Spatiotemporal Drivers of Small Mammal Community Structure in the afroalpine Ecosystems of the central Ethiopian Highlands.

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This preprint studied how habitat type and season independently and interactively shape small mammal community structure in the Guassa Menz Community Conservation Area in the central Ethiopian highlands, using live-trapping across five habitats (Festuca grassland, Helichrysum shrubland, swamp grassland, Erica heather, and juniper plantation forest) during three seasons. Abundance (Capture Success Index) was analyzed with ANOVA, while alpha diversity (Shannon diversity and richness) and community composition were assessed with NMDS, PERMANOVA, and SIMPER. The authors found that capture success peaked in the Wet season, with Helichrysum producing the highest capture rates, whereas Shannon diversity depended on habitat but not season and species richness stayed stable across spatiotemporal scales; community composition was primarily structured by habitat, with swamp grassland dominated by the wetland specialist Otomys typus. The paper does not explicitly discuss endometriosis or adenomyosis; it was included in the corpus via a keyword match in the upstream search index.

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Abstract

Small mammal communities are sensitive indicators of ecosystem processes, with their structure in tropical montane systems like the Ethiopian highlands shaped by strong habitat and seasonal gradients. A critical lack of integrated analyses examining multiple community facets simultaneously limits our understanding of the relative importance of these spatiotemporal drivers. This study quantified the independent and interactive effects of habitat type and season on community abundance, alpha diversity, and composition in the Guassa menz community conservation area. We conducted live-trapping surveys across five habitats—Festuca grassland, Helichrysum shrubland, Swamp grassland, Erica heather, and Juniper plantation forest—over three seasons (Dry, Early Dry, Wet). Data were analyzed using ANOVA for univariate metrics (Capture Success Index, CSI; Shannon Diversity, H′; Richness) and multivariate methods (NMDS, PERMANOVA, SIMPER) for community composition. Our results revealed that overall abundance (CSI) peaked significantly during the Wet season, with the Helichrysum habitat yielding the highest capture rates. In contrast, alpha diversity (H′) was significantly influenced by habitat type but not by season, while species richness remained stable across all spatiotemporal scales. Multivariate analyses demonstrated that community composition was structured primarily by habitat, with the Swamp Grassland supporting a starkly distinct assemblage driven by the wetland specialist Otomys typus, while compositional differences among upland habitats were attributed to abundance shifts of generalists like Lophuromys flavopunctatus. We conclude that community abundance, diversity, and composition are decoupled in their response to environmental drivers, with fine-scale habitat heterogeneity acting as the paramount filter for community assembly. Conservation strategies must therefore prioritize the protection of habitat mosaics, particularly diversity hotspots like the Helichrysum and unique ecological realms like the Swamp Grassland, to safeguard regional beta-diversity in this vulnerable Afroalpine ecosystem.
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Spatiotemporal Drivers of Small Mammal Community Structure in the afroalpine Ecosystems of the central Ethiopian Highlands. | Authorea try { document.documentElement.classList.add('js'); } catch (e) { } var _gaq = _gaq || []; _gaq.push(['_setAccount', 'G-8VDV14Y67G']); _gaq.push(['_trackPageview']); (function() { var ga = document.createElement('script'); ga.type = 'text/javascript'; ga.async = true; ga.src = ('https:' == document.location.protocol ? 'https://ssl' : 'http://www') + '.google-analytics.com/ga.js'; var s = document.getElementsByTagName('script')[0]; s.parentNode.insertBefore(ga, s); })(); Skip to main content Preprints Collections Wiley Open Research IET Open Research Ecological Society of Japan All Collections About About Authorea FAQs Contact Us Quick Search anywhere Search for preprint articles, keywords, etc. Search Search ADVANCED SEARCH SCROLL This is a preprint and has not been peer reviewed. Data may be preliminary. 29 November 2025 V1 Latest version Share on Spatiotemporal Drivers of Small Mammal Community Structure in the afroalpine Ecosystems of the central Ethiopian Highlands. Authors : ABRAHAM DESALEGN 0000-0002-7954-7716 [email protected] , Abdul Katakweba , and Alfan Rija Authors Info & Affiliations https://doi.org/10.22541/au.176442683.35816788/v1 205 views 152 downloads Contents Abstract Supplementary Material Information & Authors Metrics & Citations View Options References Figures Tables Media Share Abstract Small mammal communities are sensitive indicators of ecosystem processes, with their structure in tropical montane systems like the Ethiopian highlands shaped by strong habitat and seasonal gradients. A critical lack of integrated analyses examining multiple community facets simultaneously limits our understanding of the relative importance of these spatiotemporal drivers. This study quantified the independent and interactive effects of habitat type and season on community abundance, alpha diversity, and composition in the Guassa menz community conservation area. We conducted live-trapping surveys across five habitats—Festuca grassland, Helichrysum shrubland, Swamp grassland, Erica heather, and Juniper plantation forest—over three seasons (Dry, Early Dry, Wet). Data were analyzed using ANOVA for univariate metrics (Capture Success Index, CSI; Shannon Diversity, H′; Richness) and multivariate methods (NMDS, PERMANOVA, SIMPER) for community composition. Our results revealed that overall abundance (CSI) peaked significantly during the Wet season, with the Helichrysum habitat yielding the highest capture rates. In contrast, alpha diversity (H′) was significantly influenced by habitat type but not by season, while species richness remained stable across all spatiotemporal scales. Multivariate analyses demonstrated that community composition was structured primarily by habitat, with the Swamp Grassland supporting a starkly distinct assemblage driven by the wetland specialist Otomys typus, while compositional differences among upland habitats were attributed to abundance shifts of generalists like Lophuromys flavopunctatus. We conclude that community abundance, diversity, and composition are decoupled in their response to environmental drivers, with fine-scale habitat heterogeneity acting as the paramount filter for community assembly. Conservation strategies must therefore prioritize the protection of habitat mosaics, particularly diversity hotspots like the Helichrysum and unique ecological realms like the Swamp Grassland, to safeguard regional beta-diversity in this vulnerable Afroalpine ecosystem. 1. Introduction Understanding the mechanisms that structure ecological communities remains a central pursuit in ecology. Small mammal assemblages are particularly instructive model systems, as their composition, diversity, and abundance are sensitive indicators of habitat quality, resource partitioning, and biotic interactions (Happold, 2013; Jones & Safi, 2011). These community-level metrics are not merely descriptive; they are fundamental to ecosystem functioning, influencing seed dispersal, soil aeration, nutrient cycling, and predator-prey dynamics, thereby contributing to overall ecosystem stability and resilience (Stenseth et al., 2003). Consequently, quantifying the drivers of small mammal community structure is critical for predicting their responses to anthropogenic environmental change, including habitat fragmentation and climate shifts. Tropical montane ecosystems, such as the Afroalpine zones of the Ethiopian highlands, represent ideal natural laboratories for investigating community assembly processes. These regions are recognized as biodiversity hotspots characterized by steep environmental gradients, hosting a unique and often endemic fauna (Yalden & Largen, 1992). The strong abiotic gradients, most notably pronounced seasonal rainfall, and biotic gradients, comprising distinct habitat types with varying vegetation structure and resource availability, are predicted to impose powerful environmental filters (Lavorel & Garnier, 2002). These filters are expected to drive non-random patterns in community assembly, manifesting as spatial and temporal variation in alpha diversity (within a habitat) and beta-diversity (between habitats). For small mammals, which often exhibit specific microhabitat requirements and physiological constraints, the interplay between fine-scale habitat heterogeneity and seasonal resource pulses should be a primary determinant of their spatiotemporal distribution (Bekele, 1996). While it is generally accepted that both habitat type and seasonality exert influence on small mammal communities, their relative importance and interactive effects are highly context-dependent and remain poorly quantified in understudied Afroalpine systems (Kasso & Bekele, 2014). Moreover, community responses to these drivers are multi-faceted. A given habitat might profoundly influence which species are present (composition) without altering the number of species (richness), or vice-versa (Tews et al., 2004). Similarly, seasonal shifts may cause dramatic changes in overall abundance while leaving diversity metrics unchanged. A holistic and integrated understanding of community structure therefore necessitates a simultaneous examination of multiple facets—including abundance, alpha diversity, beta-diversity, and species-specific responses—an approach often missing from studies in this region. Despite the ecological significance of the Guassa Menz community conservation area, a global biodiversity hotspot, a comprehensive analysis of the simultaneous effects of fine-scale habitat heterogeneity and seasonality on the complete suite of community metrics for its small mammal fauna is conspicuously lacking. Previous studies have often focused on single aspects, such as species inventories or isolated habitat comparisons, failing to disentangle the complex spatiotemporal dynamics at play (e.g., Bekele et al., 2017). Our study fills this critical knowledge gap by employing a robust, multi-method analytical framework to provide an integrated assessment of how habitat and season interact to shape small mammal communities. The general objective of this study is to quantify the independent and interactive effects of habitat type and season on the structure of the small mammal community in the Guassa menz community conservation area, Ethiopia, with a focus on patterns of abundance, alpha diversity, and species composition. To achieve this, we pursue the following specific objectives: 1. To assess the spatiotemporal variation in overall abundance (using a Capture Success Index) and alpha diversity (Species Richness and Shannon Diversity) across distinct habitats and seasons. 2. To determine the patterns of beta-diversity and identify whether community composition is driven more by habitat type or seasonal turnover. 3. To identify the key species driving compositional differences between habitats and their association with specific environmental gradients. Research questions: 1. How do habitat type and season influence overall community abundance (capture success) and alpha diversity? 2. What is the relative importance of habitat versus season in structuring community composition (beta-diversity), and are there habitats with uniquely distinct assemblages? 3. Which species are primarily responsible for driving the observed compositional differences between habitats? Hypotheses: Hypothesis1: Habitat type will have a stronger effect on alpha diversity than season, with structurally complex habitats (e.g., Helichrysum ) supporting higher diversity, while overall abundance will peak during the Wet season across all habitats due to increased resource availability. Hypothesis2: Community composition (beta-diversity) will be significantly structured by both habitat and season, but habitat type will be the primary driver, with the Swamp Grassland supporting a functionally and taxonomically distinct community compared to the upland habitats. Hypothesis3: The distinctiveness of the Swamp Grassland community will be driven by the significant association of a wetland habitat specialist (e.g., Otomys typus ), while differences among upland habitats will be driven by abundance shifts of generalist species (e.g., Lophuromys flavopunctatus and Stenocephalemys spp.). To address the aforementioned knowledge gap, our research provides a comprehensive assessment of how spatiotemporal drivers structure small mammal communities. We simultaneously examine multiple community facets—abundance, alpha diversity, and species composition—to quantify the relative influence of fine-scale habitat heterogeneity and seasonality in the Afroalpine ecosystem of the Guassa Menz community conservation area. 2. Methods 2.1. Description of the Study Area This research was conducted in the Guassa Menz Community Conservation Area (GMCCA), located in the Central Ethiopian Highlands.( Figure 1). The area is characterized by an Afroalpine ecosystem, with a bimodal rainfall pattern consisting of a primary wet season (June–September) and a dry season (October–February), with a short ”Early Dry” period transitioning between them (Ashenafi et al., 2012). The landscape is a mosaic of five distinct habitat types: Erica Heather (dense, shrubby areas), Festuca Grassland (open plains), Helichrysum Habitat (open shrubland), Juniper Plantation Forest (planted stands), and Swamp Grassland (permanently moist graminoid habitat). Figure 1: Figure showing map of the study area 2.2. Study Design The research employed a fully factorial design to investigate the spatiotemporal dynamics of small mammal communities. Sampling was carried out across five distinct habitat types, selected to represent the major vegetation formations of the study area (Miehe & Miehe, 1994), and repeated across three distinct seasons: the Dry season (Jan-Mar), the Early Dry season (Oct-Dec), and the Wet season (Jun-Sep). This design was chosen as it is ideally suited to partition the variance in community structure into its spatial (habitat) and temporal (seasonal) components. 2.4. Small Mammal Trapping and Data Collection Small mammals were sampled using Sherman traps along 100 traps transects (with one trap at every 10 m interval) in each of the five habitats (Thomas, et al, 2020). A single Sherman live-trap (8 cm x 9 cm x 23 cm) baited with a mixture of peanut butter and crushed maize was placed at each station. Trapping was conducted for three consecutive nights during each seasonal sampling session, resulting in a total of 147 trap nights per habitat per season, and 2,205 trap nights for the entire study. Traps were checked each morning and evening. Upon capture, each individual was identified to species using standard taxonomic keys (Lavrenchenko & Bekele, 2017). For each individual, we recorded species type, sex, age class (adult or juvenile based on body size and pelage colour), reproductive condition, and body mass (to the nearest 0.5 g). All animals were released at their point of capture after processing. The primary data used for community-level analyses was the raw count of each species captured per habitat per season, which was compiled into a Species Abundance Matrix (SAM). 2.5. Community Variables Key community metrics were derived from the Species Abundance Matrix (SAM) for each habitat-season combination to capture different facets of community structure (Magurran, 2004): 2.5.1. Capture Success Index (CSI) This metric of relative abundance was calculated for each habitat and season as: CSI = (Total Number of Captures / Total Trap Nights) × 100. This index is a standard measure for comparing trappability and relative abundance across sampling sessions (Merritt et al., 2019). 2.5.2.Alpha Diversity We calculated two complementary metrics to describe within-habitat diversity: 1) Species Richness (S), defined as the total number of species captured per habitat per season, and 2) the Shannon Diversity Index (H′) to account for both species richness and evenness (Magurran, 2004). The Shannon index is calculated as H′ = -Σpi ln(pi), where pi is the proportion of individuals belonging to species i. 2.5.3. Community Composition The full Species Abundance Matrix, comprising all species counts across all habitat-by-season replicates, served as the basis for all multivariate analyses of beta-diversity and temporal turnover (Anderson et al., 2011). 2.6.Data Analysis All statistical analyses were performed in R software version 4.2.1 (R Core Team, 2022). 2.6.1. Capture Success and Alpha Diversity: The effects of Habitat (fixed factor, 5 levels), Season (fixed factor, 3 levels), and their interaction on CSI, Shannon Diversity (H′), and Species Richness were tested using separate Two-way Analysis of Variance (ANOVA). Assumptions of normality and homoscedasticity were verified using Shapiro-Wilk and Levene’s tests, respectively. 2.6.2. Beta-Diversity and Community Composition: Patterns in community composition were visualized using Non-Metric Multidimensional Scaling (NMDS) based on a Bray-Curtis dissimilarity matrix of the species abundance data. The statistical significance of the factors Habitat and Season on overall community composition was tested using a Permutational Multivariate Analysis of Variance (PERMANOVA) with 999 permutations, as implemented in the adonis2 function of the vegan package (Oksanen et al., 2022). 2.6.3. Drivers of Compositional Differences: To identify the species that contributed most to the observed dissimilarities between specific habitat types, a Similarity Percentage (SIMPER) analysis was conducted. Furthermore, to model and visualize the relationship between the entire community composition and the environmental predictors (Habitat and Season), a Redundancy Analysis (RDA) was performed. The significance of the global model and individual terms was assessed using permutation tests. 2.6.4. Sampling Adequacy: To confirm that our trapping effort was sufficient to characterize the local small mammal species pool, a Species Accumulation Curve (SAC) was constructed using the specaccum function in the vegan package. For all analyses, results were considered statistically significant at α = 0.05. 3. Results 3.1. Species Abundance and Capture Success The raw counts from the Species Abundance Matrix (SAM) established Lophuromys flavopunctatus as the unequivocally dominant species across all sampling events. Analysis of the Capture Success Index (CSI) revealed significant spatiotemporal heterogeneity, with detailed results summarized in Table 1 . The Helichrysum Habitat consistently yielded the highest CSI values, peaking at 4.44% during the Wet season, identifying it as the most critical habitat. A powerful seasonal effect was evident, with the Wet season universally recording the highest capture rates across all five habitats, while the Dry season marked the population lowest. Table 1: Capture Success Index Data Summary Habitat Season Total Captures Trap Nights CSI (%) Helichrysum Habitat Wet 100 2250 4.44 Helichrysum Habitat Early dry 71 2250 3.16 Erica Heather Wet 70 2250 3.11 Festuca Grassland Wet 68 2250 3.02 Festuca Grassland Early dry 47 2250 2.09 Swamp Grassland Wet 45 2250 2 Erica Heather Early dry 40 2250 1.78 Helichrysum Habitat Dry 30 2250 1.33 Swamp Grassland Early dry 30 2250 1.33 Juniper Plantation Forest Wet 25 2250 1.11 Festuca Grassland Dry 23 2250 1.02 Erica Heather Dry 22 2250 0.98 Juniper Plantation Forest Early dry 15 2250 0.67 Juniper Plantation Forest Dry 10 2250 0.44 Swamp Grassland Dry 10 2250 0.44 3.2. Patterns of Alpha Diversity Shannon Diversity (H′) varied across habitats and seasons, ranging from 1.26 to 1.52, as visualized in Figure 2 . A Two-way ANOVA confirmed that Habitat type was a highly significant factor influencing diversity (F = 11.23, p = 0.003), while Season and the Habitat: Season interaction were non-significant ( Table 2 ). In contrast, an ANOVA on Species Richness found no significant effects of Habitat, Season, or their interaction, indicating that the number of species (4 or 5) remained stable across the spatiotemporal matrix. Figure 2: Effect of Habitat and Season on Shannon Diversity (H’) Table2: ANOVA Results for Shannon Diversity Source Degrees of Freedom (Df) Sum of Squares (Sum Sq) Mean Square (Mean Sq) F value Habitat 4 0.817 0.2043 11.23 0.003 Season 2 0.057 0.0285 1.56 0.257 Habitat: Season 8 0.174 0.0218 1.19 0.413 Residuals 10 0.182 0.0182 3.3. Community Composition and Beta-Diversity Non-Metric Multidimensional Scaling (NMDS) ordination revealed a striking pattern. The communities from the four upland habitats (Erica Heather, Festuca Grassland, Helichrysum, Juniper Plantation Forest) formed a tight cluster, indicating highly similar species composition. In sharp contrast, all Swamp Grassland sites were widely separated, and the Wet season Swamp Grassland community was further segregated, indicating a distinct and seasonally dynamic assemblage. This observed turnover was confirmed by a highly significant PERMANOVA, which found that the combined effect of Habitat and Season explained 75.87% of the variation in community composition (F = 4.19, p = 0.005), with full results provided in Table 3 . Table 3: PERMANOVA Results Source Degrees of Freedom (Df) Sum of Squares (SumOfSqs) R2 F value Pr(>F) Model (Habitat + Season) 6 1.07018 0.75874 4.1933 0.005 Residual 8 0.34028 0.24126 Total 14 1.41046 1 3.4. Drivers of Community Structure: SIMPER and RDA Similarity Percentage (SIMPER) analysis identified the species driving compositional differences between habitats. Significant contrasts showed that the Helichrysum habitat’s distinctiveness from the Juniper Plantation Forest was driven by higher abundances of L. flavopunctatus (p = 0.035) and S. griseicauda (p = 0.023). Furthermore, the Swamp Grassland was characterized by a significant association with the wetland specialist Otomys typus (p = 0.026). Canonical Redundancy Analysis (RDA) quantified these patterns, with the global model explaining 78.31% of the variance in species data ( Table 4 ). Both Habitat (p = 0.016) and Season (p = 0.003) were significant predictors. The RDA biplot ( Figure 3 ) clearly visualized two primary gradients: RDA1, a season-abundance gradient associated with Helichrysum and the Wet season, and RDA2, a moisture gradient strongly associating the Swamp Grassland with O. typus and Arvicanthis abyssinicus . Table 4: Global and Marginal Tests for RDA Effect Degree of freedom Variance F-value p-value Global Model 6 3.9155 4.8138 0.003 Habitat 4 2.1004 3.8734 0.016 Season 2 1.8151 6.6946 0.003 Residual 8 1.0845 – – Figure 3. RDA Biplot, Redundancy analysis (RDA) of Small mammal communities. 3.5. Sampling Adequacy The Species Accumulation Curve (SAC) reached a clear asymptote at five species ( Figure 4 ), confirming that our sampling effort was sufficient to capture the local small mammal species pool. Figure 4: Species accumulation curve 4. Discussion Our integrated, multi-faceted analysis reveals a nuanced hierarchy of spatiotemporal drivers structuring the small mammal community in the Afroalpine ecosystem of Guassa Menz. A central finding is the fundamental decoupling of community metrics in their response to environmental drivers: overall abundance was a pulse-tracker, closely following seasonal resource availability ( Figure 1, Table 1 ), while patterns of diversity and composition were shaped by the more persistent template of fine-scale habitat heterogeneity. This decoupling underscores that a holistic understanding of community assembly requires moving beyond single metrics, as the factors determining ’how many individuals’ differ from those governing ’which species, and in what proportions’. The powerful seasonal signal in abundance, peaking uniformly across all habitats during the wet season ( Figure 1 ), aligns with the well-established paradigm of resource pulsing in tropical and alpine systems. This pattern is likely a consequence of a bottom-up increase in carrying capacity, where elevated primary productivity boosts the availability of critical resources such as seeds and invertebrates, thereby supporting higher population densities across the majority of species. In stark contrast to this dynamic abundance, we observed remarkable stability in species richness across both spatial and temporal gradients ( Table 1 ). This stability suggests that the presence or absence of species within a habitat is governed by longer-term habitat characteristics and the constraints of the regional species pool, rather than short-term seasonal fluctuations. However, alpha diversity, as captured by the Shannon index, was significantly influenced by habitat type, revealing a more subtle layer of community organization ( Figure2 ). The higher diversity in the structurally complex Helichrysum shrubland, without a concomitant increase in richness, indicates that this habitat primarily promotes greater community evenness. Its complex three-dimensional architecture likely provides a wider array of niches, refugia from predators, and microclimatic variation, facilitating the coexistence of species at more equitable abundances. Conversely, the simplified structure of the Juniper Plantation Forest supported a similar number of species but with lower evenness ( Figure 2, Table 1 ), demonstrating that a reliance on species counts alone can mask critical aspects of community organization and function. Crucially, our investigation identified a sharp ecological discontinuity that defines the beta-diversity of this system. While the four upland habitats formed a tight cluster in ordination space - implying a high degree of species interchange and shared membership—the Swamp Grassland was consistently and significantly segregated ( Figure 3 ). This stark compositional divergence is a hallmark of a potent environmental filter, where specific abiotic conditions, such as waterlogged soils and distinct hydrology, act as a powerful selective force, permitting only a specialist assemblage to persist. The further segregation of the Swamp Grassland community during the wet season reveals a dynamic interaction, suggesting that seasonal intensification of flooding or vegetation growth strengthens this filter, rendering the habitat even more distinct for part of the year ( Figure 3 ). This establishes the wetland-upland ecotone not as a zone of gradual transition, but as a sharp boundary generating significant beta-diversity. Our multivariate analyses move beyond these community-level patterns to pinpoint the specific taxa and ecological gradients driving the observed assemblies ( Figure 4, Table 2 ). The distinctiveness of the Swamp Grassland was mechanistically explained by its strong association with Otomys typus , a known wetland specialist ( Figure 4 ). This provides a clear, species-level link between a key habitat filter and a taxon uniquely adapted to exploit it. Within the interconnected upland habitat complex, community differentiation was primarily driven by differential abundance responses of generalist species ( Table 2 ). The significant role of Lophuromys flavopunctatus and Stenocephalemys griseicauda in distinguishing the resource rich Helichrysum shrubland from the Juniper plantation underscores that even within the generalist guild, strong habitat preferences exist, likely tied to food resources and cover ( Figure 4, Table 2 ). The Redundancy Analysis effectively synthesized these complex relationships, revealing two primary, independent axes of community variation: a primary gradient of seasonal abundance and resource phenology, and a secondary gradient of soil moisture that cleanly isolates the Swamp Grassland and its specialist fauna ( Figure 4 ). Collectively, our findings provide strong, nuanced support for our initial hypotheses. We confirmed that habitat type is a primary governor of diversity and composition ( Figures 2, 3 ), while season dictates overall abundance ( Figure 1 ). The empirical validation of the specialist-generalist dynamic—with Otomys typus defining the unique Swamp Grassland realm and generalist abundance shifts fine-tuning community structure within the uplands ( Figure 4, Table 2 ) offers a mechanistic understanding of the assembly rules at play. The primacy of habitat in structuring composition is consistent with montane ecology globally. However, the stability of species richness across our heterogeneous landscape ( Table 1 ) presents an interesting contrast to studies in more fragmented or degraded systems, suggesting that the natural habitats of the Bale Mountains may maintain sufficient connectivity and resource breadth to support a consistent local species pool at this scale. Our work thus provides a critical baseline for this biodiversity hotspot, quantifying the dynamics that underpin community resilience. Notwithstanding these insights, our study robustly identifies spatiotemporal patterns through correlative inference, and the underlying mechanisms warrant direct quantification. Future research should aim to directly measure the drivers we propose, such as food resource availability, fine-scale vegetation structure derived from LiDAR, and predation pressure assessed via camera trapping. Furthermore, mark-recapture studies would be a powerful next step, allowing estimation of vital rates to move beyond abundance correlations and understand the demographic underpinnings of community assembly. Finally, as our study was confined to a single annual cycle, it represents a snapshot of a dynamic system. Long-term monitoring is imperative to understand inter-annual variability and assess the resilience of these unique communities to escalating pressures from climate change and anthropogenic land use. By establishing the current hierarchical structure of drivers and identifying the critical ecological role of the Swamp Grassland as a filter, our study provides the essential framework upon which such future mechanistic and long-term research can be built. 5. Conclusions This study set out to disentangle the complex spatiotemporal drivers structuring the small mammal community in the afroalpine ecosystem of the Bale Mountains. By employing an integrated, multi-method approach, we demonstrated that the factors governing community structure are not monolithic but are instead metric-dependent. Our investigation yielded three fundamental insights: first, overall community abundance was highly sensitive to seasonal resource pulses, peaking in the Wet season ( Figure 1, Table 1 ), while patterns of alpha diversity were primarily a function of habitat type ( Figure 2 ), with species richness remaining notably stable ( Table 1 ). Second, habitat filtering emerged as the paramount force structuring beta-diversity, creating a stark compositional divide between the unique assemblage of the Swamp Grassland and the cluster of more similar upland habitats ( Figure 3 ). Third, these community-level patterns were driven by a clear interplay between habitat specialists, which define distinct ecological realms, and abundance shifts of widespread generalists, which fine-tune community structure within broader habitat types ( Figure 4, Table 2 ). The broader significance of these findings is twofold. Theoretically, our work underscores the critical importance of assessing multiple facets of community structure simultaneously. A study focusing solely on abundance would have concluded that season is the dominant force ( Figure 1 ), while one focused only on richness would have reported remarkable stability ( Table 1 ). It is only by examining abundance, alpha diversity, and beta-diversity in concert ( Figures 1-4, Tables 1, 2 ) that the full, nuanced picture of community assembly emerges (Bekele, 2022; Rickart et al., 2011). From a conservation perspective, our results provide actionable insights. The identification of the Helichrysum habitat as a consistent hotspot of capture success and diversity ( Figure 2 ), and the Swamp Grassland as a unique reservoir for the specialist Otomys typus ( Figures 3, 4 ), highlights that these habitats are of paramount importance. This suggests that effective conservation strategies cannot focus on a single habitat type but must instead protect a mosaic of habitats to maintain overall community integrity, functional diversity, and the high beta-diversity that characterizes this landscape. In final reflection, the main insight from this study is that a comprehensive understanding of ecological communities requires an integrated approach that captures the different rhythms and rules governing their various components. This research provides a foundational benchmark against which future changes can be measured. The vulnerable Afroalpine ecosystem is facing increasing pressure from climate change and anthropogenic activity. By elucidating how community structure is shaped by the dual pillars of habitat heterogeneity ( Figures 2, 3 ) and seasonal pulsing ( Figure 1 ), our findings provide a predictive framework. We can anticipate that forces which homogenize habitats or disrupt the timing and intensity of seasonal cycles—such as land-use conversion or climate shift—will fundamentally erode the diversity, compositional uniqueness, and ecological resilience of these remarkable small mammal communities. Conflict of Interest Statement The authors declare no conflicts of interest, financial or otherwise, that could influence the design, execution, or interpretation of this study. The funding sources had no role in the collection, analysis, or presentation of the data, nor in the decision to submit the manuscript for publication. Data Availability Statement All the required data are uploaded as supplementary material ACKNOWLEDGEMENTS I would like to extend my heartfelt gratitude to the African Center of Excellence for Innovative Rodent Pest Management and Biosensor Technology Development (ACE IRPM&BTD) for their generous sponsorship of this research project. My profound appreciation goes to my advisors, Prof. Alfan Abeid Rija and Prof. Abdul A.S. Katakweba, for their invaluable guidance and unwavering support throughout this endeavor. I also wish to honor the memory and contributions of the late Prof. Loth. S. Mulungu and extend my thanks to Dr. Meheretu Yonas of the Swedish University of Agriculture for their insightful collaboration. This work was greatly facilitated by the kind cooperation of Mekelle University, Ethiopia. We are also indebted to the Ethiopian Wildlife Conservation Authority (EWCA) for granting the research permit that allowed us to carry out our fieldwork seamlessly. A special note of thanks must go to the staff and local community of the Guassa Menz Community Conservation Area, whose hospitality and support were indispensable to this study. Finally, I am deeply grateful for the tremendous field assistance provided by Mr. Getabalew Assefa, Mr. Abebe Gossim, Mr. Demeke, and Mr. Demis Mamo, whose hard work and local expertise were vital to our efforts. Compliance with ethical standards All animal capture and handling procedures were reviewed and approved by the Ethiopian Wildlife Conservation Authority (Permit No: EWCA 31 /249/210). Necessary research and collection permits were also obtained from Sokoine University of Agriculture and the Guassa menz community conservation area. All methods were performed in accordance with the American Society of Mammalogists’ guidelines for the use of wild mammals in research. 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Supplementary Material File (list of figures for aje submision.docx) Download 1.80 MB File (list of tables for aje submision.docx) Download 13.40 KB File (raw data.pdf) Download 192.93 KB Information & Authors Information Version history V1 Version 1 29 November 2025 Copyright This work is licensed under a Non Exclusive No Reuse License. Keywords community ecology ecological experiment ecosystem terrestrial vertebrate Authors Affiliations ABRAHAM DESALEGN 0000-0002-7954-7716 [email protected] Mekelle University View all articles by this author Abdul Katakweba Sokoine University of Agriculture College of Veterinary Medicine and Biomedical Sciences View all articles by this author Alfan Rija Sokoine University of Agriculture View all articles by this author Metrics & Citations Metrics Article Usage 205 views 152 downloads .FvxKWukQNSOunydq8rnd { width: 100px; } Citations Download citation ABRAHAM DESALEGN, Abdul Katakweba, Alfan Rija. 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