Applied Soil Ecology 210 (2025) 106063

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Applied Soil Ecology

journal homepage: www.elsevier.com/locate/apsoil

Short Communication

Relationships between earthworm community, bioturbation and soil

detachment: A one-year outdoor experiment
Q.V. Pham a,* , Y. Capowiez b, P. Jouquet c,d , A.D. Nguyen e , J.L. Janeau c , T.M. Tran f,
N. Bottinelli c,f
a
Faculty of Environmental Sciences, University of Science, Vietnam National University, Hanoi (VNU), 334 Nguyen Trai, Thanh Xuan, Hanoi, Viet Nam
b
INRAE, Site Agroparc, Avignon cedex 09, 84914 Paris, France
c
IRD, CNRS, Sorbonne Universit!e, Univ Paris Est Creteil, INRAe, Institute of Ecology and Environmental Sciences (iEES-Paris), F-75005 Paris, France
d
Research and Innovation Center, Faculty of Hydrology and Water Resources Engineering, Institute of Technology of Cambodia (ITC), Phnom Penh, Cambodia
e
Institute of Ecology and Biological Resources, Vietnam Academy of Science and Technology, 18, Hoang Quoc Viet, Hanoi, Viet Nam
f
Department of Soil sciences, Soils and Fertilizers Institute (SFI), Hanoi, Viet Nam

A R T I C L E I N F O A B S T R A C T

Keywords: We conducted a one-year outdoor experiment to evaluate how the functional diversity of earthworms influences
Macrofauna soil erosion. Three species with contrasting bioturbating behaviors were selected: the polyhumic endogeic
Biostructures Pontoscolex corethrurus (feeding and casting on the soil surface and constructing shallow burrows), the endogeic
Hydrological properties
Amynthas alluxus (geophagous and permanently living in the subsoil), and the anecic Amynthas zenkevichi
(feeding and casting on the soil surface and constructing deep burrows). A total of 27 mesocosms (1 m3 each)
planted with grass were inoculated with none to all three species at two biomass levels (30 and 60 g m→2). Soil
detachment, water runoff, drainage, soil water potential, and grass biomass were monitored throughout the
whole experiment, while burrow volume, water infiltration rate, surface casts, and earthworm communities were
measured at the end. A. zenkevichi was the only surviving species and colonized all the mesocosms. Consequently,
the impact of each species and their interactions on soil detachment could not be assessed. Nevertheless, mes-
ocosms initially inoculated with earthworms showed, on average, a 7.7-fold decrease in soil detachment
compared to the control mesocosm, where no earthworms were introduced at the beginning but was later
colonized by dispersed earthworms. Structural equation modeling explained 77 % of the variance in soil
detachment. It revealed that both surface casts and burrows (measured by X-ray tomography) indirectly reduced
soil detachment by increasing water infiltration and reducing water runoff. However, surface casts also promoted
water runoff, partially counteracting these benefits. This study highlights the challenges of managing earthworm
communities in long-term mesocosm experiments under natural conditions. Despite these limitations, our
findings emphasize the crucial role of anecic earthworms in reducing soil detachment.

1. Introduction degradation can be mitigated by activity of earthworms (Le Bayon and

Binet, 2001; Jouquet et al., 2008, 2012; Bottinelli et al., 2021), which
Soil erosion, driven by water runoff, represents a significant chal- are known to influence soil porosity, water stability of aggregates and
lenge to sustainable agriculture, particularly under intense rainfall water infiltration (Shipitalo and Le Bayon, 2004; Blouin et al., 2013).
events currently observed in tropical regions (Anache et al., 2017; The mechanisms by which earthworms affect soil erosion are well-
Ahmad et al., 2020). The impact of raindrops with a kinetic energy documented (Shipitalo and Le Bayon, 2004). A consensus in the litera-
greater than the soil strength breaks down soil aggregates (Wang et al., ture suggests that the presence of earthworms reduces runoff by
2014). This process generates sediment transport, accelerates surface enhancing soil water infiltration. This occurs primarily through the
seal formation, decreases infiltration, increases overland flow, and leads presence of burrow networks especially when they maintain open
to an overall increase in soil erosion (Lal, 2001). This physical pathways for water at the soil surface (Andriuzzi et al., 2015; Le Mer

* Corresponding author.
E-mail address: phamvanquang@vnu.edu.vn (Q.V. Pham).

https://doi.org/10.1016/j.apsoil.2025.106063
Received 14 January 2025; Received in revised form 25 March 2025; Accepted 26 March 2025
Available online 29 March 2025
0929-1393/© 2025 Elsevier B.V. All rights are reserved, including those for text and data mining, AI training, and similar technologies.
Q.V. Pham et al. Applied Soil Ecology 210 (2025) 106063

et al., 2021). Furthermore, earthworms’ surface casting activities sieved through a 2 mm mesh and manually compacted in 10 cm layers to
improve soil surface roughness, reducing soil crusting and facilitating achieve a bulk density of 1.1 g cm→3, resulting in a total soil height of 75
water infiltration (Kladivko et al., 1986; Le Bayon and Binet, 2001). cm. To provide a food source for the earthworms, the top 25 cm of soil
Nevertheless, under certain conditions, surface casts can also contribute was amended with dried buffalo dung to achieve a soil organic carbon
to soil erosion (Sharpley et al., 1979; Wen et al., 2022). For instance, content of 1.5 %. After refilling, the soil surface was shaped with a 5↑
casts can be vulnerable to splash erosion (Nooren et al., 1995; Mariani slope, which was connected to a runoff water collection system. One
et al., 2007), and their fine soil materials can easily be detached and month after planting Brachiaria mutica grass, earthworms were intro-
transported during rainfall events. duced following the axial simplex sampling design (Supplementary
Since anecic, epigeic and endogeic earthworm species exhibit con- Fig. S1c). Communities were set up as either (i) monocultures, (ii)
trasting burrowing and casting activities (Capowiez et al., 2024), their centroids, consisting of equal amounts of each species, (iii) binary
effects on soil resistance to erosion are likely dependant on their mixtures, consisting of equal biomass of two species or (iv) complete
ecological category. Anecic species are highly effective at reducing mixtures consisting of species, where the proportional contributions of
erosion, as their deep, surface-connected burrows facilitate infiltration the species sum to one (Supplementary Fig. S1b). The experiment was
and their cast, enriched with organic matter stabilize soil structure. carried out at two biomass levels, 30 and 60 g fresh weight per square
While previous studies have documented anecic impacts on soil erosion meter, yielding a total of 26 earthworm communities. The biomass of
(Jouquet et al., 2008; Bottinelli et al., 2021; Wen et al., 2022), the effects earthworms was decided upon biomass reported for grasslands in Viet-
of other ecological categories is comparatively ignored. Moreover, field nam (Mulia et al., 2021). One mesocosm without earthworms served as a
research involving earthworm communities with several species could control. The experiment ran for 12 months, from November 2021 to
not disentangled the specific effects of the different species (Sharpley November 2022.
et al., 1979; Pitk”anen and Nuutinen, 1998; Le Bayon and Binet, 2001).
Consequently, how different species with distinct bioturbation behav- 2.3. Data collection
iors affect soil erosion dynamics remains unknown.
To address this, we conducted an outdoor mesocosm experiment in An automatic weather station measuring precipitation every minute
northern Vietnam, designed to assess the effects of single-species and was located in proximity to the mesocosms. Watermark sensors were
mixed-species assemblages of three earthworm species, Pontoscolex buried at 10 and 40 cm depth to continuously monitor soil water tension
corethrurus, Amynthas alluxus (endogeic species) and Amynthas zen- in each mesocosm. The sensor values ranged from 0 to 254 centibars
kevichi (anecic species). We tested species assemblages at two biomass (cb), representing soil moisture levels from wet to dry. Grass was cut
levels (30 vs. 60 g m→2) over one years. We measured soil detachment, every two months, leaving a 3 cm stub, and the air-dry biomass of the cut
water runoff and infiltration and bioturbation. We hypothesized that: (i) grass was recorded. After each rainfall event, the volumes of runoff and
earthworm species with distinct burrowing behaviors would differen- drainage water were measured, and soil particles lost by detachment
tially influence soil detachment, with anecic species exerting greater were quantified by filtering the samples through 0.45 μm membrane
effects than the two endogeic species and (ii) higher earthworm biomass filters. The water runoff coefficient is the surface water collected divided
would amplify bioturbation, enhancing infiltration and reducing erosion by daily rainfall. After one year of experimentation, we measured the
compared to lower biomass level. weight of surface casts, water infiltration rate, the volume of earthworm
burrows and the community structure of earthworms. Surface casts were
2. Materials and methods collected, air-dried and weighted. The Beerkan test was used to measure
water infiltration rate in the center of each mesocosm. Ten 330 mL
2.1. Earthworms and soil volumes of water were poured into a 19 cm diameter cylinder placed at
2–3 cm depth. The infiltration rate was then determined by the slope of
Three earthworm species from northern Vietnam were sampled for the linear regression relating the water volume to the time at soil satu-
this study: P. corethrurus, A. alluxus, and A. zenkevichi. They were ration (Capowiez et al., 2015).
assigned into ecological categories according to their morpho- The percentage of earthworm burrows ω5 mm3, found at 0–10 cm
anatomical traits (Pham et al., 2023). P. corethrurus and A. alluxus are depth, was measured using X-ray computed tomography in soil columns
endogeic species, whereas A. zenkevichi is an anecic species. However, (10 cm in height, 12.5 cm in diameter, one per mesocosm) with the Fiji/
the bioturbation behavior of the two endogeic species revealed signifi- ImageJ plugin “SoilJ” (Koestel, 2018). Finally, earthworms were hand-
cant differences: P. corethrurus exhibits surface activity, producing sorted from each mesocosm, rinsed with water, weighed for fresh
substantial amounts of casts, whereas A. alluxus shows no surface ac- biomass, and counted for abundance.
tivity and constructs deeper burrows (Pham et al., 2023, 2024).
A. alluxus (average individual fresh weight of 0.9 g) was sampled in a 2.4. Statistical analysis
forest located 50 km from Hanoi (20↑ 56′ N, 105↑ 29′ E) in soil with a
pHKCl of 4.2 and an organic carbon content of 1.0 %. P. corethrurus Throughout the experiment, earthworm communities changed
(average individual fresh weight of 0.4 g) was collected from a crop field dramatically, making it impossible to compare the effects of different
near the forest where A. alluxus was sampled, with a soil pHKCl of 4.1 and assemblages on soil detachment. Consequently, we used t-tests to
an organic carbon content of 1.2 %. A. zenkevichi (average individual compare mesocosms with varying initial earthworm biomasses (0, 30,
fresh weight of 2.4 g) were collected from a crop field (20↑ 44′ N, 106↑ and 60 g m→2, with sample sizes of n ↓ 1, 13 and 13, respectively).
17′ E) in soil with a pHKCl of 3.6 and an organic content of 1.3 %. The soil Before conducting the t-tests, normality was assessed using the Shapir-
used for the experiment was obtained in large outdoor mesocosms o–Wilk test, and homogeneity of variance was evaluated with Levene’s
previously used for growing maize and had the following properties: 22 test. These analyses were conducted using R software version 4.2.0 (R
% clay, 35 % silt, 43 % sand, pHwater of 5.3 and soil organic carbon Core Team, 2024). To further explore the relationships between soil
content of 0.3 % (Doan et al., 2015). erosion and other variables including grass biomass, earthworm abun-
dance and activity and soil properties, Partial Least Squares Structural
2.2. Outdoor mesocosms and experimental design Equation Modeling (PLS-SEM) was performed using SmartPLS software
(Ringle et al., 2024). Model fit was assessed using the Standardized Root
The experiment was conducted at the Soil and Fertilizer Institute in Mean Square Residual (SRMR) with a value below 0.1 considered
Hanoi, Vietnam (21↑ 04′ N, 105↑ 46′ E), using outdoor mesocosms (1 m3) indicative of a good model fit. Statistical significance was determined at
constructed from concrete (Supplementary Fig. S1a–b). The soil was p ε 0.05.

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Q.V. Pham et al. Applied Soil Ecology 210 (2025) 106063

3. Results interactions on soil detachment could not be assessed. The absence of

the two endogeic species indicates mortality, which can be explained by
3.1. Earthworm abundance and bioturbation several factors: (i) the rapid mineralization of buffalo dung may have
depleted soil organic carbon, leaving insufficient resources for the
A. zenkevichi was the only earthworm species found in the meso- endogeic species, which depend mainly on soil organic matter (Curry
cosms after one year. Its abundance was relatively consistent across the and Schmidt, 2007); (ii) the saturated soils caused by heavy rainfall
mesocosms, with an average density of 16.1 individuals per square events could have been detrimental to these less mobile species, which
meter, corresponding to a biomass of 14.6 g m→2 (Supplementary may have faced anaerobic conditions within the mesocosms. Indeed,
Fig. S2). However, it was impossible to determine when the two other earthworm species vary regarding their optimal soil moisture range and
species disappeared, as well as when and at what speed the anecic not all species can tolerate inundated soils (Curry, 2004; Zorn et al.,
species colonized the mesocosms. We therefore compared the meso- 2008). In contrast, A. zenkevichi, an anecic species that feeds on surface
cosms based on the introduced biomasses (b30 ↓ 30 or b60 ↓ 60 g m→2) organic matter and migrates across soil depths (Curry and Schmidt,
with the sole mesocosm initially without earthworms b0, even though by 2007), demonstrated a greater adaptability to these challenges. Its
the end of the experiment, the biomass of A. zenkevichi in this mesocosm colonization of all mesocosms suggests resource-seeking behavior or
was comparable to the biomass observed in the other mesocosms. avoidance of flooding conditions. However, the uniformly low biomass
The mass of surface casts varied between 2.5 and 4.1 kg m→2 and was at the end of the experiment indicates that intraspecific competitions
significantly lower in mesocosms b30 or b60 compared to the mesocosm likely limited their abundance. Finally, it cannot be excluded that
b0 (Table 1). The volume percentage of earthworm burrows ranged from A. zenkevichi, which colonized all the mesocosms, induced competitive
0.4 to 0.8 % and was significantly higher in mesocosms b30 or b60 interactions that significantly impaired the coexistence of the two
compared to the mesocosm b0 (Table 1). endogeic species.
Despite the mortality of the endogeic species, mesocosms inoculated
3.2. Hydrological properties and grass biomass with earthworms exhibited significant improvements in soil hydraulic
properties compared to the control mesocosm. Water infiltration and
The experiment recorded 1801 mm of rainfall in one year, including drainage increased by 2.3 and 6.6-fold, respectively, while water runoff
17 events with intensities exceeding 20 mm h→1. Soil water potential and soil detachment were substantially reduced by 1.6 and 7.7-fold,
ranged from 80 to 120 cb in December and January, and from 0 to 25 cb respectively. Our results are consistent with numerous studies that
during the rest of the year, with no differences between treatments highlight the positive impacts of anecic species on water infiltration
(Supplementary Fig. S3). The yearly runoff surface coefficient (4–30 %) under saturated conditions (Andriuzzi et al., 2015; Le Mer et al., 2021)
and soil detachment (0.06–0.1 kg m→2) were significantly lower in and the reduction of water runoff (Jouquet et al., 2008; Wen et al., 2022)
mesocosms b30 or b60 compared to the mesocosm b0 (Table 1). Addi- and erosion (Jouquet et al., 2008; Bottinelli et al., 2021).
tionally, water infiltration rate (103–236 mm h→1) and drainage The application of PLS-SEM provided additional insights into the
(96–634 mm) were also significantly reduced, while grass biomass indirect pathways through which A. zenkevichi influenced soil detach-
(1.45–1.64 kg m→2) was significantly higher in mesocosms b30 or b60 ment. An increase in burrow volume enhanced water infiltration and
compared to the mesocosm b0 (Table 1). drainage, reducing surface runoff and soil detachment. Regarding sur-
face casts, our findings revealed a dual role: on one hand, casts were
3.3. Causal relationships associated with soil detachment positively correlated to water infiltration, and thus to reduced runoff
and erosion. On the other hand, under our specific conditions, they
The PLS-SEM model could explain 77 % of the variance in soil contributed to increased water runoff, partially offsetting their erosion-
detachment and showed an acceptable fit with a SRMR of 0.068 (Fig. 1). mitigating effects. These observations are in agreement with previous
Surface casts were indirectly positively linked to soil detachment by studies demonstrating that surface casting are indeed correlated to
increasing surface water runoff. Both surface casts and earthworm burrows open at the soil surface, facilitating water infiltration, and that
burrows were indirectly negatively linked to soil detachment by casts can also act as barriers to surface runoff (Le Bayon and Binet, 2001;
increasing water infiltration which in turn decreased water runoff. Grass Jouquet et al., 2008; Bottinelli et al., 2021). The lack of a direct rela-
biomass, earthworm abundance and biomass did not influence soil tionship between surface casts and soil detachment suggests that rain-
detachment. degraded casts are transported only over very short distances and they
were mixed with the surface soil matrix which can lead to the formation
4. Discussion of crusts and then increase surface runoff.

After one year, A. zenkevichi was the only species present in the 5. Conclusions
mesocosms, with a relatively similar biomass across mesocosms, aver-
aging 16.1 g m→2, even in the one where earthworms had not been Despite the mortality of the two endogeic species, the persistence and
inoculated. As a result, the effects of the different species and their colonization of the anecic A. zenkevichi across all mesocosms illustrated

Table 1
Means ↔ standard deviations of grass biomass, earthworm biostructures and hydrological properties in mesocosms initially without earthworms (b0, n ↓ 1), and
earthworm biomass of 30 g m→2 (b30, n ↓ 13) and 60 g m→2 (b60, n ↓ 13). Comparisons between different earthworm densities were tested with t-tests. The number of
asterisks indicates the significance level, with ***p ε 0.001 and *p ε 0.05.
Mesocosm property Earthworm biomass p value

b0 b30 b60 b0 vs. b30 b0 vs. b60 b30 vs. b60

Grass biomass (kg m→2) 1.45 1.63 (0.19) 1.64 (0.19) *** *** ns
Surface cast (kg m→2) 4.1 2.9 (0.7) 2.5 (0.5) *** *** ns
Burrows (%) 0.4 0.7 (0.3) 0.8 (0.3) *** *** ns
Runoff coefficient (%) 29.1 17.5 (6.2) 18.8 (6.6) *** *** ns
Infiltration rate (mm h→1) 103 236 (192) 236 (181) * * ns
Water drainage (mm) 96 648 (388) 624 (379) *** *** ns
Soil detachment (kg m→2) 0.1 0.06 (0.03) 0.06 (0.03) *** *** ns

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Q.V. Pham et al. Applied Soil Ecology 210 (2025) 106063

Fig. 1. Structural equation model illustrating relationships among variables describing earthworm abundance, surface cast weight, grass biomass, gallery volume,
water characteristics (infiltration rate, water runoff, water drainage, and soil water tension), and soil detachment. Solid arrows indicate significant pathways, while
dotted arrows represent non-significant relationships. Path coefficients (standardized values ranging from →1 to 1) are displayed next to the arrows with significance
levels denoted by asterisks (*p ε 0.05, **p ε 0.01, ***p ε 0.001). Model R2 values are presented within the circles.

its significant role in mitigating soil detachment through its bioturbating Validation, Supervision. P. Jouquet: Writing – review & editing, Vali-
activity. We confirmed that earthworm casts played a dual role in soil dation, Supervision. A.D. Nguyen: Supervision. J.L. Janeau: Supervi-
detachment. On the one hand, they reduced water runoff velocity, sion. T.M. Tran: Project administration. N. Bottinelli: Writing – review
enhancing water infiltration. On the other hand, they were prone to & editing, Visualization, Validation, Supervision, Funding acquisition.
degradation, which increased water runoff. However, in this one-year
outdoor experiment, the effect of earthworm activity was considered
Declaration of competing interest
as being rather beneficial in terms of soil erosion and the positive effects
of casts and burrows clearly counterbalanced the negative ones. Fa-
The authors declare that they have no known competing financial
voring anecic earthworms is therefore an interesting strategy to decrease
interests or personal relationships that could have appeared to influence
soil erosion in tropical ecosystems. Future studies should focus on
the work reported in this paper.
optimizing experimental conditions to support earthworm survival,
while investigating species-specific interactions and their effects on soil
processes. Acknowledgements
Supplementary data to this article can be found online at https://doi.
org/10.1016/j.apsoil.2025.106063. This project was financially supported by the ANR project PRECIOUS
(ANR-19-CE01-0003). Pham Van Quang was funded by the French
CRediT authorship contribution statement Research Institute for Development (IRD) through “Allocations de
recherche pour une th#ese au Sud” (ARTS) and the Postdoctoral Schol-
Q.V. Pham: Writing – review & editing, Writing – original draft, arship Programme of Vingroup Innovation Foundation (VINIF), code
Methodology, Investigation. Y. Capowiez: Writing – review & editing, VINIF.2024.STS.31. We thank Jean Luc Maeght and Thierry Henry-des-
Tureaux for their assistance in setting up soil moisture sensors in

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