Post-fire recolonization of dry deciduous forests by lemurs in northwestern Madagascar

Post-fire recolonization of dry deciduous forests by lemurs in northwestern Madagascar | 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 Post-fire recolonization of dry deciduous forests by lemurs in northwestern Madagascar Naina Ratsimba Rabemananjara, Misa Miaritiana Rasolozaka, Marie Odile Ravolanirina, and 6 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-5735404/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 06 Oct, 2025 Read the published version in Biodiversity and Conservation → Version 1 posted 11 You are reading this latest preprint version Abstract Wildfires significantly threaten biodiversity, especially in tropical regions like Madagascar, where unique ecosystems face ongoing habitat loss and degradation. This study investigated the effects of forest fires on lemur abundance, species richness, and their ability to recolonize burnt vegetation in Ankarafantsika National Park (ANP), the largest protected dry deciduous forest in northwestern Madagascar. ANP hosts eight lemur species with one diurnal ( Propithecus coquereli ), two cathemeral ( Eulemur mongoz , E. fulvus ), and five nocturnal species ( Avahi occidentalis , Lepilemur edwardsi , Cheirogaleus medius , Microcebus murinus , and M. ravelobensis ). Eighteen sites with varying fire histories (1 to > 35 years post-fire) and adjacent unburnt forest parts were surveyed using diurnal and nocturnal distance sampling. Transects included burnt (700 m) and unburnt (500 m) sections. Generalized linear mixed models (GLMMs) assessed the effect of fire variables such as time since the last fire, number of fires, intervals between fires, and fire severity on lemur abundance and species richness. A full lemur community was observed only in unburnt forests and areas with extended post-fire recovery (≥ 23 years). Fires negatively impacted E. fulvus and L. edwardsi , while they did not significantly affect the abundance of small nocturnal species ( C. medius , Microcebus spp.). Lemur species richness was higher in unburnt zones and decreased with an increasing number of fires. These findings reveal the need for long recovery periods for lemur communities post-fire, suggest species-specific fire vulnerabilities, and demonstrate significant faunal impacts of this destructive driver of landscape transformation. lemur abundance species richness forest fire recolonization Ankarafantsika adaptability Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Figure 8 Introduction Fires have a long history of shaping environments with evidence dating back to the late Carboniferous (Scott 2000 ). Before humans started to use fire routinely, lightning was likely the main natural source of wildfire (Roberts 2000 ). Consequently, most terrestrial ecosystems have evolved alongside low frequencies of fire for millions of years. Meta-analyses have revealed that more than 50% of the global terrestrial habitats even need occasional fires to maintain viable environmental conditions (Pausas and Keeley 2009 ). Moreover, it was suggested that fires may have played an important role in the origins and distribution of some particular ecosystems, specifically those with rather open vegetation formations (e.g., savanna, grasslands) (Bond and Keeley 2005 ; Shlisky et al. 2007 ). However, even though some ecosystems may have evolved adaptations to fire, modern changes in the fire regime (i.e., anthropogenic increase in fire frequency, extent, or intensity) may impose severe challenges due to shorter time windows for regrowth and regeneration. Shlisky et al. ( 2007 ) showed that approximately 60% of the world's land-based habitats have undergone recent alterations in their fire regime. At least 20% of these are fire-sensitive habitats, where most species have not evolved traits to survive or regenerate after fires and are therefore highly susceptible to damage from wildfires (e.g., tropical moist forests). Despite covering only 6% of the Earth's surface, tropical forests harbor more than 80% of the planet's terrestrial biodiversity (Gardner et al. 2009 ). However, anthropogenic activities increasingly threaten this exceptional biodiversity, particularly through landscape degradation and habitat fragmentation (Wilson et al. 2015 ). A primary contributor to these disturbances is the intentional use of fire for land conversion, notably in slash-and-burn agriculture, where fire serves to clear land for agricultural expansion or to improve soil fertility (Heinimann et al. 2017 ; Curtis et al. 2018 ). Such practices accelerate deforestation and destabilize these ecosystems, further contributing to their degradation (Foley et al. 2005 ; Curtis et al. 2018 ). Many ecosystems, such as tropical rainforests, are particularly vulnerable to fire due to their lack of adaptations to frequent fire events, making them highly susceptible to shifts in fire regimes driven by human activities (Dwyer et al. 1998 ; Bond and Keeley 2005 ; Shlisky et al. 2007 ; Dantas and Pausas 2013). In contrast to fire-adapted, temperate biomes, plant species in these environments typically lack protective traits such as thick bark or deep rooting systems, leaving them susceptible to fire-induced mortality (Bond and Keeley 2005 ). Severe forest fires can destroy tree canopies, lead to the loss of understory vegetation, increase soil exposure, and enhance erosion (Bond and Keeley 2005 ). In the next step, invasive, fire-tolerant species can establish themselves in these burnt and degraded forests, and these pioneer species can outcompete native vegetation and alter ecosystem structure substantially (Bond and Keeley 2005 ). A central question in the study of fire ecology is how fires affect the composition and integrity of wildlife populations. While adaptive responses in fire-prone environments have been widely studied in plants, animal responses have received comparatively less attention (Pausas and Parr 2018 ). Animals are generally less likely to survive in the flame zone of a fire, but many are mobile enough to flee the immediate fire zone and seek shelter in fire-protected microhabitats (Pausas and Parr 2018 ; Nimmo et al. 2018). For example, the southern brown bandicoot ( Isoodon obesulus ) digs burrows for shelter in a fire-prone environment (Long 2009 ). Certain animals even modify their surroundings to protect themselves, such as some termite species (Macrotermes spp. ) which construct mounds that buffer against heat (Korb and Linsenmair 1999 ). Some birds like the malleefowl (Leipoa ocellata) construct large nesting mounds made of soil and organic material like leaf litter that reduce litter fuel loads, leading to decreased fire intensity and potentially creating fire refuges (Smith et al. 2017 ). Larger species such as elephants (Loxodonta africana and Elephas maximus) indirectly reduce the fire risk and alter their environment by consuming potential fuel materials (Holdo 2007 ). Although some species show adaptations and responses to fire, fire can still affect their abundances and species richness. These effects largely depend on the species-specific environmental plasticity and the different components of the fire regime (e.g. frequency, history, severity) (González et al. 2022 ). Fires can reduce animal abundance and diversity by destroying habitats and causing direct mortality, especially in species lacking adaptations to escape or withstand fire. Amphibians such as the spotted salamander (Ambystoma maculatum) (Fontaine and Kennedy 2012 ), which depend on moist environments, experience significant population declines in regions frequently exposed to fire. On the other hand, fires can create a mosaic of heterogeneous habitats, enhancing food availability and shelter for species (Russell et al. 1999 ). In grasslands and savannas, for example, fires promoted the growth of early successional plants, attracting herbivores like white-tailed deer (Odocoileus virginianus) and the eastern cottontail rabbit (Sylvilagus floridanus) , which thrive on fresh vegetation regrowth (Russell et al. 1999 ). Fires can also open the forest canopy, boosting insect populations and benefiting bird species like downy woodpeckers (Picoides pubescens) (Nimmo et al. 2018). Furthermore, quickly recolonizing post-fire habitats may reduce feeding competition in less populated areas and thereby convey energetic benefits for flexible individuals (Pausas and Parr 2018 ). Post-fire wildlife dynamics and thereby the recolonization potential of different animal species are still understudied particularly in the tropics and on longer time scale, but may offer key insights to understanding co-evolutionary processes in fire-prone ecosystems (González et al. 2022 ). The island of Madagascar is known for its high levels of endemism (Myers et al. 2000 ) along with the high rate of habitat degradation due to human activities, which pose a major threat to its biodiversity. Consistent with other tropical regions of the world, anthropogenic fires are a pervasive driver of landscape change and habitat degradation across Madagascar (Kull 2000 , Phelps et al. 2022 ). Fires are particularly prominent in the western, drier parts of the island (Phelps et al. 2022 ) and were described as a destructive force in modern-day Madagascar, the “isle of fire” (Kull 2000 ; Frappier-Brinton and Lehman 2022 ). However, fires may already have played a role during the formation of its ecosystems and species assemblages, as indicated by historical evidence of fires predating human arrival (Burney 1987 ; Gasse and Van Campo 2001 ; Teixeira et al. 2021 ). The ancestors of extant lemurs arrived on the island about 60 million years ago and evolved into more than 100 different species (Herrera and Dávalos 2016 ). However, the diversification of many extant genera (e.g., Microcebus ) occurred quite rapidly and rather recently within the last few million years (Poelstra et al. 2021 ; Van Elst et al. 2024 ). It is possible that adaptations to fire evolved over these time spans in habitats also prone to fire today. This hypothesis gains support from an analysis of the charcoal stratigraphy of three sediment cores from central Madagascar, which showed that late Pleistocene and early- to mid-Holocene sediments deposited before human settlement often contained more charcoal than post-settlement and modern sediments (Burney 1987 ). Lemurs are a particularly suitable and important group of study species to investigate the effects of forest fires on wildlife, as they have largely different ecologies, perform key ecosystem functions (e.g., pollination, seed dispersal, food web interactions), and the presence of a complete lemur assemblage indicates intact forest habitats (Muldoon and Goodman 2015 ). Finally, lemurs are important flagship species for conservation and ecotourism in Madagascar (Mittermeier et al. 2023 ). To date, there is insufficient information on whether lemurs or other wildlife living in the dry forests of western Madagascar possess at least some resilience against fires that are increasingly threatening and affecting the dry forest habitats of the island. The Ankarafantsika National Park (ANP) is located in northwestern Madagascar and serves as a perfect model area for studying the effects of fire on wildlife. In recent years, increasing fire intensity and severe burning episodes in the region threaten the remaining forest habitats and wildlife (Schüßler et al. 2022). ANP is inhabited by eight lemur species that differ in daily activity, body mass, and also in their IUCN conservation status, ranging from ‘Least Concern’ ( Microcebus murinus ) to ‘Critically Endangered’ ( Propithecus coquereli , Eulemur mongoz , Table 1 ). Previous work also suggested that the natural topography of ANP impacts at least the abundance of the two mouse lemurs differentially (Rakotondravony and Radespiel 2007 ). Table 1 Lemur species occurring in Ankarafantsika National Park with activity, body mass (Mittermeier et al. 2023 ), and IUCN conservation status (IUCN 2020 ). LC = Least Concern, VU = Vulnerable, EN = Endangered, CR = Critically Endangered. *Hibernates from May to mid-September Species Vernacular name Activity Body mass (g) IUCN status Propithecus coquereli Coquerel’s sifaka diurnal 3,700-4,300 CR Eulemur mongoz Mongoose lemur cathemeral 1,100-1,600 CR Eulemur fulvus Common brown lemur cathemeral 1,700-2,100 VU Avahi occidentalis Western woolly lemur nocturnal 800-1,100 VU Lepilemur edwardsi Milne-Edwards sportive lemur nocturnal 854-1,200 EN Cheirogaleus medius* Fat-tailed dwarf lemur nocturnal 120–270 VU Microcebus murinus Gray mouse lemur nocturnal 58–67 LC Microcebus ravelobensis Golden-brown mouse lemur nocturnal 56–87 VU We aim to investigate how past fires affected the different lemur species in the dry deciduous forests of ANP in northwestern Madagascar. By studying their ability and timespan for re-colonizing areas that have been previously burnt, we aim to shed light on the resilience of lemurs to forest fires. Specifically, we investigate the impacts of (1) the time since the last forest fire, (2) the maximum past fire intensity, (3) the number of past forest fires, and (4) the minimum interval between recurrent fires on lemur abundance and species richness. The relevant fire history for all study sites was provided by a parallel study for a period of 35 years (1988–2022) for which yearly remote sensing was available (Rasolozaka et al. 2024 ). To test for additional confounding effects, the influence of terrain and elevation for lemur abundance and lemur species richness is also evaluated. Materials and Methods Study area We conducted our study in Ankarafantsika National Park (ANP), which is the largest protected remaining dry forest tract in northwestern Madagascar (-16°08'60"S, 46°57'0" E) and covers an area of 1,350 km² (Fig. 1 ). It possesses some topographical complexity, as it includes river valleys at about 50 m above sea level (a.s.l.), slopes, and a rising topography from north to south where a calcitic plateau reaches peak elevations of up to 350 m a.s.l. and forms cliffs in many places in the east and south (Alonso et al. 2002 ). ANP is crucial for the conservation of numerous endangered species, such as the Critically Endangered Coquerel’s sifaka (Propithecus coquereli) , the endangered largest terrestrial carnivore of Madagascar (Cryptoprocta ferox) , and the Critically Endangered Malagasy fish eagle (Haliaeetus vociferoides) (Schwitzer et al. 2013 ; Barcala 2009 ; Razafimanjato et al. 2014 ). ANP features a seasonally dry tropical forest that includes a mix of dry deciduous forests and natural dry thickets at higher elevations, moist riverine forests in upstream valleys and around lakes at lower elevations, as well as Raphia swamp forests in downstream valleys (Goodman et al. 2021 ). Forest vegetation can be found on the dry plateau, slopes, and valleys. The average annual rainfall ranges from 1,000 to 1,500 mm, with most rain falling during the rainy season which lasts from November to April, while the almost rainless dry season covers the period from May to October (Alonso et al. 2002 ). Use of remote sensing data for study site selection We used remote sensing data to identify 18 suitable study sites with different fire histories across ANP prior to the start of the fieldwork. Rasolozaka et al. ( 2024 ) detailed the necessary working steps. In brief, this process involved screening annual Landsat satellite images (30 x 30m resolution) from 1988–2023, from which specific fire and vegetation indices were calculated to quantify the occurrence and extent of fires across the years in all study sites. M. Rasolozaka used the Normalized Burn Ratio (Key and Benson 2006 ) to identify fires, i.e., whether pixels were burnt or unburnt in a given year. By subtracting the pre- and post-fire NBR values (dNBR), M. Rasolozaka calculated the fire severity for all study sites and years (Keeley 2009 ). The rationale for our study site selection was to identify areas where an unburnt area (i.e., never burnt across the 35 years) bordered on a burnt area with a variable number of years since the last fire and a variable total number of fires experienced over the previous 35 years. Such sites were chosen to provide a mixed design for subsequent data analyses while always controlling for site-specific ecological characteristics. After completing the fieldwork, all remote sensing data were re-analyzed and thoroughly evaluated alongside field data, providing ground truthing for the final reconstruction of the site-specific fire history. This re-evaluation of the remote sensing data together with field-based records of past fire traces (e.g., fire scorches on trees and charcoal traces in soil, Rasolozaka et al., 2024 ) revealed that the prior classification of some study sites needed to be revised. Specifically, parts of the unburnt areas had indeed burnt in some sites (sites 5, 4, 13, 2, 12, 15), and some sites had experienced a heterogeneous fire history in which not all parts of the study transect were burnt during the same years (details in Supplementary Table S1 ). Study sites and spatial arrangement of fieldwork Prior to fieldwork, we selected 18 study sites that varied in their specific fire history (Table 2 ). Six sites were visited in 2022 (September - December) and a further 12 sites were visited in 2023 (May - November). The sites varied in the number of years since the last fire (1 to > 35 years), in the number of fire occurrences in each site over the last 35 years (1–7 fires), in the minimum interval between two fires (1–31 years), and in the maximum burn severity that each site experienced in the past 35 years (Table 2 ). A 1.2 km-long transect with a paired design was established in each site, which was flagged and mapped with a GPS device (Garmin GPSMAP 64) every 10 m. It consisted of a stretch of 500 m in the burnt forest (burnt zone), running perpendicularly to the fire edge, and a complementary stretch of 500 m in the unburnt forest (unburnt zone), also pointing away from the fire edge, but shifted with the burnt transect by 200 m. We connected these two transect parts by a transect of 200 m length running in parallel to the fire edge, about 50 m into the burnt forest, i.e., in the transition zone between burnt and unburnt forest. Lemur abundance survey We conducted three diurnal and three nocturnal systematic distance sampling surveys per site to monitor the 1.2km transect for sightings of any lemur species (Buckland et al. 2001 ). With four observers, we walked each transect quietly and slowly at a speed of approximately 0.5km/h (Rakotondravony and Radespiel 2007 ). Each transect was visited over three days in the morning (06h30-08h30) and evening (18h15–20h30). We used headlamps and flashlights during night surveys to identify and differentiate the eight species. Upon each lemur encounter, we recorded the number of individuals and determined the species and the position along the transect to the closest 10 m. We calculated the abundance of each species (dependent variable) separately for each partition of the transect that had a different fire history and initially also distinguished between the burnt zone and the transition zone (Table S6). Encounter rates were too small to determine population densities for any of the species for any partition of the transect (Buckland et al. 2001 ). Therefore, and to be able to subsequently model variations in abundances from different study sites, we calculated mean encounter rates per 100m transect as a proxy for abundance for each transect partition with specific fire history and for each species separately with the following formula: $$\:\text{A}\text{b}\text{u}\text{n}\text{d}\text{a}\text{n}\text{c}\text{e}=\frac{N}{L*n}*100$$ Where N: Total number of individuals encountered during all three surveys along transect partition with specific fire history L: Total length of transect partition (in meters) n: Number of surveys We further determined the lemur species richness (dependent variable) as the count of the total number of species observed in each transect partition across the three successive surveys. Modeling the effect of topographic and fire-related variables on lemur abundance and lemur species richness Due to the limited number of sightings, it was not possible to generate statistical models for Propithecus coquereli and Eulemur mongoz . For these two species, we only present descriptive data on the relationship between the species' presence and the time that elapsed since the last fire. Cheirogaleus medius hibernates during large parts of the dry season from May to mid-September (Müller and Thalmann 2002 ). Therefore, the species was only included in the abundance analysis for sites visited between September to December. All modeling approaches with species-specific abundances and lemur species richness were performed in R v. 4.4.0 (R Core Team 2024) using RStudio v.4.1.748 (RStudio team 2020 ) interface. First, we assessed the normality of all continuous variables using the Shapiro–Wilks test (Shapiro and Wilk 1965). If a variable was not normally distributed, we applied a logarithmic or square root transformation. We visually inspected the Q–Q plots before and after the transformation to ensure improvement. We modeled the impact of two topographic and five fire-related fixed factors on species-specific abundances and lemur species richness. First, we analyzed the effect of terrain (valley, slope, and plateau) and elevation (m above sea level) on lemur abundance and lemur species richness. Second, we modeled the impact of five fire-related parameters on the abundance of six lemur species and on lemur species richness. Fire-related parameters were the forest zone (unburnt/burnt), the total number of fires, the year since the last fire, the minimum interval between two fires, and the maximum fire severity in each transect partition. When fitting multivariate models, we often experienced convergence problems (Finch and Finch 2017 ; Zhang et al. 2017 ). This was likely due to the limited data set (66 partitions in 18 study sites, Table S1 ) and the resulting lack of statistical power when fitting several variables together. Therefore, we decided to calculate only univariate models for all parts of the analysis. To identify the determinants of species-specific abundances (dependent variables), we fitted Generalized Linear Mixed Models (GLMMs) using the package glmmTMB v.1.1.10 (Brooks et al. 2017 ) which allows for the implementation of different model families according to data structure and distribution. With the package DHARMa v.0.4.7 (Hartig and Hartig 2022 ) we assessed the quality of each model by testing for over- and underdispersion, outliers, and zero inflation of the residuals. In case of a zero inflation, we added the command “ziformula” to the model; in the case of overdispersion and underdispersion, we added the command “dispformula” to the model (Brooks et al. 2017 ). We selected the Tweedie family for modeling in all cases. Its regression models can accomodate highly right-skewed, as well as symmetric and heavily tailed data distributions (Ma et al. 2018 ). As the lemur species richness (dependent variable) is always an integer number with an upper observed limit of six species, we first used the cbind () function to relate the number of observed lemur species to the number of not-observed species in each partition. For this variable, we fitted Generalized Linear Mixed Effect models (GLMMs) using the package lme4 (Bates et al. 2015 ) with the binomial family, and the package DHARMa (Hartig and Hartig 2022 ) to assess the model residuals (model dispersion, outliers). We included study site and month as independent random factors when modeling species abundances, and study site as well as transect partition length as independent random factors when modeling the lemur species richness. We verified the relevance of significant variables by testing whether introducing a specific variable explained significantly more variation in the dataset compared to a null model that only contained random factors and no explanatory variable. For this comparison, we used the ANOVA function in R with a threshold p-value of less than 0.05 indicating significance. During all species-specific analyses of the impact of forest zones (burnt, transition, and unburnt), we found no significant difference between burnt and transition zones, even when applying the Tukey post-hoc test with the R package emmeans v.1.5.1 (Lenth et al. 2021 ). Given these results, we categorized both transition and burnt transect zones as burnt throughout. To correct for potential error accumulation by multiple testing, we used the Holm-Bonferroni Method to adjust the threshold for significance (Holm 1979 ). We applied this correction separately to each species and the respective topographic and fire-related variable set. We then compared the adjusted alpha with the observed p-values (Holm 1979 ) and reported the results before and after this correction. We considered p = 0.05 as the level of significance before correction. Analyses were based on two sets of data. The first set included the full dataset with all transect partitions in all sites. This dataset was used to test the impact of the topographic variables and two of the fire-related variables (zones and number of fires). The second dataset included only the dataset for the burnt partitions and was used for modeling the remaining three fire variables (time since the last fire, minimum interval between two fires, and maximum fire intensity). We generated figures using Microsoft Excel and the R packages ggplot2 v.3.4.0 (Wickham et al. 2016 ) and ggpubr v.0.6.0 (Kassambara 2018 ). Results The impact of terrain and elevation on lemur abundance and species richness During surveys, we encountered all eight lemur species present in the ANP, but only six of them had sufficient sightings for modeling abundances (Table S2). After correcting for multiple testing, the large Eulemur fulvus had significantly higher abundance in valleys than on the plateau (Estimate valley = 2.354, p = 0.007; Fig. 2 a; Table S3), but the abundance on slopes did not differ from that in valleys or on the plateau (Tukey test: Estimate slope−valley = -3.011, p = 0.115; Estimate plateau−slope = 0.657, p = 0.897; Table S5). Elevation also affected the abundance negatively (Estimate = -1.807, p = 0.025), i.e., we found less E. fulvus in areas of higher elevation, even after correcting for multiple testing (Fig. 2 b; Table S3). The abundance of the middle-sized Lepilemur edwardsi was not significantly impacted by terrain or elevation (Table S3) while the middle-sized Avahi occidentalis showed a statistical trend for lower abundance in the valley (Estimate valley = -2.046, p = 0.077; Table S3), but no longer after correcting multiple testing. Terrain and elevation did not influence the abundance of Cheirogaleus medius (Table S3). In contrast, the golden-brown mouse lemur, Microcebus ravelobensis , was significantly more abundant in valleys than on the plateau (Estimate valley = 0.920, p < 0.0001; Fig. 3 , Table S3). The post-hoc Tukey test revealed that the abundance in the valleys was also significantly higher than on slopes (Estimate slope−valley = -0.524, p = 0.042; Table S5), although there was no significant difference between the abundance on plateau and slopes (Estimate plateau−slope = -0.318, p = 0.265; Fig. 3 , Table S5). There was also a statistical trend for a lower abundance at higher altitudes (Estimate = -0.060, p = 0.067; see Table S3), but not after correcting for multiple testing. In the case of the grey mouse lemur, Microcebus murinus , neither terrain nor elevation showed a significant impact on abundance, and a statistical trend for a lower abundance in the valley than on the plateau (Estimate valley = -1.221, p = 0.053, Table S3) did not persist after correcting for multiple testing. Finally, the lemur species richness was also not statistically impacted by terrain or elevation (Table S3). The influence of time since the last fire on lemur presence/absence The time since the most recent fire in the forest ranged from one to over 35 years (Fig. 4 ). All species except Eulemur mongoz were recorded along transects in forest that had not been burned during that period (unburnt, Fig. 4 ). The three largest species Propithecus coquereli , E. mongoz , and E. fulvus were almost only observed in unburnt zones or burnt zones with fires older than 23 years. P. coquereli was only sighted along transects in four sites altogether (sites 1, 8, 11, 13) but was encountered outside the study transects in these same sites and in four others (sites 2, 15, 21, 24). Only one individual of P. coquereli was encountered in the transition zone (burnt) in one of our study sites with a very recent fire (1 year old). E. mongoz was sighted along transects in only one site and outside the study transects in six more sites. In contrast, the medium-sized lemurs Avahi occidentalis and Lepilemur edwardsi were observed in 69.2% and 76.9% of the unburnt zones, respectively (Fig. 4 ). They were also recorded in 37.0% (n = 10) and 33.3% (n = 9) of the burnt zones with different fire ages, respectively, even in those that experienced fires less than six years ago. Likewise, Cheirogaleus medius was observed in 50% of the unburnt zones as well as in 62.5% of the burnt zones, sometimes in zones of relatively young fires (2–6 years old) but also in old fire zones (Fig. 4 ). The two smallest lemur species, Microcebus ravelobensis and Microcebus murinus , were found in almost all unburnt zones ( M. ravelobensis : 100%, M. murinus : 76.9%) and in almost all burnt zones ( M. ravelobensis : 96.3%, M. murinus : 66.7%) ranging from the most recently burned (1 year post-fire) to those with older fires. The impact of fire-related variables on lemur abundance and lemur species richness We detected a negative impact of the number of fires on the abundance of Eulemur fulvus (Estimate = -1.267, p = 0.041, Fig. 5 , Table S4), but only before Holm-Bonferroni correction. Qualitatively, observations of E. fulvus dropped to almost zero, whenever there have been more than three fires in a site (Fig. 5 ). We also noted a statistical trend for the positive impact of the number of years since the last fire on E. fulvus abundance (Estimate = 0.133, p = 0.051, Table S4), but only before Holm-Bonferroni correction. The statistical modeling did not reveal any other impact of fire-related variables on the abundance of E. fulvus . The abundance of the medium-sized Avahi occidentalis in the unburnt zone was higher than in the burnt zones and negatively associated with the number of fires but was rejected after we applied the Holm-Bonferroni correction (Table S4). Qualitatively, we noted that this species was absent from sites that burned more than three times and whenever the minimum interval between two fires was less than eight years (Table S2). The abundance of the medium-sized Lepilemur edwardsi was impacted by the burn status (burnt vs. unburnt), the number of fires in a site, the minimum time interval between fires, and the maximum fire intensity (Table S4) even after correcting for multiple testing. Specifically, the species was significantly more abundant in unburnt compared to burnt areas (Estimate Unburnt = 0.923, p = 0.020; Fig. 6 a, Table S4). Moreover, the number of fires negatively influenced the abundance of L. edwardsi (Estimate = -1.131, p = 0.003, Fig. 6 b) to the point that the species was no longer observed at sites that had experienced more than three fires. Additionally, larger minimum intervals between fires lead to increased abundance of L. edwardsi , and this species was not found in burnt zones with minimum fire intervals of less than eight years (Estimate = 1.113, p = 0.012; Fig. 6 c). Lastly, higher maximum fire severities negatively affected the abundance of L. edwardsi (Estimate = -21.338, p = 0.012) leading to an absence of observations from sites with very high maximum severity (Severity > 0.12, Fig. 6 d). The other variables had no impact on L. edwardsi abundance (Table S4). The abundance of Cheirogaleus medius was significantly and positively impacted by the year since the last fire (Estimate = 0.358, p = 0.041) but this effect was no longer significant after Holm-Bonferroni correction. No other fire-related variable showed any significant effect on the abundance of C. medius (Table S4). The abundance of Microcebus murinus showed a statistical trend for a positive impact of the minimum interval between two fires (Estimate = 0.681, p = 0.071) but only before Holm-Bonferroni correction. No other fire-related variable showed a statistical effect on this species (Table S4). Finally, no fire-related variable showed any statistical effect on the abundance of Microcebus ravelobensis . (Table S4). The burn status and the number of fires (Fig. 7 a, b; Table S4) statistically influenced lemur species richness. Specifically, we found a significantly higher lemur species richness in the unburnt zone compared to the burnt zones (Estimate Unburn = 1.140, p < 0.0001; Fig. 7 a) even after Holm-Bonferroni correction. In addition, the number of fires negatively influenced lemur species richness (Estimate = -1.175, p < 0.0001, Fig. 7 b), also persisting after Holm-Bonferroni correction. The year since the last fire showed a positive statistical trend (Estimate = 0.172, p = 0.058; Table S4) for an influence on lemur species richness, likewise, the maximum fire severity showed a negative statistical trend (Estimate = -3.583, p = 0.054; Table S4), but both trends were rejected after correcting for multiple testing. The minimum interval between fires showed no statistical effect. Discussion Effects of forest fires on large-sized lemur species Due to the rarity of sightings, we could not conduct statistical analyses on two of the largest lemur species in Ankarafantsika National Park (ANP), the Critically Endangered Propithecus coquereli and Eulemur mongoz . P. coquereli was only encountered in four of 18 sites (22%) along transects and was then mainly observed in unburnt forests. As the largest lemur species in ANP that predominantly moves by vertical clinging and leaping, P. coquereli relies on robust, vertical substrates for movement, such as trunks and branches of large trees (Mittermeier et al. 2023 ) which were shown to be diminished in numbers and connectivity at least in one recently burnt part of the park (Percival et al. 2024 ). The importance of suitable substrates for locomotion is also suggested by the observation that this species was never observed on the ground inside the forest (Rabemananjara and Radespiel, unpubl. Data), although such observations were made in a non-forest matrix in another study outside ANP (Ramilison et al. 2021 ). Previous research by Kun-Rodrigues et al. ( 2014 ) revealed that P. coquereli is mostly found in the forest core and avoids forest edges, suggesting sensitivity to disturbance or possibly edge-related ecological changes (Steffens and Lehman 2018 ). Ecological changes like those at forest edges can also be expected as a result of forest fires, as canopy cover then diminishes which likely drives cascading effects on microclimate, vegetation structure, and floristic composition (Barlow and Peres 2008 ; Percival et al. 2024 ). Another explanation for the fire sensitivity of P. coquereli may be related to its feeding habits. P. coquereli has a predominantly folivorous diet (Mittermeier et al. 2023 ), and plant diversity tends to be higher in the forest interior than at the edge (McGoogan 2011 ). As forest fires very likely also lead to a reduction of plant diversity (Agus et al. 2018), burnt zones may not provide sufficient food resources for P. coquereli . Such a loss of floristic diversity may be particularly expressed in forests that experience recurrent fires that likely lead to floristic shifts in entire plant communities towards pioneer, fire-resistant species (Da Silva et al. 2018 ). Indeed, P. coquereli was never encountered in burnt zones that had experienced multiple fires over the last 35 years. However, we did once encounter a group of P. coquereli in the transition zone of a one-year-old fire and sighted another group twice in a 24-years post-fire zone. While the sightings in the transition zone may indicate a rather exploratory use of the recently burnt zone, the presence in the older fire zone suggests that this species may eventually return to previously burnt areas once conditions become favorable again. However, our current study does not provide conclusive evidence of such recolonization events yet. Further research and data collection, including longer behavioral observations of P. coquereli living close to burnt forest vegetation are needed to illuminate its habitat and resource use and thereby potential adaptations to fire-affected areas. Eulemur mongoz was encountered only twice during our entire study (once in the burnt and once in the unburnt zone of the same study site) when systematically surveying transects. One of the reasons for this rarity of sightings could be its seasonally changing activity rhythm that switches between largely diurnal activity during the rainy season to largely nocturnal activity during much of the dry season i.e., during our study period (Curtis 2006 ). This activity change to nocturnality could complicate their detection, in particular, since E. mongoz has a rather weakly developed tapetum lucidum (Martin 1995 ) and forms pairs or small family groups (Curtis and Zaramody 1999 ) that are less conspicuous than, for example, larger social groups formed by Eulemur fulvu s (Mittermeier et al. 2023 ). However, one of the two sightings during our surveys was made during daytime in the middle of the dry season (July), and E. mongoz was furthermore confirmed near six other study sites (six times seen during daytime, one time seen during nighttime), but not along the survey transects (Rabemananjara and Radespiel, unpubl. results). These opportunistic sightings suggest that this species was indeed rare along the study transects but not impossible to find inside ANP during the dry season. One explanation for their rarity in ANP could be their diet, which was suggested to be highly specialized, encompassing nectar, fruits, flowers, and leaves from a rather limited number of plant species (Sussman and Tattersall 1976 ). If those food plants were not available in the entire park, but only in some favorable forest parts, such a dietary specialization could make them extremely vulnerable to disturbances including fires. However, our sighting of E. mongoz in one forest zone that had burnt 24 years ago also shows some flexibility and resilience at least given older fires and their long-term effects. Nevertheless, the overall low number of encounters with E. mongoz aligns with other studies. Steffens et al. ( 2020 ) reported no sightings of the Critically Endangered E. mongoz during 22.15 km of survey effort in the continuous forest of ANP, while Steffens and Lehman ( 2018 ) found none over 468.18 km of surveys in 42 forest fragments in western ANP. The abundance of Eulemur fulvus appeared to be negatively influenced by the number of fires. However, the statistical support for this parameter was relatively weak which may be due to a limited statistical power or to a plastic fire response of this species. Such a plasticity may be plausible, as this species was described as a flexible generalist (Sato 2013 ) which is also suggested by its wide distribution that includes dry forests in northwestern Madagascar as well as humid rainforest habitats in central eastern Madagascar (Mittermeier et al. 2023 ). Our ecological modeling results also indicated a certain affinity to humid conditions, suggesting higher abundances of E. fulvus at lower elevations and therefore under more humid conditions (valleys) than in drier localities at higher elevations (plateau). However, some sensitivity of E. fulvus to fires was suggested by two other findings of our study: First, E. fulvus was rarely sighted in areas that had experienced more than three fires. Multiple fires were already shown to lead to increased levels of dry forest degradation, and it was hypothesized that such degraded patches may undergo very long-term or even irreversible vegetation shifts and ecosystem changes (Percival et al. 2024 ) and thereby may become permanently uninhabitable for E. fulvus . Second, E. fulvus was only sighted in burnt zones of fires that were more than 23 years old suggesting that recently burnt dry forests also do not provide suitable habitat for E. fulvus . Whether they lack essential food resources, suitable substrates for locomotion, and/or cover from predators cannot be answered at present but these questions certainly ask for more specific future research. Compared to other non-human primates, the three large lemurs in ANP require longer periods to recolonize burnt dry deciduous forests. For instance, siamangs in Sumatra ( Symphalangus syndactylus ) were reported to return to burnt areas after approximately 18 years (Lappan et al., 2021 ). Similarly, leaf-eating monkeys ( Presbytis spp.) in Indonesia recolonized burnt forests within six years post-fire, although at low densities (Chapman and Onderdonk, 1998 ). This difference could indicate that large lemurs may be more sensitive to habitat disturbances than other primate taxa, but it may also be the result of much slower vegetation regrowth and regeneration of the dry forests of Madagascar (Percival et al. 2024 ) compared to other more humid tropical forest ecosystems on the island or in other parts of the world. Percival et al. ( 2024 ) already suggested that the dry forests of ANP may have very low resilience to fire compared to fire-adapted dry forests in other parts of the world (Cavender-Bares et al. 2004 ; Mcmichael et al. 2004 ; Lozano et al. 2012 ). However, our data do not support their additional hypothesis that even one fire in this fire-sensitive forest may be sufficient to initiate a permanent transformation of dry forest into a degraded savanna/grassland vegetation. The sightings of the large lemur species in burnt forest parts of ANP after more than 23 years post-fire rather suggest that post-fire regrowth and regeneration are possible in these dry forests, even though it may take several decades. Effects of forest fires on medium-sized lemur species Modeling revealed no significant impact of fires on the abundance of the medium-sized Avahi occidentalis . This species was even recorded in some areas affected by recent fires (1–5 years post-fire), although it was recorded in burnt partitions of only 50% of the study sites. However, it was also observed in only 69% of the unburnt zones, and these mixed sighting results suggest that A. occidentalis may have specific resource requirements (Thalmann 2001 ) that are not uniformly met in ANP, but that may not be necessarily impaired by forest fires per se. Our findings also suggested that A. occidentalis may be sensitive to multiple fires. Specifically, we noted that A. occidentalis was never sighted in burnt zones that experienced more than three fires or whenever the minimum interval between consecutive fires was less than eight years. Similar to the argument made for large lemur species, multiple fires likely cause significant forest degradation (Percival et al. 2024 ). Such degradation may render the habitat unsuitable or permanently uninhabitable for this medium-sized lemur species. Given that A. occidentalis relies on vertical clinging to tree trunks or large branches and leaping between such positions in large trees (Warren and Crompton 1997 ), the availability of suitable arboreal substrates for this species-specific mode of locomotion may be an essential component of suitable habitats, which may be permanently compromised after multiple fires. The medium-sized Lepilemur edwardsi exhibited an even stronger negative response to fires than A. occidentalis . Although L. edwardsi was also occasionally sighted in recently burnt areas (1–5 years post-fire) and in burnt partitions of 38% (n = 7) of the study sites, it was significantly more abundant in unburnt than in burnt zones. L. edwardsi was also absent in areas with more than three fires or when the interval between fires was shorter than eight years. Additionally, L. edwardsi responded negatively to increasing fire severity. We hypothesize that severe fires, particularly the degradation caused by repeated fires (Percival et al., 2024 ), reduce the availability of large trees that serve as substrates for vertical clingers and leapers (Warren and Crompton, 1997 ). Additionally, these fires destroy large trees provide shelters, such as tree holes, which this species uses for daily resting (Rasoloharijaona et al., 2003 ). Sleeping holes of L. edwardsi in ANP were previously described to lie at about 3–8 m height above ground, and sleeping holes had a depth of more than 80 cm and an average wall thickness of 11 cm (Rasoloharijaona et al. 2003 ), which can clearly only be provided by large trees. If this important resource, which makes up 92% of the used shelters, is no longer available, it can be expected that the suitability and possibly the carrying capacity of a degraded post-fire forests drops considerably for L. edwardsi . Post-fire food availability may also play a role in limiting the abundance of L. edwardsi , although further research is needed to investigate this hypothesis. Effects of forest fires on small-sized lemur species Fires did not significantly influence the abundance of the three small-sized nocturnal lemurs like Cheirogaleus medius, Microcebus murinus and M. ravelobensis . All three species were recorded in both recently burned forests (1–5 years post-fire) and in forests affected by older fires (Fig. 4 ). All three species can also be categorized as omnivores, as they can consume flowers, fruits, seeds, invertebrates, insect secretions and even occasionally small vertebrates (Fietz and Ganzhorn 1999 ; Radespiel et al. 2006 ; Joly-Radko and Zimmermann 2010 ; Thorén et al. 2011 ). This dietary flexibility likely allows them to explore burned forests of various ages. Increased sun exposure in these areas may alter temperature and radiation, leading to more insect productivity and changes in fruit types and availability (Hamer 1996 ; Keane et al. 2008 ; Pausas and Keeley 2019 ; Thompson et al. 2022 ). C. medius was already stated to be quite resilient to environmental changes (Rakotoniaina et al. 2016 ). Its habit to hibernate from May to Mid-September in ANP (Müller 1999 ) which partially overlaps with the fire season (July to October), very likely helps to avoid resource shortage during periods of food scarcity, and this ability may further contribute to its resilience in highly modified post-fire zones. However, some studies indicated that C. medius tends to occur at lower densities in disturbed than in undisturbed habitats (Rakotoniaina et al. 2016 ; Hending 2021 ). This aligns with our findings that this species only reached abundances that are comparable to those in unburnt forests when the burnt areas reached a certain age post-fire (> 23 years) (Table S2), suggesting that the availability of suitable resources and habitat structure improves and regenerates with time following a fire. If this preliminary observation would be confirmed in future studies on a larger number of study sites, our lack of statistical support for negative fire effects may also be based on the small sample size (i.e, limited statistical power) with only 11 sites that could be studied after the end of the yearly hibernation period in C. medius . The two small-sized mouse lemurs, on the other hand, were present in zones of very recent to old fires (one to > 35 years old), and a negative impacts of fires on their abundance could not be detected. In other words, Microcebus murinus and M. ravelobensis showed rather similar abundances in recently burnt and older burnt zones, which underlines their high ecological flexibility and confirms previous findings that they inhabit various habitat types of different stages of degradation including secondary forest (Rakotondravony and Radespiel 2009 ; Steffens and Lehman 2018 ; Hending 2021 ; Steffens et al. 2022 ). However, they are not reacting equally to habitat fragmentation (Andriatsitohaina et al. 2020 ; Steffens et al. 2022 ). While M. murinus can even be found in small to very small forest fragments that are embedded in a non-forest savanna matrix, M. ravelobensis is usually found only in larger fragments or in continuous stretches of larger forests (Andriatsitohaina et al. 2020 ). However, in the case of geographically limited burnings in the continuous forest of ANP, the burnt forest usually borders somewhere upon stretches of unburnt forest or forest parts that underwent fires in other years. This may facilitate recolonization by both of these omnivorous generalists quickly after a fire is extinguished (Ramsay et al. 2020 ). The high resilience of both species to fire may also result from their high flexibility when selecting day shelters, which can be liana tangles, shrubs, dead wood or bark, or self-built nests located between the ground and the canopy (Radespiel et al. 1998 ; Radespiel et al. 2003 ; Thorén et al. 2010 ). These shelters are likely available even in moderately degraded post-fire landscapes and may provide critical protection under high risk of predation by raptors, snakes and carnivores (Goodman et al. 1993 ; Karpanty and Wright 2007 ). Finally, mouse lemurs are also able to change their activity patterns and enter daily torpor in case of environmental challenges (Ortmann et al. 1997 ; Schmid 2000 ; Thorén et al. 2011 ) which may contribute further to energetic resilience in post-fire landscapes. Even though this general resilience to fire has thereby been confirmed for the small, nocturnal lemurs in ANP, future research should still investigate the impact of varying fire severity and fire frequency on resource availability, reproductive output, predation risk, space use, and the resulting population sizes of these rather flexible, seasonal species in burnt zones. Even if small lemurs may use burnt forests to some extent, altered food availability and predation risk might still constrain population sizes and modulate stochastic extinction risks over time (James et al. 1997 ). Effects of forest fires on lemur species richness Habitat alteration can significantly affect species richness, particularly for primates, by altering vegetation structure, food availability, and ecosystem stability (Schwitzer et al. 2011 ). Our study has shown that the species richness of lemurs was higher in the unburnt than in the burnt forests. Studies in other tropical forests worldwide have provided similar evidence for a variety of different taxa such as in small and large mammal communities (Santosa and Kwatrina 2020 ), bird communities (Adeney et al. 2006 ; Davis et al. 2016 ; Mardiastuti 2020 ), and in herpetofaunal communities (Russell et al. 1999 ). For instance, unburned forests that maintain an intact canopy cover and have undisturbed food sources such as fruits and leaves have been shown to support a higher diversity of primates in the Kibale National Park, Uganda (Chapman and Onderdonk 1998 ). Burned areas often host primarily generalist species that can adaptively respond to fire-modified landscapes due to their dietary flexibility (omnivory) and variable locomotion that allows mitigating fire-related vegetation changes (Santos et al. 2014). Such generalists in our study were only the smallest lemur species, specifically M. ravelobensis and M. murinus . They are particularly adept at taking advantage of resources that may quickly become available again after a fire, such as insects or seeds that remain in the ground (Radespiel et al. 2006 ; Dammhahn and Kappeler 2008 ; Pruetz and Herzog 2017 ). Cumulative negative impacts of fire on lemur species richness were noted in particular after fires occurred repeatedly in sites. Recurrent fires in ANP were shown to negatively affect large lemurs, medium-sized lemurs, as well as C. medius (see above). Multiple fires are very likely to hinder forest recovery and thereby disturb resource availability (Hermosilla et al. 2019 ), but also may increase the predation risk by elevated exposure to predators in the degraded vegetation (Shanee et al. 2023 ). Moreover, large-scale fire-related vegetation shifts may lead to fragmentation of the remaining inhabitable forest patches that will likely disrupt and constrain movements and dispersal within and between the remaining sub-populations (Nimmo et al. 2019 ; Doherty et al. 2022 ). Finally, all these challenges very likely increase levels of intraspecific competition, may require individual and population-level niche shifts, generate elevated stress levels that may result in decreased reproductive success (Bolnick et al. 2010 ; Martínez-Blancas and Martorell 2020 ). Such effects can pose a significant threat to primate diversity and ecosystem resilience, especially in tropical dry forests (Van Holstein et al. 2024 ). Conservation implications Our study suggests that most lemur species do not recolonize burnt forests quickly. In fact, the lemur community in ANP only reached the level of unburnt forests when the fire happened more than 23 years ago. This slow recolonization may be attributed to various factors, including a lack of key food resources in burnt zones, insufficient substrates such as large trees needed for species-specific locomotion (e.g., clinging and leaping by Avahi and Lepilemur ) or as shelters ( Lepilemur ), and increased risks from predators or human activities in disturbed habitats. Future analyses are needed to investigate the main reasons for the species-specific fire sensitivity in more detail. Given this rather long time span that seems to be necessary for a full recovery of the lemur community after a fire, this study raises serious concerns about the amount of suitable forest habitat remaining for all lemur species in ANP. To obtain a first rough estimate of this value, we determined the area of the forest within ANP that had not experienced any fires in the last 35 years together with the area that was burned 24 years or longer ago (Fig. 8 ). Together, both partitions added up to approximately 466.6 km² (45%) of the vegetation inside ANP that should be potentially suitable for the entire lemur community consisting of eight species in the park. However, species like L. edwardsi, E. fulvus, E. mongoz and P. coquereli face dual threats: habitat degradation and hunting, which has been documented even within protected areas like the Ankarafantsika National Park (Garcia and Goodman 2003 ; Borgerson et al. 2019 ; Sato et al. 2021 ). The Critically Endangered E. mongoz was found to be the rarest of all lemur species in our study, emphasizing the need to study the habitat requirements and environmental constraints of these lemurs in more depth. Their ability or inability to adapt to environmental changes may play a key role in determining their long-term survival in highly altered landscapes. Forest fires significantly influenced lemur communities and may therefore threaten the ecosystem services that are provided by lemurs. Larger lemurs, for instance, play a crucial role in seed dispersal in intact forests (Ganzhorn et al. 1999 ; Lahann 2007 ; Sato 2012 ; Valenta et al. 2013 ). This predominant role of primate seed dispersers contrasts with other regions in the world where birds and bats also contribute substantially to these services (Howe 1986 ; Medellin and Gaona 1999 ; Herrera 2002 ). After multiple fires in ANP, only the two smallest species, the omnivorous, nocturnal mouse lemurs, seemed to be resilient and adapt to the corresponding degradation in post-fire habitats. However, their seed dispersal capacity is limited due to their small body size (Ramananjato 2024 ) which likely biases forest regeneration toward smaller-fruited plants. A recent study suggested that the Cheirogaleidae might only disperse seeds of up to 15mm in size (Ramananjato 2024 ; Ganzhorn et al. 2024). While this may help to prevent the long-term conversion of severely burnt forests into savannas, and fire-adapted lianas, for example, provide shade for early successional seedlings of species with smaller seeds, a full forest regeneration that also includes large-seeded tree species, requires a step-wise return of major seed dispersers of all sizes. Future studies are needed to compare seed sizes of plant communities in various stages of post-fire regeneration to test this hypothesis. Given the severity of fire impacts, we recommend several urgent conservation actions. First, the conservation status of the medium-sized lemurs ( A. occidentalis, L. edwardsi) and in particular that of the Endangered L. edwardsi (IUCN 2020 ) should be reassessed given that they are heavily impacted by fire (this study) and hunting (Craul et al. 2009 ; Sato et al. 2021 ), and forest fires are very prominent threats for the remaining dry forests (Frappier-Brinton and Lehman 2022 ). Second, conservation planning should focus on safeguarding the remaining unburnt forests, as they are vital for the survival of existing sub-populations and can act as source habitats facilitating the recolonization of fire-affected areas. Additionally, long-term protection strategies must be adopted to prevent burnt areas from experiencing subsequent fires, as repeated fires can significantly hinder forest recovery (Percival et al. 2024 ) and further endanger lemur populations and lemur species richness (this study). Third, the conservation efforts of the park management should be complemented by educational and socioeconomic programs to raise awareness in the rural communities, improve livelihoods, and reduce the lemur hunting pressure to protect these endangered species and facilitate long-term coexistence of humans and wildlife near and inside the park. Declarations Supplementary Information The online version contains supplementary material available at ….. Author contributions UR and DS conceptualized the study and developed the methodology; UR acquired the funding; NRR, MOR, RM, and SHR were responsible for data collection; MR provided the fire history of study sites via remote sensing work; NRR conducted the data analyses; UR, RR and HR supervised the work; NRR and UR took the lead in writing the manuscript. All authors participated in editing the manuscript drafts and approved the final version for submission. Funding This study was financially supported by Madagascar National Parks under contract number 20/DG/DOP/DAFRH/CONV/2022, as well as by the Cologne Zoo, Germany. Data availability The datasets generated and analyzed during the current study are provided in the supplementary material. Declaration of competing interests The authors declare no competing interests. Acknowledgement We want to express our gratitude to the Ministère de l’Environnement et du Développement Durable and the Direction Régionale de l’Environnement (DREDD) Boeny Betsiboka for granting us permission to conduct this research through the following research permits: № 275/22/MEDD/SG/DGGE/DAPRNE/SCBE.Re, № 070/23/MEDD/SG/DGGE/DAPRNE/SCBE.Re, and № 263/23/MEDD/SG/DGGE/DAPRNE/SCBE.Re. We sincerely thank Madagascar National Parks for their unwavering administrative and financial support, especially to the General Directors, the financial department, and Madame Lalatiana Odile Randriamiharisoa. Our appreciation also goes to the former and current Directors of Ankarafantsika National Park, Mandimby Heriniaina Andriambololona and Charles Andriamaniry, as well as the Chef du Volet Opération, logistical staff, Chefs Secteurs, and park agents for their support during our fieldwork. We are grateful to the Mention Anthropologie et Développement Durable from the Faculty of Sciences at the University of Antananarivo for facilitating all our research permits. Additionally, we thank our park guides, Jhonny Kennedy and Estoria Una, as well as the local population (CLP and Cook) for their assistance during our site visits. We also appreciate the support of Marlene Böhm, Johnny Randriafenontsoa, and Fenohery Andriantsitohaina during the data collection process in the field. This project was funded by Madagascar National Parks and supported by the Kreditanstalt für Wiederaufbau (KFW) under contract number 20/DG/DOP/DAFRH/CONV/2022. The Cologne Zoo, Germany, also provided funding for part of the fieldwork. References Adeney JM, Ginsberg JR, Russell GJ, and Kinnaird MF (2006) Effects of an ENSO-related fire on birds of a lowland tropical forest in Sumatra. Anim Conserv 9: 292-301. https://doi.org/10.1111/j.1469-1795.2006. 00035.x Agus C, et al. 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Sci Rep 3: 2424. https://doi.org/10.1038/srep02424 van Elst T, Sgarlata GM, Schüßler D, Tiley GP, Poelstra JW, Scheumann M, Blanco MB, Aleixo-Pais IG, Rina Evasoa M, Ganzhorn JU, Goodman SM, Hasiniaina AF, Hending D, Hohenlohe PA, Ibouroi MT, Iribar A, Jan F, Kappeler PM, Le Pors B, Manzi S, Olivieri G, Rakotonanahary AN, Rakotondranary SJ, Rakotondravony R, Ralison JM, Ranaivoarisoa JF, Randrianambinina B, Rasoloarison RM, Rasoloharijaona S, Rasolondraibe E, Teixeira H, Zaonarivelo JR, Louis EE, Yoder AD, Chikhi L, Radespiel U, and Salmona J (2024) Integrative taxonomy clarifies the evolution of a cryptic primate clade. Nat Ecol Evol 8: 1-16. https://doi.org/10.1038/s41559-024-02547-w van Holstein LA, McKay HD, Pimiento C, and Koops K (2024) Multidimensional primate niche space sheds light on interspecific competition in primate evolution. Commun Biol 7: 647. https://doi.org/10.1038/s42003-024-06324-0 Warren RD, and Crompton RH (1997) Locomotor ecology of Lepilemur edwardsi and Avahi occidentalis . Am J Phys Anthropol 104: 471-486. https://doi.org/10.1002/(SICI)1096-8644(199712)104:4 3.0.CO;2-V Wickham H, Chang W, and Wickham MH (2016) Package ‘ggplot2’. Create elegant data visualizations using the grammar of graphics. Version 2. https://ggplot2.tidyverse.org Wilson M, Chen XY, Corlett RT, Didham RK, Ding P, Holt RD, Holyoak M, Hu G, Hughes AC, Jiang L, Laurance WF, Liu J, Pimm SL, Robinson SK, Russo SE, Si X, Wilcove DS, Wu J, and Yu M (2015) Habitat fragmentation and biodiversity conservation: key findings and future challenges. Landsc Ecol 31: 219–227. https://doi.org/10.1007/s10980-015-0312-3 Zhang Y, Zhou H, Zhou J, and Sun W (2017) Regression Models for Multivariate Count Data. J Comput Graph Stat 26: 1-13. https://doi.org/10.1080/10618600.2016.1154063 Table 2 Table 2 is available in the Supplementary Files section. Additional Declarations No competing interests reported. Supplementary Files Supplementaryinformations.docx Table2.docx Cite Share Download PDF Status: Published Journal Publication published 06 Oct, 2025 Read the published version in Biodiversity and Conservation → Version 1 posted Editorial decision: Revision requested 29 Apr, 2025 Reviews received at journal 18 Mar, 2025 Reviewers agreed at journal 03 Mar, 2025 Reviews received at journal 03 Mar, 2025 Reviewers agreed at journal 03 Mar, 2025 Reviewers agreed at journal 02 Mar, 2025 Reviewers agreed at journal 01 Mar, 2025 Reviewers invited by journal 26 Feb, 2025 Editor assigned by journal 03 Jan, 2025 Submission checks completed at journal 02 Jan, 2025 First submitted to journal 30 Dec, 2024 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. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-5735404","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":397447061,"identity":"2aee7716-d3a0-4d46-aeae-f6808c247794","order_by":0,"name":"Naina Ratsimba Rabemananjara","email":"","orcid":"","institution":"University of Veterinary Medicine Hannover","correspondingAuthor":false,"prefix":"","firstName":"Naina","middleName":"Ratsimba","lastName":"Rabemananjara","suffix":""},{"id":397447062,"identity":"c85b66e9-aa45-4431-a458-8c39bc2398ef","order_by":1,"name":"Misa Miaritiana Rasolozaka","email":"","orcid":"","institution":"University of Veterinary Medicine Hannover","correspondingAuthor":false,"prefix":"","firstName":"Misa","middleName":"Miaritiana","lastName":"Rasolozaka","suffix":""},{"id":397447063,"identity":"c0befb2a-e5eb-40d7-8a04-30485f638d9d","order_by":2,"name":"Marie Odile Ravolanirina","email":"","orcid":"","institution":"University of Mahajanga","correspondingAuthor":false,"prefix":"","firstName":"Marie","middleName":"Odile","lastName":"Ravolanirina","suffix":""},{"id":397447064,"identity":"0efba370-d2e0-4275-90a8-30017f509700","order_by":3,"name":"Rogula Marivola","email":"","orcid":"","institution":"University of Mahajanga","correspondingAuthor":false,"prefix":"","firstName":"Rogula","middleName":"","lastName":"Marivola","suffix":""},{"id":397447066,"identity":"febb7c24-c50d-4063-872a-70e8075a083e","order_by":4,"name":"Seheno Harilala Randriamiarantsoa","email":"","orcid":"","institution":"University of Antananarivo","correspondingAuthor":false,"prefix":"","firstName":"Seheno","middleName":"Harilala","lastName":"Randriamiarantsoa","suffix":""},{"id":397447067,"identity":"60102101-3117-410f-ba77-f7b1f050d7b5","order_by":5,"name":"Romule Rakotondravony","email":"","orcid":"","institution":"University of Mahajanga","correspondingAuthor":false,"prefix":"","firstName":"Romule","middleName":"","lastName":"Rakotondravony","suffix":""},{"id":397447069,"identity":"a5b75311-189d-4c21-b5c5-4223f253f788","order_by":6,"name":"Hanta Razafindraibe","email":"","orcid":"","institution":"University of Antananarivo","correspondingAuthor":false,"prefix":"","firstName":"Hanta","middleName":"","lastName":"Razafindraibe","suffix":""},{"id":397447070,"identity":"9702ea0b-894a-4d2a-950b-7f27654d9ad2","order_by":7,"name":"Dominik Schüßler","email":"","orcid":"","institution":"University of Hildesheim","correspondingAuthor":false,"prefix":"","firstName":"Dominik","middleName":"","lastName":"Schüßler","suffix":""},{"id":397447071,"identity":"92f8a856-7695-4cde-94dd-03377705f205","order_by":8,"name":"Ute Radespiel","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAABLUlEQVRIie3QsUrDQBjA8e8IXJbUrgnV9BW+kkGkg6+SW9rRtUPBK4HrYnWtNNhX6CSOKQfpcg/QoWC6dKqQqVTQ4kUHhaSIm8P94W757sdxB2Ay/d8svQiHrAcOAtDvgVNxmP4kocI/ER0TCL8SfI4Sq9dv+81JNMjYw9vpuc1p9vq08nExymDbK5OUhkSl3QBX8wjZIzoXN4ndGqlNgGqBJFYVxEHCqWQzlwm3ILi82rk1Idn9uANWTVSQek74QV5Px2y4Z3FBQuq9F2S60eRQdQuQgZAhLJkAxr9Io7jlzqWa8BLx0g7OB7fd1ky/xQ3TwEGV0MaZkEHd6cA8TkvkRMr1mu/azeZkuM7zvn+JC069FyF9aqck2/bLv6xLPnf3+OhYlcRkMplMug9q621X/EePvwAAAABJRU5ErkJggg==","orcid":"","institution":"University of Veterinary Medicine Hannover","correspondingAuthor":true,"prefix":"","firstName":"Ute","middleName":"","lastName":"Radespiel","suffix":""}],"badges":[],"createdAt":"2024-12-30 12:10:36","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-5735404/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-5735404/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s10531-025-03167-x","type":"published","date":"2025-10-06T15:57:32+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":73018233,"identity":"2a394d35-e9dd-43b1-b8eb-b2e44d2e5d1c","added_by":"auto","created_at":"2025-01-06 03:12:16","extension":"jpg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":4416496,"visible":true,"origin":"","legend":"\u003cp\u003eMap of Ankarafantsika National Park located in northwestern Madagascar (inlay map) with the 18 study sites selected based on prior remote sensing work (Source: Landsat 09 image in August 2023 with true color composite (Band 04, Band 03, Band 02); Shapefile of Madagascar: https://data.humdata.org/dataset/cod-ab-mdg; Villages and roads: Madagascar National Parks).\u003c/p\u003e","description":"","filename":"Fig1.tiff.jpg","url":"https://assets-eu.researchsquare.com/files/rs-5735404/v1/d6e5fc5a0380fe69d6b5d9e9.jpg"},{"id":73018251,"identity":"37b0f15c-b337-4017-aab8-d84e8a3aa233","added_by":"auto","created_at":"2025-01-06 03:12:18","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":457813,"visible":true,"origin":"","legend":"\u003cp\u003eRelationships between the abundance of \u003cem\u003eE. fulvus\u003c/em\u003e per 100 m and the topographic variables with significant effects\u003cstrong\u003e (a) \u003c/strong\u003eBox plots showing\u003cstrong\u003e \u003c/strong\u003eabundance/100m of \u003cem\u003eEulemur fulvus\u003c/em\u003e on plateau, slopes, and in valleys with individual data points being displayed as jitter. Boxplots display the median in a horizontal line with interquartile range (box) and minimum and maximum values (whiskers) without outliers. \u003cstrong\u003e(b)\u003c/strong\u003eAbundance/100 m of \u003cem\u003eE. fulvus\u003c/em\u003e in relation to elevation (in m a.s.l.). Red line: linear regression line, size of black dots encodes for the number of data points showing the same value.\u003c/p\u003e","description":"","filename":"Fig2.tiff.jpg","url":"https://assets-eu.researchsquare.com/files/rs-5735404/v1/0aff0b740c980d646e173ede.jpg"},{"id":73018455,"identity":"a7765e13-fb20-4e31-a832-0807815a3eeb","added_by":"auto","created_at":"2025-01-06 03:20:18","extension":"jpg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":589121,"visible":true,"origin":"","legend":"\u003cp\u003eComparison of the abundance of \u003cem\u003eMicrocebus ravelobensis\u003c/em\u003e on plateau, slopes, and in valleys. Individual data points are displayed as jitter. Boxplots display the median in a horizontal line with an interquartile range (box) andthe minimum and maximum values (whiskers) without outliers.\u003c/p\u003e","description":"","filename":"Fig3.tiff.jpg","url":"https://assets-eu.researchsquare.com/files/rs-5735404/v1/2934ec0eb244581d0d3776f3.jpg"},{"id":73018447,"identity":"f037c96b-54ed-4598-85ea-61fbe92e7a3a","added_by":"auto","created_at":"2025-01-06 03:20:17","extension":"jpg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":112508,"visible":true,"origin":"","legend":"\u003cp\u003ePresence and absence of eight lemur species in burnt zones, categorized by the time since the most recent fire (1 to \u0026gt;35 years) and compared to unburnt zones. Colored segments represent the presence of each species within a specific fire-age category. \"NA\" indicates a site where no data was available due to the hibernation period of \u003cem\u003eCheirogaleus medius\u003c/em\u003e. n: The number of sites containing partitions surveyed within each fire-age category. Numbers inside the segments indicate the number of sites where the species was documented for each category.\u003c/p\u003e","description":"","filename":"Fig4.tiff.jpg","url":"https://assets-eu.researchsquare.com/files/rs-5735404/v1/1c8ca447138b1cb86c60744b.jpg"},{"id":73018264,"identity":"23bd4c2e-c5a5-4211-8884-279591b9fcc2","added_by":"auto","created_at":"2025-01-06 03:12:18","extension":"jpg","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":457105,"visible":true,"origin":"","legend":"\u003cp\u003eRelationship between the number of fires and the abundance of \u003cem\u003eE. fulvus\u003c/em\u003e per 100 m. The size of the black dots encodes for the number of data points showing the same value. The red line represents the linear regression line.\u003c/p\u003e","description":"","filename":"Fig5.tiff.jpg","url":"https://assets-eu.researchsquare.com/files/rs-5735404/v1/42f0d5926733a7b7e63da1be.jpg"},{"id":73018452,"identity":"70f65cd0-a6ad-435b-8345-dd6ddd00d878","added_by":"auto","created_at":"2025-01-06 03:20:17","extension":"jpg","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":1098131,"visible":true,"origin":"","legend":"\u003cp\u003eRelationships between the abundance of \u003cem\u003eL. edwardsi\u003c/em\u003e per 100 m and the fire-related variables with significant effects. \u003cstrong\u003e(a)\u003c/strong\u003eComparison of \u003cem\u003eL. edwardsi\u003c/em\u003e abundance in burnt versus unburnt zones. Individual data points are displayed as jitter. Boxplots display the median in a horizontal line with an interquartile range and the minimum and maximum values (whiskers) without outliers. \u003cstrong\u003e(b)\u003c/strong\u003e Relationship between the abundance of \u003cem\u003eL. edwardsi\u003c/em\u003e and the number of fires experienced per transect partition. \u003cstrong\u003e(c)\u003c/strong\u003eRelationship between the minimum interval between two fires and the abundance of \u003cem\u003eL. edwardsi\u003c/em\u003e. \u003cstrong\u003e(d)\u003c/strong\u003e Relationship between the maximum fire severity and the abundance of \u003cem\u003eL. edwardsi\u003c/em\u003e. The sizes of the black dots in \u003cstrong\u003e(b)\u003c/strong\u003e, \u003cstrong\u003e(c)\u003c/strong\u003e, and \u003cstrong\u003e(d) \u003c/strong\u003eencode for the number of data points showing the same value. The red line represents the linear regression line.\u003c/p\u003e","description":"","filename":"Fig6.tiff.jpg","url":"https://assets-eu.researchsquare.com/files/rs-5735404/v1/cd14de2a963a60cf3879ccdb.jpg"},{"id":73018250,"identity":"2e28cb65-ac60-4a71-95d4-c19e3198a836","added_by":"auto","created_at":"2025-01-06 03:12:18","extension":"jpg","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":402831,"visible":true,"origin":"","legend":"\u003cp\u003eImpact of burn status and the number of fires on lemur species richness. \u003cstrong\u003e(a)\u003c/strong\u003e Comparison of lemur species richness in burnt and unburnt zones. Individual data points are displayed as jitter. Boxplots display the median in a horizontal line with an interquartile range and the minimum and maximum values (whiskers) without outliers. \u003cstrong\u003e(b)\u003c/strong\u003e Relationship between the number of fires and lemur species richness. The sizes of the black dots encode the number of data points showing the same value. The linear regression line is shown in red.\u003c/p\u003e","description":"","filename":"Fig7.tiff.jpg","url":"https://assets-eu.researchsquare.com/files/rs-5735404/v1/a4dcea9ce5764d5e3d84e835.jpg"},{"id":73018236,"identity":"eb205a22-297e-4502-8e3d-ba1a1b831d47","added_by":"auto","created_at":"2025-01-06 03:12:16","extension":"jpg","order_by":8,"title":"Figure 8","display":"","copyAsset":false,"role":"figure","size":2988334,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eFire history and vegetation map of Ankarafantsika National Park (1988–2022) \u003c/strong\u003eshowing the distribution of fire-affected and unburnt areas within the park. Green areas represent zones that never burnt between 1988 – 2022. Gray areas indicate regions with fire histories less than 23 years old, and yellow areas indicate regions with a more than 23-year old fire history.\u003c/p\u003e","description":"","filename":"Fig8.tiff.jpg","url":"https://assets-eu.researchsquare.com/files/rs-5735404/v1/ce55e27df86b6c830fe5618c.jpg"},{"id":93419821,"identity":"93ffc79c-ecf9-4696-97d9-5d0ed94667a7","added_by":"auto","created_at":"2025-10-13 16:08:03","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":11774419,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-5735404/v1/13958a65-9766-40dd-8db6-c5e10d5a7da9.pdf"},{"id":73018234,"identity":"ae029ab2-d4c6-4a30-9daa-65bd5063ea2f","added_by":"auto","created_at":"2025-01-06 03:12:16","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":85910,"visible":true,"origin":"","legend":"","description":"","filename":"Supplementaryinformations.docx","url":"https://assets-eu.researchsquare.com/files/rs-5735404/v1/753232d7a700e6fda5bc0356.docx"},{"id":73018232,"identity":"b6e70047-bdaa-4fe9-bb96-36a8c8b9f582","added_by":"auto","created_at":"2025-01-06 03:12:16","extension":"docx","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":23710,"visible":true,"origin":"","legend":"","description":"","filename":"Table2.docx","url":"https://assets-eu.researchsquare.com/files/rs-5735404/v1/52caf85944171877706f58b4.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Post-fire recolonization of dry deciduous forests by lemurs in northwestern Madagascar","fulltext":[{"header":"Introduction","content":"\u003cp\u003eFires have a long history of shaping environments with evidence dating back to the late Carboniferous (Scott \u003cspan citationid=\"CR111\" class=\"CitationRef\"\u003e2000\u003c/span\u003e). Before humans started to use fire routinely, lightning was likely the main natural source of wildfire (Roberts \u003cspan citationid=\"CR100\" class=\"CitationRef\"\u003e2000\u003c/span\u003e). Consequently, most terrestrial ecosystems have evolved alongside low frequencies of fire for millions of years. Meta-analyses have revealed that more than 50% of the global terrestrial habitats even need occasional fires to maintain viable environmental conditions (Pausas and Keeley \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). Moreover, it was suggested that fires may have played an important role in the origins and distribution of some particular ecosystems, specifically those with rather open vegetation formations (e.g., savanna, grasslands) (Bond and Keeley \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Shlisky et al. \u003cspan citationid=\"CR113\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). However, even though some ecosystems may have evolved adaptations to fire, modern changes in the fire regime (i.e., anthropogenic increase in fire frequency, extent, or intensity) may impose severe challenges due to shorter time windows for regrowth and regeneration. Shlisky et al. (\u003cspan citationid=\"CR113\" class=\"CitationRef\"\u003e2007\u003c/span\u003e) showed that approximately 60% of the world's land-based habitats have undergone recent alterations in their fire regime. At least 20% of these are fire-sensitive habitats, where most species have not evolved traits to survive or regenerate after fires and are therefore highly susceptible to damage from wildfires (e.g., tropical moist forests).\u003c/p\u003e \u003cp\u003eDespite covering only 6% of the Earth's surface, tropical forests harbor more than 80% of the planet's terrestrial biodiversity (Gardner et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). However, anthropogenic activities increasingly threaten this exceptional biodiversity, particularly through landscape degradation and habitat fragmentation (Wilson et al. \u003cspan citationid=\"CR129\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). A primary contributor to these disturbances is the intentional use of fire for land conversion, notably in slash-and-burn agriculture, where fire serves to clear land for agricultural expansion or to improve soil fertility (Heinimann et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Curtis et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Such practices accelerate deforestation and destabilize these ecosystems, further contributing to their degradation (Foley et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Curtis et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2018\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eMany ecosystems, such as tropical rainforests, are particularly vulnerable to fire due to their lack of adaptations to frequent fire events, making them highly susceptible to shifts in fire regimes driven by human activities (Dwyer et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Bond and Keeley \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Shlisky et al. \u003cspan citationid=\"CR113\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Dantas and Pausas 2013). In contrast to fire-adapted, temperate biomes, plant species in these environments typically lack protective traits such as thick bark or deep rooting systems, leaving them susceptible to fire-induced mortality (Bond and Keeley \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). Severe forest fires can destroy tree canopies, lead to the loss of understory vegetation, increase soil exposure, and enhance erosion (Bond and Keeley \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). In the next step, invasive, fire-tolerant species can establish themselves in these burnt and degraded forests, and these pioneer species can outcompete native vegetation and alter ecosystem structure substantially (Bond and Keeley \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2005\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eA central question in the study of fire ecology is how fires affect the composition and integrity of wildlife populations. While adaptive responses in fire-prone environments have been widely studied in plants, animal responses have received comparatively less attention (Pausas and Parr \u003cspan citationid=\"CR82\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Animals are generally less likely to survive in the flame zone of a fire, but many are mobile enough to flee the immediate fire zone and seek shelter in fire-protected microhabitats (Pausas and Parr \u003cspan citationid=\"CR82\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Nimmo et al. 2018). For example, the southern brown bandicoot (\u003cem\u003eIsoodon obesulus\u003c/em\u003e) digs burrows for shelter in a fire-prone environment (Long \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). Certain animals even modify their surroundings to protect themselves, such as some termite species \u003cem\u003e(Macrotermes\u003c/em\u003e spp.\u003cem\u003e)\u003c/em\u003e which construct mounds that buffer against heat (Korb and Linsenmair \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e1999\u003c/span\u003e). Some birds like the malleefowl \u003cem\u003e(Leipoa ocellata)\u003c/em\u003e construct large nesting mounds made of soil and organic material like leaf litter that reduce litter fuel loads, leading to decreased fire intensity and potentially creating fire refuges (Smith et al. \u003cspan citationid=\"CR114\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Larger species such as elephants \u003cem\u003e(Loxodonta africana and Elephas maximus)\u003c/em\u003e indirectly reduce the fire risk and alter their environment by consuming potential fuel materials (Holdo \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2007\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eAlthough some species show adaptations and responses to fire, fire can still affect their abundances and species richness. These effects largely depend on the species-specific environmental plasticity and the different components of the fire regime (e.g. frequency, history, severity) (Gonz\u0026aacute;lez et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Fires can reduce animal abundance and diversity by destroying habitats and causing direct mortality, especially in species lacking adaptations to escape or withstand fire. Amphibians such as the spotted salamander \u003cem\u003e(Ambystoma maculatum)\u003c/em\u003e (Fontaine and Kennedy \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2012\u003c/span\u003e), which depend on moist environments, experience significant population declines in regions frequently exposed to fire. On the other hand, fires can create a mosaic of heterogeneous habitats, enhancing food availability and shelter for species (Russell et al. \u003cspan citationid=\"CR102\" class=\"CitationRef\"\u003e1999\u003c/span\u003e). In grasslands and savannas, for example, fires promoted the growth of early successional plants, attracting herbivores like white-tailed deer \u003cem\u003e(Odocoileus virginianus)\u003c/em\u003e and the eastern cottontail rabbit \u003cem\u003e(Sylvilagus floridanus)\u003c/em\u003e, which thrive on fresh vegetation regrowth (Russell et al. \u003cspan citationid=\"CR102\" class=\"CitationRef\"\u003e1999\u003c/span\u003e). Fires can also open the forest canopy, boosting insect populations and benefiting bird species like downy woodpeckers \u003cem\u003e(Picoides pubescens)\u003c/em\u003e (Nimmo et al. 2018). Furthermore, quickly recolonizing post-fire habitats may reduce feeding competition in less populated areas and thereby convey energetic benefits for flexible individuals (Pausas and Parr \u003cspan citationid=\"CR82\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Post-fire wildlife dynamics and thereby the recolonization potential of different animal species are still understudied particularly in the tropics and on longer time scale, but may offer key insights to understanding co-evolutionary processes in fire-prone ecosystems (Gonz\u0026aacute;lez et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe island of Madagascar is known for its high levels of endemism (Myers et al. \u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2000\u003c/span\u003e) along with the high rate of habitat degradation due to human activities, which pose a major threat to its biodiversity. Consistent with other tropical regions of the world, anthropogenic fires are a pervasive driver of landscape change and habitat degradation across Madagascar (Kull \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2000\u003c/span\u003e, Phelps et al. \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Fires are particularly prominent in the western, drier parts of the island (Phelps et al. \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) and were described as a destructive force in modern-day Madagascar, the \u0026ldquo;isle of fire\u0026rdquo; (Kull \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Frappier-Brinton and Lehman \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). However, fires may already have played a role during the formation of its ecosystems and species assemblages, as indicated by historical evidence of fires predating human arrival (Burney \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e1987\u003c/span\u003e; Gasse and Van Campo \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Teixeira et al. \u003cspan citationid=\"CR119\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe ancestors of extant lemurs arrived on the island about 60\u0026nbsp;million years ago and evolved into more than 100 different species (Herrera and D\u0026aacute;valos \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). However, the diversification of many extant genera (e.g., \u003cem\u003eMicrocebus\u003c/em\u003e) occurred quite rapidly and rather recently within the last few million years (Poelstra et al. \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Van Elst et al. \u003cspan citationid=\"CR125\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). It is possible that adaptations to fire evolved over these time spans in habitats also prone to fire today. This hypothesis gains support from an analysis of the charcoal stratigraphy of three sediment cores from central Madagascar, which showed that late Pleistocene and early- to mid-Holocene sediments deposited before human settlement often contained more charcoal than post-settlement and modern sediments (Burney \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e1987\u003c/span\u003e). Lemurs are a particularly suitable and important group of study species to investigate the effects of forest fires on wildlife, as they have largely different ecologies, perform key ecosystem functions (e.g., pollination, seed dispersal, food web interactions), and the presence of a complete lemur assemblage indicates intact forest habitats (Muldoon and Goodman \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Finally, lemurs are important flagship species for conservation and ecotourism in Madagascar (Mittermeier et al. \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). To date, there is insufficient information on whether lemurs or other wildlife living in the dry forests of western Madagascar possess at least some resilience against fires that are increasingly threatening and affecting the dry forest habitats of the island.\u003c/p\u003e \u003cp\u003eThe Ankarafantsika National Park (ANP) is located in northwestern Madagascar and serves as a perfect model area for studying the effects of fire on wildlife. In recent years, increasing fire intensity and severe burning episodes in the region threaten the remaining forest habitats and wildlife (Sch\u0026uuml;\u0026szlig;ler et al. 2022). ANP is inhabited by eight lemur species that differ in daily activity, body mass, and also in their IUCN conservation status, ranging from \u0026lsquo;Least Concern\u0026rsquo; (\u003cem\u003eMicrocebus murinus\u003c/em\u003e) to \u0026lsquo;Critically Endangered\u0026rsquo; (\u003cem\u003ePropithecus coquereli\u003c/em\u003e, \u003cem\u003eEulemur mongoz\u003c/em\u003e, Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Previous work also suggested that the natural topography of ANP impacts at least the abundance of the two mouse lemurs differentially (Rakotondravony and Radespiel \u003cspan citationid=\"CR91\" class=\"CitationRef\"\u003e2007\u003c/span\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\u003eLemur species occurring in Ankarafantsika National Park with activity, body mass (Mittermeier et al. \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), and IUCN conservation status (IUCN \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). LC\u0026thinsp;=\u0026thinsp;Least Concern, VU\u0026thinsp;=\u0026thinsp;Vulnerable, EN\u0026thinsp;=\u0026thinsp;Endangered, CR\u0026thinsp;=\u0026thinsp;Critically Endangered. *Hibernates from May to mid-September\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"5\"\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 \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSpecies\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eVernacular name\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eActivity\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eBody mass (g)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eIUCN status\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003ePropithecus coquereli\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCoquerel\u0026rsquo;s sifaka\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003ediurnal\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3,700-4,300\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eCR\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eEulemur mongoz\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMongoose lemur\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003ecathemeral\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1,100-1,600\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eCR\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eEulemur fulvus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCommon brown lemur\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003ecathemeral\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1,700-2,100\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eVU\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAvahi occidentalis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eWestern woolly lemur\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003enocturnal\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e800-1,100\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eVU\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eLepilemur edwardsi\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMilne-Edwards sportive lemur\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003enocturnal\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e854-1,200\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eEN\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eCheirogaleus medius*\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eFat-tailed dwarf lemur\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003enocturnal\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e120\u0026ndash;270\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eVU\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eMicrocebus murinus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eGray mouse lemur\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003enocturnal\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e58\u0026ndash;67\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eLC\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eMicrocebus ravelobensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eGolden-brown mouse lemur\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003enocturnal\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e56\u0026ndash;87\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eVU\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\u003eWe aim to investigate how past fires affected the different lemur species in the dry deciduous forests of ANP in northwestern Madagascar. By studying their ability and timespan for re-colonizing areas that have been previously burnt, we aim to shed light on the resilience of lemurs to forest fires. Specifically, we investigate the impacts of (1) the time since the last forest fire, (2) the maximum past fire intensity, (3) the number of past forest fires, and (4) the minimum interval between recurrent fires on lemur abundance and species richness. The relevant fire history for all study sites was provided by a parallel study for a period of 35 years (1988\u0026ndash;2022) for which yearly remote sensing was available (Rasolozaka et al. \u003cspan citationid=\"CR98\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). To test for additional confounding effects, the influence of terrain and elevation for lemur abundance and lemur species richness is also evaluated.\u003c/p\u003e"},{"header":"Materials and Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eStudy area\u003c/h2\u003e \u003cp\u003eWe conducted our study in Ankarafantsika National Park (ANP), which is the largest protected remaining dry forest tract in northwestern Madagascar (-16\u0026deg;08'60\"S, 46\u0026deg;57'0\" E) and covers an area of 1,350 km\u0026sup2; (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). It possesses some topographical complexity, as it includes river valleys at about 50 m above sea level (a.s.l.), slopes, and a rising topography from north to south where a calcitic plateau reaches peak elevations of up to 350 m a.s.l. and forms cliffs in many places in the east and south (Alonso et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2002\u003c/span\u003e). ANP is crucial for the conservation of numerous endangered species, such as the Critically Endangered Coquerel\u0026rsquo;s sifaka \u003cem\u003e(Propithecus coquereli)\u003c/em\u003e, the endangered largest terrestrial carnivore of Madagascar \u003cem\u003e(Cryptoprocta ferox)\u003c/em\u003e, and the Critically Endangered Malagasy fish eagle \u003cem\u003e(Haliaeetus vociferoides)\u003c/em\u003e (Schwitzer et al. \u003cspan citationid=\"CR109\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Barcala \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Razafimanjato et al. \u003cspan citationid=\"CR99\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). ANP features a seasonally dry tropical forest that includes a mix of dry deciduous forests and natural dry thickets at higher elevations, moist riverine forests in upstream valleys and around lakes at lower elevations, as well as \u003cem\u003eRaphia\u003c/em\u003e swamp forests in downstream valleys (Goodman et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Forest vegetation can be found on the dry plateau, slopes, and valleys. The average annual rainfall ranges from 1,000 to 1,500 mm, with most rain falling during the rainy season which lasts from November to April, while the almost rainless dry season covers the period from May to October (Alonso et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2002\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eUse of remote sensing data for study site selection\u003c/h3\u003e\n\u003cp\u003eWe used remote sensing data to identify 18 suitable study sites with different fire histories across ANP prior to the start of the fieldwork. Rasolozaka et al. (\u003cspan citationid=\"CR98\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) detailed the necessary working steps. In brief, this process involved screening annual Landsat satellite images (30 x 30m resolution) from 1988\u0026ndash;2023, from which specific fire and vegetation indices were calculated to quantify the occurrence and extent of fires across the years in all study sites. M. Rasolozaka used the Normalized Burn Ratio (Key and Benson \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2006\u003c/span\u003e) to identify fires, i.e., whether pixels were burnt or unburnt in a given year. By subtracting the pre- and post-fire NBR values (dNBR), M. Rasolozaka calculated the fire severity for all study sites and years (Keeley \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). The rationale for our study site selection was to identify areas where an unburnt area (i.e., never burnt across the 35 years) bordered on a burnt area with a variable number of years since the last fire and a variable total number of fires experienced over the previous 35 years. Such sites were chosen to provide a mixed design for subsequent data analyses while always controlling for site-specific ecological characteristics. After completing the fieldwork, all remote sensing data were re-analyzed and thoroughly evaluated alongside field data, providing ground truthing for the final reconstruction of the site-specific fire history. This re-evaluation of the remote sensing data together with field-based records of past fire traces (e.g., fire scorches on trees and charcoal traces in soil, Rasolozaka et al., \u003cspan citationid=\"CR98\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) revealed that the prior classification of some study sites needed to be revised. Specifically, parts of the unburnt areas had indeed burnt in some sites (sites 5, 4, 13, 2, 12, 15), and some sites had experienced a heterogeneous fire history in which not all parts of the study transect were burnt during the same years (details in Supplementary Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e).\u003c/p\u003e\n\u003ch3\u003eStudy sites and spatial arrangement of fieldwork\u003c/h3\u003e\n\u003cp\u003ePrior to fieldwork, we selected 18 study sites that varied in their specific fire history (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Six sites were visited in 2022 (September - December) and a further 12 sites were visited in 2023 (May - November). The sites varied in the number of years since the last fire (1 to \u0026gt;\u0026thinsp;35 years), in the number of fire occurrences in each site over the last 35 years (1\u0026ndash;7 fires), in the minimum interval between two fires (1\u0026ndash;31 years), and in the maximum burn severity that each site experienced in the past 35 years (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). A 1.2 km-long transect with a paired design was established in each site, which was flagged and mapped with a GPS device (Garmin GPSMAP 64) every 10 m. It consisted of a stretch of 500 m in the burnt forest (burnt zone), running perpendicularly to the fire edge, and a complementary stretch of 500 m in the unburnt forest (unburnt zone), also pointing away from the fire edge, but shifted with the burnt transect by 200 m. We connected these two transect parts by a transect of 200 m length running in parallel to the fire edge, about 50 m into the burnt forest, i.e., in the transition zone between burnt and unburnt forest.\u003c/p\u003e\n\u003ch3\u003eLemur abundance survey\u003c/h3\u003e\n\u003cp\u003eWe conducted three diurnal and three nocturnal systematic distance sampling surveys per site to monitor the 1.2km transect for sightings of any lemur species (Buckland et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2001\u003c/span\u003e). With four observers, we walked each transect quietly and slowly at a speed of approximately 0.5km/h (Rakotondravony and Radespiel \u003cspan citationid=\"CR91\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). Each transect was visited over three days in the morning (06h30-08h30) and evening (18h15\u0026ndash;20h30). We used headlamps and flashlights during night surveys to identify and differentiate the eight species. Upon each lemur encounter, we recorded the number of individuals and determined the species and the position along the transect to the closest 10 m.\u003c/p\u003e \u003cp\u003eWe calculated the abundance of each species (dependent variable) separately for each partition of the transect that had a different fire history and initially also distinguished between the burnt zone and the transition zone (Table S6). Encounter rates were too small to determine population densities for any of the species for any partition of the transect (Buckland et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2001\u003c/span\u003e). Therefore, and to be able to subsequently model variations in abundances from different study sites, we calculated mean encounter rates per 100m transect as a proxy for abundance for each transect partition with specific fire history and for each species separately with the following formula:\u003cdiv id=\"Equa\" class=\"Equation\"\u003e\u003cdiv format=\"TEX\" class=\"mathdisplay\" id=\"FileID_Equa\" name=\"EquationSource\"\u003e\n$$\\:\\text{A}\\text{b}\\text{u}\\text{n}\\text{d}\\text{a}\\text{n}\\text{c}\\text{e}=\\frac{N}{L*n}*100$$\u003c/div\u003e\u003c/div\u003e\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eWhere\u003c/p\u003e \u003cp\u003eN: Total number of individuals encountered during all three surveys along transect partition with specific fire history\u003c/p\u003e \u003cp\u003eL: Total length of transect partition (in meters)\u003c/p\u003e \u003cp\u003en: Number of surveys\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003cp\u003eWe further determined the lemur species richness (dependent variable) as the count of the total number of species observed in each transect partition across the three successive surveys.\u003c/p\u003e\n\u003ch3\u003eModeling the effect of topographic and fire-related variables on lemur abundance and lemur species richness\u003c/h3\u003e\n\u003cp\u003eDue to the limited number of sightings, it was not possible to generate statistical models for \u003cem\u003ePropithecus coquereli\u003c/em\u003e and \u003cem\u003eEulemur mongoz\u003c/em\u003e. For these two species, we only present descriptive data on the relationship between the species' presence and the time that elapsed since the last fire. \u003cem\u003eCheirogaleus medius\u003c/em\u003e hibernates during large parts of the dry season from May to mid-September (M\u0026uuml;ller and Thalmann \u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e2002\u003c/span\u003e). Therefore, the species was only included in the abundance analysis for sites visited between September to December.\u003c/p\u003e \u003cp\u003eAll modeling approaches with species-specific abundances and lemur species richness were performed in R v. 4.4.0 (R Core Team 2024) using RStudio v.4.1.748 (RStudio team \u003cspan citationid=\"CR101\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) interface. First, we assessed the normality of all continuous variables using the Shapiro\u0026ndash;Wilks test (Shapiro and Wilk 1965). If a variable was not normally distributed, we applied a logarithmic or square root transformation. We visually inspected the Q\u0026ndash;Q plots before and after the transformation to ensure improvement.\u003c/p\u003e \u003cp\u003eWe modeled the impact of two topographic and five fire-related fixed factors on species-specific abundances and lemur species richness. First, we analyzed the effect of terrain (valley, slope, and plateau) and elevation (m above sea level) on lemur abundance and lemur species richness. Second, we modeled the impact of five fire-related parameters on the abundance of six lemur species and on lemur species richness. Fire-related parameters were the forest zone (unburnt/burnt), the total number of fires, the year since the last fire, the minimum interval between two fires, and the maximum fire severity in each transect partition. When fitting multivariate models, we often experienced convergence problems (Finch and Finch \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Zhang et al. \u003cspan citationid=\"CR130\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). This was likely due to the limited data set (66 partitions in 18 study sites, Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e) and the resulting lack of statistical power when fitting several variables together. Therefore, we decided to calculate only univariate models for all parts of the analysis.\u003c/p\u003e \u003cp\u003eTo identify the determinants of species-specific abundances (dependent variables), we fitted Generalized Linear Mixed Models (GLMMs) using the package glmmTMB v.1.1.10 (Brooks et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) which allows for the implementation of different model families according to data structure and distribution. With the package DHARMa v.0.4.7 (Hartig and Hartig \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) we assessed the quality of each model by testing for over- and underdispersion, outliers, and zero inflation of the residuals. In case of a zero inflation, we added the command \u0026ldquo;ziformula\u0026rdquo; to the model; in the case of overdispersion and underdispersion, we added the command \u0026ldquo;dispformula\u0026rdquo; to the model (Brooks et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). We selected the Tweedie family for modeling in all cases. Its regression models can accomodate highly right-skewed, as well as symmetric and heavily tailed data distributions (Ma et al. \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2018\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eAs the lemur species richness (dependent variable) is always an integer number with an upper observed limit of six species, we first used the \u003cem\u003ecbind\u003c/em\u003e() function to relate the number of observed lemur species to the number of not-observed species in each partition. For this variable, we fitted Generalized Linear Mixed Effect models (GLMMs) using the package lme4 (Bates et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2015\u003c/span\u003e) with the binomial family, and the package DHARMa (Hartig and Hartig \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) to assess the model residuals (model dispersion, outliers).\u003c/p\u003e \u003cp\u003eWe included study site and month as independent random factors when modeling species abundances, and study site as well as transect partition length as independent random factors when modeling the lemur species richness. We verified the relevance of significant variables by testing whether introducing a specific variable explained significantly more variation in the dataset compared to a null model that only contained random factors and no explanatory variable. For this comparison, we used the ANOVA function in R with a threshold p-value of less than 0.05 indicating significance. During all species-specific analyses of the impact of forest zones (burnt, transition, and unburnt), we found no significant difference between burnt and transition zones, even when applying the Tukey post-hoc test with the R package emmeans v.1.5.1 (Lenth et al. \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Given these results, we categorized both transition and burnt transect zones as burnt throughout.\u003c/p\u003e \u003cp\u003eTo correct for potential error accumulation by multiple testing, we used the Holm-Bonferroni Method to adjust the threshold for significance (Holm \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e1979\u003c/span\u003e). We applied this correction separately to each species and the respective topographic and fire-related variable set. We then compared the adjusted alpha with the observed p-values (Holm \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e1979\u003c/span\u003e) and reported the results before and after this correction. We considered p\u0026thinsp;=\u0026thinsp;0.05 as the level of significance before correction.\u003c/p\u003e \u003cp\u003eAnalyses were based on two sets of data. The first set included the full dataset with all transect partitions in all sites. This dataset was used to test the impact of the topographic variables and two of the fire-related variables (zones and number of fires). The second dataset included only the dataset for the burnt partitions and was used for modeling the remaining three fire variables (time since the last fire, minimum interval between two fires, and maximum fire intensity). We generated figures using Microsoft Excel and the R packages ggplot2 v.3.4.0 (Wickham et al. \u003cspan citationid=\"CR128\" class=\"CitationRef\"\u003e2016\u003c/span\u003e) and ggpubr v.0.6.0 (Kassambara \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2018\u003c/span\u003e).\u003c/p\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec9\" class=\"Section2\"\u003e \u003ch2\u003eThe impact of terrain and elevation on lemur abundance and species richness\u003c/h2\u003e \u003cp\u003eDuring surveys, we encountered all eight lemur species present in the ANP, but only six of them had sufficient sightings for modeling abundances (Table S2). After correcting for multiple testing, the large \u003cem\u003eEulemur fulvus\u003c/em\u003e had significantly higher abundance in valleys than on the plateau (Estimate\u003csub\u003evalley\u003c/sub\u003e = 2.354, p\u0026thinsp;=\u0026thinsp;0.007; Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ea; Table S3), but the abundance on slopes did not differ from that in valleys or on the plateau (Tukey test: Estimate\u003csub\u003eslope\u0026minus;valley\u003c/sub\u003e = -3.011, p\u0026thinsp;=\u0026thinsp;0.115; Estimate\u003csub\u003eplateau\u0026minus;slope\u003c/sub\u003e = 0.657, p\u0026thinsp;=\u0026thinsp;0.897; Table S5). Elevation also affected the abundance negatively (Estimate = -1.807, p\u0026thinsp;=\u0026thinsp;0.025), i.e., we found less \u003cem\u003eE. fulvus\u003c/em\u003e in areas of higher elevation, even after correcting for multiple testing (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eb; Table S3). The abundance of the middle-sized \u003cem\u003eLepilemur edwardsi\u003c/em\u003e was not significantly impacted by terrain or elevation (Table S3) while the middle-sized \u003cem\u003eAvahi occidentalis\u003c/em\u003e showed a statistical trend for lower abundance in the valley (Estimate\u003csub\u003evalley\u003c/sub\u003e = -2.046, p\u0026thinsp;=\u0026thinsp;0.077; Table S3), but no longer after correcting multiple testing. Terrain and elevation did not influence the abundance of \u003cem\u003eCheirogaleus medius\u003c/em\u003e (Table S3). In contrast, the golden-brown mouse lemur, \u003cem\u003eMicrocebus ravelobensis\u003c/em\u003e, was significantly more abundant in valleys than on the plateau (Estimate\u003csub\u003evalley\u003c/sub\u003e = 0.920, p\u0026thinsp;\u0026lt;\u0026thinsp;0.0001; Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e, Table S3). The post-hoc Tukey test revealed that the abundance in the valleys was also significantly higher than on slopes (Estimate\u003csub\u003eslope\u0026minus;valley\u003c/sub\u003e = -0.524, p\u0026thinsp;=\u0026thinsp;0.042; Table S5), although there was no significant difference between the abundance on plateau and slopes (Estimate\u003csub\u003eplateau\u0026minus;slope\u003c/sub\u003e= -0.318, p\u0026thinsp;=\u0026thinsp;0.265; Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e, Table S5). There was also a statistical trend for a lower abundance at higher altitudes (Estimate = -0.060, p\u0026thinsp;=\u0026thinsp;0.067; see Table S3), but not after correcting for multiple testing. In the case of the grey mouse lemur, \u003cem\u003eMicrocebus murinus\u003c/em\u003e, neither terrain nor elevation showed a significant impact on abundance, and a statistical trend for a lower abundance in the valley than on the plateau (Estimate\u003csub\u003evalley\u003c/sub\u003e = -1.221, p\u0026thinsp;=\u0026thinsp;0.053, Table S3) did not persist after correcting for multiple testing. Finally, the lemur species richness was also not statistically impacted by terrain or elevation (Table S3).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eThe influence of time since the last fire on lemur presence/absence\u003c/h3\u003e\n\u003cp\u003eThe time since the most recent fire in the forest ranged from one to over 35 years (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). All species except \u003cem\u003eEulemur mongoz\u003c/em\u003e were recorded along transects in forest that had not been burned during that period (unburnt, Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). The three largest species \u003cem\u003ePropithecus coquereli\u003c/em\u003e, \u003cem\u003eE. mongoz\u003c/em\u003e, and \u003cem\u003eE. fulvus\u003c/em\u003e were almost only observed in unburnt zones or burnt zones with fires older than 23 years. \u003cem\u003eP. coquereli\u003c/em\u003e was only sighted along transects in four sites altogether (sites 1, 8, 11, 13) but was encountered outside the study transects in these same sites and in four others (sites 2, 15, 21, 24). Only one individual of \u003cem\u003eP. coquereli\u003c/em\u003e was encountered in the transition zone (burnt) in one of our study sites with a very recent fire (1\u0026nbsp;year old). \u003cem\u003eE. mongoz\u003c/em\u003e was sighted along transects in only one site and outside the study transects in six more sites. In contrast, the medium-sized lemurs \u003cem\u003eAvahi occidentalis\u003c/em\u003e and \u003cem\u003eLepilemur edwardsi\u003c/em\u003e were observed in 69.2% and 76.9% of the unburnt zones, respectively (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). They were also recorded in 37.0% (n\u0026thinsp;=\u0026thinsp;10) and 33.3% (n\u0026thinsp;=\u0026thinsp;9) of the burnt zones with different fire ages, respectively, even in those that experienced fires less than six years ago. Likewise, \u003cem\u003eCheirogaleus medius\u003c/em\u003e was observed in 50% of the unburnt zones as well as in 62.5% of the burnt zones, sometimes in zones of relatively young fires (2\u0026ndash;6 years old) but also in old fire zones (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). The two smallest lemur species, \u003cem\u003eMicrocebus ravelobensis\u003c/em\u003e and \u003cem\u003eMicrocebus murinus\u003c/em\u003e, were found in almost all unburnt zones (\u003cem\u003eM. ravelobensis\u003c/em\u003e: 100%, \u003cem\u003eM. murinus\u003c/em\u003e: 76.9%) and in almost all burnt zones (\u003cem\u003eM. ravelobensis\u003c/em\u003e: 96.3%, \u003cem\u003eM. murinus\u003c/em\u003e: 66.7%) ranging from the most recently burned (1\u0026nbsp;year post-fire) to those with older fires.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003eThe impact of fire-related variables on lemur abundance and lemur species richness\u003c/h2\u003e \u003cp\u003eWe detected a negative impact of the number of fires on the abundance of \u003cem\u003eEulemur fulvus\u003c/em\u003e (Estimate = -1.267, p\u0026thinsp;=\u0026thinsp;0.041, Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e, Table S4), but only before Holm-Bonferroni correction. Qualitatively, observations of \u003cem\u003eE. fulvus\u003c/em\u003e dropped to almost zero, whenever there have been more than three fires in a site (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e). We also noted a statistical trend for the positive impact of the number of years since the last fire on \u003cem\u003eE. fulvus\u003c/em\u003e abundance (Estimate\u0026thinsp;=\u0026thinsp;0.133, p\u0026thinsp;=\u0026thinsp;0.051, Table S4), but only before Holm-Bonferroni correction. The statistical modeling did not reveal any other impact of fire-related variables on the abundance of \u003cem\u003eE. fulvus\u003c/em\u003e.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eThe abundance of the medium-sized \u003cem\u003eAvahi occidentalis\u003c/em\u003e in the unburnt zone was higher than in the burnt zones and negatively associated with the number of fires but was rejected after we applied the Holm-Bonferroni correction (Table S4). Qualitatively, we noted that this species was absent from sites that burned more than three times and whenever the minimum interval between two fires was less than eight years (Table S2). The abundance of the medium-sized \u003cem\u003eLepilemur edwardsi\u003c/em\u003e was impacted by the burn status (burnt vs. unburnt), the number of fires in a site, the minimum time interval between fires, and the maximum fire intensity (Table S4) even after correcting for multiple testing. Specifically, the species was significantly more abundant in unburnt compared to burnt areas (Estimate\u003csub\u003eUnburnt\u003c/sub\u003e = 0.923, p\u0026thinsp;=\u0026thinsp;0.020; Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003ea, Table S4). Moreover, the number of fires negatively influenced the abundance of \u003cem\u003eL. edwardsi\u003c/em\u003e (Estimate = -1.131, p\u0026thinsp;=\u0026thinsp;0.003, Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003eb) to the point that the species was no longer observed at sites that had experienced more than three fires. Additionally, larger minimum intervals between fires lead to increased abundance of \u003cem\u003eL. edwardsi\u003c/em\u003e, and this species was not found in burnt zones with minimum fire intervals of less than eight years (Estimate\u0026thinsp;=\u0026thinsp;1.113, p\u0026thinsp;=\u0026thinsp;0.012; Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003ec). Lastly, higher maximum fire severities negatively affected the abundance of \u003cem\u003eL. edwardsi\u003c/em\u003e (Estimate = -21.338, p\u0026thinsp;=\u0026thinsp;0.012) leading to an absence of observations from sites with very high maximum severity (Severity\u0026thinsp;\u0026gt;\u0026thinsp;0.12, Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003ed). The other variables had no impact on \u003cem\u003eL. edwardsi\u003c/em\u003e abundance (Table S4).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eThe abundance of \u003cem\u003eCheirogaleus medius\u003c/em\u003e was significantly and positively impacted by the year since the last fire (Estimate\u0026thinsp;=\u0026thinsp;0.358, p\u0026thinsp;=\u0026thinsp;0.041) but this effect was no longer significant after Holm-Bonferroni correction. No other fire-related variable showed any significant effect on the abundance of \u003cem\u003eC. medius\u003c/em\u003e (Table S4). The abundance of \u003cem\u003eMicrocebus murinus\u003c/em\u003e showed a statistical trend for a positive impact of the minimum interval between two fires (Estimate\u0026thinsp;=\u0026thinsp;0.681, p\u0026thinsp;=\u0026thinsp;0.071) but only before Holm-Bonferroni correction. No other fire-related variable showed a statistical effect on this species (Table S4). Finally, no fire-related variable showed any statistical effect on the abundance of \u003cem\u003eMicrocebus ravelobensis\u003c/em\u003e. (Table S4).\u003c/p\u003e \u003cp\u003eThe burn status and the number of fires (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003ea, b; Table S4) statistically influenced lemur species richness. Specifically, we found a significantly higher lemur species richness in the unburnt zone compared to the burnt zones (Estimate\u003csub\u003eUnburn\u003c/sub\u003e = 1.140, p\u0026thinsp;\u0026lt;\u0026thinsp;0.0001; Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003ea) even after Holm-Bonferroni correction. In addition, the number of fires negatively influenced lemur species richness (Estimate = -1.175, p\u0026thinsp;\u0026lt;\u0026thinsp;0.0001, Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003eb), also persisting after Holm-Bonferroni correction. The year since the last fire showed a positive statistical trend (Estimate\u0026thinsp;=\u0026thinsp;0.172, p\u0026thinsp;=\u0026thinsp;0.058; Table S4) for an influence on lemur species richness, likewise, the maximum fire severity showed a negative statistical trend (Estimate = -3.583, p\u0026thinsp;=\u0026thinsp;0.054; Table S4), but both trends were rejected after correcting for multiple testing. The minimum interval between fires showed no statistical effect.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e"},{"header":"Discussion","content":"\u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003eEffects of forest fires on large-sized lemur species\u003c/h2\u003e \u003cp\u003eDue to the rarity of sightings, we could not conduct statistical analyses on two of the largest lemur species in Ankarafantsika National Park (ANP), the Critically Endangered \u003cem\u003ePropithecus coquereli\u003c/em\u003e and \u003cem\u003eEulemur mongoz\u003c/em\u003e. \u003cem\u003eP. coquereli\u003c/em\u003e was only encountered in four of 18 sites (22%) along transects and was then mainly observed in unburnt forests. As the largest lemur species in ANP that predominantly moves by vertical clinging and leaping, \u003cem\u003eP. coquereli\u003c/em\u003e relies on robust, vertical substrates for movement, such as trunks and branches of large trees (Mittermeier et al. \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) which were shown to be diminished in numbers and connectivity at least in one recently burnt part of the park (Percival et al. \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). The importance of suitable substrates for locomotion is also suggested by the observation that this species was never observed on the ground inside the forest (Rabemananjara and Radespiel, unpubl. Data), although such observations were made in a non-forest matrix in another study outside ANP (Ramilison et al. \u003cspan citationid=\"CR95\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Previous research by Kun-Rodrigues et al. (\u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2014\u003c/span\u003e) revealed that \u003cem\u003eP. coquereli\u003c/em\u003e is mostly found in the forest core and avoids forest edges, suggesting sensitivity to disturbance or possibly edge-related ecological changes (Steffens and Lehman \u003cspan citationid=\"CR115\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Ecological changes like those at forest edges can also be expected as a result of forest fires, as canopy cover then diminishes which likely drives cascading effects on microclimate, vegetation structure, and floristic composition (Barlow and Peres \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Percival et al. \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Another explanation for the fire sensitivity of \u003cem\u003eP. coquereli\u003c/em\u003e may be related to its feeding habits. \u003cem\u003eP. coquereli\u003c/em\u003e has a predominantly folivorous diet (Mittermeier et al. \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), and plant diversity tends to be higher in the forest interior than at the edge (McGoogan \u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). As forest fires very likely also lead to a reduction of plant diversity (Agus et al. 2018), burnt zones may not provide sufficient food resources for \u003cem\u003eP. coquereli\u003c/em\u003e. Such a loss of floristic diversity may be particularly expressed in forests that experience recurrent fires that likely lead to floristic shifts in entire plant communities towards pioneer, fire-resistant species (Da Silva et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Indeed, \u003cem\u003eP. coquereli\u003c/em\u003e was never encountered in burnt zones that had experienced multiple fires over the last 35 years. However, we did once encounter a group of \u003cem\u003eP. coquereli\u003c/em\u003e in the transition zone of a one-year-old fire and sighted another group twice in a 24-years post-fire zone. While the sightings in the transition zone may indicate a rather exploratory use of the recently burnt zone, the presence in the older fire zone suggests that this species may eventually return to previously burnt areas once conditions become favorable again. However, our current study does not provide conclusive evidence of such recolonization events yet. Further research and data collection, including longer behavioral observations of \u003cem\u003eP. coquereli\u003c/em\u003e living close to burnt forest vegetation are needed to illuminate its habitat and resource use and thereby potential adaptations to fire-affected areas.\u003c/p\u003e \u003cp\u003e \u003cem\u003eEulemur mongoz\u003c/em\u003e was encountered only twice during our entire study (once in the burnt and once in the unburnt zone of the same study site) when systematically surveying transects. One of the reasons for this rarity of sightings could be its seasonally changing activity rhythm that switches between largely diurnal activity during the rainy season to largely nocturnal activity during much of the dry season i.e., during our study period (Curtis \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). This activity change to nocturnality could complicate their detection, in particular, since \u003cem\u003eE. mongoz\u003c/em\u003e has a rather weakly developed \u003cem\u003etapetum lucidum\u003c/em\u003e (Martin \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e1995\u003c/span\u003e) and forms pairs or small family groups (Curtis and Zaramody \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e1999\u003c/span\u003e) that are less conspicuous than, for example, larger social groups formed by \u003cem\u003eEulemur fulvu\u003c/em\u003es (Mittermeier et al. \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). However, one of the two sightings during our surveys was made during daytime in the middle of the dry season (July), and \u003cem\u003eE. mongoz\u003c/em\u003e was furthermore confirmed near six other study sites (six times seen during daytime, one time seen during nighttime), but not along the survey transects (Rabemananjara and Radespiel, unpubl. results). These opportunistic sightings suggest that this species was indeed rare along the study transects but not impossible to find inside ANP during the dry season. One explanation for their rarity in ANP could be their diet, which was suggested to be highly specialized, encompassing nectar, fruits, flowers, and leaves from a rather limited number of plant species (Sussman and Tattersall \u003cspan citationid=\"CR118\" class=\"CitationRef\"\u003e1976\u003c/span\u003e). If those food plants were not available in the entire park, but only in some favorable forest parts, such a dietary specialization could make them extremely vulnerable to disturbances including fires. However, our sighting of \u003cem\u003eE. mongoz\u003c/em\u003e in one forest zone that had burnt 24 years ago also shows some flexibility and resilience at least given older fires and their long-term effects. Nevertheless, the overall low number of encounters with \u003cem\u003eE. mongoz\u003c/em\u003e aligns with other studies. Steffens et al. (\u003cspan citationid=\"CR116\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) reported no sightings of the Critically Endangered \u003cem\u003eE. mongoz\u003c/em\u003e during 22.15 km of survey effort in the continuous forest of ANP, while Steffens and Lehman (\u003cspan citationid=\"CR115\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) found none over 468.18 km of surveys in 42 forest fragments in western ANP.\u003c/p\u003e \u003cp\u003eThe abundance of \u003cem\u003eEulemur fulvus\u003c/em\u003e appeared to be negatively influenced by the number of fires. However, the statistical support for this parameter was relatively weak which may be due to a limited statistical power or to a plastic fire response of this species. Such a plasticity may be plausible, as this species was described as a flexible generalist (Sato \u003cspan citationid=\"CR105\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) which is also suggested by its wide distribution that includes dry forests in northwestern Madagascar as well as humid rainforest habitats in central eastern Madagascar (Mittermeier et al. \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Our ecological modeling results also indicated a certain affinity to humid conditions, suggesting higher abundances of \u003cem\u003eE. fulvus\u003c/em\u003e at lower elevations and therefore under more humid conditions (valleys) than in drier localities at higher elevations (plateau). However, some sensitivity of \u003cem\u003eE. fulvus\u003c/em\u003e to fires was suggested by two other findings of our study: First, \u003cem\u003eE. fulvus\u003c/em\u003e was rarely sighted in areas that had experienced more than three fires. Multiple fires were already shown to lead to increased levels of dry forest degradation, and it was hypothesized that such degraded patches may undergo very long-term or even irreversible vegetation shifts and ecosystem changes (Percival et al. \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) and thereby may become permanently uninhabitable for \u003cem\u003eE. fulvus\u003c/em\u003e. Second, \u003cem\u003eE. fulvus\u003c/em\u003e was only sighted in burnt zones of fires that were more than 23 years old suggesting that recently burnt dry forests also do not provide suitable habitat for \u003cem\u003eE. fulvus\u003c/em\u003e. Whether they lack essential food resources, suitable substrates for locomotion, and/or cover from predators cannot be answered at present but these questions certainly ask for more specific future research.\u003c/p\u003e \u003cp\u003eCompared to other non-human primates, the three large lemurs in ANP require longer periods to recolonize burnt dry deciduous forests. For instance, siamangs in Sumatra (\u003cem\u003eSymphalangus syndactylus\u003c/em\u003e) were reported to return to burnt areas after approximately 18 years (Lappan et al., \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Similarly, leaf-eating monkeys (\u003cem\u003ePresbytis\u003c/em\u003e spp.) in Indonesia recolonized burnt forests within six years post-fire, although at low densities (Chapman and Onderdonk, \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e1998\u003c/span\u003e). This difference could indicate that large lemurs may be more sensitive to habitat disturbances than other primate taxa, but it may also be the result of much slower vegetation regrowth and regeneration of the dry forests of Madagascar (Percival et al. \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) compared to other more humid tropical forest ecosystems on the island or in other parts of the world. Percival et al. (\u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) already suggested that the dry forests of ANP may have very low resilience to fire compared to fire-adapted dry forests in other parts of the world (Cavender-Bares et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Mcmichael et al. \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Lozano et al. \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). However, our data do not support their additional hypothesis that even one fire in this fire-sensitive forest may be sufficient to initiate a permanent transformation of dry forest into a degraded savanna/grassland vegetation. The sightings of the large lemur species in burnt forest parts of ANP after more than 23 years post-fire rather suggest that post-fire regrowth and regeneration are possible in these dry forests, even though it may take several decades.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec14\" class=\"Section2\"\u003e \u003ch2\u003eEffects of forest fires on medium-sized lemur species\u003c/h2\u003e \u003cp\u003eModeling revealed no significant impact of fires on the abundance of the medium-sized \u003cem\u003eAvahi occidentalis\u003c/em\u003e. This species was even recorded in some areas affected by recent fires (1\u0026ndash;5 years post-fire), although it was recorded in burnt partitions of only 50% of the study sites. However, it was also observed in only 69% of the unburnt zones, and these mixed sighting results suggest that \u003cem\u003eA. occidentalis\u003c/em\u003e may have specific resource requirements (Thalmann \u003cspan citationid=\"CR120\" class=\"CitationRef\"\u003e2001\u003c/span\u003e) that are not uniformly met in ANP, but that may not be necessarily impaired by forest fires per se. Our findings also suggested that \u003cem\u003eA. occidentalis\u003c/em\u003e may be sensitive to multiple fires. Specifically, we noted that \u003cem\u003eA. occidentalis\u003c/em\u003e was never sighted in burnt zones that experienced more than three fires or whenever the minimum interval between consecutive fires was less than eight years. Similar to the argument made for large lemur species, multiple fires likely cause significant forest degradation (Percival et al. \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Such degradation may render the habitat unsuitable or permanently uninhabitable for this medium-sized lemur species. Given that \u003cem\u003eA. occidentalis\u003c/em\u003e relies on vertical clinging to tree trunks or large branches and leaping between such positions in large trees (Warren and Crompton \u003cspan citationid=\"CR127\" class=\"CitationRef\"\u003e1997\u003c/span\u003e), the availability of suitable arboreal substrates for this species-specific mode of locomotion may be an essential component of suitable habitats, which may be permanently compromised after multiple fires.\u003c/p\u003e \u003cp\u003eThe medium-sized \u003cem\u003eLepilemur edwardsi\u003c/em\u003e exhibited an even stronger negative response to fires than \u003cem\u003eA. occidentalis\u003c/em\u003e. Although \u003cem\u003eL. edwardsi\u003c/em\u003e was also occasionally sighted in recently burnt areas (1\u0026ndash;5 years post-fire) and in burnt partitions of 38% (n\u0026thinsp;=\u0026thinsp;7) of the study sites, it was significantly more abundant in unburnt than in burnt zones. \u003cem\u003eL. edwardsi\u003c/em\u003e was also absent in areas with more than three fires or when the interval between fires was shorter than eight years. Additionally, \u003cem\u003eL. edwardsi\u003c/em\u003e responded negatively to increasing fire severity. We hypothesize that severe fires, particularly the degradation caused by repeated fires (Percival et al., \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), reduce the availability of large trees that serve as substrates for vertical clingers and leapers (Warren and Crompton, \u003cspan citationid=\"CR127\" class=\"CitationRef\"\u003e1997\u003c/span\u003e). Additionally, these fires destroy large trees provide shelters, such as tree holes, which this species uses for daily resting (Rasoloharijaona et al., \u003cspan citationid=\"CR97\" class=\"CitationRef\"\u003e2003\u003c/span\u003e). Sleeping holes of \u003cem\u003eL. edwardsi\u003c/em\u003e in ANP were previously described to lie at about 3\u0026ndash;8 m height above ground, and sleeping holes had a depth of more than 80 cm and an average wall thickness of 11 cm (Rasoloharijaona et al. \u003cspan citationid=\"CR97\" class=\"CitationRef\"\u003e2003\u003c/span\u003e), which can clearly only be provided by large trees. If this important resource, which makes up 92% of the used shelters, is no longer available, it can be expected that the suitability and possibly the carrying capacity of a degraded post-fire forests drops considerably for \u003cem\u003eL. edwardsi\u003c/em\u003e. Post-fire food availability may also play a role in limiting the abundance of \u003cem\u003eL. edwardsi\u003c/em\u003e, although further research is needed to investigate this hypothesis.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec15\" class=\"Section2\"\u003e \u003ch2\u003eEffects of forest fires on small-sized lemur species\u003c/h2\u003e \u003cp\u003eFires did not significantly influence the abundance of the three small-sized nocturnal lemurs like \u003cem\u003eCheirogaleus medius, Microcebus murinus\u003c/em\u003e and \u003cem\u003eM. ravelobensis\u003c/em\u003e. All three species were recorded in both recently burned forests (1\u0026ndash;5 years post-fire) and in forests affected by older fires (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). All three species can also be categorized as omnivores, as they can consume flowers, fruits, seeds, invertebrates, insect secretions and even occasionally small vertebrates (Fietz and Ganzhorn \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Radespiel et al. \u003cspan citationid=\"CR90\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Joly-Radko and Zimmermann \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Thor\u0026eacute;n et al. \u003cspan citationid=\"CR122\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). This dietary flexibility likely allows them to explore burned forests of various ages. Increased sun exposure in these areas may alter temperature and radiation, leading to more insect productivity and changes in fruit types and availability (Hamer \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e1996\u003c/span\u003e; Keane et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Pausas and Keeley \u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Thompson et al. \u003cspan citationid=\"CR121\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). \u003cem\u003eC. medius\u003c/em\u003e was already stated to be quite resilient to environmental changes (Rakotoniaina et al. \u003cspan citationid=\"CR93\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Its habit to hibernate from May to Mid-September in ANP (M\u0026uuml;ller \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e1999\u003c/span\u003e) which partially overlaps with the fire season (July to October), very likely helps to avoid resource shortage during periods of food scarcity, and this ability may further contribute to its resilience in highly modified post-fire zones. However, some studies indicated that \u003cem\u003eC. medius\u003c/em\u003e tends to occur at lower densities in disturbed than in undisturbed habitats (Rakotoniaina et al. \u003cspan citationid=\"CR93\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Hending \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). This aligns with our findings that this species only reached abundances that are comparable to those in unburnt forests when the burnt areas reached a certain age post-fire (\u0026gt;\u0026thinsp;23 years) (Table S2), suggesting that the availability of suitable resources and habitat structure improves and regenerates with time following a fire. If this preliminary observation would be confirmed in future studies on a larger number of study sites, our lack of statistical support for negative fire effects may also be based on the small sample size (i.e, limited statistical power) with only 11 sites that could be studied after the end of the yearly hibernation period in \u003cem\u003eC. medius\u003c/em\u003e.\u003c/p\u003e \u003cp\u003eThe two small-sized mouse lemurs, on the other hand, were present in zones of very recent to old fires (one to \u0026gt;\u0026thinsp;35 years old), and a negative impacts of fires on their abundance could not be detected. In other words, \u003cem\u003eMicrocebus murinus\u003c/em\u003e and \u003cem\u003eM. ravelobensis\u003c/em\u003e showed rather similar abundances in recently burnt and older burnt zones, which underlines their high ecological flexibility and confirms previous findings that they inhabit various habitat types of different stages of degradation including secondary forest (Rakotondravony and Radespiel \u003cspan citationid=\"CR92\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Steffens and Lehman \u003cspan citationid=\"CR115\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Hending \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Steffens et al. \u003cspan citationid=\"CR117\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). However, they are not reacting equally to habitat fragmentation (Andriatsitohaina et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Steffens et al. \u003cspan citationid=\"CR117\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). While \u003cem\u003eM. murinus\u003c/em\u003e can even be found in small to very small forest fragments that are embedded in a non-forest savanna matrix, \u003cem\u003eM. ravelobensis\u003c/em\u003e is usually found only in larger fragments or in continuous stretches of larger forests (Andriatsitohaina et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). However, in the case of geographically limited burnings in the continuous forest of ANP, the burnt forest usually borders somewhere upon stretches of unburnt forest or forest parts that underwent fires in other years. This may facilitate recolonization by both of these omnivorous generalists quickly after a fire is extinguished (Ramsay et al. \u003cspan citationid=\"CR96\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The high resilience of both species to fire may also result from their high flexibility when selecting day shelters, which can be liana tangles, shrubs, dead wood or bark, or self-built nests located between the ground and the canopy (Radespiel et al. \u003cspan citationid=\"CR88\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Radespiel et al. \u003cspan citationid=\"CR89\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Thor\u0026eacute;n et al. \u003cspan citationid=\"CR123\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). These shelters are likely available even in moderately degraded post-fire landscapes and may provide critical protection under high risk of predation by raptors, snakes and carnivores (Goodman et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e1993\u003c/span\u003e; Karpanty and Wright \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). Finally, mouse lemurs are also able to change their activity patterns and enter daily torpor in case of environmental challenges (Ortmann et al. \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; Schmid \u003cspan citationid=\"CR107\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Thor\u0026eacute;n et al. \u003cspan citationid=\"CR122\" class=\"CitationRef\"\u003e2011\u003c/span\u003e) which may contribute further to energetic resilience in post-fire landscapes.\u003c/p\u003e \u003cp\u003eEven though this general resilience to fire has thereby been confirmed for the small, nocturnal lemurs in ANP, future research should still investigate the impact of varying fire severity and fire frequency on resource availability, reproductive output, predation risk, space use, and the resulting population sizes of these rather flexible, seasonal species in burnt zones. Even if small lemurs may use burnt forests to some extent, altered food availability and predation risk might still constrain population sizes and modulate stochastic extinction risks over time (James et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e1997\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec16\" class=\"Section2\"\u003e \u003ch2\u003eEffects of forest fires on lemur species richness\u003c/h2\u003e \u003cp\u003eHabitat alteration can significantly affect species richness, particularly for primates, by altering vegetation structure, food availability, and ecosystem stability (Schwitzer et al. \u003cspan citationid=\"CR110\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). Our study has shown that the species richness of lemurs was higher in the unburnt than in the burnt forests. Studies in other tropical forests worldwide have provided similar evidence for a variety of different taxa such as in small and large mammal communities (Santosa and Kwatrina \u003cspan citationid=\"CR103\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), bird communities (Adeney et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Davis et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Mardiastuti \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), and in herpetofaunal communities (Russell et al. \u003cspan citationid=\"CR102\" class=\"CitationRef\"\u003e1999\u003c/span\u003e). For instance, unburned forests that maintain an intact canopy cover and have undisturbed food sources such as fruits and leaves have been shown to support a higher diversity of primates in the Kibale National Park, Uganda (Chapman and Onderdonk \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e1998\u003c/span\u003e). Burned areas often host primarily generalist species that can adaptively respond to fire-modified landscapes due to their dietary flexibility (omnivory) and variable locomotion that allows mitigating fire-related vegetation changes (Santos et al. 2014). Such generalists in our study were only the smallest lemur species, specifically \u003cem\u003eM. ravelobensis\u003c/em\u003e and \u003cem\u003eM. murinus\u003c/em\u003e. They are particularly adept at taking advantage of resources that may quickly become available again after a fire, such as insects or seeds that remain in the ground (Radespiel et al. \u003cspan citationid=\"CR90\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Dammhahn and Kappeler \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Pruetz and Herzog \u003cspan citationid=\"CR86\" class=\"CitationRef\"\u003e2017\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eCumulative negative impacts of fire on lemur species richness were noted in particular after fires occurred repeatedly in sites. Recurrent fires in ANP were shown to negatively affect large lemurs, medium-sized lemurs, as well as \u003cem\u003eC. medius\u003c/em\u003e (see above). Multiple fires are very likely to hinder forest recovery and thereby disturb resource availability (Hermosilla et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2019\u003c/span\u003e), but also may increase the predation risk by elevated exposure to predators in the degraded vegetation (Shanee et al. \u003cspan citationid=\"CR112\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Moreover, large-scale fire-related vegetation shifts may lead to fragmentation of the remaining inhabitable forest patches that will likely disrupt and constrain movements and dispersal within and between the remaining sub-populations (Nimmo et al. \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Doherty et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Finally, all these challenges very likely increase levels of intraspecific competition, may require individual and population-level niche shifts, generate elevated stress levels that may result in decreased reproductive success (Bolnick et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Mart\u0026iacute;nez-Blancas and Martorell \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Such effects can pose a significant threat to primate diversity and ecosystem resilience, especially in tropical dry forests (Van Holstein et al. \u003cspan citationid=\"CR126\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec17\" class=\"Section2\"\u003e \u003ch2\u003eConservation implications\u003c/h2\u003e \u003cp\u003eOur study suggests that most lemur species do not recolonize burnt forests quickly. In fact, the lemur community in ANP only reached the level of unburnt forests when the fire happened more than 23 years ago. This slow recolonization may be attributed to various factors, including a lack of key food resources in burnt zones, insufficient substrates such as large trees needed for species-specific locomotion (e.g., clinging and leaping by \u003cem\u003eAvahi\u003c/em\u003e and \u003cem\u003eLepilemur\u003c/em\u003e) or as shelters (\u003cem\u003eLepilemur\u003c/em\u003e), and increased risks from predators or human activities in disturbed habitats. Future analyses are needed to investigate the main reasons for the species-specific fire sensitivity in more detail.\u003c/p\u003e \u003cp\u003eGiven this rather long time span that seems to be necessary for a full recovery of the lemur community after a fire, this study raises serious concerns about the amount of suitable forest habitat remaining for all lemur species in ANP. To obtain a first rough estimate of this value, we determined the area of the forest within ANP that had not experienced any fires in the last 35 years together with the area that was burned 24 years or longer ago (Fig.\u0026nbsp;\u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e8\u003c/span\u003e). Together, both partitions added up to approximately 466.6 km\u0026sup2; (45%) of the vegetation inside ANP that should be potentially suitable for the entire lemur community consisting of eight species in the park. However, species like \u003cem\u003eL. edwardsi, E. fulvus, E. mongoz\u003c/em\u003e and \u003cem\u003eP. coquereli\u003c/em\u003e face dual threats: habitat degradation and hunting, which has been documented even within protected areas like the Ankarafantsika National Park (Garcia and Goodman \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Borgerson et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Sato et al. \u003cspan citationid=\"CR106\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). The Critically Endangered \u003cem\u003eE. mongoz\u003c/em\u003e was found to be the rarest of all lemur species in our study, emphasizing the need to study the habitat requirements and environmental constraints of these lemurs in more depth. Their ability or inability to adapt to environmental changes may play a key role in determining their long-term survival in highly altered landscapes.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eForest fires significantly influenced lemur communities and may therefore threaten the ecosystem services that are provided by lemurs. Larger lemurs, for instance, play a crucial role in seed dispersal in intact forests (Ganzhorn et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Lahann \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Sato \u003cspan citationid=\"CR104\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Valenta et al. \u003cspan citationid=\"CR124\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). This predominant role of primate seed dispersers contrasts with other regions in the world where birds and bats also contribute substantially to these services (Howe \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e1986\u003c/span\u003e; Medellin and Gaona \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Herrera \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2002\u003c/span\u003e). After multiple fires in ANP, only the two smallest species, the omnivorous, nocturnal mouse lemurs, seemed to be resilient and adapt to the corresponding degradation in post-fire habitats. However, their seed dispersal capacity is limited due to their small body size (Ramananjato \u003cspan citationid=\"CR94\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) which likely biases forest regeneration toward smaller-fruited plants. A recent study suggested that the Cheirogaleidae might only disperse seeds of up to 15mm in size (Ramananjato \u003cspan citationid=\"CR94\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Ganzhorn et al. 2024). While this may help to prevent the long-term conversion of severely burnt forests into savannas, and fire-adapted lianas, for example, provide shade for early successional seedlings of species with smaller seeds, a full forest regeneration that also includes large-seeded tree species, requires a step-wise return of major seed dispersers of all sizes. Future studies are needed to compare seed sizes of plant communities in various stages of post-fire regeneration to test this hypothesis.\u003c/p\u003e \u003cp\u003eGiven the severity of fire impacts, we recommend several urgent conservation actions. First, the conservation status of the medium-sized lemurs (\u003cem\u003eA. occidentalis, L. edwardsi)\u003c/em\u003e and in particular that of the Endangered \u003cem\u003eL. edwardsi\u003c/em\u003e (IUCN \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) should be reassessed given that they are heavily impacted by fire (this study) and hunting (Craul et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Sato et al. \u003cspan citationid=\"CR106\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), and forest fires are very prominent threats for the remaining dry forests (Frappier-Brinton and Lehman \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Second, conservation planning should focus on safeguarding the remaining unburnt forests, as they are vital for the survival of existing sub-populations and can act as source habitats facilitating the recolonization of fire-affected areas. Additionally, long-term protection strategies must be adopted to prevent burnt areas from experiencing subsequent fires, as repeated fires can significantly hinder forest recovery (Percival et al. \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) and further endanger lemur populations and lemur species richness (this study). Third, the conservation efforts of the park management should be complemented by educational and socioeconomic programs to raise awareness in the rural communities, improve livelihoods, and reduce the lemur hunting pressure to protect these endangered species and facilitate long-term coexistence of humans and wildlife near and inside the park.\u003c/p\u003e \u003c/div\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eSupplementary Information\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003eThe online version contains supplementary material available at \u0026hellip;..\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor contributions\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eUR and DS conceptualized the study and developed the methodology; UR acquired the funding; NRR, MOR, RM, and SHR were responsible for data collection; MR provided the fire history of study sites via remote sensing work; NRR conducted the data analyses; UR, RR and HR supervised the work; NRR and UR took the lead in writing the manuscript. All authors participated in editing the manuscript drafts and approved the final version for submission.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis study was financially supported by Madagascar National Parks under contract number 20/DG/DOP/DAFRH/CONV/2022, as well as by the Cologne Zoo, Germany.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eData availability\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe datasets generated and analyzed during the current study are provided in the supplementary material.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eDeclaration of\u003c/strong\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003cstrong\u003ecompeting interests\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;The authors declare no competing interests.\u003c/p\u003e\u003ch2\u003eAcknowledgement\u003c/h2\u003e\u003cp\u003e We want to express our gratitude to the Minist\u0026egrave;re de l\u0026rsquo;Environnement et du D\u0026eacute;veloppement Durable and the Direction R\u0026eacute;gionale de l\u0026rsquo;Environnement (DREDD) Boeny Betsiboka for granting us permission to conduct this research through the following research permits: № 275/22/MEDD/SG/DGGE/DAPRNE/SCBE.Re, № 070/23/MEDD/SG/DGGE/DAPRNE/SCBE.Re, and № 263/23/MEDD/SG/DGGE/DAPRNE/SCBE.Re. We sincerely thank Madagascar National Parks for their unwavering administrative and financial support, especially to the General Directors, the financial department, and Madame Lalatiana Odile Randriamiharisoa. Our appreciation also goes to the former and current Directors of Ankarafantsika National Park, Mandimby Heriniaina Andriambololona and Charles Andriamaniry, as well as the Chef du Volet Op\u0026eacute;ration, logistical staff, Chefs Secteurs, and park agents for their support during our fieldwork. We are grateful to the Mention Anthropologie et D\u0026eacute;veloppement Durable from the Faculty of Sciences at the University of Antananarivo for facilitating all our research permits. Additionally, we thank our park guides, Jhonny Kennedy and Estoria Una, as well as the local population (CLP and Cook) for their assistance during our site visits. We also appreciate the support of Marlene B\u0026ouml;hm, Johnny Randriafenontsoa, and Fenohery Andriantsitohaina during the data collection process in the field. This project was funded by Madagascar National Parks and supported by the Kreditanstalt f\u0026uuml;r Wiederaufbau (KFW) under contract number 20/DG/DOP/DAFRH/CONV/2022. The Cologne Zoo, Germany, also provided funding for part of the fieldwork.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAdeney JM, Ginsberg JR, Russell GJ, and Kinnaird MF (2006) Effects of an ENSO-related fire on birds of a lowland tropical forest in Sumatra. Anim Conserv 9: 292-301. https://doi.org/10.1111/j.1469-1795.2006. 00035.x\u003c/li\u003e\n\u003cli\u003eAgus C, et al. (2019) The impact of forest fire on the biodiversity and the soil characteristics of tropical peatland. In: Leal Filho W, Barbir J, Preziosi R (eds) Handbook of climate change and biodiversity. Climate Change Management. 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Nat Ecol Evol 8: 1-16. https://doi.org/10.1038/s41559-024-02547-w\u003c/li\u003e\n\u003cli\u003evan Holstein LA, McKay HD, Pimiento C, and Koops K (2024) Multidimensional primate niche space sheds light on interspecific competition in primate evolution. Commun Biol 7: 647. https://doi.org/10.1038/s42003-024-06324-0\u003c/li\u003e\n\u003cli\u003eWarren RD, and Crompton RH (1997) Locomotor ecology of \u003cem\u003eLepilemur edwardsi \u003c/em\u003eand \u003cem\u003eAvahi occidentalis\u003c/em\u003e. Am J Phys Anthropol 104: 471-486. https://doi.org/10.1002/(SICI)1096-8644(199712)104:4\u0026lt;471::AID-AJPA 4\u0026gt; 3.0.CO;2-V\u003c/li\u003e\n\u003cli\u003eWickham H, Chang W, and Wickham MH (2016) Package \u0026lsquo;ggplot2\u0026rsquo;. Create elegant data visualizations using the grammar of graphics. Version 2. https://ggplot2.tidyverse.org\u003c/li\u003e\n\u003cli\u003eWilson M, Chen XY, Corlett RT, Didham RK, Ding P, Holt RD, Holyoak M, Hu G, Hughes AC, Jiang L, Laurance WF, Liu J, Pimm SL, Robinson SK, Russo SE, Si X, Wilcove DS, Wu J, and Yu M (2015) Habitat fragmentation and biodiversity conservation: key findings and future challenges. Landsc Ecol 31: 219\u0026ndash;227. https://doi.org/10.1007/s10980-015-0312-3\u003c/li\u003e\n\u003cli\u003eZhang Y, Zhou H, Zhou J, and Sun W (2017) Regression Models for Multivariate Count Data. J Comput Graph Stat 26: 1-13. https://doi.org/10.1080/10618600.2016.1154063\u003c/li\u003e\n\u003c/ol\u003e"},{"header":"Table 2","content":"\u003cp\u003eTable 2 is available in the Supplementary Files section.\u003c/p\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"biodiversity-and-conservation","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"bioc","sideBox":"Learn more about [Biodiversity and Conservation](https://www.springer.com/journal/10531)","snPcode":"10531","submissionUrl":"https://submission.nature.com/new-submission/10531/3","title":"Biodiversity and Conservation","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"lemur abundance, species richness, forest fire, recolonization, Ankarafantsika, adaptability","lastPublishedDoi":"10.21203/rs.3.rs-5735404/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-5735404/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eWildfires significantly threaten biodiversity, especially in tropical regions like Madagascar, where unique ecosystems face ongoing habitat loss and degradation. This study investigated the effects of forest fires on lemur abundance, species richness, and their ability to recolonize burnt vegetation in Ankarafantsika National Park (ANP), the largest protected dry deciduous forest in northwestern Madagascar. ANP hosts eight lemur species with one diurnal (\u003cem\u003ePropithecus coquereli\u003c/em\u003e), two cathemeral (\u003cem\u003eEulemur mongoz\u003c/em\u003e, \u003cem\u003eE. fulvus\u003c/em\u003e), and five nocturnal species (\u003cem\u003eAvahi occidentalis\u003c/em\u003e, \u003cem\u003eLepilemur edwardsi\u003c/em\u003e, \u003cem\u003eCheirogaleus medius\u003c/em\u003e, \u003cem\u003eMicrocebus murinus\u003c/em\u003e, and \u003cem\u003eM. ravelobensis\u003c/em\u003e). Eighteen sites with varying fire histories (1 to \u0026gt;\u0026thinsp;35 years post-fire) and adjacent unburnt forest parts were surveyed using diurnal and nocturnal distance sampling. Transects included burnt (700 m) and unburnt (500 m) sections. Generalized linear mixed models (GLMMs) assessed the effect of fire variables such as time since the last fire, number of fires, intervals between fires, and fire severity on lemur abundance and species richness. A full lemur community was observed only in unburnt forests and areas with extended post-fire recovery (\u0026ge;\u0026thinsp;23 years). Fires negatively impacted \u003cem\u003eE. fulvus\u003c/em\u003e and \u003cem\u003eL. edwardsi\u003c/em\u003e, while they did not significantly affect the abundance of small nocturnal species (\u003cem\u003eC. medius\u003c/em\u003e, \u003cem\u003eMicrocebus\u003c/em\u003e spp.). Lemur species richness was higher in unburnt zones and decreased with an increasing number of fires. These findings reveal the need for long recovery periods for lemur communities post-fire, suggest species-specific fire vulnerabilities, and demonstrate significant faunal impacts of this destructive driver of landscape transformation.\u003c/p\u003e","manuscriptTitle":"Post-fire recolonization of dry deciduous forests by lemurs in northwestern Madagascar","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-01-06 03:12:11","doi":"10.21203/rs.3.rs-5735404/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2025-04-29T19:38:52+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-03-18T22:50:00+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"35592504465031205272378673838782600113","date":"2025-03-03T15:25:24+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-03-03T11:41:50+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"205132387367970156892451016487970957481","date":"2025-03-03T10:18:30+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"138829162103460869530333127347701997718","date":"2025-03-03T00:54:49+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"47308230851516312401391296591189192220","date":"2025-03-01T07:22:25+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-02-26T17:03:30+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-01-03T19:46:54+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-01-02T14:51:02+00:00","index":"","fulltext":""},{"type":"submitted","content":"Biodiversity and Conservation","date":"2024-12-30T12:00:10+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"biodiversity-and-conservation","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"bioc","sideBox":"Learn more about [Biodiversity and Conservation](https://www.springer.com/journal/10531)","snPcode":"10531","submissionUrl":"https://submission.nature.com/new-submission/10531/3","title":"Biodiversity and Conservation","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"30162554-1d66-4252-aae1-f6dca47201c2","owner":[],"postedDate":"January 6th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2025-10-13T16:03:43+00:00","versionOfRecord":{"articleIdentity":"rs-5735404","link":"https://doi.org/10.1007/s10531-025-03167-x","journal":{"identity":"biodiversity-and-conservation","isVorOnly":false,"title":"Biodiversity and Conservation"},"publishedOn":"2025-10-06 15:57:32","publishedOnDateReadable":"October 6th, 2025"},"versionCreatedAt":"2025-01-06 03:12:11","video":"","vorDoi":"10.1007/s10531-025-03167-x","vorDoiUrl":"https://doi.org/10.1007/s10531-025-03167-x","workflowStages":[]},"version":"v1","identity":"rs-5735404","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-5735404","identity":"rs-5735404","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}