Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e64043, enero-diciembre 2025 (Publicado Mar. 03, 2025)

Does mesofaunal abundance and composition influence

mangrove leaf litter decomposition?

Margarita Loría-Naranjo1,2*; https://orcid.org/0000-0003-4396-7388

Jeffrey A. Sibaja-Cordero1,2; https://orcid.org/0000-0001-5323-356X

Jorge Cortés1,2; https://orcid.org/0000-0001-7004-8649

1. Centro de Investigación en Ciencias del Mar y Limnología (CIMAR) Ciudad de la Investigación, Universidad de Costa

Rica, San Pedro; maggie.loria@gmail.com; jorge.cortes@ucr.ac.cr; jeffrey.sibaja@ucr.ac.cr

2. Escuela de Biología, Universidad de Costa Rica, San Pedro.

Received 04-IX-2024. Corrected 18-XI-2024. Accepted 05-II-2025.

ABSTRACT

Introduction: Litter decomposition in mangrove forest is a process in which organic matter produced as leaves

in the trees is later transferred to the sediment. Several invertebrates participate in the fragmentation of leaves,

and they might have a great impact in the later stages of decomposition of mangrove leaves.

Objective: The study aims to characterize mesofauna inhabiting leaf litter of Rhizophora racemosa, in the South

Pacific of Costa Rica during the dry and rainy season.

Methods: Five pre-weighted litter bag samples were set out during each season. Each sample consisted of

specimens adhered to the leaves after 0, 15, 30, 60, 90 and 120 days, from two sites in Térraba-Sierpe National

Wetland. The samples were washed in a sieve (0.5 mm) and the organisms were separated and identified. Taxa

were assigned to trophic groups to evaluate their influence in the decomposition of the leaves. Additionally, leaf

detritus samples retained in the sieve were dried in an oven at 70 °C to calculate the remaining vegetal matter

for each site and season.

Results: The most abundant taxa were: Bivalvia (26 %), Nematoda (19.45 %), Gastropoda (14.78 %), Nemertea

(14.61 %), and Ostracoda (8.20 %). Based on the trophic classification of the mesofauna found, it is expected that

most of them are indirect consumers of R. racemosa litter. Correlation analysis shows that, depending on the site

and the season, the mass loss of leaf litter was greater with a greater mesofauna abundance. A combined influ-

ence of water salinity, temperature and dissolved oxygen explains 25 % of the changes observed in the mesofauna

found in leaf litter samples throughout the sampling period.

Conclusions: Leaf litter decomposition process evidences the presence of an abundant and diverse mesofauna

community. The abundance of this mesofauna seems to have a positive influence on the decomposition of the

leaves. It is a vital issue to comprehend the role of mesofauna as potential consumers of mangrove leaf litter, in

order to conservate biodiversity in this ecosystem.

Key words: Rhizophora racemosa; estuarine invertebrates; feeding guilds; nutrient recycling.

RESUMEN

¿La abundancia y composición de la mesofauna influye en la descomposición

de la hojarasca de manglar?

Introducción: La descomposición de hojarasca del manglar es un proceso donde la materia orgánica producida

como hojas se transfiere posteriormente al sedimento. Varios invertebrados participan en la fragmentación y

posiblemente tienen impacto en etapas posteriores de descomposición de las hojas.

https://doi.org/10.15517/rev.biol.trop..v73iS1.64043

SUPPLEMENT

2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e64043, enero-diciembre 2025 (Publicado Mar. 03, 2025)

INTRODUCTION

Litter decomposition is a process in which

organic matter produced by mangrove forests

is transferred to the sediment. This process

involves at least four basic steps: 1) the removal

of soluble compounds from leaves by water, 2)

colonization by microorganisms, 3) consump-

tion of plant matter by herbivores, and 4) the

combined action of physical forces such as

water movement, deposition, and sediment

supply (Ashton et al., 1999; Dewiyanti, 2010;

Kathiresan & Bingham, 2001; Palit et al., 2022;

Swift et al., 1979). To study litter decomposi-

tion in mangroves, it is important to consider

factors such as dissolved oxygen in sediments,

soil salinity, forest type, and the composition

of litter material (Galeano et al., 2010; Lugo &

Snedaker, 1974; Matheri et al., 2024; Singh et

al., 1993; Twilley & Day, 1999).

Several invertebrates play a crucial role

in this process, especially in the later stages of

decomposition of mangrove leaves. This group

of organisms is called mesofauna, which ranges

in size from 0.2 mm to 2 mm (Matheri et al.,

2024). Mesofauna includes annelids, peracari-

dans, microarthropods, mollusks, nematodes,

gastrotrichs, and others (Dittmann, 2001; Gray

& Elliott, 2009; Maggenti & Maggenti, 2005).

When considering other functional groups,

such as microbes and macrofauna, it has been

found that mesofauna contributes between 23

% and 27 % to the overall litter decomposition

process in mangroves (Islam et al., 2024).

D’Croz et al. (1989) studied the mesofauna

community associated with the leaf litter of

Rhizophora mangle during the decomposition

process in Panama Bay. In Brazil, de Oliveira et

al. (2011) identified and quantified the benthic

macrofauna present during the leaf decomposi-

tion of R. mangle and Laguncularia racemosa.

Gladstone-Gallagher (2012) examined the ben-

thic community associated with the detritus of

Avicennia marina in two sites of a temperate

mangrove forest in New Zealand. When meso-

fauna is excluded, the rate of litter decomposi-

tion can be reduced by up to 50 %, suggesting

that these organisms play a significant role in

explaining patterns of leaf litter decomposition

(Powers et al., 2009). Therefore, the abundance

and composition of mesofauna serve as bioin-

dicators of the environmental quality of man-

grove systems (Xiping et al., 2015).

In Panama, the most abundant meso-

fauna groups found in litter samples of R.

mangle included polychaetes, bivalves, gas-

tropods, crabs, and shrimps (D’Croz et al.,

Objetivo: Este estudio tiene como objetivo caracterizar la mesofauna que habita la hojarasca de Rhizophora race-

mosa, en el Pacífico Sur de Costa Rica, durante las épocas seca y lluviosa.

Métodos: Se colocaron cinco bolsas de hojas pre-pesadas durante cada época. Cada conjunto se usó para estu-

diar los especímenes adheridos a las hojas después de 0, 15, 30, 60, 90 y 120 días, en dos sitios del Humedal

Nacional Térraba-Sierpe. Las muestras se lavaron en un tamiz (0.5 mm) y los organismos se separaron e identi-

ficaron. Los taxones se asignaron a grupos tróficos para evaluar su influencia en la descomposición de las hojas.

Adicionalmente, las muestras de detritos de hojas retenidas en el tamiz se secaron a 70 °C para calcular la materia

vegetal remanente por sitio y época.

Resultados: Los taxones más abundantes fueron: Bivalvia (26 %), Nematoda (19.45 %), Gastropoda (14.78 %),

Nemertea (14.61 %), and Ostracoda (8.20 %). Según la clasificación trófica de la mesofauna, se encontró que la

mayoría eran consumidores indirectos de la hojarasca. El análisis de correlación mostró que, dependiendo del

sitio y la estación, la pérdida de masa fue mayor con una mayor cantidad de mesofauna. Una influencia com-

binada de la salinidad, temperatura y oxígeno disuelto del agua explica el 25 % de los cambios observados en la

mesofauna de la hojarasca a lo largo del tiempo estudiado.

Conclusiones: Una abundante y diversa comunidad de mesofauna participa en el proceso de descomposición

de la hojarasca en el manglar de Térraba-Sierpe. La abundancia de esta mesofauna aparenta tener una influencia

positiva en la descomposición de las hojas. La comprensión del papel de esta mesofauna como consumidora de la

hojarasca es vital para la conservación de la biodiversidad en este ecosistema.

Palabras clave: Rhizophora racemosa; invertebrados estuarinos; gremios alimenticios; reciclaje de nutrientes.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e64043, enero-diciembre 2025 (Publicado Mar. 03, 2025)

1989). In a mangrove forest in the State of

Bahía, Brazil, annelids (Oligochaeta and Poly-

chaeta) were the most abundant organisms

during the decomposition of R. mangle leaves

(de Oliveira et al., 2011).

In Costa Rica, Díaz-Navarro (1993) stud-

ied the decomposition of Rhizophora sp. leaves

at Punta Morales in the Gulf of Nicoya. Recent-

ly, Loría-Naranjo et al. (2018) researched the

decomposition process of Rhizophora racemosa

leaf litter in the Térraba-Sierpe National Wet-

land (TSNW). The present study aims to deter-

mine the variation in mesofauna abundance

and composition at different sites, seasons, and

stages of leaf litter decomposition for the man-

grove tree R. racemosa.

MATERIALS AND METHODS

The present study was conducted in the

Térraba-Sierpe National Wetland (TSNW),

which is the largest mangrove forest in Costa

Rica, covering an area of 120.33 km² (Quesada-

Alpízar et al., 2006). Rhizophora racemosa is

the most common mangrove species along

the Pacific coast of Costa Rica (Jiménez, 1994;

Silva-Benavides, 2009). This species typically

inhabits the inner regions of the forest (Bar-

rantes-Leiva & Cerdas-Salas, 2015; Jiménez,

1988; Jiménez, 1994) and is the most abundant

mangrove species within TSNW, occupying an

area of 76.70 km² (Barrantes-Leiva & Cerdas-

Salas, 2015).

Study Site: TSNW is in Osa, Puntarenas,

Costa Rica, at coordinates 8°47’–9°03’ N and

83°29’–83°38’ W. The area is nourished by

two major rivers: the Grande de Térraba and

the Sierpe. TSNW encompasses approximately

40 % of Costa Rica’s total mangrove cover-

age (Windevoxhel-Lora, 1998). In 2013, the

annual precipitation was recorded at 3 301 mm,

with a mean atmospheric temperature of 26.6

°C (Instituto Meteorológico Nacional [IMN],

2014). The Talamanca Cordillera blocks the

trade winds, leading to an extended rainy sea-

son and a limited dry season (Jiménez, 1999).

Two sampling sites within TSNW were selected:

Estero Caballo (Site 1, located at 8°53’11.11” N

and 83°34’2.23” W) in the Sierpe basin, and

Boca Nueva (Site 2, located at 8°59’16.46” N

and 83°37’30.28” W) in the Grande de Térraba

basin. At both sites, R. racemosa was the pre-

dominant species found along the riverbanks.

Methodology: Ten field trips were con-

ducted during 2013, five during the dry season

(from January to April), and five during the

rainy season (from July to November). This

resulted in two experimental periods. Each

experiment period consisted of litter decompo-

sition data after 0, 15, 30, 60, 90, and 120 days.

At each site, senescent leaves of R. rac-

emosa were collected from the forest floor. Only

leaves with a yellowish color but no evidence

of any physical damage (holes or bites) were

collected, to guarantee a similar initial level of

decay. Litter bags of 10 x 20 cm and a mesh size

of 1 mm (Roberts et al., 1987) were used. For

both sites (Estero Caballo and Boca Nueva), 50

litter bags with 10 g of litter were attached to

roots or trunks, so that they were exposed to

wave action and different environmental condi-

tions depending on the season. Five litter bags

from each site were retrieved on each field trip.

In the laboratory, the samples contained in

the bags were washed using a sieve with a pore

size of 0.5 mm. The organisms retained into the

sieve were fixed in 70 % ethanol. The organisms

adhered to five leaf litter samples from each

site were quantified and identified. For this

purpose, a stereoscope along with taxonomic

keys (de León-González, 2009; Giere, 1993;

Glockner-Fagetti, 2009; Glockner-Fagetti &

Egremi-Valdéz, 2009; Hernández-Alcántara &

Solís-Weiss, 2009; Roldán, 1988; Salazar-Vallejo

& Rizo, 2009) were used. Individuals were

identified at the level of phylum, class, or order,

depending on their characteristics and the level

of detail achievable with the stereoscope. This

methodology provided a chronological record

of the types of fauna colonizing the samples

over various decomposition periods (0, 15,

30, 60, 90, and 120 days). Additionally, the

taxa were classified according to the different

4Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e64043, enero-diciembre 2025 (Publicado Mar. 03, 2025)

trophic groups they belong to, allowing to eval-

uate their influence on leaf decomposition.

Environmental data, including salinity

(UPS), temperature (°C), and dissolved oxygen

(mg O2/l), were recorded at both sites using a

multiparametric sensor during each field trip.

Precipitation (mm) and atmospheric tempera-

ture (°C) data were obtained from the closest

meteorological station in Palmar Norte, Pun-

tarenas (IMN, 2014).

Leaf mass loss: To estimate mass loss,

dry leaf litter samples were analyzed. The leaf

detritus collected in the sieve was dried in an

oven at 70° C until it reached a constant weight.

The remaining mass of leaves after 15, 30, 60,

90, and 120 days of decomposition was used to

determine the Spearman correlation with total

mesofauna abundance for each site and season.

Mesofauna community: To assess the

similarity in the composition of taxa that colo-

nized the samples, taxa abundance data were

transformed using log (x + 1) and the Bray-

Curtis similarity index was applied. To visual-

ize differences in taxa composition between

sample types (particularly those collected from

Site 1 or Site 2, during dry or rainy season), a

Multidimensional Scaling analysis (MDS) of

the pooled samples was performed, consider-

ing decomposition time, site and season, using

PRIMER 6.0.

Two-way similarity analysis (ANOSIM)

were conducted on two occasions with the

sample matrix (Clarke & Warwick, 1994) in

PAST software. The first analysis included the

time factor (15, 30, 60, 90 and 120 days) and

the site factor (Estero Caballo or Boca Nueva).

The second analysis considered the time fac-

tor alongside season (dry or rainy). An R

value of ANOSIM with P < 0.05 indicated the

degree of differentiation between levels of the

analyzed factors. This value ranges from 0 to

1, where 1 represents a complete difference in

taxa composition (Clarke & Warwick, 1994).

The probabilities obtained in these tests were

corrected using Bonferroni method, as the

same data matrix was utilized by both analyzes.

In addition, a posteriori comparisons were

performed to determine the R values between

sampling dates for each site according to the

season (Clarke & Warwick, 1994). To identify

which taxa contributed most to the difference

observed between the levels of each factor (site,

season, and time), three SIMPER tests were

conducted in the PAST software. Similar ANO-

SIM and SIMPER tests were also executed for

the trophic groups matrix of the invertebrates.

A BIO-ENV analysis was conducted to

identify the environmental variables—such as

water salinity, dissolved oxygen in water, water

temperature, environmental temperature, and

precipitation—that best explain the changes in

taxa composition under each sampling condi-

tion (Clarke & Warwick, 1994). Additionally,

Mantel tests were performed to assess whether

there is a correlation between each individual

environmental variable and the composition of

the taxa.

RESULTS

Mesofauna abundance: A total of 17 875

individuals were identified and classified in 46

taxa of mesofauna. The most abundant taxa

included Bivalvia (4 648 ind.), Nematoda (3 478

ind.), Gastropoda (2 642 ind.), Nemertea (2 612

ind.), and Ostracoda (1 466 ind.) (Fig. 1). A

complete list of organisms found in each sample

can be found in Appendix 1.

The total abundance of mesofauna was not

associated with the proportion of remaining

leaf litter mass in Site 1 during the dry season

(r = -0.3, P = 0.520) or in Site 2 during the rainy

season (r = -0.66, P = 0.136). However, in Site

1 during the rainy season (r = -0.94, P = 0.003)

and in Site 2 during the dry season (r = -0.94,

P = 0.003), there was a significant relationship

between mesofauna abundance and the propor-

tional loss of mass: a greater number of organ-

isms in the samples corresponded to a greater

loss of mass over the decomposition period.

Taxonomic composition: The mesofauna

composition varied significantly across sites,

showing a 41 % difference (ANOSIM, R = 0.41,

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e64043, enero-diciembre 2025 (Publicado Mar. 03, 2025)

P < 0.001). Additionally, the composition dif-

fered by season (dry or rainy), with a 16 %

variation noted (ANOSIM, R = 0.16, P < 0.001).

Samples from Site 1 in both seasons exhibited

greater similarity than those from Site 2. Nota-

bly, the mesofauna composition at Site 2 during

the rainy season was the most distinct when

compared to all other treatments (Fig. 2).

Significant differences in taxa composi-

tion, ranging from 14 % to 26 %, were observed

throughout the leaf litter decomposition pro-

cess (ANOSIM, R = 0.14 - 0.26, P < 0.001).

During the dry season at Site 1, although the

dates appeared relatively clustered on the MDS

plot (Fig. 2), significant differences in taxa

composition were found between different days

(P < 0.05), with a complete 100 % difference

observed between day 30 and day 120. In con-

trast, Site 2 exhibited a high level of differentia-

tion in mesofauna composition between days

during the dry season (Fig. 2).

Similarly, although the dates of the rainy

season at Site 1 were closer to one another com-

pared to other sites, all a posteriori comparisons

indicated significant changes in taxa composi-

tion. Conversely, at Site 2, only the comparisons

between days 90 and 120 showed significant

differences in mesofauna composition when

compared to earlier days (Fig. 2).

According to the SIMPER analysis, the taxa

that primarily contributed to the differences in

decomposition time and seasons were Bivalvia,

Nematoda, Gastropoda, Ostracoda, and Gam-

maridea. The populations of Gastropoda and

Gammaridea peaked at 60 days, while Nema-

toda reached its peak at 90 days. Bivalvia and

Ostracoda presented their highest abundance at

120 days (Table 1). Furthermore, Gastropoda,

Nematoda, Bivalvia, Ostracoda, and Harpacti-

coida were found to be more abundant in Site

Fig. 2. Multidimensional Scaling analysis (MDS),

showing the distance of similarity (Bray-Curtis) based on

composition of mesofauna (pooled samples) between days

of decomposition process, according to site and season.

TSNW, Puntarenas, Costa Rica.

Fig. 1. Number of individuals from the most abundant taxa found in leaf litter samples according to decomposition time and

season. TSNW, Puntarenas, Costa Rica.

6Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e64043, enero-diciembre 2025 (Publicado Mar. 03, 2025)

1 compared to Site 2 (Table 2). Additionally,

Bivalvia and Ostracoda were more prevalent

during the dry season, whereas Nematoda,

Gastropoda, and Gammaridea were more com-

mon during the rainy season (Table 3).

Composition by Trophic Groups: The

classification of taxa into different trophic

groups is displayed in Table 4. This classifica-

tion includes multiple trophic groups for the

same taxa to ensure that all potential repre-

sentatives are considered. The differentiation

in the composition of trophic groups between

sites was found to be 37 % (ANOSIM, R = 0.37,

P < 0.001). Additionally, the differentiation

in trophic group composition between the

dry and rainy seasons was 12 % (R = 0.12, P

< 0.001). In terms of decomposition time, the

differentiation ranged from 10 % to 18 % (R =

0.10-0.18, P < 0.001). Notably, the most signifi-

cant differences in the composition of trophic

groups occurred between the different sites.

The most abundant taxa, which were the

same that mainly contributed to differenc-

es for decomposition time, belonged to the

trophic groups: suspensor feeders (Bivalvia),

herbivore-carnivore (Nemertea, Gastropoda),

detritivore-herbivore (Ostracoda, Harpacticoi-

da, Gammaridea), and detritivore-herbivore-

carnivore-omnivore (Nematoda), respectively

Table 1

SIMPER test for the time factor, showing the five most influential taxa for composition differences and their mean abundance

between sampling decomposition days. TSNW, Puntarenas, Costa Rica.

Taxa Contribution to

difference (%)

Mean abundance

(15 days)

Mean abundance

(30 days)

Mean abundance

(60 days)

Mean abundance

(90 days)

Mean abundance

(120 days)

Bivalvia 10.68 5.1 13.6 57.3 46.2 110

Nematoda 9.73 5.5 10.9 21.4 79 57.1

Gastropoda 9.31 11.7 42.1 56 7.6 14.8

Ostracoda 7.11 2.95 6.2 4.35 6 53.8

Gammaridea 5.62 1.1 5.1 11.3 1.2 3.55

Table 3

SIMPER test for season factor, showing the five most influential taxa for composition differences and their mean abundance

between sampling decomposition days. TSNW, Puntarenas, Costa Rica.

Taxa Contribution to difference (%) Mean abundance for dry season Mean abundance for rainy season

Bivalvia 10.52 60.2 32.8

Nematoda 9.49 22.8 46.8

Gastropoda 9.38 21.8 31

Ostracoda 6.91 22.8 6.48

Gammaridea 5.52 1.98 6.94

Table 2

SIMPER test for site factor, showing the five most influential taxa for composition differences and their mean abundance

between sites and seasons. TSNW, Puntarenas, Costa Rica.

Taxa Contribution to difference (%) Mean abundance for Site 1 Mean abundance for Site 2

Gastropoda 11.69 51 1.84

Nematoda 10.7 63.7 5.86

Bivalvia 10.38 54 39

Ostracoda 7.32 11.3 18

Harparticoida 6.17 6.42 0.8

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e64043, enero-diciembre 2025 (Publicado Mar. 03, 2025)

(Table 5). The herbivore-carnivore and suspen-

sor feeders, were the groups that contributed

the most to the differences of mesofaunal com-

position for site and season (Table 6).

Association with environmental vari-

ables: Water salinity and precipitation were

notably different between seasons (Table 7).

Precipitation (Mantel, P = 0.610) and envi-

ronmental temperature (Mantel, P = 0.090) do

not influence mesofauna composition. Indi-

vidually, salinity (Mantel, P = 0.134), water

temperature (Mantel, P = 0.988) and dissolved

oxygen in water (Mantel, P = 0.331) fail to

explain the changes observed in mesofaunal

composition. However, together these three

variables can contribute in part to explain the

changes observed in mesofauna composition

in litter samples over time (BIO-ENV, r = 0.25,

P = 0.039).

Table 4

Classification by trophic groups for the taxa found in the samples and their mean total abundance according to season and

site. TSNW, Puntarenas, Costa Rica.

Site Season

TotalDry Rainy

Trophic group Taxa 1 2 1 2

Carnivore Hydrozoa; Platyhelminthes; Polychaeta:

Chrysopetallidae, Nereididae, Phyllodocidae,

Polynoidae, Syllidae, Halacaridae; Diptera:

Ceratopogonidae; Chordata: Gobiidae

214 213 291 84 802

Detritivore Kinorhyncha; Oligochaeta; Sipuncula; Polychaeta:

Ampharetidae, Maldanidae, Orbiniidae;

Amphipoda: Hyperiidea; Decapoda: Anomura;

Diptera: Chironomidae

54 51 109 10 224

Herbivore Tanaidacea; Insecta: Collembola, Lepidoptera 134 17 205 83 439

Omnivore Cnidaria: Actinaria; Decapoda: Alpheidae;

Diptera: Dolichopododidae

2 4 11 5 22

Sediment feeder Polychaeta: Capitellidae, Cirratulidae, Paraonidae 24 16 8 0 48

Suspensor feeder Polychaeta: Chaetopteridae; Cirripedia; Bivalvia 1 124 1 904 1 592 164 4 784

Detritivore, Herbivore Foraminifera; Ostracoda; Copepoda:

Harpacticoida; Amphipoda: Gammaroidea;

Insecta: Hydrophilidae

562 1 058 969 67 2 656

Herbivore, Carnivore Nemertea; Gastropoda 1 199 89 3 961 55 254

Herbivore, Omnivore Isopoda; Decapoda: Brachyura 8 19 51 63 141

Detritivore, Herbivorous, Carnivore, Omnivore Nematoda 932 207 2 253 86 3 478

Detritivore, Carnivore, Omnivore Polychaeta: Pilargidae 10001

Table 5

SIMPER test for time factor, showing the five most influential trophic groups for the difference composition and their mean

abundance between sampling dates. TSNW, Puntarenas, Costa Rica.

Trophic group

Difference

contribution

(%)

Mean

Abundance

(15 days)

Mean

Abundance

(30 days)

Mean

Abundance

(60 days)

Mean

Abundance

(90 days)

Mean

Abundance

(120 days)

Suspensor feeder 26.76 5.65 15.2 60 47.1 111

Herbivore, Carnivore 24.35 11.7 42.1 57 105 47

Detritivore, Herbivore, Carnivore, Omnivore 18.05 5.5 10.9 21.4 79 57.1

Detritivore, Herbivore 15.17 7.9 16.2 24.9 16.5 67.3

Carnivore 6.33 7.75 5.6 7.85 6.15 12.8

8Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e64043, enero-diciembre 2025 (Publicado Mar. 03, 2025)

DISCUSSION

Correlations done in this investigation

show that with a greater amount of mesofauna

in the litter, the mass losses are greater, depend-

ing on the site and the season. However, a more

specific identification of mesofauna must find

another possible relation for the treatments that

did not relate to mesofauna abundance (as in

the case of Site 1 during the dry season and Site

2 during the rainy season). Melo et al. (2013)

stated that mesofauna abundance and taxa

diversity could depend on freshwater discharge

from river flows, and, in this investigation, the

biggest difference in mesofaunal abundance

and composition was between sites, each one

located in a different river basin (Sierpe and

Grande de Térraba).

Regions of TSNW located in the Sierpe

basin, such as Estero Caballo (Site 1) tend to

have a major marine influence than regions

located in Grande de Térraba, such as Boca

Nueva (Site 2) (Picado, 2016). A combined

action of water salinity, water temperature, and

dissolved oxygen of water is necessary to achieve

25 % of the changes observed in the composi-

tion of mesofauna found in leaf litter samples

throughout the sampling time, and this fact

highlights a synergistic effect of these environ-

mental variables on mesofauna composition.

Other environmental variables not considered

in this study, such as flood frequency, type of

sediment, or microbial activity, can explain part

of the changes observed in mesofauna commu-

nity composition (Palit et al., 2022).

From the total number of individuals

quantified and identified throughout this study,

microbivalves were found to be the most abun-

dant taxa in juvenile or adult forms. Although

bivalves can be found adhering to the surface

of the leaves (Hogarth, 2007), they feed mainly

on suspended matter (Brusca & Brusca, 2005;

Hogarth, 2007). Water characteristics such as

salinity, dissolved oxygen, and temperature

influence the presence of bivalves and other

filter feeders on the mangrove litter (Vereycken

& Aldridge, 2023). The degradation of man-

grove leaves produces particulate organic mat-

ter, which serves as a crucial food source for

these filter feeders (Wang et al., 2015).

Table 6

SIMPER test for site and season factor, showing the five most influence trophic groups for the difference composition and

their mean abundance between sites and seasons. TSNW, Puntarenas, Costa Rica.

Trophic groups Difference

contribution (%)

Mean abundance per site Difference

contribution (%)

Mean abundance per season

Site 1 Site 2 Dry season Rainy season

Herbivore, Carnivore 30.91 103 1.88 24.83 25.8 79.3

Suspensor feeder 22.96 54.3 41.4 26.53 60.6 35.1

Detritivore, Herbivore,

Carnivore, Omnivore 19.71 63.7 5.86 17.64 22.8 46.8

Detritivore, Herbivore 14.41 30.6 22.5 15.15 32.4 20.7

Carnivore 4.95 10.1 5.94 6.40 8.54 7.5

Table 7

Mean value and standard deviation for the environmental parameters. TSNW, Puntarenas, Costa Rica.

Season Site Dissolved oxygen

(mg/l)

Water salinity

(ppm)

Water temperature

(ºC)

Precipitation

(mm)

Environmental

temperature (ºC)

Dry 14.46 ± 0.84 22.76 ± 2.29 29.83 ± 0.17 4.64 ± 1.08 26.72 ± 0.95

26.81 ± 0.96 27.39 ± 3.32 30.08 ± 0.33 4.64 ± 1.08 26.72 ± 0.95

Rainy 14.80 ± 0.29 9.62 ± 3.74 28.68 ± 0.76 11.01 ± 0.95 26.16 ± 0.14

26.84 ± 1.65 2.41 ± 1.85 27.10 ± 0.30 11.01 ± 0.95 26.16 ± 0.14

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e64043, enero-diciembre 2025 (Publicado Mar. 03, 2025)

The abundance of mesofauna in leaf litter

samples was linked to mass loss, and conse-

quently, to leaf litter decomposition at Site 1

during the rainy season and at Site 2 during the

dry season. This indicates that the decomposi-

tion of leaf litter could be influenced by the

presence of mesofauna, which varies according

to the specific conditions of each site and sea-

son. These findings are consistent with Powers

et al. (2009), who found that higher decompo-

sition rates occurred in leaf litter samples that

contained mesofauna.

In general, litter degradation tends to pro-

ceed more rapidly in tropical regions compared

to temperate environments. This pattern has

been attributed to lower biological activity from

invertebrates in temperate mangroves (D’Croz

et al., 1989; Gladstone-Gallagher, 2012). The

absence of mesofauna can reduce the decom-

position rate by as much as fifty percent (Pow-

ers et al., 2009). For the species R. racemosa in

TSNW, Loría-Naranjo et al. (2018) reported

a leaf litter decomposition rate of k = 0.012,

which aligns with other values found in tropical

regions and is relatively fast (Kulal et al., 2008).

However, it is important to note that excluding

certain functional groups of fauna using litter

mesh bags can significantly affect the decay rate

(Islam et al., 2024; Vinh et al., 2020).

Variations in the composition of meso-

fauna communities during different days of

leaf litter decomposition can be partially attrib-

uted to changes in specific organisms or condi-

tions within the leaves throughout this process.

Matheri et al. (2024) established that meso-

fauna communities respond significantly to the

duration of decomposition. Treplin & Zimmer

(2012) noted that crabs tend to prefer aged,

decaying leaves due to the decrease of solu-

ble leaf compounds over time, which means

these crabs do not necessarily play a crucial

role in leaf breakdown during the early stages

of decomposition.

In a mangrove ecosystem in southwestern

Puerto Rico, Torres-Pratts & Schizas (2007)

observed that the abundance of nematodes in

the litter increases progressively as decomposi-

tion advances. For the leaf litter of R. racemosa

in the TSNW, a significant abundance of nema-

todes was recorded at the end of the experi-

mental period during the dry season. A peak

in nematode abundance was observed after 90

days of decomposition during the rainy sea-

son. Many nematode species feed on the algae,

fungi, and bacteria that colonize decomposing

organic matter in estuarine habitats (Moens &

Vincx, 1997; Sánchez-Monge & Cortés, 2024).

Therefore, it is likely that the peak of nematodes

corresponds to an increase in these microor-

ganisms, which serve as food sources for the

nematodes present in the leaf litter.

There was also a noticeable seasonal change

in the mesofauna. Selviani et al. (2024) high-

lighted that climate and microclimate influ-

ence the presence of microbiota, which in turn

affects the decomposition rate of mangrove

litter. In this study, a total of 67 individuals of

Brachyura (crabs from various families) were

observed, with 57 of them identified during the

rainy season. However, the brachyuran crabs

were not among the most abundant taxa and

did not significantly contribute to the differ-

ences observed between samples. Crabs are

considered key organisms in the ecological pro-

cesses of mangrove forests, as they are known to

consume detritus from leaf litter, which influ-

ences leaf decomposition and nutrient cycling

(Hogarth, 2007; Jiménez, 1994; Lacerda, 2002).

These organisms play a vital role in the frag-

mentation of leaves by scraping the surface to

consume algae and other epibionts.

Based on the trophic classification of the

mesofauna present, it is expected that most

organisms are indirect consumers of R. rac-

emosa litter. For example, rest of fungi, diatoms

and bacteria have been found in the stomach

contents of springtails (Castaño-Meneses et

al., 2004), and nematodes. This suggests that

these organisms likely benefit from the decom-

position process of the litter by feeding on

their epibionts. On the decaying leaf litter of

R. racemosa, taxa of aquatic insects were also

observed, including Trichoptera, Lepidoptera

larvae and Diptera families (Chironomidae,

Ceratopogonidae and Dolichopodidae). Rincón

& Covich (2014) noted that Trichoptera species

10 Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e64043, enero-diciembre 2025 (Publicado Mar. 03, 2025)

Phylloicus pulchrus plays a significant role in

litter decomposition despite its low density, as

it scrapes the surface and consumes the fibrous

components of the leaves. Similarly, Lepidop-

tera larvae also consume leaves in mangrove

ecosystems (Lacerda, 2002). Diptera were the

most abundant insects found in the samples

analyzed in this study, which is consistent

with findings from other mangrove ecosystems

(Hogarth, 2007; Lacerda, 2002). Their mouth-

parts are adapted for licking or sucking other

plant fluids (Brusca & Brusca, 2005).

In this investigation, some of the most

abundant taxa found in leaf litter samples

belonged to trophic groups that consume

decayed leaves, including Nemertea, Gastrop-

oda, Ostracoda, Harpacticoida, Gammaridea,

and Nematoda. Among these, gastropods were

particularly prominent (Hogarth, 2007) and

represented the most abundant taxa in the leaf

litter samples. Although some gastropods feed

directly on leaf litter, such as Terebralia palustris

(Potaminidae), which primarily consumes leaf

litter and is one of the most significant leaf litter

removers in mangrove forests (Hogarth, 2007),

Melo et al. (2013) also identified another small

gastropod, Olivella minuta, as one of the most

abundant taxa of benthic macrofauna in Brazil-

ian mangroves.

Thirteen polychaete families were identi-

fied (Ampharetidae, Capitellidae, Cirratulidae,

Chaetopteridae, Chrysopetallidae, Maldanidae,

Nereididae, Orbiniidae, Paraonidae, Pilargidae,

Phyllodocidae, Polynoidea, Sylidae). Families

Capitellidae, Chaetopteridae, Nereididae, Orbi-

niidae and Phyllodocidae in previous studies

on polychaete communities in mangroves of

tropical estuaries were described as herbivores

or detritivores (López et al., 2002; Sarkar et

al., 2005). The family Nereididae (many of its

representatives are carnivorous or omnivorous)

was the largest among the polychaete fami-

lies found in the litter samples of R. racemosa

in TSNW. Although López et al. (2002) and

Sarkar et al. (2005) studied the community

of polychaetes present in the sediment of the

mangrove, these organisms can also act as her-

bivores or detritivores, turning into potential

litter consumers, and therefore into collabora-

tors of the decomposition process.

Ostracods are microcrustaceans that rank

among the five most abundant taxa found in

the litter of R. racemosa. Vargas (1987) noted

that ostracods are one of the most prevalent

groups of benthic fauna in the mangrove for-

est of Punta Morales, Costa Rica. In Puerto

Rico, however, ostracods have been reported to

occur in low quantities in the litter of R. mangle

(Torres-Pratts & Schizas, 2007).

Copepods from the order Harpacticoida

were not among the most abundant taxa, but

they significantly contributed to the differ-

ences in composition between the sampling

sites. Harpacticoid copepods, particularly those

from the Darcythomsoniidae family, are known

for their exclusive consumption of mangrove

litter (Nagelkerken et al., 2008). In this study,

Harpacticoida were classified as detritivores

and herbivores; however, they were not linked

to the rate of leaf litter decomposition.

There are notable changes in the com-

position of the mesofauna community that

colonizes the samples over the decomposition

period, as well as differences observed between

sites and seasons. However, these changes do

not appear to be directly linked to the loss of

litter mass during decomposition, regardless of

the site or season. Consequently, the diversity

of taxa and trophic groups found in the samples

does not have a direct relationship with the leaf

litter decomposition rate. Instead of the varia-

tions in mesofauna composition, it is the abun-

dance of this mesofauna that influences the leaf

litter decomposition of R. racemosa.

The study of mesofauna presence dur-

ing the leaf litter decomposition process of R.

racemosa in TSNW mangrove evidences an

abundant mesofauna community composed of

diverse taxa that still needs deeper study efforts

to achieve their total role in nutrient recycling,

soil formation, primary productivity, and ecol-

ogy of mangrove forests. The increase in the

comprehension of these processes is vital for

the conservation and sustainability of the eco-

system. Therefore, the research addressed here

represents a contribution to this understanding.

Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e64043, enero-diciembre 2025 (Publicado Mar. 03, 2025)

Ethical statement: the authors declare that

they all agree with this publication and made

significant contributions; that there is no con-

flict of interest of any kind; and that we fol-

lowed all pertinent ethical and legal procedures

and requirements. All financial sources are fully

and clearly stated in the acknowledgments sec-

tion. A signed document has been filed in the

journal archives.

See supplementary material

a11v73s1-suppl1

ACKNOWLEDGMENTS

Authors thank Jorge Picado, Carlos and

Leo of the Biology Team of the Instituto Costar-

ricense de Electricidad (ICE) at Buenos Aires,

Puntarenas, for their support in the field. To

Álvaro Morales and Emma Segura of the Cen-

tro de Investigación en Ciencias del Mar y

Limnología (CIMAR) at the University of Costa

Rica, for administrative support. To the Insti-

tuto Metereológico Nacional (IMN) which pro-

vided the temperature and rainfall data. Thanks

to Juan Carlos Azofeifa, Marco Corrales, Vik-

toria Bogantes, Rita Vargas, Paola Campos and

Kimberly García who helped with organism’s

quantification and identification.

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