites, pseudoscorpions, and diplurans. Population densities and species diversity were consistently highest in the litter layer and declined with increasing soil depth, with clear seasonal peaks observed during the study period. Ants dominated the macrofauna, while mites and springtails were the most diverse mesofaunal taxa. The study revealed clear seasonal and vertical patterns in the density and diversity of soil faunal communities in the Hyrcanian forest ecosystem. These findings provide baseline information for biodiversity conservation and future monitoring of soil fauna in Hyrcanian forests. Thereby, conserving litter quality, maintaining habitat integrity, and monitoring soil faunal diversity are critical for sustaining forest ecosystem functions in such a globally valuable and inherited region.

Graphical Abstract

Macrofauna and mesofauna in Hyrcanian Protected Forest: Seasonal patterns of the arthropod diversity and community structure across soil depths

Keywords

Abundance

Community composition

Conservation

Faunal diversity

Iran

Leaf litter

Subjects

Biodiversity

Adiningrat, D.P., Siegenthaler, A., Schlund, M., Wang, T., Skidmore, A.K., Rousseau, M. & Heurich, M. (2025) Effect of forest structural attributes on soil microbial diversity in mixed temperate forests. Plant Soil, 517, 867–885. https://doi.org/10.1007/s11104-025-07907-4

Akhani, H., Djamali, M., Ghorbanalizadeh, A. & Ramezani, E. (2010) Plant biodiversity of Hyrcanian relict forests, N Iran: An overview of the flora, vegetation, palaeoecology and conservation. Pakistan Journal of Botany, 42, 231–258.

Anthony, M.A., Bender, S.F. & van der Heijden, M.G.A. (2023) Enumerating soil biodiversity. Proceedings of the National Academy of Sciences of the United States of America, 120 (33), e2304663120. https://doi.org/10.1073/pnas.2304663120

Angst, G., Potapov, A., Joly, F.X., Angst, Š., Frouz, J., Ganault, P. & Eisenhauer, N. (2024) Conceptualizing soil fauna effects on labile and stabilized soil organic matter. Nature Communications, 15, 5005. https://doi.org/10.1038/s41467-024-49240-x

Ardakani, M.R. (2007) Ecology. University of Tehran Press, Tehran. 340 pp.

Berg, M.P. (2012) Patterns of biodiversity at fine and small spatial scales. In: Wall, D.H., Bardgett, R.D., Behan-Pelletier, V., Herrick, J.E., Jones, T.H., Ritz, K., Six, J., Strong, D.R. & van der Putten, W.H. (eds) Soil Ecology and Ecosystem Services. Oxford University Press, UK, pp. 136–152. https://doi.org/10.1093/acprof:oso/9780199575923.003.0014

Berry, G., Tanruean, K., Napiroon, T., Saetang, W., Penpo, M., Duangrod, S. & Poolprasert, P. (2020) The community structure of beneficial and harmful arthropod fauna in rice field ecosystems in lower Northern Thailand. The Journal of Applied Science, 19 (1), 1–12. https://doi.org/10.14416/j.appsci.2020.01.001

Bretfeld, G. (1999) Synopses on Palaearctic Collembola. V. 2. Symphypleona. Abhandlungen und Berichte des Naturkundemuseums, Gӧrlitz, 71 (1), 1‒318.

Burton, V.J. & Eggleton, P. (2016) Microhabitat heterogeneity enhances soil macrofauna and plant species diversity in an ash - field maple woodland. European Journal of Soil Biology, 75, 97–106. https://doi.org/10.1016/j.ejsobi.2016.04.012.

Cayuela, M.L., Clause, J., Frouz, J. & Baveye, P.C. (2020) Interactive Feedbacks between soil fauna and soil processes. Frontiers in Environmental Science, 8, 14, https://doi.org/10.3389/fenvs.2020.00014

Chahartaghi, M., Langel, R., Scheu, S. & Ruess, L. (2005) Feeding guilds in Collembola based on nitrogen stable isotope ratios. Soil Biology and Biochemistry, 37 (9), 1718–1725. https://doi.org/10.1016/j.soilbio.2005.02.006

Chahartaghi, M., Scheu, S. & Ruess, L. (2006) Sex ratio and mode of reproduction in Collembola of an oak-beech forest. Pedobiologia, 50 (4), 331–340. https://doi.org/10.1016/j.pedobi.2006.06.001

Chahartaghi, M., Maraun, M., Scheu, S. & Domes, K. (2009) Resource depletion and colonization: A comparison between parthenogenetic and sexual Collembola species. Pedobiologia, 52 (3), 181–189. https://doi.org/10.1016/j.pedobi.2008.08.003

Coleman, D.C., Geisen, S. & Wall, D.H. (2023) Soil fauna: occurrence, biodiversity, and roles in ecosystem function. In: Paul, E.A. & Frey, S.D. (eds) Soil Microbiology, Ecology and Biochemistry, 5th ed. Elsevier, Amsterdam, Netherlands, pp. 131–159. https://doi.org/10.1016/B978-0-12-822941-5.00005-3

de Pedro, L., Perera-Fernández, L.G., López-Gallego, E., Pérez-Marcos, M. & Sanchez, J.A. (2020) The effect of cover crops on the biodiversity and abundance of ground-dwelling arthropods in a Mediterranean pear orchard. Agronomy, 10 (4), 580. https://doi.org/10.3390/agronomy10040580

Decaëns, T., Jiménez, J.J., Gioia, C., Measey, G.J. & Lavelle, P. (2006) The values of soil animals for conservation biology. European Journal of Soil Biology, 42 (1), 23–38. https://doi.org/10.1016/j.ejsobi.2006.07.001

D'Haese, C., Shayanmehr, M., Yoosefi Lafooraki, E. & Tully, T. (2025) An initial assessment of the Collembola (Hexapoda) fauna of the Bula Hyrcanian Forest (Iran) at the interface of two biodiversity hotspots. Soil Organisms, 97, 211–245. https://doi.org/10.25674/451

Domene, X. (2016) A critical analysis of meso- and macrofauna effects following biochar supplementation. In: Ralebitso-Senior, K.R. & Orr, C.H. (eds) Biochar Application: Essential Soil Microbial Ecology. Elsevier Inc., Amsterdam, pp. 268–292. https://doi.org/10.1016/b978-0-12-803433-0.00011-4

Domínguez, M.V.S., González, E., Fabián, D., Salvo, A. & Fenoglio, M.S. (2020) Arthropod diversity and ecological processes on green roofs in a semi-rural area of Argentina: Similarity to neighbor ground habitats and landscape effects. Landscape and Urban Planning, 199, 103816. https://doi.org/10.1016/j.landurbplan.2020.103816

Eckert, M., Gaigher, R., Pryke, J.S., Janion-Scheepers, C. & Samways, M.J. (2023) High trait diversity among soil arthropods allows their survival in a heterogeneous eco-agricultural mosaic. Soil Biology and Biochemistry, 186, 109180. https://doi.org/10.1016/j.soilbio.2023.109180

Fjellberg, A. (2007) The Collembola of Fennoscandia and Denmark. Part II: Entomobryomorpha and Symphypleona. Fauna Entomologica Scandinavica, 42, 1–264. https://doi.org/10.1163/ej.9789004157705.i-265

Frouz, J., Ali, A., Frouzova, J. & Lobinske, R.J. (2004) Horizontal and vertical distribution of soil macroarthropods along a spatio-temporal moisture gradient in subtropical central Florida. Environmental Entomology, 33 (5), 1282–1295. https://doi.org/10.1603/0046-225X-33.5.1282

Gagnarli, E., Goggioli, D., Tarchi, F., Guidi, S., Nannelli, R., Vignozzi, N., Valboa, G., Lottero, M.R., Corino, L. & Simoni, S. (2015) Case study of microarthropod communities to assess soil quality in different managed vineyards. Soil, 1, 527–536. https://doi.org/10.5194/soil-1-527-2015

García-Segura, D., Castillo-Murrieta, I.M., Martínez-Rabelo, F., Gomez-Anaya, A., Rodríguez-Campos, J., Hernández-Castellanos, B., Contreras-Ramos, S.M. & Barois, I. (2018) Macrofauna and mesofauna from soil contaminated by oil extraction. Geoderma, 332, 180–189. http://dx.doi.org/10.1016/j.geoderma.2017.06.013

Ghiglieno, I., Simonetto, A., Orlando, F., Donna, P., Tonni, M., Valenti, L. & Gilioli, G. (2020) Response of the arthropod community to soil characteristics and management in the Franciacorta Viticultural area (Lombardy, Italy). Agronomy, 10 (5), 740. https://doi.org/10.3390/agronomy10050740

Gholizadeh, H., Naqinezhad, A. & Chytrý, M. (2019) Hyrcanian Forest Vegetation Database. Phytocoenologia, 49 (2), 209–210. https://doi.org/10.1127/phyto/2018/0315

Ghomi-Avili, A., Akbarinia, M., Hosseini, S.M., Talebian, M.H. & Knapp, H.D. (2020) Fuzzy and Boolean operation based modelling for evaluation of ecological capability in the Hyrcanian Forests. Journal of Forest Science, 66 (4), 170–184. https://doi.org/10.17221/130/2019-JFS

Ghorbanalizadeh, A. & Akhani, H. (2022) Plant diversity of Hyrcanian relict forests: An annotated checklist, chorology and threat categories of endemic and near endemic vascular plant species. Plant Diversity, 44 (1), 39–69. https://doi.org/10.1016/j.pld.2021.07.005

Gomyo, M. & Kuraji, K. (2016) Effect of the litter layer on runoff and evapotranspiration using the paired watershed method. Journal of Forestry Research, 21 (6), 306–313. https://doi.org/10.1007/s10310-016-0542-5

Gonçalves, F., Nunes, C., Carlos, C., López, A., Oliveira, I., Crespí, A., Teixeira, B., Pinto, R., Costa, C.A. & Torres, L. (2020) Do soil management practices affect the activity density, diversity, and stability of soil arthropods in vineyards? Agriculture, Ecosystems and Environment, 294, 106863. https://doi.org/10.1016/j.agee.2020.106863

Guo, X., Bian, Z., Wang, S., Wang, Q., Zhang, Y., Zhou, J. & Lin, L. (2020) Prediction of the spatial distribution of soil arthropods using a random forest model: A case study in Changtu County, Northeast China. Agriculture, Ecosystems & Environment, 292, 106818. https://doi.org/10.1016/j.agee.2020.106818

Haghdoost, N., Akbarinia, M., Hosseini, S.M. & Kooch, Y. (2011) Conversion of Hyrcanian degraded forests to plantations: Effects on soil C and N stocks. Annals of Biological Research, 2 (5), 385–399. https://www.researchgate.net/publication/275333499

Hasin, S. & Booncher, K. (2020) Change in ground-dwelling arthropod communities in different agroecosystems in Wang Nam Khiao, Nakhon Ratchasima province, Thailand. Agriculture and Natural Resources, 54 (2), 139–149. https://doi.org/10.34044/j.anres.2020.54.2.04

Hébert, C. (2023) Forest arthropod diversity. In: Allison, J.D., Paine, T.D., Slippers, B. & Wingfield, M.J. (eds) Forest Entomology and Pathology. Springer, Cham, pp. 45–90. https://doi.org/10.1007/978-3-031-11553-0_3

Hector, A. & Bagchi, R. (2007) Biodiversity and ecosystem multifunctionality. Nature, 448 (7150), 188–190. https://doi.org/10.1038/nature05947

Heydari, M., Eslaminejad, P., Kakhki, F.V., Mirab-balou, M., Omidipour, R., Prévosto, B., Kooch, Y. & Lucas-Borja, M.E. (2020) Soil quality and mesofauna diversity relationship are modulated by woody species and seasonality in semiarid oak forest. Forest Ecology and Management, 473, 118332. https://doi.org/10.1016/j.foreco.2020.118332

Homami Totmaj, L., Ramezani, E., Alizadeh, K. & Behling, H. (2021) Four millennia of vegetation and environmental history above the Hyrcanian forest, northern Iran. Vegetation History and Archaeobotany, 30, 611–621. https://doi.org/10.1007/s00334-020-00813-y

Hosseini, S.M. (2006) Death of Elm trees in the Hyrcanian forests of Iran. In: IUFRO Working Party 7.03.10. Proceedings of the Workshop 2006. Gmunden, Austria Vienna, pp. 183–186.

Jordana, R. (2012) Capbryinae and Entomobryini. In: Dunger, W. & Burkhardt, U. (eds) Synopses on Palearctic Collembola. Soil Organisms, 84 (1), 1–391.

Jourgholami, M., Labelle, E.R. & Feghhi, J. (2019) Efficacy of leaf litter mulch to mitigate runoff and sediment yield following mechanized operations in the Hyrcanian mixed forests. Journal of Soils and Sediments, 19, 2076–2088. https://doi.org/10.1007/s11368-018-2194-x

Kelleher, E.M. & Choi, Y.D. (2020) Role of plant diversity on arthropod communities in a restored tallgrass prairie of the U.S. Midwest. Restoration Ecology, 28 (6), 1464–1474. https://doi.org/10.1111/rec.13238

Kooch, Y. & Noghre, N. (2020) The effect of shrubland and grassland vegetation types on soil fauna and flora activities in a mountainous semi-arid landscape of Iran. Science of the Total Environment, 703, 135497. https://doi.org/10.1016/j.scitotenv.2019.135497

Kooch, Y., Moghimian, N. & Kolb, S. (2019) Microbial hotspot areas of C and N cycles in old-growth Hyrcanian forests top soils. Forest Ecology and Management, 446, 93–104. https://doi.org/10.1016/j.foreco.2019.05.022

Kooch, Y., Ehsani, S. & Akbarinia, M. (2020) Stratification of soil organic matter and biota dynamics in natural and anthropogenic ecosystems. Soil & Tillage Research, 200, 104621. https://doi.org/10.1016/j.still.2020.104621

Kounda-Kiki, C., Vaçulik, A., Ponge, J.F. & Sarthou, C. (2004) Soil arthropods in a developmental succession on the Nouragues inselberg (French Guiana). Biology and Fertility of Soils, 40, 119–127. https://doi.org/10.1007/s00374-004-0749-z

Lavelle, P., Bignell, D., Lepage, M., Wolters, V., Roger, P., Ineson, P., Heal, O.W. & Dhillion, S. (1997) Soil function in a changing world: the role of invertebrate ecosystem engineers. European Journal of Soil Biology, 33 (4), 159–193.

Li, X., Xiao, Q., Niu, J., Dymond, S., McPherson, E.G., van Doorn, N., Yu, X., Xie, B., Zhang, K. & Li, J. (2017) Rainfall interception by tree crown and leaf litter: an interactive process. Hydrological Processes, 31, 3533–3542. https://doi.org/10.1002/hyp.11275

Liang, K. (2024) The role of leaf litter in forest soil fertility and microbial diversity. Molecular Soil Biology, 15 (3), 99–108. https://doi.org/10.5376/msb.2024.15.0011

Lin, D., Wang, F., Fanin, N., Pang, M., Dou, P., Wang, H., Qian, S., Zhao, L., Yang, Y., Mi, X. & Ma, K. (2019) Soil fauna promote litter decomposition but do not alter the relationship between leaf economics spectrum and litter decomposability. Soil Biology and Biochemistry, 136, 107519. https://doi.org/10.1016/j.soilbio.2019.107519

Ma, C., Yin, X. & Wang, H. (2019) Soil fauna effect on Dryas octopetala litter decomposition in an Alpine tundra of the Changbai Mountains, China. Alpine Botany, 129, 53–62. https://doi.org/10.1007/s00035-018-0215-4

Marsden, C., Martin-Chave, A., Cortet, J., Hedde, M. & Capowiez, Y. (2020) How agroforestry systems influence soil fauna and their functions - a review. Plant Soil, 453, 29–44. https://doi.org/10.1007/s11104-019-04322-4

Mehrafrooz Mayvan, M. & Shayanmehr, M. (2016) An investigation of soil Chilopoda from Semeskandeh forest (Sari, Mazandaran, Iran). Taxonomy and Biosystematics, 7 (25), 1–12. https://dor.isc.ac/dor/20.1001.1.20088906.1394.7.25.2.2

Mehrafrooz Mayvan, M., Shayanmehr, M. & Scheu, S. (2015) Depth distribution and inter-annual fluctuations in density and diversity of Collembola in an Iranian Hyrcanian forest. Soil Organisms, 87 (3), 239–247. https://soil-organisms.org/SO/article/view/293

Menta, C. & Remelli, S. (2020) Soil health and arthropods: from complex system to worthwhile investigation. Insects, 11 (1), 54. https://doi.org/10.3390/insects11010054

Mohammadi, J., Shataee, S., Namiranian, M. & Næsset, E. (2017) Modeling biophysical properties of broad-leaved stands in the hyrcanian forests of Iran using fused airborne laser scanner data and ultraCam-D images. International Journal of Applied Earth Observation and Geoinformation, 61, 32–45. http://dx.doi.org/10.1016/j.jag.2017.05.003

Moradi, H., Naqinezhad, A., Siadati, S., Yousefi, Y., Attar, F., Etemad, V. & Reif, A. (2016) Elevational gradient and vegetation environmental relationships in the central Hyrcanian forests of northern Iran. Nordic Journal of Botany, 34, 1–14. https://doi.org/10.1111/njb.00535

Moradi, J., John, J., Vicentini, F., Vesela, H., Vicena, J., Ardestani, M.M. & Frouz, J. (2020) Vertical distribution of soil fauna and microbial community under two contrasting post mining chronosequences: Sites reclaimed by alder plantation and unreclaimed regrowth. Global Ecology and Conservation, 23, e01165. https://doi.org/10.1016/j.gecco.2020.e01165

Nacif, M.E., Kitzberger, T. & Garibaldi, L.A. (2020) Positive outcomes between herbivore diversity and tree survival: Responses to management intensity in a Patagonian forest. Forest Ecology and Management, 458, 117738. https://doi.org/10.1016/j.foreco.2019.117738

Nakhutsrishvili, G., Zazanashvili, N., Batsatsashvili, K. & Montalvo Mancheno, C.S. (2015) Colchic and Hyrcanian forests of the Caucasus: similarities, differences and conservation status. Flora Mediterranea, 25 (Special Issue), 185–192. https://doi.org/10.7320/FlMedit25SI.185

O’Donnell, P.P. & Wright, W. (2021) Assessing the conservation and enhancement value of revegetated strips on arthropod assemblages in a pasture landscape. Journal of Environmental Management, 278, 111522. https://doi.org/10.1016/j.jenvman.2020.111522

Palacios-Vargas, J., Castaño-Meneses, G., Gómez-Anaya, J., Martínez-Yrizar, A., Mejía-Recamier, B.E., & Martínez-Sánchez, J. (2007) Litter and soil arthropods diversity and density in a tropical dry forest ecosystem in Western Mexico. Biodiversity and Conservation, 16, 3703–3717. https://doi.org/10.1007/s10531-006-9109-7

Pequeno, P.A.C.L., Franklin, E. & Norton, R.A. (2020) Determinants of intra-annual population dynamics in a tropical soil arthropod. Biotropica, 52, 129–138. https://doi.org/10.1111/btp.12731

Pereira, J. de M., Segat, J.C., Baretta, D., Vasconcellos, R.L. de F., Baretta, C.R.D.M. & Cardoso, E.J.B.N. (2017) Soil macrofauna as a soil quality indicator in native and replanted Araucaria angustifolia forests. Revista Brasileira de Ciênciado Solo, 41, 1–15. https://doi.org/10.1590/18069657rbcs20160261

Pestana, L.F.A., de Souza, A.L.T., Tanaka, M.O., Labarque, F.M. & Soares, J.A.H. (2020) Interactive effects between vegetation structure and soil fertility on tropical ground-dwelling arthropod assemblages. Applied Soil Ecology, 155, 103624. https://doi.org/10.1016/j.apsoil.2020.103624

Pielou, E.C. (1969) An Introduction to Mathematical Ecology. Wiley and Sons, New, 300 pp.

Potapov, M. (2001) Isotomidae. In: Dunger, W. (ed.) Synopses on Palaeartic Collembola. Vol. 3. Abhandlungen und Berichte des Naturkundemuseums Görlitz, 73 (2), pp. 1–603.

Sánchez-Montoya, M.M., Tockner, K., von Schiller, D., Miñano, J., Catarineu, C., Lencina, J.L., Barberá, G.G. & Ruhi, A. (2020) Dynamics of ground-dwelling arthropod metacommunities in intermittent streams: The key role of dry riverbeds. Biological Conservation, 241, 108328. https://doi.org/10.1016/j.biocon.2019.108328

Shayanmehr, M., Mehrafrooz Mayvan, M., Yoosefi Lafooraki, E., Kahrarian, M., Yahyapour, E., Daghighi, E., Jordana, R., Arbea, J., Potapov, M., D’Haese, C., Tully, T., Skarżyński, D., Smolis, A., Galli, L., Sendra, A., Gioia Cipola, N., Molero, R. & Kováč, L. (2025) Arthropod diversity in Iran: Subphylum Hexapoda Latreille, 1825, Basal Hexapod Taxa. Journal of Insect Biodiversity and Systematics, 11 (4), 1085–1114. https://doi.org/10.48311/jibs.11.4.1085

Sibly, R.M. & Hone, J. (2002) Population growth rate and its determinants: An overview. Philosophical Transactions of the Royal Society B: Biological Sciences, 357 (1425), 1153–1170. https://doi.org/10.1098/rstb.2002.1117

Stritih, A., Lecina-Diaz, J., Mohr, J., Schattenberg, C., Ammer, C., Anselmetto, N., Bauhus, J., Bončina, A., Garbarino, M., Hlásny, T., Lindner, M., Lingua, E., Knific, E., Krüger, K., Marangon, D., Morresi, D., Mulec, K., Nagel, T.A., Potterf, M., Stritih, J., Thom, D., Welling, D. & Seidl, R. (2026) Can “Closer-to-Nature” forest management sustain biodiversity and ecosystem services in an uncertain future? lessons from Central Europe. Current Forestry Reports, 12, 1. https://doi.org/10.1007/s40725-025-00264-6

Takeda, H. & Abe, T. (2001) Templates of food-habitat resources for the organization of soil animals in temperate and tropical forests. Ecological Research, 16 (5), 961–973. https://doi.org/10.1046/j.1440-1703.2001.00450.x

Thibaud, J.M., Schulz, H.J. & da Gama Assalino, M.M. (2004) Hypogastruridae. In: Dunger, W. (eds) Synopses on Palaearctic Collembola. Vol. 4. Abhandlungen und Berichte des Naturkundemuseums Görlitz, 75 (2), pp. 1–287.

Tilman, D., Fargione, J., Wolff, B., D’Antonio, C., Dobson, A., Howarth, R., Schindler, D., Schlesinger, W.H., Simberloff, D. & Swackhamer, D. (2001) Forecasting Agriculturally Driven Global Environmental Change. Science, 292 (5515), 281–284. https://doi.org/10.1126/science.1057544

Tresch, S., Frey, D., Le Bayon, R.C., Zanetta, A., Rasche, F., Fliessbach, A. & Moretti, M. (2019) Litter decomposition driven by soil fauna, plant diversity and soil management in urban gardens. Science of the Total Environment, 658, 1614–1629. https://doi.org/10.1016/j.scitotenv.2018.12.235

Truffer, O., Lieberherr, E., Van Ruymbeke, K., Chimisso, C., Rowe, T., Vranken, L. & Ohmura, T. (2025) Mapping the Research of Forest Ecosystem Services in Europe: A Review. Forest Policy and Economics, 181, 103663. https://doi.org/10.1016/j.forpol.2025.103663

Tscharntke, T., Klein, A.M., Kruess, A., Steffan-Dewenter, I. & Thies, C. (2005) Landscape perspectives on agricultural intensification and biodiversity-Ecosystem service management. Ecology Letters, 8, 857–874. https://doi.org/10.1111/j.1461-0248.2005.00782.x

Tully, T., Shayanmehr, M., Yoosefi Lafooraki, E., Ghajar Sepanlou, M. & D’Haese, C. (2025) Diversity and community structure of Collembola (Hexapoda) across sites and microhabitats in the Bula protected area, a UNESCO-listed Hyrcanian forest at the crossroads of two biodiversity hotspots (Iran). bioRxiv 2025.05.20.650270. https://doi.org/10.1101/2025.05.20.650270

Utami, S., Kurniawan, A. & Muslimin, I. (2019) Arthropod diversity on forest stands in South Sumatra, Indonesia. IOP Conf. Series: Earth and Environmental Science, 394, 012003. https://doi.org/10.1088/1755-1315/394/1/012003

Valencia-Cuevas, L., Rodríguez-Domínguez, A., Mussali-Galante, P., Ramos-Quintana, F. & Tovar-Sánchez, E. (2020) Influence of edaphic factors along an altitudinal gradient on a litter arthropod community in an Abies-Quercus forest in Mexico. Acta Oecologica, 108, 103609. https://doi.org/10.1016/j.actao.2020.103609

van Straalen, N.M. (1998) Evaluation of bioindicator systems derived from soil arthropod communities. Applied Soil Ecology, 9 (1–3), 429–437. https://doi.org/10.1016/S0929-1393(98)00101-2

Venuste, N. (2020) Impacts of forest and agricultural land use on soil and litter arthropod assemblages in Southern province of Rwanda. Rwanda Journal of Engineering, Science, Technology and Environment, 3 (1), 1–16. https://dx.doi.org/10.4314/rjeste.v3i1.7

Waez-Mousavi, S.M. (2017) Humus systems in the Caspian Hyrcanian temperate forests. Applied Soil Ecology, 123, 664–667. http://dx.doi.org/10.1016/j.apsoil.2017.09.022

Wang, B., Verheyen, K., Baeten, L. & De Smedt, P. (2021) Herb litter mediates tree litter decomposition and soil fauna composition. Soil Biology and Biochemistry, 152, 108063. https://doi.org/10.1016/j.soilbio.2020.108063

Wiwatwitaya, D. & Takeda, H. (2005) Seasonal changes in soil arthropod abundance in the dry evergreen forest of north-east Thailand, with special reference to collembolan communities. Ecological Research, 20, 59–70. https://doi.org/10.1007/s11284-004-0013-x

Wu, P. & Wang, C. (2019) Differences in spatiotemporal dynamics between soil macrofauna and mesofauna communities in forest ecosystems: The significance for soil fauna diversity monitoring. Geoderma, 337, 266–272. https://doi.org/10.1016/j.geoderma.2018.09.031

Xin, W.D., Yin, X.Q. & Song, B. (2012) Contribution of soil fauna to litter decomposition in Songnen sandy lands in northeastern China. Journal of Arid Environments, 77, 90–95, https://doi.org/10.1016/j.jaridenv.2011.10.001

Xu, G., Zhang, Y., Zhang, S. & Ma, K. (2020) Biodiversity associations of soil fauna and plants depend on plant life form and are accounted for by rare taxa along an elevational gradient. Soil Biology and Biochemistry, 140, 107640. https://doi.org/10.1016/j.soilbio.2019.107640

Yang, X., Shao, M. & Li, T. (2020) Effects of terrestrial isopods on soil nutrients during litter decomposition. Geoderma, 376, 114546. https://doi.org/10.1016/j.geoderma.2020.114546

Zagatto, M.R.G., Niva, C.C., Thomazini, M.J., Baretta, D., Santos, A., Nadolny, H., Cardoso, G.B.X. & Brown, G.G. (2017) Soil Invertebrates in different land use systems: How Integrated production systems and seasonality affect soil mesofauna communities. Journal of Agricultural Science and Technology, 7, 158–169. https://doi.org/10.17265/2161-6264/2017.03.003

Zagatto, M.R.G., Filho, L.C.I.O., Pompeo, P.N., Niva, C.C., Baretta, D. & Cardoso, E.J.B.N. (2020) Mesofauna and macrofauna in soil and litter of mixed plantations. In: Bran Nogueira Cardoso, E., Gonçalves, J., Balieiro, F. & Franco, A. (eds) Mixed Plantations of Eucalyptus and Leguminous Trees. Springer, Cham, pp. 155–171. https://doi.org/10.1007/978-3-030-32365-3_8

Zahriban Hesari, M., Shataee, S., Maghsoudi, Y., Mohammadi, J., Fransson, J.E.S. & Persson, H.J. (2020) Forest variable estimations using TanDEM-X Data in Hyrcanian Forests. Canadian Journal of Remote Sensing, 46 (2), 166–176. https://doi.org/10.1080/07038992.2020.1763790

Zan, P., Mao, Z. & Sun, T. (2022) Effects of soil fauna on litter decomposition in Chinese forests: a meta-analysis. PeerJ, 10, e12747. http://doi.org/10.7717/peerj.12747

Zhang, L., Liu, J., Yin, R., Xu, Z., You, C., Li, H., Wang, L., Liu, S., Xu, H., Xu, L., Liu, Y., Wang, Y. & Tan, B. (2023) Soil fauna accelerated litter C and N release by improving litter quality across an elevational gradient. Ecological Processes, 12, 47. https://doi.org/10.1186/s13717-023-00459-4

Zhang, Z.Q. (2011) Animal biodiversity: An introduction to higher-level classification and taxonomic richness. Zootaxa, 3148 (1), 7–12. https://doi.org/10.11646/zootaxa.3148.1.3