Interactive effects of litter diversity, functional traits, and macrofauna on decomposition of litter mixtures from lianas and trees

Denis Mburu Njoroge; Yanlin Zhu; Jean Evans Israel Codjia; Juan Zuo; Gbadamassi G.O. Dossa; Yali Song; Douglas Schaefer; Mareike Roeder; Rhett D. Harrison; Johannes H.C. Cornelissen

doi:10.1016/j.soilbio.2026.110163

Interactive effects of litter diversity, functional traits, and macrofauna on decomposition of litter mixtures from lianas and trees

Denis Mburu Njoroge
, Yanlin Zhu
, Jean Evans Israel Codjia
, Juan Zuo^*
, Gbadamassi G.O. Dossa^*
, Yali Song^*
, Douglas Schaefer
, Mareike Roeder
, Rhett D. Harrison
, Johannes H.C. Cornelissen

^*Corresponding author for this work

Systems Ecology

Research output: Contribution to Journal › Article › Academic › peer-review

Abstract

Understanding the effect of litter mixing on litter decomposition rates is a prerequisite for predicting global carbon and nutrient cycling. However, the complex interactions associated with the decomposition of litter mixtures hinder their inclusion in global models. Soil fauna play an important role in the decomposition of litter mixtures, and their body size affects their ability to decompose litter. However, how different groups of soil fauna interact with litter traits to drive litter mixture decomposition remains poorly known. Here, through a field leaf litterbag decomposition experiment, we tested how soil fauna size and litter traits affect the decomposition of litter mixtures. We hypothesized that: (H1) The rate of decomposition and the magnitude of the litter mixing effect are greater in the presence of macrofauna. (H2) The effect of litter traits on the litter decomposition rate and the litter mixing effect decreases as the size of soil fauna increases. We used litter from two plant functional groups (trees and lianas) and incubated 27 litter treatments, including 12 single species, six 2-species mixtures, and nine 4-species mixtures. Each litter treatment was incubated using three litterbag mesh size treatments representing access to fauna of increasing body size (0.07 mm, 2.0 mm, and 5.0 mm) for 365 days. Both H1 and H2 were only partially supported. Macrofauna access accelerated decomposition and increased the litter mixing effect for tree-only mixtures. Initial litter traits predicted mixture decomposition rates in the presence of microbes and mesofauna, but not in the presence of macrofauna. Surprisingly, initial traits did not predict the litter mixing effect. This study highlights that litter traits may vary in their importance to different groups of soil fauna. In nature, therefore, where fauna communities (micro-, meso-, and macro-fauna) co-occur, and litter mixtures vary spatially, it may be difficult to predict decomposition rates.

Original language	English
Article number	110163
Pages (from-to)	1-10
Number of pages	10
Journal	Soil Biology and Biochemistry
Volume	220
Early online date	17 Apr 2026
DOIs	https://doi.org/10.1016/j.soilbio.2026.110163
Publication status	E-pub ahead of print - 17 Apr 2026

Bibliographical note

Publisher Copyright:
© 2026 Elsevier Ltd.

Funding

This work was funded by National Natural Science Foundation of China (NSFC) ( W2432022 , 31850410488 ), Yunnan Province Government for Talents Program ( E1YN101B01 ), Zhi Hui Yunnan Program ( 202203AM140024 ), and young international staff Chinese Academy of Sciences (CAS) president international fellowship initiative (PIFI) grant ( 2019FYB0001 ). This work also benefited from the Open Funding from CAS Key Laboratory of Tropical Forest Ecology, Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences ( 22-CAS-TFE-06 ), and from Southeast Asia Biodiversity Research Institute, Chinese Academy of Sciences (CAS-SEABRI: #Y4ZK111B05). The study was supported by the International Partnership Program of the Chinese Academy of Sciences , Grant No. 067GJHZ2024082FN . DMN was supported by funding from the European Union (ERC, BIOCOMP, GA 101075426). We appreciate Jinfa Dao for his help during fieldwork and sample processing. In addition, we are grateful to the members of the Institutional Center for Shared Technologies and Facilities of Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, for their assistance with analyzing the initial litter chemical traits. We are grateful to the editorial board and the anonymous reviewers for their suggestions that enabled us to improve substantially the quality of the manuscript.

Funders	Funder number
Xishuangbanna Tropical Botanical Garden
European Commission
CAS Key Laboratory of Tropical Forest Ecology
Yunnan Province Government for Talents Program	E1YN101B01
Southeast Asia Biodiversity Research Institute	067GJHZ2024082FN
Chinese Academy of Sciences	22-CAS-TFE-06
European Research Council	GA 101075426
Zhi Hui Yunnan Program	2019FYB0001, 202203AM140024
National Natural Science Foundation of China	31850410488, W2432022

Keywords

Biogeochemical cycling
Diversity effect
Ecosystem functioning
Functional traits
Liana afterlives
Litter mixing
Non-additive effects

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10.1016/j.soilbio.2026.110163

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Njoroge, D. M., Zhu, Y., Codjia, J. E. I., Zuo, J., Dossa, G. G. O., Song, Y., Schaefer, D., Roeder, M., Harrison, R. D., & Cornelissen, J. H. C. (2026). Interactive effects of litter diversity, functional traits, and macrofauna on decomposition of litter mixtures from lianas and trees. Soil Biology and Biochemistry, 220, 1-10. Article 110163. Advance online publication. https://doi.org/10.1016/j.soilbio.2026.110163

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title = "Interactive effects of litter diversity, functional traits, and macrofauna on decomposition of litter mixtures from lianas and trees",

abstract = "Understanding the effect of litter mixing on litter decomposition rates is a prerequisite for predicting global carbon and nutrient cycling. However, the complex interactions associated with the decomposition of litter mixtures hinder their inclusion in global models. Soil fauna play an important role in the decomposition of litter mixtures, and their body size affects their ability to decompose litter. However, how different groups of soil fauna interact with litter traits to drive litter mixture decomposition remains poorly known. Here, through a field leaf litterbag decomposition experiment, we tested how soil fauna size and litter traits affect the decomposition of litter mixtures. We hypothesized that: (H1) The rate of decomposition and the magnitude of the litter mixing effect are greater in the presence of macrofauna. (H2) The effect of litter traits on the litter decomposition rate and the litter mixing effect decreases as the size of soil fauna increases. We used litter from two plant functional groups (trees and lianas) and incubated 27 litter treatments, including 12 single species, six 2-species mixtures, and nine 4-species mixtures. Each litter treatment was incubated using three litterbag mesh size treatments representing access to fauna of increasing body size (0.07 mm, 2.0 mm, and 5.0 mm) for 365 days. Both H1 and H2 were only partially supported. Macrofauna access accelerated decomposition and increased the litter mixing effect for tree-only mixtures. Initial litter traits predicted mixture decomposition rates in the presence of microbes and mesofauna, but not in the presence of macrofauna. Surprisingly, initial traits did not predict the litter mixing effect. This study highlights that litter traits may vary in their importance to different groups of soil fauna. In nature, therefore, where fauna communities (micro-, meso-, and macro-fauna) co-occur, and litter mixtures vary spatially, it may be difficult to predict decomposition rates.",

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AU - Njoroge, Denis Mburu

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AU - Codjia, Jean Evans Israel

AU - Zuo, Juan

AU - Dossa, Gbadamassi G.O.

AU - Song, Yali

AU - Schaefer, Douglas

AU - Roeder, Mareike

AU - Harrison, Rhett D.

AU - Cornelissen, Johannes H.C.

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N2 - Understanding the effect of litter mixing on litter decomposition rates is a prerequisite for predicting global carbon and nutrient cycling. However, the complex interactions associated with the decomposition of litter mixtures hinder their inclusion in global models. Soil fauna play an important role in the decomposition of litter mixtures, and their body size affects their ability to decompose litter. However, how different groups of soil fauna interact with litter traits to drive litter mixture decomposition remains poorly known. Here, through a field leaf litterbag decomposition experiment, we tested how soil fauna size and litter traits affect the decomposition of litter mixtures. We hypothesized that: (H1) The rate of decomposition and the magnitude of the litter mixing effect are greater in the presence of macrofauna. (H2) The effect of litter traits on the litter decomposition rate and the litter mixing effect decreases as the size of soil fauna increases. We used litter from two plant functional groups (trees and lianas) and incubated 27 litter treatments, including 12 single species, six 2-species mixtures, and nine 4-species mixtures. Each litter treatment was incubated using three litterbag mesh size treatments representing access to fauna of increasing body size (0.07 mm, 2.0 mm, and 5.0 mm) for 365 days. Both H1 and H2 were only partially supported. Macrofauna access accelerated decomposition and increased the litter mixing effect for tree-only mixtures. Initial litter traits predicted mixture decomposition rates in the presence of microbes and mesofauna, but not in the presence of macrofauna. Surprisingly, initial traits did not predict the litter mixing effect. This study highlights that litter traits may vary in their importance to different groups of soil fauna. In nature, therefore, where fauna communities (micro-, meso-, and macro-fauna) co-occur, and litter mixtures vary spatially, it may be difficult to predict decomposition rates.

AB - Understanding the effect of litter mixing on litter decomposition rates is a prerequisite for predicting global carbon and nutrient cycling. However, the complex interactions associated with the decomposition of litter mixtures hinder their inclusion in global models. Soil fauna play an important role in the decomposition of litter mixtures, and their body size affects their ability to decompose litter. However, how different groups of soil fauna interact with litter traits to drive litter mixture decomposition remains poorly known. Here, through a field leaf litterbag decomposition experiment, we tested how soil fauna size and litter traits affect the decomposition of litter mixtures. We hypothesized that: (H1) The rate of decomposition and the magnitude of the litter mixing effect are greater in the presence of macrofauna. (H2) The effect of litter traits on the litter decomposition rate and the litter mixing effect decreases as the size of soil fauna increases. We used litter from two plant functional groups (trees and lianas) and incubated 27 litter treatments, including 12 single species, six 2-species mixtures, and nine 4-species mixtures. Each litter treatment was incubated using three litterbag mesh size treatments representing access to fauna of increasing body size (0.07 mm, 2.0 mm, and 5.0 mm) for 365 days. Both H1 and H2 were only partially supported. Macrofauna access accelerated decomposition and increased the litter mixing effect for tree-only mixtures. Initial litter traits predicted mixture decomposition rates in the presence of microbes and mesofauna, but not in the presence of macrofauna. Surprisingly, initial traits did not predict the litter mixing effect. This study highlights that litter traits may vary in their importance to different groups of soil fauna. In nature, therefore, where fauna communities (micro-, meso-, and macro-fauna) co-occur, and litter mixtures vary spatially, it may be difficult to predict decomposition rates.

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KW - Ecosystem functioning

KW - Functional traits

KW - Liana afterlives

KW - Litter mixing

KW - Non-additive effects

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M1 - 110163

ER -

Interactive effects of litter diversity, functional traits, and macrofauna on decomposition of litter mixtures from lianas and trees

Abstract

Bibliographical note

Funding

Keywords

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