Burrowing crabs as potential bioindicators of sediment carbon storage in mangroves of the Colombian Pacific with different degrees of anthropogenic disturbance.

Burrowing crabs as potential bioindicators of sediment carbon storage in mangroves of the Colombian Pacific with different degrees of anthropogenic disturbance. | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Burrowing crabs as potential bioindicators of sediment carbon storage in mangroves of the Colombian Pacific with different degrees of anthropogenic disturbance. LUIS ALEJANDRO SANDOVAL, DALLIAM FELIPE CHACÓN, ADRIANA MARTÍNEZ, and 2 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-7095077/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Little is known about the indicator potential of sediment carbon storage of mangrove crabs (as ecosystem engineers) for blue carbon sequestration assessment. This study aimed to examine the crab trophic guild’s structure and composition in relation to mangroves with different degrees of anthropogenic disturbance index and climate seasons in the Colombian Pacific mangrove forests. Crab trophic guilds were also linked to soil organic carbon content (SOC) and other soil environmental variables in each ecosystem. Ocypodids and deposit feeders were the most representative crabs in terms of species numbers. Mangrove forests with the highest anthropogenic intervention (Pingüita, San Pedro y Rompido) have a high abundance of the deposit feeders, Uca spp., Minuca spp., and Omnivores species, as well as the lowest SOC values. Meanwhile, forests with low and moderate intervention (El Morro and Bocagrande) have a high abundance of deposit feeders, Leptuca spp., and the highest SOC. Crab burrows's abundance of Leptuca spp. could be good bioindicators of higher sediment carbon storage and mangrove ecosystem quality, whereas Uca spp., Minuca spp., and Omnivores species could be used as potential indicators for disturbed systems and lower sediment carbon storage. Macrobenthos ecosystem engineers anthropogenic intervention trophic guild Uca Leptuca Minuca Figures Figure 1 Figure 2 Figure 3 Introduction 'Blue Carbon' refers to the carbon captured by the world's ocean or coastal vegetated ecosystems. Since the term was coined about a decade ago (McLeod et al. 2011 ), our knowledge about blue carbon sequestration has rapidly evolved. In mangrove forests, sequestration rates and standing C stocks are exceptionally high and exceed those of comparable terrestrial ecosystems (e.g., tropical rainforests) (Donato et al. 2011 ; Mcleod et al. 2011 ). There are local and global scale estimates of sediment carbon in mangrove forests (e.i., Atwood et al. 2017; Jennerjahn 2020 ; Ouyang and Lee 2020 ; Wang et al. 2021 ). Thus, the potential of mangrove forests to sequester and store large amounts of carbon has been relatively well documented (Santos et al. 2021 ). Nevertheless, the role of the macrobenthos, as ecosystem engineers, in sediment biogeochemical cycles is still inconclusive (Qiu et al. 2019 ). For instance, crabs have been considered good indicators of the environmental habitat quality of mangrove ecosystems (e.i., Amaral et al. 2009 ; Retnaningdyah et al. 2022 ) but little is known about the indicator potential of sediment carbon storage of mangrove crabs for blue carbon sequestration assessment. Ecosystem engineers are organisms that directly or indirectly affect other organisms' composition and critical biological processes by changing abiotic environments (e.g., Jones et al. 1994 ; Wright and Jones 2004 ). Sesarmid and ocypodid crabs are the mangrove fauna's major bioturbating and bioengineering components through their feeding and digging activities (Cannicci et al. 2008 ; Kristensen et al. 2008 ; Lee 1998 ). They can play an essential role in retaining organic matter (OM), as some species drag leaf litter and propagules into their burrows, restricting the export through tidal flushing (Ashton 2002 ; Lee 2008 ). Through their burrow, other species enhance oxygen flux into the waterlogged sediment, facilitating oxidation and enhancing nutrient availability (Kristensen and Alongi 2006 ). Sesarmids are herbivorous crabs that can consume up to 81% of mangrove litter production (Nordhaus et al. 2006 ). They can dig burrows down to 2 meters in depth and store leaf litter, mangrove propagules, and other organic material from different sources, enhancing the retention of OC within the forest (Kristensen et al. 2008 ; Andreetta et al. 2013). Conversely, a recent study showed that the sesarmid crab Ucides cordatus feeding and digging activities would not increase the organic matter contents in the sediments since their burrows would expose deeper parts of the sediment to aerobic conditions, thus facilitating the decomposition of initially preserved organic matter (Katzer 2023 ). In this way, the differences in OM content between mangrove forests could be explained by hydro-geomorphic differences and less by crab or crab burrow-related effects. This includes the tidal amplitude, implications, and riverine influences (Bouillon et al. 2003 ). Ocypodid crabs are filter feeders that can exploit food sources from the sediment, such as bacteria, benthic microalgae, and meiofauna (Reinsel 2004 ). As filter feeders, they feed on the sediment surface, never burying organic material, as typical for sesarmids (Andreetta et al. 2013). Burrows of the pantropical genus Uca are the most abundant and conspicuous, and they are typically less deep than sesarmids (Michaels and Zieman 2013 ). Some studies have shown that the presence of Uca burrows aerates sediments, thus facilitating the decomposition of OM (e.i., Katz 1980 ; Montague 1982 ). Conversely, an experimental study showed that their burrows have little effect on surrounding sediment oxygen concentrations. Thus, the differences in the decomposition of OM would depend significantly on the characteristics of the sediment in which the burrows are located (Michael and Zieman 2013). Likewise, a recent study showed that crab burrowing could trap and intercept detritus, consequently promoting the retention and accumulation of soil carbon instead of facilitating the oxidation of OM (Qiu et al. 2019 ). Mangroves of the Tropical Eastern Pacific (TEP) are the second-largest mangrove area in the Neotropics (FAO 2007 ). They are prevalent coastal ecosystems along the Pacific coasts of Colombia, with several structural features and service provisions that make them important regionally and globally (Castellanos et al. 2020). As part of the Eastern Pacific Basin, this area is affected by a large tidal range and is one of the rainiest regions of the world; this set of conditions converges in mangrove forests with unique characteristics (Blanco and Cantera 2001; Medina-Contreras et al. 2022 ). However, large mangrove areas in the Colombian Pacific remain poorly studied (Castellanos et al. 2020), and they could be affected by multiple anthropogenic stressors such as deforestation; wood production for fuel, coal, and construction materials; urban and industrial development (Palacios et al. 2019; Rodríguez-Rodríguez et al. 2021 ; FAO 2023 ). Recently, Gómez-García, et al. ( 2024 ), found a relationship between the degree of anthropogenic intervention and carbon reserves in deltaic mangroves of the region, where systems with the highest anthropogenic disturbance (measured through the anthropogenic disturbance index) registered the lowest values in blue carbon. In this study, field investigations were conducted to examine the link of burrowing crabs (as key bioturbators) to soil organic matter dynamics in mangrove forests of the Colombian Pacific with different anthropogenic and environmental stressors. We used the different disturbance degrees in mangroves of the Colombian Pacific by Gómez-García et al. ( 2024 ), and grouped burrowing crabs by trophic guilds to link crabs’ abundance and soil organic carbon content (SOC) in each ecosystem. The abundance of some crab trophic guilds was expected to correlate with the SOC to evaluate crabs as bioindicators of sediment carbon storage. It was also expected that mangrove forests with fewer anthropogenic stressors would have higher SOC concentrations, as shown recently by Gómez-García et al. ( 2024 ). Material and methods Study area The Pacific Coast of Colombia is one of the wettest regions in America (mean humidity between 88 and 100%) with one of the highest annual precipitations for the coastal areas of the Western Hemisphere (precipitation > 500 cm yr − 1 ), with a wet period from January to April and a very wet period from May to December (Castellanos-Galindo et al. 2013 ). Temperatures above 24°C. (Rangel and Arellano 2004 ), and average air temperature of 25°C (Cantera and Blanco 2001). Five mangrove systems (Fig. 1 ) with different levels of anthropogenic intervention were selected: (1) El Morro, a near pristine mangrove located in the internal zone of the Bahía Málaga estuary (4°02'58 " N, 77°11'28" W), where there is a Natural National Park and low anthropogenic disturbance index (ADI = 0) (Gómez-García et al. 2024 ). (2) San Pedro (03° 50′ 11" N, 77° 15′ 30" W) and (3) Piangüita (03° 50′ 32" N, 77° 12′ 23" W) mangroves within Buenaventura Bay, which are near to the main port on the Colombian and have a high anthropogenic disturbance index (ADI = 5 y 8, respectively) (Gómez-García et al. 2024 ). (4) Bocagrande (1°47'30"N, 8°52'18"W), and (5) Rompido (1°48'45"N, 78°50'16"W) mangroves set south of Tumaco Bay, which are near to Tumaco municipal (200000 inhabitants) and have a moderate level (ADI = 2) and a high anthropogenic intervention (ADI = 7), respectively (Gómez-García et al. 2024 ). Relevant characteristics of the five systems are shown in Table 1 . Table 1 Main features of the five mangrove forests studied in the Colombian pacific coast. Data taken from Gómez-García et al. ( 2024 ). * Unpublish data. Feature/sample site El Morro San Pedro Piangüita Bocagrande Rompido Anthropogenic disturbance index (ADI) 0 5 8 2 7 Dominant mangrove species Rhizophora mangle Mora oleifera Pelliciera rhizophorae Rhizophora mangle Laguncularia racemosa Mora oleifera Pelliciera rhizophorae Rhizophora mangle Pelliciera rhizophorae Laguncularia racemosa Rhizophora mangle Pelliciera rhizophorae Rhizophora mangle Avicennia germinans Abundance (ind/ha) 783.3 ± 110.80 1283.3 ± 492.27 1333.3 ± 320.06 500.0 ± 112.54 983.3 ± 110.80 Basal Area (m 2 /ha) 27.36 ± 8.01 15.08 ± 228 1620 ± 2,93 43.85 ± 18.93 1540 ± 179 Diameter at breast height (cm) 17.02 ± 1.91 12.95 ± 0.92 10.91 ± 0.88 29.83 ± 3.07 13.47 ± 0.79 *Apparent density (g/cm 3 ) 0.4 0.9 0.6 0.7 0.9 Field sampling Sampling took place in 2023. The density of crabs was estimated during low tides during both the rainy and less rainy seasons. In each of the five different study ecosystems, five 2 × 2 m random quadrats were sampled to assess the density of crab populations. Individuals were first counted visually at each site to assess the relative frequencies of the populations present in the quadrat (Skov and Hartnoll 2001 ; Cannicci et al. 2009 ). Then, crabs were collected manually to avoid underestimating specimens not active on the surface during the visual crab counts. Samples were collected from the selected quadrats using a Russian drill hole soil sampler (0–50 cm) to determine soil components and organic carbon content. The samples were analyzed to determine the percentages of sand, silt, clay, bulk density, and organic carbon (SOC). Temperature, pH, and conductivity from interstitial water were registered in situ using a soil pH-moisture meter. Soil analyses Bulk samples were air-dried, sieved (< 2 mm) and homogenized. The Bouyoucos-hydrometer method was used to analyze particle size (Bouyoucos, 1962 ). To determine soil bulk density, samples with a known volume metal corer were oven-dried at 60°C for 48 h and weighed. The bulk density was calculated as the ratio of the dry mass of the soil sample to its volume. Soil organic carbon was determined by the Walkley and Black method (Andreetta et al. 2014 ). Soil organic C stored in the soil was calculated by the following formula: $$\:{SOC}_{tot}={\sum\:}_{1}^{n}(OC\:\times\:\:DB\:\times\:\:Dh)$$ where SOC is the soil organic carbon stock (kg m − 2 ), OC is the organic carbon content for each depth interval (mg g − 1), BD is the bulk density (g cm − 3 ), and Dh is the soil thickness interval. Statistical analysis Crabs were classified into five trophic categories: herbivores, omnivores, carnivores, deposit feeders, and filter feeders. Feeding guilds were assigned according to information found in local literature (Table 2 ). Before multivariate analysis, trophic guild assemblage data were transformed by the 4th root to reduce the influence of abundant species relative to rare ones (Clarke and Warwick 2001). With these data, Bray-Curtis similarity matrices were calculated and used as input for non-parametric permutational multivariate analysis of variances (PERMANOVA) to test for ecosystem and season differences in crab community structure and composition. The P-values were then obtained using 9999 permutations. A posteriori pairwise comparison with the PERMANOVA t-statistics was conducted to elucidate spatial differences within mangrove ecosystems. To reveal the relationships between the abundance of crab trophic guilds and soil properties, a Canonical Correspondence Analysis (CCA) was carried out. Finally, we performed a Spearman rank correlation (SRC) analysis of the abundance of crab trophic guilds and soil properties between explanatory variables to demonstrate correlations. Table 2 Crabs’ species from mangrove forests of the Colombian Pacific and their trophic guild regarding local sources. Family/Subfamily Species Trophic guild Abbreviation Source Ocypodidae Rafinesque, 1815 Ocypodinae Rafinesque, 1815 Uca sp . Latreille, 1819 Deposit feeders DEP1 Medina-Contreras et al. 2022 Uca (Petruca) panamensis (Stimpson, 1859) Uca heteropleura (Smith, 1870) Uca stylifera (H. Milne Edwards, 1852) Gelasiminae Miers, 1886 Leptuca sp Bott, 1973 Deposit feeders DEP2 Medina-Contreras et al. 2020 Leptuca inaequalis (Rathbun, 1935) Leptuca tenuipedis (Crane, 1941) Leptuca batuenta (Crane, 1941) Leptuca tomentosa (Crane, 1941) Minuca sp . Bott, 1954 Deposit feeders DEP3 Minuca umbratila (Crane, 1941) Minuca brevifron s (Stimpson, 1860) Ucidinae Števcic, 2005 Ucides occidentalis (Ortmann, 1897) Herbivores HER Medina-Contreras et al. 2020 Grapsidae MacLeay, 1838 Pachygrapsus sp . Randall, 1840 Omnivores OMN Medina-Contreras et al. 2020 Pachygrapsus socius Stimpson, 1871 Goniopsis pulchra (Lockington, 1877) Panopeidae Ortmann, 1893 Panopeus sp H. Milne Edwards, 1834 Carnivores CAR Medina-Contreras et al. 2020 Panopeus chilensis H. Milne Edwards & Lucas, 1843 Panopeus purpureus Lockington, 1877 Eurypanopeus transversus (Stimpson, 1860) Eurypanopeus planus (Smith, 1869) Porcellanidae Haworth, 1825 Petrolisthes zacae Haig, 1968 Filter feeders FIL Medina-Contreras et al. 2020 Results In total, 22 species from 4 families were captured (Table 2 ). The most dominant families in terms of species numbers per family were Ocypodidae (12 spp.) and Panopeidae (5 spp.). Deposit feeders were the most representative guild in terms of species numbers, which were grouped by gender at Deposit feeders 1 ( Uca spp.), Deposit feeders 2 ( Leptuca spp.), and Deposit feeders 3 ( Minuca spp.) (Table 2 ). PERMANOVA analysis revealed significant differences in crab community structure between mangrove forests (p 0.05). Posteriori pairwise comparisons did not show differences between San Pedro and Piangüita and between Rompido and Piangüita, ecosystems with a high ADI (Table 3 ). Table 3 Non-parametric pairwise comparisons of 4th-root transformed trophic guilds crabs’ assemblage. In bold significant differences. San Pedro Pingüita El Morro Bocagrade Rompido San Pedro 0.0903 0.0076 0.0167 0.0076 Piangüita 0.0071 0.0145 0.0527 El Morro 0.0075 0.0077 Bocagrande 0.0075 Rompido Deposit feeder 1 ( Uca spp .) and 3 ( Minuca spp .) were the most abundant species in San Pedro; deposit feeder 2 ( Leptuca spp .) were the most abundant species in El Morro; carnivores were the most abundant species in Rompido (Table 2 , Fig. 2 ). There was little abundance of herbivores and filters; for this reason, former trophic guilds were excluded from the CCA. The canonical correlation describes the principal tendencies in the relationship between crabs' trophic guilds and their environment (Fig. 3 ). The projections of a species on this axis show its preference for high or low values of this environmental gradient. The first and second ordination axes represent this variation best and showed eigenvalues of 0.116 and 0.035, respectively, which amounted to 71.2% and 21.6% of the variation in crab abundance (Table 4 ). Axis 1 separated the ecosystems with a low and moderate anthropogenic disturbance indexon the right side, which are characterized by the highest SOC, clay, BD, and Cond, in opposition to the ecosystems with a high ADI on the left side, with lowest SOC, clay, BD and Cond, and higher sand characterize. The DEP 2 ( Leptuca spp.) strongly correlated to sites in the middle estuary ecosystems with low and moderate ADI. The DEP 1 ( Uca spp.), DEP 3 ( Minuca spp.), and Omnivores species are more closely related to sites with a high ADI (Fig. 3 ). Table 4 Summary of canonical correspondence analysis for abundance data of crab’s trophic guilds with mangrove ecosystems and soil environmental variables. Axis Eigenvalue % Axis Eigenvalue p 1 0.11597 71.38 1 0.116 0.001 2 0.035053 21.58 2 0.03505 0.037 3 0.011023 6.785 3 0.01102 0.120 4 0.00042 0.2585 4 0.00042 0.721 Since DEP 1 and 3 showed similar correlations with the soil environmental variable, they were analyzed as a group at the SRC analysis. Thus, DEP 1 + 3 ( Uca spp . + Minuca spp. ) showed a significant and negative correlation with SOC and a positive correlation with sand (Table 5 ). Omnivores were negatively associated with SOC and clay, while DEP 2 ( Leptuca spp. ) was positively associated with SOC and clay (Table 5 ). Table 5 Spearman’s rank correlation matrix for abundance data of crab’s trophic guilds with soil environmental variables. In bold significant result. SOC Sand Clay Leptuca spp (DEP2) DEP 1 + 3 Om SOC 1.97E-05 9.39E-08 3.26E-05 0.023937 0.01842 Sand -0.74462 0.00014203 0.0023252 0.032543 0.085356 Clay 0.84692 -0.68846 0.0011255 0.054947 0.021035 Leptuca spp (DEP2) 0.7315 -0.58095 0.61287 0.74411 0.034515 ADD 1 + 3 -0.45018 0.4286 -0.38852 -0.068724 0.59903 Om -0.46761 0.35101 -0.45889 -0.42431 -0.11049 Table 6 Crab contribution to sediment carbon storage of four mangrove systems compared. Authors Country Approach Analysis Trophic guild/ Group assemblage Crab species Crab contribution Proportion of contribution Andreetta et al. 2014 Kenya Relationships between crab assemblage and SOC Distance-based linear model Deposit feeders Uca annulipes, Uca inversa, Uca chlorophthalmus, Uca urvillei Increase 0,55 Burrowing Sesarmid Neosarmatium africanum, Neosarmatium smithi Increase 0,56 Non-Burrowing Sesarmid Chiromantes ortmanni, Chiromantes eulimene, Perisesarma guttatum , Parasesarma leptosoma Increase 0,42 Katzer, 2023 Colombia Field experiment with crab-inhabited and field investigations crab-free burrows Kruskal-Wallis test Burrowing Sesarmid Ucides cordatus Decrease 0,14 Qiu et al. 2019 China Comparison between areas with few or no crab burrows and areas with high density crab burrows One-way analysis of variance Burrowing crabs, mainly herbivorous Helice tientsinensis as the dominating species Increase N.A This study Colombia Correlation between abundance data of crab’s trophic guilds with soil environmental variables Spearman’s rank Deposit feeders Leptuca spp. Increase 0,73 Deposit feeders Uca spp., Minuca spp. Decrease -0,45 Omnivores Pachygrapsus spp., Goniopsis pulchra Decrease -0,47 Discussion This study compared the crab trophic guild’s structure and composition in relation to mangroves with different degrees of anthropogenic intervention levels and climate seasons. The systems included a near pristine mangrove (El Morro) with a low anthropogenic intervention level, three sites with a high level (San Pedro, Piangüita, and Rompido), and a system with a moderate level (Bocagrande). These systems also have differences regarding soil apparent density, reenforcing anthropogenic intervention levels. Crab trophic guilds were also linked to soil organic carbon content (SOC) in each ecosystem, showing they can be used as potential bioindicators for evaluating blue carbon sequestration as discussed below. Mangrove habitats generally have lower species richness of macrobenthos (e.g., 17–85 species) than adjoining habitats such as seagrass meadows and open sand/mudflats. However, brachyuran crabs constitute one of the most diverse invertebrate groups in the mangrove ecosystem (Lee 2008 ). Crab richness in the mangrove forests of the Colombian Pacific (22 species from 4 families) was lower than in mangroves of the Indo-Pacific region, where there is higher mangrove species richness, such as the Indian coast, where the East coast hosts 127 crab species and the West Coast 83 species (Sathish et al. 2023 ). Crab species richness in the Colombian Pacific would be higher than in the Colombian Caribbean coast, where, for instance, Sandoval (2023) found a total of 10 macrobenthos species. Higher mangrove species richness and organic carbon availability in the soil have been positively correlated with higher crab species richness and abundance (Sathish et al. 2023 ). The environmental conditions, such as salinity, depth, and sea surface temperature, are relevant drivers of the distribution of mangrove crabs worldwide (Sharifian et al. 2021 ). The abundance of crab species belonging to the Ocypodidae family is relatively high in the Colombian Pacific, as in other mangrove ecosystems (Lee 2008 ). Deposit feeders were the most representative trophic guild in terms of species numbers, most of them belonging to the Ocypodidae family. There was no difference in crab community structure between rainy seasons. This would be explained because this region has one of the highest annual precipitations for the coastal areas of the western hemisphere, where there is a rainy season (May to December, average 746 mm/month) and the less rainy season (January to April, average 422 mm7month) (Castellanos-Galindo et al. 2013 ). However, we found significant differences in crab community structure between mangrove forests, where San Pedro and Piangüita, ecosystems with the highest ADI, did not show differences in their community structure but showed remarkable differences regarding the other three forests (see below), which suggests that anthropogenic intervention conditions would drive the distribution of mangrove crabs in this region. The canonical correlation describes the principal trends in the relationship between crab’s trophic guilds and their environment, supporting that their distribution would be linked to anthropogenic intervention. Axis 1 separated the ecosystems on the right side with low and moderate anthropogenic intervention levels (El Morro and Bocagrande). On the left, it separated the ecosystems with the highest intervention (Pingüita, San Pedro, and Rompido). Former ecosystems have a high abundance of Uca spp. (DEP 1), Minuca spp. (DEP 3), and omnivore species, as well as the lowest SOC values. While El Morro and Bocagrande have a high abundance of Leptuca spp. (DEP 2) and the highest SOC. Which suggests that Leptuca spp. would be a good bio-indicator of higher sediment carbon storage and the mangrove ecosystem's quality in the Colombian Pacific mangrove ecosystems. Whereas Uca spp., Minuca spp., and omnivores crabs could be used as potential indicators for disturbed systems and lower sediment carbon storage supporting our hypothesis. The above would be explained since mangrove crabs are sensitive to changes in mangrove cover area and changes in their community structure are useful tools to detect mangrove habitat status (Sathish et al. 2023 ). Some studies have shown that mangrove degradation negatively affects crab biodiversity and communities’ structure (e.i., Ashton et al. 2003 ; Maia and Coutinho 2013 ). Here, we showed significant differences in crab trophic guild’s structure and composition in relation to mangroves with different degrees of anthropogenic intervention. Likewise, Leptuca spp. crabs, dominated in low and moderate anthropogenic intervention levels forests (with the highest SOC values), whereas Uca spp., Minuca spp., and omnivores crabs were more abundant in the degraded forests (with lowest SOC values). The field investigation indicated that mangrove soils in the region vary considerably according to anthropogenic intervention as showed recently by Gómez-García et al. ( 2024 ). Our results suggest that ecosystems with high mangrove degradation, such as San Pedro and Pingüita within Buenaventura Bay, which have been intensively degraded for domestic use, such as firewood and house building (Palacios and Cantera 2017 ), have lost their whole capacity for soil carbon storage. Uca spp., Minuca spp., and omnivores crabs could be used as potential indicators for these disturbed systems. This is supported by findings of other studies that have reported ocypodid crabs, mainly Uca species, dominating in disturbed, degraded, or young rehabilitation sites (Macintosh et al. 2002 ; Ashton et al. 2003 ). The effect of burrowing crabs on organic carbon storage varies according to anthropogenic intervention. In ecosystems with a high mangrove degradation, crab burrows ( Uca spp., Minuca spp. and omnivore species) could expose sediment parts to aerobic conditions, thus facilitating the decomposition of initially preserved organic matter, as found in another study (Katzer, 2023 ). To explain this, we can hypothesize that crab burrows in ecosystems with high mangrove degradation could not trap enough organic matter to improve the contents of SOC since they have less carbon accumulated in aerial biomass (Gómez-García et al. 2024 ). Consequently, they have a smaller amount of detritus available. Conversely, in crab burrows of Leptuca spp. in non-degraded systems, detritus could be trapped and intercepted, especially plant detritus, which could be an important mechanism for promoting carbon retention and accumulation, consistent with other studies (Table 5 ). However, we acknowledge that other factors as vegetation and inundation times are recognized as controls of SOC in mangrove ecosystems (Andreetta et al. 2014 ). Therefore, a more detailed understanding of the influence of burrowing crabs on SOC is required. In summary, the most dominant family in species numbers per family was Ocypodidae. Deposit feeders were the most representative trophic guild. The field investigation indicated that anthropogenic intervention conditions would drive the distribution of mangrove crabs in this region. Likewise, the effect of burrowing crabs on organic carbon storage varies according to anthropogenic intervention. Ecosystems with the highest anthropogenic intervention (Pingüita, San Pedro y Rompido) have a high abundance of Uca spp., Minuca spp., and omnivores species and the lowest SOC values. Meanwhile, systems with low and moderate intervention (El Morro and Bocagrande) have a high abundance of Leptuca spp. and the highest SOC. Burrowing crabs' abundance could be a good bioindicator of sediment carbon storage and the mangrove ecosystem's quality on the Colombian Pacific coasts. Declarations This study was funded by the Sistema General de Regalías (FCTel-SGR: BPIN 2020000100054). Ethics and Consent to Participate declarations: not applicable. Author Contribution SLA, MJE and GAI developed the concept and designed the study. CDF, MA and GAI conducted the field study. SLA, MJE and GAI analysed the data. SLA wrote the original draft of the manuscript. SLA, MJE and GAI interpreted results. All authors discussed the results, commented on and improved various versions of the manuscript. 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J Sea Res 59:16–29. https://doi.org/10.1016/j.seares.2007.05.002 Lee SY (1998) Ecological role of grapsid crabs in mangrove ecosystems: a review. Mar Freshw Res 49:335–343 Macintosh DJ, Ashton EC, Havanon S (2002) Mangrove rehabilitation and intertidal biodiversity: a study in the Ranong mangrove ecosystem, Thailand. Est Coast Shelf Sci 55:331–345. https://doi.org/10.1006/ecss.2001.0896 McLeod E, Chmura GL, Bouillon S, Salm R, Björk M, Duarte CM, Lovelock CE, Schlesinger WH, Silliman BR (2011) A blueprint for blue carbon: toward an improved understanding of the role of vegetated coastal habitats in sequestering CO2. Front Ecol Environ 9:552–560. https://doi.org/10.1890/110004 Maia RC, Coutinho R (2013) The influence of mangrove structure on the spatial distribution of Melampus coffeus (Gastropoda: Ellobiidae) in Brazilian estuaries. Pan-Am J Aquat Sci 8:21–29 Medina-Contreras D, Arenas F, Cantera-Kintz J, Sánchez A, Lázarus JF (2022) Carbon sources supporting macrobenthic crustaceans in tropical eastern pacific mangroves. Food Webs 30. https://doi.org/10.1016/j.fooweb.2022.e00219 Michaels RA, Zieman JC (2013) Fiddler crab (Uca spp.) burrows have little effect on surrounding sediment oxygen concentrations. J Exp Mar Biol Ecol 448:104–113. http://dx.doi.org/10.1016/j.jembe.2013.06.020 Montague CL (1982) The influence of fiddler crab burrows and burrowing on metabolic processes in salt marsh sediments. In: Kennedy VS (ed) Estuarine Comparisons. Academic, New York, pp 283–301 Nellemann C, Corcoran E, Duarte C, Valdes L, Young C, Fonseca L, Grimsditch G (2009) Blue Carbon - the Role of Healthy Oceans in Binding Carbon. UNEP, Arendal Nordhaus I, Wolff M, Diele K (2006) Litter processing and population food intake of the mangrove crab Ucides cordatus in a high intertidal forest in northern Brazil. Est Coast Shelf Sci 67:239–250. https://doi.org/10.1016/j.ecss.2005.11.022 Ouyang X, Lee SY (2020) Improved estimates on global carbon stock and carbon pools in tidal wetlands. Nat Commun 11:317. https://doi.org/10.1038/s41467-019-14120-2 Palacios ML, Cantera JR (2017) Mangrove timber use as an ecosystem service in the Colombian Pacific. Hydrobiologia 803:345–358. https://doi.org/10.1007/s10750-017-3309-x Palacios-Peñaranda ML, Cantera JR, Peña-Salamanca EJ (2019) Carbon stocks in mangrove forests of the Colombian Pacific. Est Coast Shelf Sci 227:106299. https://doi.org/10.1016/j.ecss.2019.106299 Qiu D, Cui B, Yan J, Ma X, Ning Z, Wang F, Sui H, Bai J (2019) Effect of burrowing crabs on retention and accumulation of soil carbon and nitrogen in an intertidal salt marsh. J Sea Res 154:101808. https://doi.org/10.1016/j.seares.2019.101808 Rangel O, Arellano H (2004) Clima del Chocó biogeográfico/costa pacífica de Colombia. Colombia Diversidad Biótica IV: El Chocó Biogeográfico/Costa Pacífica. Universidad Nacional de Colombia Reinsel KA (2004) Impact of fiddler crab foraging and tidal inundation on an inter-tidal sandflat: season-dependent effects in one tidal cycle. J Exp Mar Biol Ecol 313:1–17. https://doi.org/10.1016/j.jembe.2004.06.003 Retnaningdyah C, Febriansyah SC, Hakim L (2022) Evaluation of the quality of mangrove ecosystems using macrozoobenthos as bioindicators in the Southern Coast of East Java, Indonesia. Biodivers J 23:1247. https://doi.org/10.13057/biodiv/d231247 Rodríguez-Rodríguez JA, Mancera-Pineda JE, Tavera H (2021) Mangrove restoration in Colombia: Trends and lessons learned. Ecol Manag 496:119414. https://doi.org/10.1016/j.foreco Sanders CJ, Santos I, Steven ADL, Lovelock CE (2017) Global patterns in mangrove soil carbon stocks and losses. Nat Clim Change 7:523. https://doi.org/10.1038/nclimate3326 Santos IR, Burdige DJ, Jennerjahn TC, Bouillon S, Cabral A, Serrano O, Wernberg T, Filbee-Dexter K, Guimond JA, Tamborski JJ (2021) The renaissance of Odum's outwelling hypothesis in 'Blue Carbon' science. Est Coast Shelf Sci 255:107361. https://doi.org/10.1016/j.ecss.2021.107361 Sathish C, Reddy DN, Bharti VS, Deshmukhe G, Jaiswar AK (2023) Diversity and distribution of mangrove associated crabs (Infraorder: Brachyura) of India and the relationship between mangroves and crabs. Thalassas J 39(2):847–866. https://doi.org/10.1007/s41208-023-00578-x Sharifian S, Kamrani E, Saeedi H (2021) Global future distributions of mangrove crabs in response to climate change. Wetlands 41(8):99. https://doi.org/10.1007/s13157-021-01503-9 Skov MW, Hartnoll RG (2001) Comparative suitability of binocular observation, burrow counting and excavation for the quantification of the mangrove fiddler crab Uca annulipes (H. Milne Edwards). Hydrobiologia 449:201–212. https://doi.org/10.1007/978-94-017-0645-2_22 Wang F, Sanders CJ, Santos IR, Tang J, Schurech M, Kirwan ML, Kopp RE, Zhu K, Li X, Yuan J, Liu W, Li Z (2021) Global blue carbon accumulation in tidal wetlands increases with climate change. Natl Sci Rev 8(9):nwaa296. https://doi.org/10.1093/nsr/nwaa296 Wright JP, Jones CG (2004) Predicting effects of ecosystem engineers on patch-scale species richness from primary productivity. Ecology 85:2071–2081. https://doi.org/10.1890/02-8018 Additional Declarations No competing interests reported. Cite Share Download PDF Status: Posted Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. 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13:40:13","extension":"xml","order_by":9,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":127187,"visible":true,"origin":"","legend":"","description":"","filename":"eda52741d65a4a6683c0d8b675b449c61structuring.xml","url":"https://assets-eu.researchsquare.com/files/rs-7095077/v1/7719079612457c66f7bb8db3.xml"},{"id":93687322,"identity":"c65848a6-f60f-4456-b5b6-de35fe896009","added_by":"auto","created_at":"2025-10-16 13:24:13","extension":"html","order_by":10,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":137476,"visible":true,"origin":"","legend":"","description":"","filename":"earlyproof.html","url":"https://assets-eu.researchsquare.com/files/rs-7095077/v1/cc9fbdc8461bc4a71bae0eba.html"},{"id":93687310,"identity":"8ad731c1-7b7d-4772-b5ee-e27946493d33","added_by":"auto","created_at":"2025-10-16 13:24:13","extension":"jpg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":245418,"visible":true,"origin":"","legend":"\u003cp\u003eMap of the Colombian Pacific coast showing the five studied mangrove ecosystems.\u003c/p\u003e","description":"","filename":"1.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7095077/v1/6ee0f35fe5ec162d4d0255c9.jpg"},{"id":93687313,"identity":"211da047-e186-4580-9d40-96219df4da26","added_by":"auto","created_at":"2025-10-16 13:24:13","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":78251,"visible":true,"origin":"","legend":"\u003cp\u003eCrab trophic guilds abundance in five mangrove forests of the Colombian Pacific. DEP: Deposit feeder.\u003c/p\u003e","description":"","filename":"2.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7095077/v1/888303aa0a6f40fda2d21aa0.jpg"},{"id":93688605,"identity":"908011b6-d2bd-4614-a484-89b8bf0910ca","added_by":"auto","created_at":"2025-10-16 13:40:13","extension":"jpg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":25820,"visible":true,"origin":"","legend":"\u003cp\u003eCanonical correspondence analysis for abundance data of crab’s trophic guilds with mangrove ecosystems and soil environmental variables represented by vectors. Each green line represents a soil environmental variable.\u003c/p\u003e","description":"","filename":"3.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7095077/v1/0ada607204e339a5a6c59324.jpg"},{"id":93689009,"identity":"27879e64-1f8e-4a43-9cf5-bb5e830be5b3","added_by":"auto","created_at":"2025-10-16 13:48:14","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1197371,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7095077/v1/ced75de1-b521-4f18-8578-849cb7b3dcb6.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Burrowing crabs as potential bioindicators of sediment carbon storage in mangroves of the Colombian Pacific with different degrees of anthropogenic disturbance.","fulltext":[{"header":"Introduction","content":"\u003cp\u003e'Blue Carbon' refers to the carbon captured by the world's ocean or coastal vegetated ecosystems. Since the term was coined about a decade ago (McLeod et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2011\u003c/span\u003e), our knowledge about blue carbon sequestration has rapidly evolved. In mangrove forests, sequestration rates and standing C stocks are exceptionally high and exceed those of comparable terrestrial ecosystems (e.g., tropical rainforests) (Donato et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Mcleod et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). There are local and global scale estimates of sediment carbon in mangrove forests (e.i., Atwood et al. 2017; Jennerjahn \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Ouyang and Lee \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Wang et al. \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Thus, the potential of mangrove forests to sequester and store large amounts of carbon has been relatively well documented (Santos et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Nevertheless, the role of the macrobenthos, as ecosystem engineers, in sediment biogeochemical cycles is still inconclusive (Qiu et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). For instance, crabs have been considered good indicators of the environmental habitat quality of mangrove ecosystems (e.i., Amaral et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Retnaningdyah et al. \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) but little is known about the indicator potential of sediment carbon storage of mangrove crabs for blue carbon sequestration assessment.\u003c/p\u003e\u003cp\u003eEcosystem engineers are organisms that directly or indirectly affect other organisms' composition and critical biological processes by changing abiotic environments (e.g., Jones et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e1994\u003c/span\u003e; Wright and Jones \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). Sesarmid and ocypodid crabs are the mangrove fauna's major bioturbating and bioengineering components through their feeding and digging activities (Cannicci et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Kristensen et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Lee \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e1998\u003c/span\u003e). They can play an essential role in retaining organic matter (OM), as some species drag leaf litter and propagules into their burrows, restricting the export through tidal flushing (Ashton \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Lee \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Through their burrow, other species enhance oxygen flux into the waterlogged sediment, facilitating oxidation and enhancing nutrient availability (Kristensen and Alongi \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2006\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eSesarmids are herbivorous crabs that can consume up to 81% of mangrove litter production (Nordhaus et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). They can dig burrows down to 2 meters in depth and store leaf litter, mangrove propagules, and other organic material from different sources, enhancing the retention of OC within the forest (Kristensen et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Andreetta et al. 2013). Conversely, a recent study showed that the sesarmid crab \u003cem\u003eUcides cordatus\u003c/em\u003e feeding and digging activities would not increase the organic matter contents in the sediments since their burrows would expose deeper parts of the sediment to aerobic conditions, thus facilitating the decomposition of initially preserved organic matter (Katzer \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). In this way, the differences in OM content between mangrove forests could be explained by hydro-geomorphic differences and less by crab or crab burrow-related effects. This includes the tidal amplitude, implications, and riverine influences (Bouillon et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2003\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eOcypodid crabs are filter feeders that can exploit food sources from the sediment, such as bacteria, benthic microalgae, and meiofauna (Reinsel \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). As filter feeders, they feed on the sediment surface, never burying organic material, as typical for sesarmids (Andreetta et al. 2013). Burrows of the pantropical genus \u003cem\u003eUca\u003c/em\u003e are the most abundant and conspicuous, and they are typically less deep than sesarmids (Michaels and Zieman \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Some studies have shown that the presence of \u003cem\u003eUca\u003c/em\u003e burrows aerates sediments, thus facilitating the decomposition of OM (e.i., Katz \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e1980\u003c/span\u003e; Montague \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e1982\u003c/span\u003e). Conversely, an experimental study showed that their burrows have little effect on surrounding sediment oxygen concentrations. Thus, the differences in the decomposition of OM would depend significantly on the characteristics of the sediment in which the burrows are located (Michael and Zieman 2013). Likewise, a recent study showed that crab burrowing could trap and intercept detritus, consequently promoting the retention and accumulation of soil carbon instead of facilitating the oxidation of OM (Qiu et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eMangroves of the Tropical Eastern Pacific (TEP) are the second-largest mangrove area in the Neotropics (FAO \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). They are prevalent coastal ecosystems along the Pacific coasts of Colombia, with several structural features and service provisions that make them important regionally and globally (Castellanos et al. 2020). As part of the Eastern Pacific Basin, this area is affected by a large tidal range and is one of the rainiest regions of the world; this set of conditions converges in mangrove forests with unique characteristics (Blanco and Cantera 2001; Medina-Contreras et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). However, large mangrove areas in the Colombian Pacific remain poorly studied (Castellanos et al. 2020), and they could be affected by multiple anthropogenic stressors such as deforestation; wood production for fuel, coal, and construction materials; urban and industrial development (Palacios et al. 2019; Rodr\u0026iacute;guez-Rodr\u0026iacute;guez et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; FAO \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Recently, G\u0026oacute;mez-Garc\u0026iacute;a, et al. (\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), found a relationship between the degree of anthropogenic intervention and carbon reserves in deltaic mangroves of the region, where systems with the highest anthropogenic disturbance (measured through the anthropogenic disturbance index) registered the lowest values in blue carbon.\u003c/p\u003e\u003cp\u003eIn this study, field investigations were conducted to examine the link of burrowing crabs (as key bioturbators) to soil organic matter dynamics in mangrove forests of the Colombian Pacific with different anthropogenic and environmental stressors. We used the different disturbance degrees in mangroves of the Colombian Pacific by G\u0026oacute;mez-Garc\u0026iacute;a et al. (\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), and grouped burrowing crabs by trophic guilds to link crabs\u0026rsquo; abundance and soil organic carbon content (SOC) in each ecosystem. The abundance of some crab trophic guilds was expected to correlate with the SOC to evaluate crabs as bioindicators of sediment carbon storage. It was also expected that mangrove forests with fewer anthropogenic stressors would have higher SOC concentrations, as shown recently by G\u0026oacute;mez-Garc\u0026iacute;a et al. (\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e"},{"header":"Material and methods","content":"\u003cp\u003eStudy area\u003c/p\u003e\u003cp\u003eThe Pacific Coast of Colombia is one of the wettest regions in America (mean humidity between 88 and 100%) with one of the highest annual precipitations for the coastal areas of the Western Hemisphere (precipitation\u0026thinsp;\u0026gt;\u0026thinsp;500 cm yr\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e), with a wet period from January to April and a very wet period from May to December (Castellanos-Galindo et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Temperatures above 24\u0026deg;C. (Rangel and Arellano \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2004\u003c/span\u003e), and average air temperature of 25\u0026deg;C (Cantera and Blanco 2001). Five mangrove systems (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e) with different levels of anthropogenic intervention were selected: (1) El Morro, a near pristine mangrove located in the internal zone of the Bah\u0026iacute;a M\u0026aacute;laga estuary (4\u0026deg;02'58 \" N, 77\u0026deg;11'28\" W), where there is a Natural National Park and low anthropogenic disturbance index (ADI\u0026thinsp;=\u0026thinsp;0) (G\u0026oacute;mez-Garc\u0026iacute;a et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). (2) San Pedro (03\u0026deg; 50\u0026prime; 11\" N, 77\u0026deg; 15\u0026prime; 30\" W) and (3) Piang\u0026uuml;ita (03\u0026deg; 50\u0026prime; 32\" N, 77\u0026deg; 12\u0026prime; 23\" W) mangroves within Buenaventura Bay, which are near to the main port on the Colombian and have a high anthropogenic disturbance index (ADI\u0026thinsp;=\u0026thinsp;5 y 8, respectively) (G\u0026oacute;mez-Garc\u0026iacute;a et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). (4) Bocagrande (1\u0026deg;47'30\"N, 8\u0026deg;52'18\"W), and (5) Rompido (1\u0026deg;48'45\"N, 78\u0026deg;50'16\"W) mangroves set south of Tumaco Bay, which are near to Tumaco municipal (200000 inhabitants) and have a moderate level (ADI\u0026thinsp;=\u0026thinsp;2) and a high anthropogenic intervention (ADI\u0026thinsp;=\u0026thinsp;7), respectively (G\u0026oacute;mez-Garc\u0026iacute;a et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Relevant characteristics of the five systems are shown in Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eMain features of the five mangrove forests studied in the Colombian pacific coast. Data taken from G\u0026oacute;mez-Garc\u0026iacute;a et al. (\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). * Unpublish data.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"6\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eFeature/sample site\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eEl Morro\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eSan Pedro\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003ePiang\u0026uuml;ita\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003eBocagrande\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003eRompido\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eAnthropogenic disturbance index (ADI)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e7\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDominant mangrove species\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eRhizophora mangle\u003c/em\u003e\u003c/p\u003e\u003cp\u003e\u003cem\u003eMora oleifera\u003c/em\u003e\u003c/p\u003e\u003cp\u003e\u003cem\u003ePelliciera rhizophorae\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e\u003cem\u003eRhizophora mangle\u003c/em\u003e\u003c/p\u003e\u003cp\u003e\u003cem\u003eLaguncularia racemosa\u003c/em\u003e\u003c/p\u003e\u003cp\u003e\u003cem\u003eMora oleifera\u003c/em\u003e\u003c/p\u003e\u003cp\u003e\u003cem\u003ePelliciera rhizophorae\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003eRhizophora mangle\u003c/em\u003e\u003c/p\u003e\u003cp\u003e\u003cem\u003ePelliciera rhizophorae\u003c/em\u003e\u003c/p\u003e\u003cp\u003e\u003cem\u003eLaguncularia racemosa\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u003cem\u003eRhizophora mangle\u003c/em\u003e\u003c/p\u003e\u003cp\u003e\u003cem\u003ePelliciera rhizophorae\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u003cem\u003eRhizophora mangle\u003c/em\u003e\u003c/p\u003e\u003cp\u003e\u003cem\u003eAvicennia germinans\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eAbundance (ind/ha)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e783.3\u0026thinsp;\u0026plusmn;\u0026thinsp;110.80\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1283.3\u0026thinsp;\u0026plusmn;\u0026thinsp;492.27\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e1333.3\u0026thinsp;\u0026plusmn;\u0026thinsp;320.06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e500.0\u0026thinsp;\u0026plusmn;\u0026thinsp;112.54\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e983.3\u0026thinsp;\u0026plusmn;\u0026thinsp;110.80\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eBasal Area (m\u003csup\u003e2\u003c/sup\u003e/ha)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e27.36\u0026thinsp;\u0026plusmn;\u0026thinsp;8.01\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e15.08\u0026thinsp;\u0026plusmn;\u0026thinsp;228\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e1620\u0026thinsp;\u0026plusmn;\u0026thinsp;2,93\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e43.85\u0026thinsp;\u0026plusmn;\u0026thinsp;18.93\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e1540\u0026thinsp;\u0026plusmn;\u0026thinsp;179\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDiameter at breast height (cm)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e17.02\u0026thinsp;\u0026plusmn;\u0026thinsp;1.91\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e12.95\u0026thinsp;\u0026plusmn;\u0026thinsp;0.92\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e10.91\u0026thinsp;\u0026plusmn;\u0026thinsp;0.88\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e29.83\u0026thinsp;\u0026plusmn;\u0026thinsp;3.07\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e13.47\u0026thinsp;\u0026plusmn;\u0026thinsp;0.79\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e*Apparent density (g/cm\u003csup\u003e3\u003c/sup\u003e)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.9\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003eField sampling\u003c/p\u003e\u003cp\u003eSampling took place in 2023. The density of crabs was estimated during low tides during both the rainy and less rainy seasons. In each of the five different study ecosystems, five 2 \u0026times; 2 m random quadrats were sampled to assess the density of crab populations. Individuals were first counted visually at each site to assess the relative frequencies of the populations present in the quadrat (Skov and Hartnoll \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Cannicci et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). Then, crabs were collected manually to avoid underestimating specimens not active on the surface during the visual crab counts.\u003c/p\u003e\u003cp\u003eSamples were collected from the selected quadrats using a Russian drill hole soil sampler (0\u0026ndash;50 cm) to determine soil components and organic carbon content. The samples were analyzed to determine the percentages of sand, silt, clay, bulk density, and organic carbon (SOC). Temperature, pH, and conductivity from interstitial water were registered \u003cem\u003ein situ\u003c/em\u003e using a soil pH-moisture meter.\u003c/p\u003e\u003cp\u003eSoil analyses\u003c/p\u003e\u003cp\u003eBulk samples were air-dried, sieved (\u0026lt;\u0026thinsp;2 mm) and homogenized. The Bouyoucos-hydrometer method was used to analyze particle size (Bouyoucos, \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e1962\u003c/span\u003e). To determine soil bulk density, samples with a known volume metal corer were oven-dried at 60\u0026deg;C for 48 h and weighed. The bulk density was calculated as the ratio of the dry mass of the soil sample to its volume. Soil organic carbon was determined by the Walkley and Black method (Andreetta et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Soil organic C stored in the soil was calculated by the following formula:\u003cdiv id=\"Equa\" class=\"Equation\"\u003e\u003cdiv format=\"TEX\" class=\"mathdisplay\" id=\"FileID_Equa\" name=\"EquationSource\"\u003e\n$$\\:{SOC}_{tot}={\\sum\\:}_{1}^{n}(OC\\:\\times\\:\\:DB\\:\\times\\:\\:Dh)$$\u003c/div\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003ewhere SOC is the soil organic carbon stock (kg m\u003csup\u003e\u0026minus;\u0026thinsp;2\u003c/sup\u003e), OC is the organic carbon content for each depth interval (mg g\u0026thinsp;\u0026minus;\u0026thinsp;1), BD is the bulk density (g cm\u003csup\u003e\u0026minus;\u0026thinsp;3\u003c/sup\u003e), and Dh is the soil thickness interval.\u003c/p\u003e\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e\u003ch2\u003eStatistical analysis\u003c/h2\u003e\u003cp\u003eCrabs were classified into five trophic categories: herbivores, omnivores, carnivores, deposit feeders, and filter feeders. Feeding guilds were assigned according to information found in local literature (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Before multivariate analysis, trophic guild assemblage data were transformed by the 4th root to reduce the influence of abundant species relative to rare ones (Clarke and Warwick 2001). With these data, Bray-Curtis similarity matrices were calculated and used as input for non-parametric permutational multivariate analysis of variances (PERMANOVA) to test for ecosystem and season differences in crab community structure and composition. The P-values were then obtained using 9999 permutations. A posteriori pairwise comparison with the PERMANOVA t-statistics was conducted to elucidate spatial differences within mangrove ecosystems. To reveal the relationships between the abundance of crab trophic guilds and soil properties, a Canonical Correspondence Analysis (CCA) was carried out. Finally, we performed a Spearman rank correlation (SRC) analysis of the abundance of crab trophic guilds and soil properties between explanatory variables to demonstrate correlations.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eCrabs\u0026rsquo; species from mangrove forests of the Colombian Pacific and their trophic guild regarding local sources.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"6\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eFamily/Subfamily\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003eSpecies\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eTrophic guild\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003eAbbreviation\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003eSource\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eOcypodidae Rafinesque, 1815\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eOcypodinae Rafinesque, 1815\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eUca sp\u003c/em\u003e. Latreille, 1819\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\" morerows=\"3\" rowspan=\"4\"\u003e\u003cp\u003eDeposit feeders\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\" morerows=\"3\" rowspan=\"4\"\u003e\u003cp\u003eDEP1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\" morerows=\"3\" rowspan=\"4\"\u003e\u003cp\u003eMedina-Contreras et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2022\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eUca (Petruca) panamensis\u003c/em\u003e (Stimpson, 1859)\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eUca heteropleura\u003c/em\u003e (Smith, 1870)\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eUca stylifera\u003c/em\u003e (H. Milne Edwards, 1852)\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eGelasiminae Miers, 1886\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eLeptuca sp\u003c/em\u003e Bott, 1973\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\" morerows=\"4\" rowspan=\"5\"\u003e\u003cp\u003eDeposit feeders\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\" morerows=\"4\" rowspan=\"5\"\u003e\u003cp\u003eDEP2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\" morerows=\"4\" rowspan=\"5\"\u003e\u003cp\u003eMedina-Contreras et al. 2020\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eLeptuca inaequalis\u003c/em\u003e (Rathbun, 1935)\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eLeptuca tenuipedis\u003c/em\u003e (Crane, 1941)\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eLeptuca batuenta\u003c/em\u003e (Crane, 1941)\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eLeptuca tomentosa\u003c/em\u003e (Crane, 1941)\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eMinuca sp\u003c/em\u003e. Bott, 1954\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003eDeposit feeders\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003eDEP3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\" morerows=\"2\" rowspan=\"3\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eMinuca umbratila\u003c/em\u003e (Crane, 1941)\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eMinuca brevifron\u003c/em\u003es (Stimpson, 1860)\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eUcidinae Števcic, 2005\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eUcides occidentalis\u003c/em\u003e (Ortmann, 1897)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eHerbivores\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eHER\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eMedina-Contreras et al. 2020\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eGrapsidae MacLeay, 1838\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003ePachygrapsus sp\u003c/em\u003e. Randall, 1840\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003eOmnivores\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003eOMN\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003eMedina-Contreras et al. 2020\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003ePachygrapsus socius\u003c/em\u003e Stimpson, 1871\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eGoniopsis pulchra\u003c/em\u003e (Lockington, 1877)\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePanopeidae Ortmann, 1893\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003ePanopeus sp\u003c/em\u003e H. Milne Edwards, 1834\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003eCarnivores\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003eCAR\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\" morerows=\"4\" rowspan=\"5\"\u003e\u003cp\u003eMedina-Contreras et al. 2020\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003ePanopeus chilensis\u003c/em\u003e H. Milne Edwards \u0026amp; Lucas, 1843\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003ePanopeus purpureus\u003c/em\u003e Lockington, 1877\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eEurypanopeus transversus\u003c/em\u003e (Stimpson, 1860)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"3\" nameend=\"c5\" namest=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003eEurypanopeus planus\u003c/em\u003e (Smith, 1869)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePorcellanidae Haworth, 1825\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003e\u003cem\u003ePetrolisthes zacae\u003c/em\u003e Haig, 1968\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eFilter feeders\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eFIL\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eMedina-Contreras et al. 2020\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003c/div\u003e"},{"header":"Results","content":"\u003cp\u003eIn total, 22 species from 4 families were captured (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). The most dominant families in terms of species numbers per family were Ocypodidae (12 spp.) and Panopeidae (5 spp.). Deposit feeders were the most representative guild in terms of species numbers, which were grouped by gender at Deposit feeders 1 (\u003cem\u003eUca\u003c/em\u003e spp.), Deposit feeders 2 (\u003cem\u003eLeptuca\u003c/em\u003e spp.), and Deposit feeders 3 (\u003cem\u003eMinuca\u003c/em\u003e spp.) (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). PERMANOVA analysis revealed significant differences in crab community structure between mangrove forests (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05). However, there was no difference between rainy seasons (p\u0026thinsp;\u0026gt;\u0026thinsp;0.05). Posteriori pairwise comparisons did not show differences between San Pedro and Piang\u0026uuml;ita and between Rompido and Piang\u0026uuml;ita, ecosystems with a high ADI (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eNon-parametric pairwise comparisons of 4th-root transformed trophic guilds crabs\u0026rsquo; assemblage. In bold significant differences.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"6\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eSan Pedro\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePing\u0026uuml;ita\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eEl Morro\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003eBocagrade\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003eRompido\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eSan Pedro\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.0903\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cb\u003e0.0076\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u003cb\u003e0.0167\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u003cb\u003e0.0076\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePiang\u0026uuml;ita\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cb\u003e0.0071\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u003cb\u003e0.0145\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.0527\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eEl Morro\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u003cb\u003e0.0075\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u003cb\u003e0.0077\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eBocagrande\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u003cb\u003e0.0075\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eRompido\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003eDeposit feeder 1 (\u003cem\u003eUca spp\u003c/em\u003e.) and 3 (\u003cem\u003eMinuca spp\u003c/em\u003e.) were the most abundant species in San Pedro; deposit feeder 2 (\u003cem\u003eLeptuca spp\u003c/em\u003e.) were the most abundant species in El Morro; carnivores were the most abundant species in Rompido (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e, Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). There was little abundance of herbivores and filters; for this reason, former trophic guilds were excluded from the CCA. The canonical correlation describes the principal tendencies in the relationship between crabs' trophic guilds and their environment (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). The projections of a species on this axis show its preference for high or low values of this environmental gradient. The first and second ordination axes represent this variation best and showed eigenvalues of 0.116 and 0.035, respectively, which amounted to 71.2% and 21.6% of the variation in crab abundance (Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). Axis 1 separated the ecosystems with a low and moderate anthropogenic disturbance indexon the right side, which are characterized by the highest SOC, clay, BD, and Cond, in opposition to the ecosystems with a high ADI on the left side, with lowest SOC, clay, BD and Cond, and higher sand characterize. The DEP 2 (\u003cem\u003eLeptuca\u003c/em\u003e spp.) strongly correlated to sites in the middle estuary ecosystems with low and moderate ADI. The DEP 1 (\u003cem\u003eUca\u003c/em\u003e spp.), DEP 3 (\u003cem\u003eMinuca\u003c/em\u003e spp.), and Omnivores species are more closely related to sites with a high ADI (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab4\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 4\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eSummary of canonical correspondence analysis for abundance data of crab\u0026rsquo;s trophic guilds with mangrove ecosystems and soil environmental variables.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"7\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eAxis\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eEigenvalue\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003e%\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003eAxis\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003eEigenvalue\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003ep\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.11597\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e71.38\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.116\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.001\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.035053\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e21.58\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.03505\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.037\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.011023\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e6.785\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.01102\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.120\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.00042\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.2585\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.00042\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.721\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003eSince DEP 1 and 3 showed similar correlations with the soil environmental variable, they were analyzed as a group at the SRC analysis. Thus, DEP 1\u0026thinsp;+\u0026thinsp;3 (\u003cem\u003eUca spp\u003c/em\u003e. + \u003cem\u003eMinuca spp.\u003c/em\u003e) showed a significant and negative correlation with SOC and a positive correlation with sand (Table\u0026nbsp;\u003cspan refid=\"Tab5\" class=\"InternalRef\"\u003e5\u003c/span\u003e). Omnivores were negatively associated with SOC and clay, while DEP 2 (\u003cem\u003eLeptuca spp.\u003c/em\u003e) was positively associated with SOC and clay (Table\u0026nbsp;\u003cspan refid=\"Tab5\" class=\"InternalRef\"\u003e5\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab5\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 5\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eSpearman\u0026rsquo;s rank correlation matrix for abundance data of crab\u0026rsquo;s trophic guilds with soil environmental variables. In bold significant result.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"7\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eSOC\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eSand\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eClay\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u003cem\u003eLeptuca spp\u003c/em\u003e (DEP2)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003eDEP 1\u0026thinsp;+\u0026thinsp;3\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003eOm\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eSOC\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1.97E-05\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e9.39E-08\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u003cb\u003e3.26E-05\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u003cb\u003e0.023937\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e\u003cb\u003e0.01842\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eSand\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-0.74462\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.00014203\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u003cb\u003e0.0023252\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u003cb\u003e0.032543\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.085356\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eClay\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.84692\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-0.68846\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u003cb\u003e0.0011255\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.054947\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e\u003cb\u003e0.021035\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eLeptuca\u003c/em\u003e spp (DEP2)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cb\u003e0.7315\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e\u003cb\u003e-0.58095\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cb\u003e0.61287\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.74411\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.034515\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eADD 1\u0026thinsp;+\u0026thinsp;3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cb\u003e-0.45018\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e\u003cb\u003e0.4286\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e-0.38852\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e-0.068724\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.59903\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eOm\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cb\u003e-0.46761\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.35101\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cb\u003e-0.45889\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e-0.42431\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e-0.11049\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab6\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 6\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eCrab contribution to sediment carbon storage of four mangrove systems compared.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"8\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eAuthors\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eCountry\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eApproach\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAnalysis\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003eTrophic guild/ Group assemblage\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003eCrab species\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003eCrab contribution\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c8\"\u003e\u003cp\u003eProportion of contribution\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003eAndreetta et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2014\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003eKenya\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003eRelationships\u003c/p\u003e\u003cp\u003ebetween crab assemblage and SOC\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003eDistance-based linear model\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDeposit feeders\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u003cem\u003eUca annulipes, Uca inversa, Uca chlorophthalmus, Uca urvillei\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eIncrease\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0,55\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eBurrowing\u003c/p\u003e\u003cp\u003eSesarmid\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u003cem\u003eNeosarmatium africanum, Neosarmatium smithi\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eIncrease\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0,56\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eNon-Burrowing Sesarmid\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u003cem\u003eChiromantes ortmanni, Chiromantes eulimene, Perisesarma guttatum\u003c/em\u003e,\u003c/p\u003e\u003cp\u003e\u003cem\u003eParasesarma leptosoma\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eIncrease\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0,42\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eKatzer, \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2023\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eColombia\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eField experiment with crab-inhabited and field investigations\u003c/p\u003e\u003cp\u003ecrab-free burrows\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eKruskal-Wallis test\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eBurrowing\u003c/p\u003e\u003cp\u003eSesarmid\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u003cem\u003eUcides cordatus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDecrease\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0,14\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eQiu et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2019\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eChina\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eComparison between areas with few or no crab burrows and areas with high density crab burrows\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eOne-way\u003c/p\u003e\u003cp\u003eanalysis of variance\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eBurrowing crabs, mainly herbivorous\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u003cem\u003eHelice tientsinensis\u003c/em\u003e as the dominating species\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eIncrease\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eN.A\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003eThis study\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003eColombia\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003eCorrelation between abundance data of crab\u0026rsquo;s trophic guilds with soil environmental variables\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\" morerows=\"2\" rowspan=\"3\"\u003e\u003cp\u003eSpearman\u0026rsquo;s rank\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDeposit feeders\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u003cem\u003eLeptuca\u003c/em\u003e spp.\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eIncrease\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0,73\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eDeposit feeders\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u003cem\u003eUca\u003c/em\u003e spp., \u003cem\u003eMinuca\u003c/em\u003e spp.\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDecrease\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e-0,45\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eOmnivores\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u003cem\u003ePachygrapsus spp., Goniopsis pulchra\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDecrease\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e-0,47\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eThis study compared the crab trophic guild\u0026rsquo;s structure and composition in relation to mangroves with different degrees of anthropogenic intervention levels and climate seasons. The systems included a near pristine mangrove (El Morro) with a low anthropogenic intervention level, three sites with a high level (San Pedro, Piang\u0026uuml;ita, and Rompido), and a system with a moderate level (Bocagrande). These systems also have differences regarding soil apparent density, reenforcing anthropogenic intervention levels. Crab trophic guilds were also linked to soil organic carbon content (SOC) in each ecosystem, showing they can be used as potential bioindicators for evaluating blue carbon sequestration as discussed below.\u003c/p\u003e\u003cp\u003eMangrove habitats generally have lower species richness of macrobenthos (e.g., 17\u0026ndash;85 species) than adjoining habitats such as seagrass meadows and open sand/mudflats. However, brachyuran crabs constitute one of the most diverse invertebrate groups in the mangrove ecosystem (Lee \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Crab richness in the mangrove forests of the Colombian Pacific (22 species from 4 families) was lower than in mangroves of the Indo-Pacific region, where there is higher mangrove species richness, such as the Indian coast, where the East coast hosts 127 crab species and the West Coast 83 species (Sathish et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Crab species richness in the Colombian Pacific would be higher than in the Colombian Caribbean coast, where, for instance, Sandoval (2023) found a total of 10 macrobenthos species. Higher mangrove species richness and organic carbon availability in the soil have been positively correlated with higher crab species richness and abundance (Sathish et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). The environmental conditions, such as salinity, depth, and sea surface temperature, are relevant drivers of the distribution of mangrove crabs worldwide (Sharifian et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eThe abundance of crab species belonging to the Ocypodidae family is relatively high in the Colombian Pacific, as in other mangrove ecosystems (Lee \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Deposit feeders were the most representative trophic guild in terms of species numbers, most of them belonging to the Ocypodidae family. There was no difference in crab community structure between rainy seasons. This would be explained because this region has one of the highest annual precipitations for the coastal areas of the western hemisphere, where there is a rainy season (May to December, average 746 mm/month) and the less rainy season (January to April, average 422 mm7month) (Castellanos-Galindo et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). However, we found significant differences in crab community structure between mangrove forests, where San Pedro and Piang\u0026uuml;ita, ecosystems with the highest ADI, did not show differences in their community structure but showed remarkable differences regarding the other three forests (see below), which suggests that anthropogenic intervention conditions would drive the distribution of mangrove crabs in this region.\u003c/p\u003e\u003cp\u003eThe canonical correlation describes the principal trends in the relationship between crab\u0026rsquo;s trophic guilds and their environment, supporting that their distribution would be linked to anthropogenic intervention. Axis 1 separated the ecosystems on the right side with low and moderate anthropogenic intervention levels (El Morro and Bocagrande). On the left, it separated the ecosystems with the highest intervention (Ping\u0026uuml;ita, San Pedro, and Rompido). Former ecosystems have a high abundance of \u003cem\u003eUca\u003c/em\u003e spp. (DEP 1), \u003cem\u003eMinuca\u003c/em\u003e spp. (DEP 3), and omnivore species, as well as the lowest SOC values. While El Morro and Bocagrande have a high abundance of \u003cem\u003eLeptuca\u003c/em\u003e spp. (DEP 2) and the highest SOC. Which suggests that \u003cem\u003eLeptuca\u003c/em\u003e spp. would be a good bio-indicator of higher sediment carbon storage and the mangrove ecosystem's quality in the Colombian Pacific mangrove ecosystems. Whereas \u003cem\u003eUca\u003c/em\u003e spp., \u003cem\u003eMinuca\u003c/em\u003e spp., and omnivores crabs could be used as potential indicators for disturbed systems and lower sediment carbon storage supporting our hypothesis.\u003c/p\u003e\u003cp\u003eThe above would be explained since mangrove crabs are sensitive to changes in mangrove cover area and changes in their community structure are useful tools to detect mangrove habitat status (Sathish et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Some studies have shown that mangrove degradation negatively affects crab biodiversity and communities\u0026rsquo; structure (e.i., Ashton et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Maia and Coutinho \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Here, we showed significant differences in crab trophic guild\u0026rsquo;s structure and composition in relation to mangroves with different degrees of anthropogenic intervention. Likewise, \u003cem\u003eLeptuca\u003c/em\u003e spp. crabs, dominated in low and moderate anthropogenic intervention levels forests (with the highest SOC values), whereas \u003cem\u003eUca\u003c/em\u003e spp., \u003cem\u003eMinuca\u003c/em\u003e spp., and omnivores crabs were more abundant in the degraded forests (with lowest SOC values).\u003c/p\u003e\u003cp\u003eThe field investigation indicated that mangrove soils in the region vary considerably according to anthropogenic intervention as showed recently by G\u0026oacute;mez-Garc\u0026iacute;a et al. (\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Our results suggest that ecosystems with high mangrove degradation, such as San Pedro and Ping\u0026uuml;ita within Buenaventura Bay, which have been intensively degraded for domestic use, such as firewood and house building (Palacios and Cantera \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), have lost their whole capacity for soil carbon storage. \u003cem\u003eUca\u003c/em\u003e spp., \u003cem\u003eMinuca\u003c/em\u003e spp., and omnivores crabs could be used as potential indicators for these disturbed systems. This is supported by findings of other studies that have reported ocypodid crabs, mainly \u003cem\u003eUca\u003c/em\u003e species, dominating in disturbed, degraded, or young rehabilitation sites (Macintosh et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Ashton et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2003\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eThe effect of burrowing crabs on organic carbon storage varies according to anthropogenic intervention. In ecosystems with a high mangrove degradation, crab burrows (\u003cem\u003eUca\u003c/em\u003e spp., \u003cem\u003eMinuca\u003c/em\u003e spp. and omnivore species) could expose sediment parts to aerobic conditions, thus facilitating the decomposition of initially preserved organic matter, as found in another study (Katzer, \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). To explain this, we can hypothesize that crab burrows in ecosystems with high mangrove degradation could not trap enough organic matter to improve the contents of SOC since they have less carbon accumulated in aerial biomass (G\u0026oacute;mez-Garc\u0026iacute;a et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Consequently, they have a smaller amount of detritus available. Conversely, in crab burrows of \u003cem\u003eLeptuca\u003c/em\u003e spp. in non-degraded systems, detritus could be trapped and intercepted, especially plant detritus, which could be an important mechanism for promoting carbon retention and accumulation, consistent with other studies (Table\u0026nbsp;\u003cspan refid=\"Tab5\" class=\"InternalRef\"\u003e5\u003c/span\u003e). However, we acknowledge that other factors as vegetation and inundation times are recognized as controls of SOC in mangrove ecosystems (Andreetta et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Therefore, a more detailed understanding of the influence of burrowing crabs on SOC is required.\u003c/p\u003e\u003cp\u003eIn summary, the most dominant family in species numbers per family was Ocypodidae. Deposit feeders were the most representative trophic guild. The field investigation indicated that anthropogenic intervention conditions would drive the distribution of mangrove crabs in this region. Likewise, the effect of burrowing crabs on organic carbon storage varies according to anthropogenic intervention. Ecosystems with the highest anthropogenic intervention (Ping\u0026uuml;ita, San Pedro y Rompido) have a high abundance of \u003cem\u003eUca\u003c/em\u003e spp., \u003cem\u003eMinuca\u003c/em\u003e spp., and omnivores species and the lowest SOC values. Meanwhile, systems with low and moderate intervention (El Morro and Bocagrande) have a high abundance of \u003cem\u003eLeptuca\u003c/em\u003e spp. and the highest SOC. Burrowing crabs' abundance could be a good bioindicator of sediment carbon storage and the mangrove ecosystem's quality on the Colombian Pacific coasts.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003eThis study was funded by the Sistema General de Regal\u0026iacute;as (FCTel-SGR: BPIN 2020000100054).\u003c/p\u003e\u003cp\u003eEthics and Consent to Participate declarations: not applicable.\u003c/p\u003e\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eSLA, MJE and GAI developed the concept and designed the study. CDF, MA and GAI conducted the field study. SLA, MJE and GAI analysed the data. SLA wrote the original draft of the manuscript. SLA, MJE and GAI interpreted results. All authors discussed the results, commented on and improved various versions of the manuscript.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAmaral V, Penha-Lopes G, Paula J (2009) RNA/DNA ratio of crabs as an indicator of mangrove habitat quality. Aquat Conserv Mar Freshw 19:S56\u0026ndash;S62. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1002/aqc.1039\u003c/span\u003e\u003cspan address=\"10.1002/aqc.1039\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003cli\u003e\u003cspan\u003eAndreetta A, Fusi M, Cameldi I, Cim\u0026ograve; F, Carnicelli S, Cannicci S (2014) Mangrove carbon sink. Do burrowing crabs contribute to sediment carbon storage? 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Ecology 85:2071\u0026ndash;2081. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1890/02-8018\u003c/span\u003e\u003cspan address=\"10.1890/02-8018\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Macrobenthos, ecosystem engineers, anthropogenic intervention, trophic guild, Uca, Leptuca, Minuca","lastPublishedDoi":"10.21203/rs.3.rs-7095077/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-7095077/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eLittle is known about the indicator potential of sediment carbon storage of mangrove crabs (as ecosystem engineers) for blue carbon sequestration assessment. This study aimed to examine the crab trophic guild\u0026rsquo;s structure and composition in relation to mangroves with different degrees of anthropogenic disturbance index and climate seasons in the Colombian Pacific mangrove forests. Crab trophic guilds were also linked to soil organic carbon content (SOC) and other soil environmental variables in each ecosystem. Ocypodids and deposit feeders were the most representative crabs in terms of species numbers. Mangrove forests with the highest anthropogenic intervention (Ping\u0026uuml;ita, San Pedro y Rompido) have a high abundance of the deposit feeders, \u003cem\u003eUca\u003c/em\u003e spp., \u003cem\u003eMinuca\u003c/em\u003e spp., and Omnivores species, as well as the lowest SOC values. Meanwhile, forests with low and moderate intervention (El Morro and Bocagrande) have a high abundance of deposit feeders, \u003cem\u003eLeptuca\u003c/em\u003e spp., and the highest SOC. Crab burrows's abundance of \u003cem\u003eLeptuca\u003c/em\u003e spp. could be good bioindicators of higher sediment carbon storage and mangrove ecosystem quality, whereas \u003cem\u003eUca\u003c/em\u003e spp., \u003cem\u003eMinuca\u003c/em\u003e spp., and Omnivores species could be used as potential indicators for disturbed systems and lower sediment carbon storage.\u003c/p\u003e","manuscriptTitle":"Burrowing crabs as potential bioindicators of sediment carbon storage in mangroves of the Colombian Pacific with different degrees of anthropogenic disturbance.","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-10-16 13:24:08","doi":"10.21203/rs.3.rs-7095077/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"c9ab20c7-fceb-4e04-a553-6d6aa86c46f2","owner":[],"postedDate":"October 16th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2025-10-16T13:24:08+00:00","versionOfRecord":[],"versionCreatedAt":"2025-10-16 13:24:08","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-7095077","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-7095077","identity":"rs-7095077","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}