Edge effects on amphibian diversity in the Yoko Forest Reserve, Democratic Republic of the Congo

Edge effects on amphibian diversity in the Yoko Forest Reserve, Democratic Republic of the Congo | Authorea try { document.documentElement.classList.add('js'); } catch (e) { } var _gaq = _gaq || []; _gaq.push(['_setAccount', 'G-8VDV14Y67G']); _gaq.push(['_trackPageview']); (function() { var ga = document.createElement('script'); ga.type = 'text/javascript'; ga.async = true; ga.src = ('https:' == document.location.protocol ? 'https://ssl' : 'http://www') + '.google-analytics.com/ga.js'; var s = document.getElementsByTagName('script')[0]; s.parentNode.insertBefore(ga, s); })(); Skip to main content Preprints Collections Wiley Open Research IET Open Research Ecological Society of Japan All Collections About About Authorea FAQs Contact Us Quick Search anywhere Search for preprint articles, keywords, etc. Search Search ADVANCED SEARCH SCROLL This is a preprint and has not been peer reviewed. Data may be preliminary. 24 April 2025 V1 Latest version Share on Edge effects on amphibian diversity in the Yoko Forest Reserve, Democratic Republic of the Congo Authors : Loving Musubaho 0000-0002-5670-7422 [email protected] , Léon Iyongo , Jean-Claude Mukinzi , Jasmin Mutahinga 0009-0003-9098-388X , Gabriel Badjedjea , Hippolyte Nshimba , and BOGAERT Jan Authors Info & Affiliations https://doi.org/10.22541/au.174549300.01640102/v1 345 views 224 downloads Contents Abstract Information & Authors Metrics & Citations View Options References Figures Tables Media Share Abstract Edges provide valuable insight into the effects of landscape structure on forest ecosystems and faunal distribution. This study assessed edge effects on amphibian diversity in the Yoko Forest Reserve, Democratic Republic of the Congo. Over twelve months (from December 2020 to November 2021), amphibians were surveyed during 24 nocturnal field campaigns using visual detection, habitat searches, and acoustic hearing of vocalizations. Twelve sites were sampled, each including five habitats: primary forest, fallow, field, and two edge types (primary forest-fallow and primary forest-field). Amphibian abundance, species richness, diversity, evenness, and relative density were compared between each edge and its adjacent habitats. A total of 5516 individuals (33 species, 17 genera, 10 families) were recorded. Edges had lower abundance, richness and density than the primary forest but higher than adjacent disturbed habitats. The forest-fallow edge maintained similar diversity to the primary forest, while the forest-field edge showed significantly lower diversity. Forest evenness was lower than at both edges. Several species, including Hylarana albolabris, Leptopelis calcaratus, L. christyi, L. millsoni, L. notatus, L. ocellatus and Sclerophrys pusilla showed intermediate abundances at the primary forest-fallow edge. Similarly, Afrixalus osorioi and Ptychadena mascareniensis showed intermediate abundances in the primary forest-field edge. Scelophrys gracilipes and S. pusilla showed a positive preference for the primary forest-field edge. Afrixalus osorioi, A. quadrivittatus, Chiromantis rufescens, Hylambates verrucosus, Hyperolius sp., H. langi, H. ocellatus, Leptopelis calcaratus, Ptychadena christyi, P. mascareniensis, P. perreti, Sclerophrys gutturalis and S. gracilipes showed a positive preference for the primary forest-fallow edge. Among these species, Chiromantis rufescens and P. mascareniensis showed a moderate but significant preference for this edge, enabling them to be considered specialists of this habitat. These results confirm the structuring role of edges in forest amphibian population’s dynamics. Edge effects on amphibian diversity in the Yoko Forest Reserve, Democratic Republic of the Congo Loving Musubaho 1,2,3* , Léon Iyongo 4,5 , Jean-Claude Mukinzi 6,7 , Jasmin Mutahinga 6,8 , Gabriel Badjedjea 9 , Hippolyte Nshimba 10 & Jan Bogaert 1* 1 Gembloux Agro-Bio Tech, Université de Liège, 2 Passage des Déportés, 5030 Gembloux, Belgium; 2 Environnement and Sustainable Development Option, Institut Supérieur de Développement Rural de Goma, Goma P.O. Box 232, North-Kivu, Democratic Republic of the Congo; 3 Research Center for Environmental Planning, Goma P.O. Box 302, North-Kivu, Democratic Republic of the Congo; 4 Faculty of Renewable Natural Resources Management, University of Kisangani, Kisangani P.O. Box 2012, Tshopo, Democratic Republic of the Congo; 5 Laboratoire d’Ecologie du Paysage et de Gestion des Ressources Fauniques, University of Kisangani, Kisangani P.O. Box 2012, Tshopo, Democratic Republic of the Congo; 6 Department of Ecology and Animal Resources Management, Faculty of Sciences, University of Kisangani, Kisangani P.O. Box 2012, Tshopo, Democratic Republic of the Congo ; 7 Laboratoire d’Ecologie et Gestion des Ressources Animales, Faculty of Sciences, University of Kisangani, Kisangani P.O. Box 2012, Tshopo, Democratic Republic of the Congo; 8 Action pour le Droit de l’Environnement et la Restauration des Ecosystèmes (ADRE-RDC), Kisangani, Tshopo, Democratic Republic of the Congo; 9 Centre de Surveillance de la Biodiversité, Department of Ecology and Aquatic Resources Biodiversity, Faculty of Sciences, University of Kisangani, Kisangani P.O. Box 2012, Tshopo, Democratic Republic of the Congo ; 10 Departement of Ecology and Plant Ressources Management, Faculty of Sciences, Université de Kisangani, Kisangani P.O. Box 2012, Tshopo, Democratic Republic of the Congo. * : Corresponding author : [email protected] (Musubaho L). Abstract : Edges provide valuable insight into the effects of landscape structure on forest ecosystems and faunal distribution. This study assessed edge effects on amphibian diversity in the Yoko Forest Reserve, Democratic Republic of the Congo. Over twelve months (from December 2020 to November 2021), amphibians were surveyed during 24 nocturnal field campaigns using visual detection, habitat searches, and acoustic hearing of vocalizations. Twelve sites were sampled, each including five habitats: primary forest, fallow, field, and two edge types (primary forest-fallow and primary forest-field). Amphibian abundance, species richness, diversity, evenness, and relative density were compared between each edge and its adjacent habitats. A total of 5516 individuals (33 species, 17 genera, 10 families) were recorded. Edges had lower abundance, richness and density than the primary forest but higher than adjacent disturbed habitats. The forest-fallow edge maintained similar diversity to the primary forest, while the forest-field edge showed significantly lower diversity. Forest evenness was lower than at both edges. Several species, including Hylarana albolabris , Leptopelis calcaratus , L. christyi , L. millsoni , L. notatus , L. ocellatus and Sclerophrys pusilla showed intermediate abundances at the primary forest-fallow edge. Similarly, Afrixalus osorioi and Ptychadena mascareniensis showed intermediate abundances in the primary forest-field edge. Scelophrys gracilipes and S. pusilla showed a positive preference for the primary forest-field edge. Afrixalus osorioi , A. quadrivittatus , Chiromantis rufescens , Hylambates verrucosus , Hyperolius sp. , H. langi , H. ocellatus , Leptopelis calcaratus , Ptychadena christyi , P. mascareniensis , P. perreti , Sclerophrys gutturalis and S. gracilipes showed a positive preference for the primary forest-fallow edge. Among these species, Chiromantis rufescens and P. mascareniensis showed a moderate but significant preference for this edge, enabling them to be considered specialists of this habitat. These results confirm the structuring role of edges in forest amphibian population’s dynamics. Keywords : amphibians, adjacent habitats, edge effects, Yoko Forest Reserve. Introduction Landscape and habitat characteristics influence fauna as well as connectivity between animal populations (Archis et al. 2018; Musubaho et al. 2024b). This also applies to amphibians, whose role as bioindicators of ecological processes is well established (Koirala et al. 2019; Widiana et al. 2019; Vagmaker et al. 2020). Because of their sensitivity to habitat modifications, these changes can affect their distribution and survival (Belasen et al. 2019; Betts et al. 2019; Anjos et al. 2020). At the landscape scale, these transformations influence not only the connectivity between these habitats, but also the intensity of edge effects (Cushman et al. 2016; Widiana et al. 2019). The loss of this connectivity can have negative consequences on the distribution of amphibians (Campos et al. 2020). Changes in landscape structure lead to the isolation of habitat fragments (Lourenço-de-Moraes et al. 2018), a phenomenon particularly prevalent in forest areas in the Democratic Republic of the Congo (DRC). The forests in and around Kisangani are particularly threatened by slash-and-burn agriculture and timber exploitation (Masimo et al. 2020; Kipute et al. 2023). This fragmentation leads to the formation of edges, which play an important role in biodiversity distribution, landscape structuring and ecosystem processes (Peyras et al. 2013; Porensky & Young, 2013; Barlow et al. 2016; Pfeifer et al. 2017; Posse-Sarmiento & Banks-Leite, 2024). To better understand the influence of edges on amphibians, the present study was carried out in the Yoko Forest Reserve (YFRE). The edge effects on amphibians have been the subject of numerous studies worldwide (Porensky & Young, 2013; Pfeifer et al. 2017), particularly in tropical America (Santos-Barrera & Urbina-Cardona, 2011; Schneider-Maunoury et al. 2016; Posse-Sarmiento & Banks-Leite, 2024), Asia (Scriven et al. 2018; Widiana et al. 2019) and Africa (Hofer et al. 2000; Lehtinen et al. 2003). In the DRC, particularly in the Kisangani region, research of this kind is still in its infancy. This analysis, the first to focus on edge effects on amphibians in this region, inspired by related works on rodents in the Masako Forest Reserve (Iyongo et al. 2009a; Iyongo et al. 2012; Meniko et al. 2020). This study verifies the following hypotheses : (i) Given that anthropization is impacting amphibians in the Kisangani region, particularly in the YFRE (Musubaho et al. 2024b), the resulting habitat modifications could affect the distribution of abundance and specific richness between edges and their adjacent habitats; (ii) It is well established that edges, due to their particular ecological conditions (Cadenasso et al. 2003a; Harper et al. 2005; Niyukuri et al. 2014), exhibit statistically higher abundances, specific richness, diversities, equitabilities and relative densities than adjacent habitats (Iyongo et al. 2012; Niyukuri et al. 2014; Magura et al. 2017; Erdös et al. 2019; Widiana et al. 2019; Meniko et al. 2020; Ho et al. 2023). Similar observations are expected in the YFRE; (iii) Variations in environmental conditions influence the distribution of species, favoring those better adapted to a given habitat type (edges or adjacent habitats), where they become more abundant (Cortés et al. 2008; Iyongo et al. 2012; Niyukuri et al. 2014; Schneider-Maunoury et al. 2016). It is likely that this phenomenon is also true in the study area; (iv) As several studies have identified certain amphibian species as primary forest specialists (Magura et al. 2017; Decena et al. 2020; Musubaho et al. 2024b), this study postulates that edge specialists may also exist in the YFRE. The study aims to compare amphibian diversity between edges (primary forest-fallow, primary forest-field) and their adjacent habitats (field, fallow and primary forest). Specifically, this research tries to (i) assess the influence of edges on the distribution of amphibian populations; (ii) compare edges with adjacent habitats in terms of the amphibian community; (iii) analyze, through their abundances, the degree of adaptation and response of species to edge conditions and (iv) identify the preferences of amphibian species according to the different habitat types present in the YFRE. As in Musubaho et al. (2024b), the field and the fallow are considered as disturbed habitats, the field being more disturbed than the fallow, while the primary forest is an undisturbed habitat serving as a control habitat. Materials et methods Study area The YFRE is located near the equator, between 0°15′ and 0°20′ N et 25°14′ and 25°20′ E (Boyemba, 2011), on the road connecting Kisangani with Ubundu, in the Tshopo Province. It is characterized by an average annual temperature of 25°C, an average annual rainfall of 1750 mm and by the absence of dry months (Picard et al. 2015). As throughout the Kisangani region, rainfall throughout the year is interrupted by two sub-dry seasons. The long rainy season runs from September to December and the short rainy season covers the months from March to June. On the other hand, the long sub-dry season extends through January and February and the short sub-dry season covers July and August (Kahindo, 2011; Sabongo, 2015). The hydrographic network is dominated by rivers, the most important of which are Yoko and Biaro streams. They receive water of numerous small stream before discharging into the Congo River. The Yoko stream, which gave the studied reserve its name, flows from west to east, dividing the study zone into two blocks, north and south. Within each block, six sites were identified (Fig. 1) and in each site, five habitats were described and sampled. These habitats are primary forest, field, fallow, primary forest-fallow and primary forest-field edges. These edges were identified on the basis of their vegetation physiognomy, which differs from that of adjacent habitats (Iyongo et al. 2009a; Iyongo et al. 2012; Widiana et al., 2019; Meniko et al. 2020). Figure 1 . Location map and structure of the sampling sites in the two blocks of the YFRE, Tshopo Province, DRC. Twelve sites were selected and within each site, five habitats were exploited: primary forest (PF), fallow (FL), field (FD), primary forest-fallow edge (PFFLE) and primary forest-field edge (PFFDE). Primary forest is dominated by Trilepisium madagascariense Dc. (Moraceae), Uapaca guineensis Mull. Arg (Euphorbiaceae), Milicia excelsa (Welw.) C.C. Berg. (Fabaceae), Chrysophyllum pruniforme Pierre ex Engler (Sapotaceae), Sterculia tragacantha Lindley (Malvaceae), Strombosia pustulata Oliver (Strombosiaceae), Microdesmis yafungana J. Léonard (Pandaceae), Turraeanthus africanus (Welw.) Pellegr. (Meliaceae), Afrostyrax lepidophyllum Mildbr (Uaceae), Megaphrynium macrostachyum (Bentham) Milne-Redh, Marantochloa mannii (Bentham) Milne-Redh (Maranthaceae). Fallow is dominated by Tetrorchidium didymostemon (Baillon) Pax et Hoffm. (Euphorbiaceae), Oncoba welwitschii Oliver, (Flacourtiaceae), Zanthoxylum gilletii (De Wild.) P.G Waterman (Rutaceae), Triumfetta cordifolia Guill., Perr. & A. Rich (Malvaceae), Pycnanthus angolensis (Welw.) Exell (Myristicaceae), Canarium schweinfurthii Engler (Burseraceae), Harungana madagascariensis Lam. ex Poiret (Hypericaceae). The species Oriza sativa L., Zea mays L. (Poaceae), Capsicum annuum L., Solanum melongena L. (Solananceae), Musa sp . (Musaceae), Manihot esculenta Crantz (Euphorbiaceae), Elaeis guineensis Jacq (Arecaceae) are among the most dominant in the field. Orphaned primary forest species such as Julbernardia seretii (De Wild.) Troupin, Pterocarpus soyauxii Taub. (Fabaceae), Petersianthus macrophylla (P. Beauv.) Liben (Lecythidaceae), Khaya anthotheca Dc. (Meliaceae), Margaritaria discoidea (Baillon) Webster (Phyllanthaceae) are also recorded in the field. Primary forest-fallow edge is dominated by Trilepisium madagascariense Dc., Treculia africana Decne. (Moraceae), Pauridiantha callicarpoides (Hiern) Bremek. (Rubiaceae), Zanthoxylum gilletii (De Wild.) P.G. Waterman (Rutaceae), Alchornea floribunda Mull. Arg. (Euphorbiaceae), Pycnanthus angolensis (Welw.) Exell (Myristicaceae), Cola bruneelii De Wild., (Malvaceae), Oncoba welwitschii Oliver (Flacourtiaceae), Hexalobus crispiflorus A. Rich. (Annonaceae), Afromomum sangineum K. Schum. (Zingiberaceae). Among dominant species in Primary forest-field edge there are Chrysophyllum africanum A. Dc. (Sapotaceae), Trema orientalis (L.) Blume (Cannabaceae), Khaya anthotheca Dc. (Meliaceae), Triumfeta cordifolia Guill., Perr & A. Rich. (Malvaceae) et Selaginella kraussiana (Kunze) A. Braun (Selaginellaceae). Methods The study covered 5516 amphibians collected using the method and techniques previously described (Musubaho et al. 2024ab), who deal with the diversity and endemism of amphibians in the YFRE, the anthropogenic effects on amphibian diversity and habitat similarity in the same forest ecosystem, respectively. All amphibians belong to the Anura order and divided into 10 families, 17 genera and 33 species (Table 1). In this article, the name Hylarana albolabris has been used in place of Amnirana albolabris in accordance with the systematic revision of this species (Frost, 2024). Table 1. Amphibian species surveyed between December 2020 and November 2021 in five habitats in the YFRE, Democratic Republic of the Congo.Ni = number of individuals collected. Arthroleptidae Mivart, 1869 Arthroleptis variabilis Matschie, 1893 20 Arthroleptis tuberosus Anderson, 1905 31 Leptopelis calcaratus Boulenger, 1906 41 Leptopelis ocellatus Mocquard, 1902 68 L. notatus Buchloz & Peters in Peters, 1875 322 Leptopelis christyi Boulenger, 1912 313 Leptopelis millsoni Boulenger, 1895 217 Cardioglossa leucomystax Boulenger, 1903 48 Bufonidae Gray, 1825 Sclerophrys gutturalis Power, 1927 13 Sclerophrys pusilla Mertens, 1937 143 Sclerophrys gracilipes Boulenger, 1899 30 Nectophryne batesii Boulenger, 1913 4 Dicroglossidae Anderson, 1871 Hoplobatrachus occipitalis Günther , 1858 9 Hyperoliidae Laurent, 1943 Hylambates verrucosus Boulenger, 1912 91 Afrixalus quadrivittatus Werner, 1908 19 Afrixalus osorioi Ferreira, 1906 60 Afrixalus equatorialis Laurent, 1941 3 Hyperolius ocellatus Günther, 1858 15 Hyperolius platyceps Boulenger, 1900 175 Hyperolius langi Noble, 1924 26 Hyperolius parallelus Günther, 1858 7 Hyperolius sp. 20 Cryptothylax greshoffi Schilthuis, 1889 6 Kassina maculosa Sternfeld, 1917 1 Phrynobatrachidae Laurent, 1941 Phrynobatrachus auritus Boulenger, 1900 731 Phrynobatrachus perpalmatus Boulenger, 1898 17 Pipidae Gray, 1825 Xenopus pygmaeus Loumont, 1986 47 Ptychadenidae Dubois, 1987 Ptychadena christyi Boulenger, 1919 207 Ptychadena perreti Guibe & Lamotte, 1958 134 Ptychadena mascareniensis Duméril & Bibron, 1841 349 Pyxicephalidae Bonaparte, 1850 Amietia nutti Boulenger, 1896 26 Ranidae Rafinesque, 1814 Hylarana (Amnirana) albolabris Hallowell, 1856 1986 Rhacophoridae Hoffman, 1932 Chiromantis rufescens Günther, 1869 337 10 families 33 species 5516 Amphibian abundance and specific richness are expected to vary across sites, and to be statistically different between each edge and its adjacent habitats. The Shannon-Wiener diversity index (Equation 1) was used to assess amphibian diversity in each habitat. This ecological measure quantifies and compares diversity expressed in biologically interpretable units translating the effective number of species (Legendre and Legendre, 2012; Marcon, 2022). \begin{equation} H^{{}^{\prime}}=-\sum_{i=1}^{S}{p_{i}\ln p_{i}}\ \ \ \ \ \ \ (1)\nonumber \\ \end{equation} where \(H^{\prime}\) is the Shannon index and pi is the proportion of individuals of species \(i\) (\(pi=ni/N\), \(ni\) equals the number of individuals of species \(i\) et \(N\) is the total number of individuals collected ). Given that edges combine the characteristics of adjacent habitats, we expect these edges to be characterized by diversity values statistically different from those of their adjacent habitats. So, to assess how individuals are distributed between species within habitats, we used Piélou’s evenness expressed by equation 2 (Magurran, 2004). \(J=\frac{H^{{}^{\prime}}}{\text{Hmax}}\) = \(\frac{H^{{}^{\prime}}}{\text{lnS}}\) (2) where \(J\) corresponds to Piélou’s evenness, \(H^{\prime}\) translates the Shannon-Wiener index or real diversity and Hmax is the maximum diversity. Considering the conditions of variation of the results of this index, values close to 1 expected in edges should be significantly different from those close to 0 expected in adjacent habitats. Amphibian relative densities were calculated using relationship (3) (Nicolas et al. 2003) and consist to the numbers of individuals per unit area. \(T\left(\%\right)=\frac{N}{\left(\text{n\ x\ t}\right)}\ \times\ 100\)(3) where T(%) is the relative density (the number of individuals per hectare), \(N\) is the number of individuals captured in the habitat, \(n\) is the number of collectors and \(t\) is the number of effective capture nights. The number of amphibians per hectare in each edge is expected to be significantly different from the number of amphibians observed in each of its adjacent habitats. As the data were not normally distributed and were unpaired, the Kruskal-Wallis test was applied to examine whether there were significant differences between edges and their adjacent habitats about the calculated indices. The Mann-Whitney test was used to compare each edge with each of its adjacent habitats. The above tests were used because the data did not follow the normal distribution. Before applying the Kruskal-Wallis and Mann-Whitney tests, the Shapiro test was applied to check the normality of the data derived from the calculated parameters. All p -values were less than 0.0001, the one for species richness being slightly higher (0.0014). Dunn’s test, using Bonferoni’s adjustment, was applied to determine which edge showed significant differences with the adjacent habitats. To assess the degree to which species adapt to habitat conditions through their abundances, this study used the typology of species responses to habitat heterogeneity (Iyongo et al. 2009a) (Table 2). To implement this typology, each species was analyzed separately. Table 2. Typology of species responses to habitat heterogeneity and conditions (Iyongo et al. 2009a) : H1 (habitat 1), H3 (habitat 3). The letters \(a\), \(b\), ab and \(c\) are assigned according to the results of the test used on the abundance of the species in each edge and its adjacent habitats. I a a a No edge effect, the species is a generalist; no difference in abundance between habitats. II a b a Edge effect : if b>a, the species prefers the edge and if ba and b>c, the species prefers the edge ; if b<a and bb) and (b>c) or (a<b) and (bb, the species avoids habitat 3 and if b>a, the species seeks habitat 3. VI ab a b No edge effects detected. [1]¿p#1 For the purposes of visualizing Figure 3, the Mann-Whitney test was performed for display on the graph. The typology of species responses to habitat heterogeneity was determined according to the relationships ”primary forest-edge-fallow” and ”primary forest-edge-field”, in order to capture the species reaction to the influence of these two edge types. Nevertheless, some of amphibian species in the YFRE are expected to be better adapted to edge conditions. To determine the ecological preference of species in relation to edges and their adjacent habitats, the point biserial correlation coefficient was calculated. This is the quantitative equivalent of the correlation phi coefficient (Borcard et al. 2018) whose values vary between [-1, 1]. De Cáceres and Legendre (2009) consider this approach to be more useful than the indicator-value approach, as it can detect negative preferences. In this article, the values of the point biserial correlation coefficient (Φ) are interpreted as follows : if Φ = 0, there is independence (there is no link) ; if |Φ| < 0,1, the preference is very weak ; if 0,1 ≤ |Φ| < 0,3, the preference is weak ; if 0,3 ≤ |Φ| < 0,5, preference is moderate ; if 0,5 ≤ |Φ| < 0,7, preference is strong ; si 0,7 ≤ |Φ| ≤ 1, preference is very strong. The significance of species preference was assessed on the basis of the confidence interval. Confidence intervals (CIs) were generated using the bootstrapping method, with 1000 iterations. Therefore, when both bounds have the same sign, the preference is considered significant; in other cases, the preference is considered insignificant (Borcard et al. 2018). Based on species adaptations, we expect to detect species that prefer edges more than their adjacent habitats (edge specialists). Conversely, we will know which species avoid edges. We consider edge specialists to be species that show a positive and at least moderate preference for edges, with a significant level of preference and statistically higher abundances than in adjacent habitats. Field data were processed in Excel 2016. All analyses were performed using R.4.4.0 (through RStudio 2024.04+764) (R Core Team, 2024). Abundance, species richness, Shannon-Wiener diversity index and Piélou equitability were calculated using the ”diversityresult” function in the ”BiodiversityR 2.16-1” package (Kindt and Coe, 2005). Amphibian density in habitats was calculated using equation 3. Normality tests were performed using the ”Shapiro.test” function, while the Kruskal-Wallis test was performed using the ”kruskal.test” function, both from the ”Stats” package, R’s default package. Dunn’s test was applied using the ”dunn_test” function from the ”rstatix 0.7.2” package (Kassambara, 2023b). The graphs constituting figures (2) and (3) were produced using the ”ggplot2 3.5.1” package (Wickham, 2016). These graphs were assembled using the ”ggarrange” function in the ”ggpubr 0.6.0” package (Kassambara, 2023a). The results of the Mann-Whitney and Kruskal-Wallis tests were visualized on the graphs in figure (3) using the ”stat_compare_means” function in the ”ggpubr 0.6.0” package. The point biserial correlation coefficient was calculated using the ”strassoc” function in the ”indicspecies 1.7.14” package (De Cáceres and Legendre, 2009). Results The data collected showed that amphibian abundance and specific richness vary between each edge and its adjacent habitats in the twelve sites exploited within the study area. The average abundance and specific richness per habitat are respectively 12.167 ± 12.104 and 3.416 ± 2.429 in the primary forest-field edge, 77.250 ± 58.865 and 13.167 ± 2,758 in the primary forest-fallow edge, 3.583 ± 3.728 and 1.500 ± 0.904 in the field, 36.833 ± 22.270 and 11.083 ± 4.078 in the fallow, 329.833 ± 40.618 and 20.083 ± 3.232 in the primary forest. These variations in abundance and specific richness (Fig. 2) confirm the first hypothesis of this study, which states that habitat conditions resulting from anthropization have an impact on the distribution of abundance and specific richness between edges and their adjacent habitats. Figure 2. Abundances (A) and specific richness (B) of amphibians captured between December 2020 and November 2021 in the YFRE. Each site was characterized by five habitats: primary forest (PF), fallow (FL), field (FD), primary forest-fallow edge (PFFLE) and primary forest-field edge (PFFDE). The abscissas represent habitats, while the ordinates correspond to the number of individuals and species respectively. The grey dots represent the values obtained per site, while the black dots represent the averages of the values obtained per habitat at the twelve sites. Primary forest-field edge provided intermediate values of abundance ( N = 146), species richness ( n = 15), diversity ( H’ = 1.678), evenness (J = 0.620) and relative density ( T (%) = 12.674 ind/ha) between the field ( N = 43; n = 6; H’ = 1.168; J = 0.652; T (%) = 3.733 ind/ha) and primary forest ( N = 3958; n = 32; H’ = 2.083; J = 0.601; T (%) = 343.576 ind/ha). Also, primary forest-fallow edge was characterized by intermediate values of the same indices ( N = 927; n = 26; H’ = 2.557; J = 0.820; T (%) = 80.469 ind/ha) between fallow ( N = 442; n = 27; H’ = 2.701; T (%) = 38.368 ind/ha) and primary forest. The results of the Mann-Whitney test showed that the primary forest-field edge did not systematically show a significant difference with the field; the same applied to the primary forest-fallow edge, which did not show a significant difference with the fallow (Fig. 3). Nevertheless, the primary forest showed significant differences, with values higher than those of the two edges surveyed, in terms of abundance, species richness and relative density. These observations partially confirm the second hypothesis of this study, according to which, due to their particular ecological conditions, the edges present statistically higher abundances, specific richness, diversity, evenness and relative densities than those of adjacent habitats. Figure 3: Comparison of abundances (A), specific richness (B), Shannon diversity (C), Piélou evenness (D) and relative densities (E) of amphibians captured in the edges and their adjacent habitats exploited between December 2020 and November 2021 in the YFRE. The abscissas represent habitats, while the ordinates indicate values for abundance, species richness, Shannon diversity, Piélou evenness and relative density, respectively. The barplots indicate the values obtained for each parameter in each habitat. The dots represent the averages of the parameters calculated for all twelve sites for each habitat. Vertical lines illustrate standard deviations; horizontal lines indicate the habitats compared and values above these horizontal lines indicate the results of the Mann-Whitney test. The results of the Kruskal-Wallis test are presented in the top left-hand corner for each parameter. Species abundances in the edges and their adjacent habitats (Table 3; Table 4, Appendix 1) indicated that: In the primary forest-fallow relationship, Hylarana albolabris , Leptopelis calcaratus , Leptopelis christyi , L. millsoni , L. notatus , L. ocellatus and Sclerophrys pusilla provided, in the primary forest-fallow edge, intermediate abundances to those of adjacent habitats (Type IV). They are therefore sensitive to edge effects. In contrast, Amietia nutti , Arthroleptis variabilis , Phrynobatrachus auritus , P. perpalmatus and Xenopus pygmaeus were better established in primary forest (high abundances), and avoided fallow and edge (low abundances) (Type V). Edge effect was not detected for Arthroleptis tuberosus , Cardioglossa leucomystax and Hyperolius platyceps by their abundances in habitat 1 which were intermediate between edge and habitat 3 (Type VI). All the remaining 18 species, apart from those mentioned, didn’t show differences in abundances between edge and its adjacent habitats (Type I) ; In the primary forest-field relationship, Afrixalus osorioi and Ptychadena mascareniensis provided, in the primary forest-field edge, abundances that were intermediate to those of adjacent habitats (Type IV). They are therefore sensitive to edge effects. Afrixalus equatorialis , A. quadrivittatus , Cryptothylax greshoffi , Hoplobatrachus occipitalis , Hyperolius sp., H. ocellatus , Kassina maculosa , Nectophryne batesii , Sclerophrys gracilipes , S. gutturalis and S. pusilla showed abundances that did not differ significantly between the edge and adjacent habitats (Type I). These species are therefore not sensitive to edge effects. All the remaining 20 species, apart from those mentioned, prefer primary forest and avoid field and edge (Type V). These observations confirm the third hypothesis of this study, according to which species best adapted to edges or adjacent habitats are particularly abundant there, due to the changing environmental conditions specific to these two types of habitat. Table 3. Species responses to edge conditions exploited between December 2020 and November 2021 in the YFRE. \(N\) = total number of individuals per species in all habitats throughout the study period; \(n\) = number of individuals of each species for each of the relationships explored: the primary forest-fallow relationship and the primary forest-field relationship . The letters \(a\), ab et \(b\)represent the results of Dunn’s test on the abundance of each species in the edge and in its adjacent habitats, with a > b. Habitats symbolized by the same letters are not statistically different in terms of species abundance, while significant differences in abundance were obtained between habitats bearing different letters. Numbers I, IV, V and VI reflect the types of species response to habitat heterogeneity and conditions, and are explained in Tables 1 and 4, Appendix 1. [1]¿p#1 n FL PFFLE PF Type n PF PFFDE FD Type Hylarana albolabris 1986 1975 b ab a IV 1789 a b b V Phrynobatrachus auritus 731 728 b b a V 660 a b b V Ptychadena mascareniensis 349 278 a a a I 163 a ab b IV Chiromantis rufescens 337 332 a a a I 93 a b b V Leptopelis notatus 322 320 b ab a IV 247 a b b V Leptopelis christyi 313 310 b ab a IV 244 a b b V Leptopelis millsoni 217 217 b ab a IV 162 a b b V Ptychadena christyi 207 206 a a a I 97 a b b V Hyperolius platyceps 175 175 ab b a VI 133 a b b V Sclerophrys pusilla 143 71 a ab b IV 77 a a a I Ptychadena perreti 134 130 a a a I 74 a b b V Hylambates verrucosus 91 91 a a a I 46 a b b V Leptopelis ocellatus 68 67 b ab a IV 54 a b b V Afrixalus osorioi 60 58 a a a I 31 a ab b IV Cardioglossa leucomystax 48 48 ab b a VI 43 a b b V Xenopus pygmaeus 47 47 b b a V 41 a b b V Leptopelis calcaratus 41 41 b ab a IV 28 a b b V Arthroleptis tuberosus 31 31 ab b a VI 22 a b b V Sclerophrys gracilipes 30 22 a a a I 15 a a a I Amietia nutti 26 26 b b a V 26 a b b V Hyperolius langi 26 26 a a a I 10 a b b V Arthroleptis variabilis 20 19 b b a V 17 a b b V Hyperolius sp. 20 20 a a a I 7 a a a I Afrixalus quadrivittatus 19 19 a a a I 11 a a a I Phrynobatrachus perpalmatus 17 17 b b a V 16 a b b V Hyperolius ocellatus 15 15 a a a I 11 a b b V Sclerophrys gutturalis 13 9 a a a I 4 a a a I Hoplobatrachus occipitalis 9 8 a a a I 7 a a a I Hyperolius parallelus 7 7 a a a I 7 a a a I Cryptothylax greshoffi 6 6 a a a I 5 a a a I Nectophryne batesii 4 4 a a a I 3 a a a I Afrixalus equatorialis 3 3 a a a I 3 a a a I Kassina maculosa 1 1 a a a I 1 a a a I The results of the point biserial correlation coefficient, used to determine species preferences in relation to habitats (Fig. 4; Table 5, Appendix 2), indicated that of the species present in the YFRE, two ( Sclerophrys gracilipes and S. pusilla ) showed a positive preference for the primary forest-field edge. Similarly, Afrixalus osorioi , A. quadrivittatus , Chiromantis rufescens , Hylambates verrucosus , Hyperolius sp., H. langi , H. ocellatus , Leptopelis calcaratus , Ptychadena christyi , P. mascareniensis , P. perreti , Sclerophrys gracilipes and S. gutturalis show a positive preference for the primary forest-fallow edge. Of these species, Chiromantis rufescens and Ptychadena mascareniensis showed a positive, moderate and significant preference for the primary forest-fallow edge, of which they can be considered specialists. All species recorded in the study area, with the exception of Sclerophrys gutturalis and S. pusilla , showed a positive preference for primary forest. However, all species showed a negative preference for the field, with the exception of Sclerophrys gutturalis , which had a positive preference for this habitat. Only A. quadrivittatus , Nectophryne batesii , Sclerophrys gracilipes , S. gutturalis and S. pusilla had a positive preference for fallow. The preferences of these two species ( Chiromantis rufescens and Ptychadena mascareniensis ) for the primary forest-fallow edge confirm the fourth hypothesis of this study, which states that apart from some species being primary forest specialists, there are other species that are edge specialists. Figure 4. Amphibian preferences for exploited habitats between December 2020 and November 2021 in the YFRE. This graphic representation is taken from the table 5, Appendix 3. Most species show a more positive preference for primary forest than for forest edges. Discussion The presence of edges in the YFRE illustrates the impact of habitat fragmentation. This can lead to a variety of consequences on amphibians (Kusrini, 2013), including low mobility (Bowne & Bowers, 2004) and increased vulnerability to mortality when moving through inhospitable environments (Carr et al. 2002). Indeed, habitat modification leads to the formation of edges, which are characterized by a distinct microclimate, great heterogeneity and fluctuating environmental conditions, independent of those of adjacent habitats (Cadenasso et al. 2003a; Harper et al. 2005). In fragmented landscapes, the role of edges varies according to their contrast and forest structure (Dodonov et al. 2013). These variations directly influence amphibian distribution (Pfeifer et al. 2017; Scriven et al. 2018; Widiana et al. 2019). The results of this study clearly demonstrate that the primary forest is by far richer in amphibians than any of the edges in the YFRE. This corroborates with other research attesting that amphibians prefer forest environments because they are characterized by cooler temperatures and high humidity, conducive to the survival of individuals during dry periods (Schneider-Maunoury et al. 2016; Widiana et al. 2019). By including the air humidity, diversity of ecological niches, availability of food resources and canopy cover that characterize primary forest, this creates a particularly favorable environment for amphibians (Jackson & Blackburn, 2010; Santos-Barrera & Urbina-Cardona, 2011). In the YFRE, the primary forest fulfils all these conditions, which explains its attractiveness for the species recorded (Musubaho et al. 2024b). Reduced litter in edges has negative effects on frog and toad litter populations, leading to loss of biodiversity, disruption of ecosystem processes (Pfeifer et al. 2017) and increased predation and mortality (Ewers & Didham, 2006). In contrast, other studies suggest that edges are key areas for biodiversity, which is in line with the edge effect theory, according to which, edges harbor the greatest number of species, compared to adjacent habitats (Niyukuri et al. 2014; Bátori et al. 2018; Erdös et al. 2019; 2023; Ho et al. 2023). Observations have confirmed this theory for terrestrial beetles (Magura et al. 2017), amphibians (Hofer et al. 2000; Widiana et al. 2019), birds (Delage et al. 2000) and rodents (Iyongo et al. 2012; Meniko et al. 2020). The differences found around amphibian specific richness between edges and their adjacent habitats could be influenced by several factors, including canopy cover, understory density, litter depth and temperature variations (Urbina-Cardona et al. 2006). Moreover, the way in which these factors modify the habitats could be responsible for the variations in species richness. However, an earlier study showed no significant difference in amphibian species richness between edges and their adjacent habitats (Dawson & Hostetler, 2008). In this study, the measurement of diversity between the different habitats reveals that the primary forest-fallow edge has a higher diversity than the primary forest. This could be explained by a better even distribution of species in this habitat. These results are in line with previous studies showing that edges are often characterized by high diversity (Widiana et al. 2019; Meniko et al., 2020; Ho et al., 2023). In line with other researches, as the primary forest-fallow edge combines the characteristics of adjacent habitats, this provides this habitat with conditions that thus make it favorable to amphibians (Ho et al., 2023). In contrast, amphibian diversity in the primary forest-fallow edge is lower than in the fallow. This high diversity in the fallow could be linked to the great adaptability of amphibians to the conditions of this habitat (Tumushimire et al. 2020; Musubaho et al. 2024b), as well as to the abundance of food resources, which plays a decisive role in the presence of species (Mukinzi et al. 2005). The shading provided by the woody species of the primary forest mitigates edge effects by facilitating the mobility of amphibians between the primary forest and the fallow via their edge (Cruz-Elizalde et al. 2016; Oda et al. 2016). However, other studies have reported that edges are characterized by lower diversity, as their environmental conditions (temperature, humidity, litter, canopy) are less favorable for amphibians, particularly frogs (Cortés et al. 2008). These factors could explain the best distribution of individuals between species in the fallow and in the primary forest-fallow edge. On the other hand, the low evenness in the primary forest could be the result of the strong adaptation to this habitat of the species Hylarana albolabris and Phrynobatrachus auritus , which together account for 61.5% of total species abundance in the primary forest, compared with 16.7% distributed between the fallow and the two edges, as these species are absent from the field. Fragmentation makes habitats unfavorable to amphibians, because their openness increases vulnerability to predation by increasing the probability of detection of individuals by their predators (Karraker et al. 2018; Musubaho et al. 2024b). It alters ecological processes in habitats and disrupts predator-prey relationships, leading to reduction in amphibian abundance and density (El Hamoumi & Himmi, 2010; Karraker et al. 2018). This reduction is one of the main factors in the gradual decline of species (Schneider-Maunoury et al. 2016). In the YFRE, these factors could explain the gradual decline in amphibian density as they move from primary forest to fallow and field, via edge zones (Figure 3). Studies have shown that the maintenance of amphibians in various habitats depends on the roles they play in trophic chains as preys or predators (Toledo et al. 2007; Vagmaker et al. 2020). Landscape quality strongly influences amphibian distribution (Musubaho et al. 2024b), and the ecological conditions of habitats influence species’ ability to adapt (Schiøtz, 1999; Howell, 2010). For example, in the primary forest-field relationship, the species Amietia nutti , Hylarana albolabris , Arthroleptis tuberosus , A. variabilis , Cardioglossa leucomystax , Chiromantis rufescens , Hylambates verrucosus , Hyperolius langi , H. ocellatus , H. platyceps , Leptopelis calcaratus , L. christyi , L. millsoni , L. notatus , L. ocellatus , Phrynobatrachus auritus , P. perpalmatus , Ptychadena christyi , P. perreti , and Xenopus pygmaeus are best represented in primary forest and avoid the field as well as the primary forest-field edge. Afrixalus osorioi and Ptychadena mascareniensis , on the other hand, show intermediate abundance in the edge. In the primary forest-field relationship, species such as Amietia nutti , Arthroleptis variabilis , Phrynobatrachus auritus , P. perpalmatus and Xenopus pygmaeus are better adapted to primary forest conditions, avoiding fallow and the primary forest-fallow edge, while Hylarana albolabris , Leptopelis calcaratus , L. christyi , L. millsoni , L. notatus , L. ocellatus and Sclerophrys pusilla show intermediate abundances in the edge. According to previous studies, amphibian adaptation to a diversity of habitats is a strategy maintaining their populations (Decout et al. 2012; Holzer et al. 2017; Figueiredo et al. 2019). The fact that only a few species adapt to disturbed habitat conditions may be a factor in the vulnerability of amphibians in the YFRE, due anthropogenic activities. For example (figure (4)), Sclerophrys gutturalis and S. pusilla showed a best preference for fallow, while Nectophryne batesii prefers both fallow and field. Fragmentation of natural habitats leads to significant changes in biodiversity (Barlow et al. 2016; Pfeifer et al. 2017; Decena et al. 2020; Belloto-Trigo et al. 2023). Amphibians are particularly sensitive to microclimatic modifications and changes in the vegetation structure of edges (Marsh & Pearman, 1997; Demaynadier & hunter, 1998). Edges differ in their specific composition (Anderson et al. 2003), being recognized as particular habitats with their own ecological conditions and species (Cadenasso et al. 2003a; Imbeau et al. 2003). Edge effects are recognized as an important process affecting individual and species behavior, native community structure and ecological interactions in the transition zone (or ecotone) between two or more contrasting habitats, particularly in fragmented landscapes (Wimp et al. 2011; Peyras et al. 2013). These effects include an increase in temperature, a reduction in soil moisture and an increase in wind speed, often with negative consequences on certain species (Laurence, 2008; Cortés et al. 2008). These impacts are particularly marked for primary forest specialist species, which are more sensitive to changes in their habitat (Mestre, 2017; Sykes et al. 2020). For example, the species Amietia nutti , Afrixalus equatorialis , Hyperolius parallelus and Kassina maculosa were observed exclusively in primary forest, while Phrynobatrachus perpalmatus and Xenopus pygmaeus were predominantly found in this habitat (94.1% and 88.2% of individuals respectively), with only a few occurrences in the primary forest-fallow edge. These six species were identified as primary forest specialists by Musubaho et al (2024b). The degree of specialization of species influences their response to edge effects: specialist species react strongly to disturbance, while generalists react very weakly or neutrally (Peyras et al. 2013). The results of this study show that fragmentation strongly influences the habitat preferences by amphibians. Most of the species present in the YFRE show a positive preference for primary forest, with the exception of Sclerophrys gutturalis , which prefers fallow, and S. pusilla , which shows an almost equal preference for fallow and the primary forest-field edge. Among species with a positive preference for primary forest, Sclerophrys gracilipes prefers fallow more than primary forest, while Chiromantis rufescens , Hyperolius sp., H. langi and Ptychadena mascareniensis prefer the primary forest-fallow edge more than primary forest. Preference analysis (Table 5, Appendix 2) shows that Chiromantis rufescens and Ptychadena mascareniensis have a moderate and significant preference for the primary forest-fallow edge. This suggests that both species can be considered specialists of this type of edge. However, the fact that these 2 species leave type I in the primary forest-fallow relationship for types IV and V respectively (Table 4, Appendix 1) shows the low appreciation by these species of the presence of field in the YFRE. This low appreciation observed in the primary forest-field relationship can also be seen by the number of species avoiding the field and the primary forest-field edge, which is much higher than that of species avoiding the fallow and the primary forest-fallow edge (Table 4, Appendix 1). All the above species were classified as primary forest generalists by Musubaho et al (2024b), Ptychadena mascareniensis having shown very high ecological plasticity, even turning over in the field. In this study, it is one of five species found in all five habitats, along with Leptopelis christyi , Ptychadena perreti , Sclerophrys gracilipes and S. pusilla . Data for Hoplobatrachus occipitalis , Hyperolius parallelus , Cryptothylax greshoffi , Nectophryne batesii , Afrixalus equatorialis and Kassina maculosa (Table 4, Appendix 2) should be treated with caution, given their low abundances in the various habitats. Studies on invertebrates have shown that natural edges can be permeable to forest specialist species, particularly for spiders, centipedes and beetles (Gallé & Torma, 2009; Lacasella et al. 2015). These edges are unfavorable to species from open environments because they prevent them from crossing over to the primary forest (Lacasella et al. 2015). On the other hand, nthropogenic edges are often impenetrable to forest specialists, obstructing their dispersal (Magura et al. 2017). However, these edges are invaded by groups from surrounding environments, including geometrid moths (Axmacher et al. 2004), ground beetles (Knapp et al. 2013), spiders and ground beetles (Matveinen-Huju et al. 2009), but are repellent to forest specialists such as bark beetles (Peltonen & Heliövaara, 1998) and some species of ground beetles (Knapp et al. 2013). Similarly, the functional connectivity of forest fragments is greatly reduced by high-contrast matrices (recent plantations) for native forest species, while soft edges (native forests and mature plantations) maintain functional connectivity (Peyras et al. 2013). We think that primary forest-field edges are subject to anthropogenic activities, making them less favorable for the dispersal of primary forest specialist amphibian species. The literature indicates that anthropogenic edges accelerate forest degradation and consequently promote biodiversity loss. Thus, their increasing presence in a landscape must be avoided (Harper et al. 2005) and their restoration is an imperative task in the management of ecosystems and biodiversity. Promoting habitat heterogeneity and reducing the contrast between these edges and adjacent habitats are important strategies in restoring and maintaining ecological connectivity (Anderson & Carter, 1987; Samways, 2007). Conclusions Landscape fragmentation has a negative impact on ecosystems, in particular by creating and multiplying edges, which have a considerable impact on amphibians, particularly in terms of abundance, specific richness, diversity, evenness and relative density. These edges have intermediate values between adjacent habitats. Thus, for the primary forest-field edge, the values observed are intermediate between those of primary forest and field, while for the primary forest-fallow edge, these values are intermediate between those of primary forest and fallow. Abundance and species richness are significantly lower in both edges than in primary forest. Primary forest, on the other hand, although low in diversity compared with fallow and the primary forest-fallow edge, remains significantly more diverse than the primary forest-field edge. Furthermore, the primary forest is characterized by the lowest evenness, which is significantly lower than that of the primary forest-fallow edge, but remains low, with no significant difference, compared to the primary forest-field edge. The differences found between each edge and its adjacent habitats clearly illustrate edge effects on amphibian distribution. These effects revealed that the species Hylarana albolabris , Leptopelis calcaratus , L. christyi , L. millsoni , L. notatus , L. ocellatus and Sclerophrys pusilla show intermediate abundances in the primary forest-fallow edge, compared the habitats it separates. In contrast, Amietia nutti , Arthroleptis variabilis , Phrynobatrachus auritus , P. perpalmatus and Xenopus pygmaeus are better established in primary forest, avoiding both the fallow and the primary forest-fallow edge. No edge effects were observed for Arthroleptis tuberosus , Cardioglossa leucomystax and Hyperolius platyceps . In addition, Afrixalus osorioi and Ptychadena mascareniensis show intermediate abundances in the primary forest-field edge compared with its adjacent habitats. Finally, the species Afrixalus equatorialis , A. quadrivittatus , Cryptothylax greshoffi , Hoplobatrachus occipitalis , Hyperolius sp., H. ocellatus , Kassina maculosa , Nectophryne batesii , Sclerophrys gracilipes , S. gutturalis and S. pusilla show no difference in abundance between primary forest, field and edge. In terms of species adaptations to habitats, Sclerophrys gracilipes and S. pusilla show a positive preference for the primary forest-field edge. Similarly, Afrixalus osorioi , A. quadrivittatus , Chiromantis rufescens , Hylambates verrucosus , Hyperolius sp., H. langi , H. ocellatus , Leptopelis calcaratus , Ptychadena christyi , P. mascareniensis , P. perreti , Sclerophrys gracilipes and S. gutturalis show a positive preference for the primary forest-fallow edge. The species C. rufescens and P. mascareniensis show a positive, moderate and significant preference for the primary forest-fallow edge, making them specialists of this habitat. The edge effects on amphibian distribution, as demonstrated in the present study, are manifested by significant differences in the parameters compared between edges and their adjacent habitats. These differences reflect the impact of fragmentation in the YFRE, where habitat heterogeneity plays a fundamental role in the diversity of amphibian responses to edges. Consequently, preserving and protecting edges, and preventing changes to their structure, composition and characteristics, as well as those of adjacent habitats, are essential to maintaining biodiversity, which depends on healthy ecosystems. Author contributions : Musubaho L . : conceptualization (Equal), methodology (Equal), investigation (Lead), data curation (Equal), formal analysis (Equal), visualisation (Equal), writing-original draft (Lead). Iyongo L . : conceptualization (Equal), methodology (Lead), project administration (Equal), supervision (Equal), validation (Equal), writing-review & editing (Equal). Mukinzi J-C . : methodology (Equal), formal analysis (Supporting), supervision (Equal), writing-review & editing (Equal). Mutahinga J . : data curation (Equal), formal analysis (Equal), visualisation (Equal), writing-review & editing (Equal). Badjedjea G . : data curation (Supporting), formal analysis (Supporting), writing-review & editing (Equal). Nshimba H . : methodology (Equal), formal analysis (Supporting), supervision (Equal), writing-review & editing (Equal). Bogaert J . : conceptualization (Lead), funding acquisition (Lead), project administration (Lead), resources (Lead), supervision (Lead), validation (Lead), writing-review & editing (Lead). Data Availability Statement : All the data used to create this publication have been uploaded as additional information. Acknowledgements : All activities relating to this research were fully funded by the University of Liège’s ”Glocal Health” project, thanks to the PhD grant awarded to Loving Musubaho. The authors express their deep gratitude to the field teams, whose courage and commitment were exemplary throughout the data collection period, particularly under night-time working conditions. They are grateful to Václav Gvozdík of the Institute of Vertebrate Biology of the Czech Academy of Sciences/Czech Republic, for his modest contribution to the identification of captured amphibians and the systematic revision of some species. The managers of the Laboratoire d’Ecologie et Gestion des Ressources Animales (LEGERA) of the Faculty of Sciences of the University of Kisangani are thanked for availability of their premises throughout the study. Sincere thanks to Christien Kimbuluma of the Laboratoire d’Ecologie et Aménagement Forestier (LECAFOR/RDC) for the botanical description of the habitats exploited. References Anderson, M.A., Carter, C.I., 1987. Shaping ride sides to benefit wild plants and butterflies. In Jardine, D.C (Ed.), Wildlife Management in Forests (pp. 66-80). Lancaster : Institute of Chartered Forests. Anderson, C.S., Cady, A.B., Meikle, D., 2003. Effects of vegetation structure and edge habitat on the density and distribution of white-footed mice ( Peromyscus leucopus ) in small and large forest patches. Canadian Journal of Zoology 81 : 897-904. Anjos, A.G., Costa, R.N., Brito, D., Solé, M., 2020. Is there an association between the ecological characteristics of anurans from the Brazilian Atlantic Forest and their extinction risk? Ethology, Ecology & Evolution , 32 (4): 336-350. Archis, J.N., Akcali, C., Stuart, B.L., Kikuchi, D., Chunco, A.J., 2018. Is the future already here? The impact of climate change on the distribution of the eastern coral snake ( Micrurus fulvius ). PeerJ 6: e4647. Axmacher, J.C., Holtmann, G., Scheurmann, L., Brehm, G., Muller-Hohenstein, K., Fieldler, K., 2004. Diversity of geometrid moths (Lepidoptera: Geometridae) along anAfrotropical elevational rainforest transect. Diversity and Distributions 10: 293-302. Barlow, J., Lennox, G.D., Ferreira, J., Berenguer, E., Lees, A.C., Nally, R.M., Thomson, J.R. et al., 2016. Anthropogenic disturbance in tropical forests can double biodiversity loss from deforestation. Nature 535 : 144-147. Bátori, Z., Erdős, L., Kelemen, A., Deák, B., Valkóé, R., Bragina, T.M., Kiss, P.J., Kröel-Dulay, G., Tölgyesi, C., 2018. Diversity patterns in sandy forest-steppes : a comparative study from the western and central Palearctic. Biodivers. Conserv . 27 : 1011-1030. Belasen, A.M., Bletz, M.C., Leite, DS., Toledo, L.F., James, T.Y., 2019. Long-term habitat fragmentation is associated with reduced MHC IIB diversity and increase infections in amphibian hosts. Front. Ecol. Evol . 6 : 236. Bellotto-Trigo, F.C.; Uezu, A.; Hatfield, J.H.; Morante-Filho, J.C.; dos Anjos, L.; Develey, P.F.; Clegg, T.; Orme, C.D.L.; Banks-Leite, C. Intraspecific variation in sensitivity to habitat fragmentation is influenced by forest cover and distance to the range edge. Biol. Conserv . 2023, 284 , 110167. Betts, M.G., Wolf, C., Pfeifer, M., Banks-Leite, C., Arroyo-Rodriguez, V., Ribeiro, D.B., Barlow, J., Eigenbrod, F., Faria, D., Fletcher, R.J.et al., 2019. Extinction filters mediate the global effects of habitat fragmentation on animals. Science 366: 1236-1239. Borcard, D., Gillet, F., Legendre, P. 2018. Numerical Ecology with R, 2 nde Edition, Springer International Publishing AG, 435. Bowne, D.R., Bowers, M.A., 2004. Interpatch movements in spatially structured populations : a literature review. Landscape Ecology 19 :1-20. Boyemba, B.F. Ecologie de Pericopsis elata (Harms) Van Meeuwen (Fabaceae), Arbre de Forêt Tropicale Africaine à Répartition Agrégée . Ph.D. Thesis, Université Libre de Bruxelles, Bruxelles, Belgique, 2011. Cadenasso, M.L., Pickett, S.T.A., Weathers, K.C., Jones, C.G., 2003a. A framework for a theory of ecological boundaries. Bioscience 53 : 750-758. Carr, L.W., Pope, S.E., Fahrig, L., 2002. Impacts of landscape transformation by road. In : Gutzwiller, K.J. (Ed.), Concepts and applications of Landscape Ecology in Biological Conservation. Springer-Verlag, New York. Campos, F.S., Lourenço-de-Moraes, R., Ruas, D.S., Mira-Mendes, C.V., Franch, M., Liorente, G.A., Solé, M., Cabral, P., 2020. Searching for networks: ecological connectivity for amphibians under climate change. Environmental Management 65: 46-61. Cortés, A.M., Ramírez-Pinilla, M.P., Suárez, H.A., Tovar, E., 2008. Edge effects on richness, abundance and diversity of frogs in Andean cloud forest fragments. South American Journal of Herpetology 3 (3): 213-222. Cruz-Elizalde, R., Berriozabal-Islas, C., Hernandez-Salinas, U., Martinez-Morales, M.A., Ramirez-Bautista, A., 2016. Amphibian species richness and diversity in a modified tropical environment of central Mexico. Tropical Ecology 57 (3): 407-417. Cushman, S.A., Elliot, N.B., Macdonald, D.W., Loveridge, A.J., 2016. A multi-scale assessment of population connectivity in african lions (Panthera leo) in response to landscape change. Landscape Ecol . 31 : 1337-1353. Dawson, D.E., Hostetler, M.E., 2008. Herpetofaunal use of edge and interior habitats in urban forest remnants. Urban Habitats 5(1), ISSN 1541-7115. De Cáceres, M., Legendre, P., 2009. Associations between species and groups of sites: indices and statistical inference. Ecology 90(12): 3566-3574. Decena, S.C.P., Avorque, C.A., Decena, I.C.P., Asis, P.D., Pacle, B., 2020. Impact of habitat alteration on amphibian diversity and species composition in a lowland tropical rainforest in Northeastern Leyte, Philippines. Scientific Reports 10: 10547. Decout, S., Manel, S., Miaud, C., Luque, S., 2012. Integrative approach for landscape-based graph connectivity analysis: a case study with the common frog ( Rana temporaria ) in human-dominated landscapes. Landscape Ecol . 27: 267-279. Delage, V., Fortin, M-J., Desroches, A., 2000. Effets de lisière et d’isolement des habitats d’oiseaux chanteurs dans les tourbières exploitées. Ecoscience 7(2): 149-156. Demaynadier, P.G., Hunter, M.L.Jr., 1998. Effects of sylvicultural edges on the distribution and abundance of amphibians in Maine. Conservation Biology 12 : 340-352. Dodonov, P., Harper, K.A., Silva-Matos, D.M., 2013. The role of edge contrast and forest structure in edge influence : vegetation and microclimate at edges in the Brazilian cerrado. Plant Ecol . Doi : 10.1007/s11258-013-0256-0. El Hamoumi, R., Himm, O., 2010. Distribution et état des lieux des peuplements d’Amphibiens dans le complexe de zones humides du bas Loukkos (Larache, Maroc). Bull. Inst. Sc. 32 (2), 95-100. Erdös, L., Krstonošić, D., Kiss, P.J., Bátori, Z., Tölgyesi, C., Škvorc, Z., 2019. Plant composition and diversity at edges in a semi-natural forest-grassland mosaic. Plant. Ecol . 220 : 279-292. Erdös, L., Ho, K.V., Bátori, Z., Kröel-Dulay, G., Ónodi, G., Tölgyesi, C., Török, P., Lengyel, A., 2023. Taxonomy, functional and phylogenetic diversity peaks do not coincide along a compositional gradient in forest-grassland mosaics. J. Ecol . 111 : 182-197. Ewers, R.M., Didham, R.K., 2006. Continous response functions for quantifying the strength of edge effects. J. Appl. Ecol . 43 : 527-536. Figueiredo, G.T., Storti, L.F., Lourenco-De-Moraes, R., Shibatta, O.A., Anjos, L., 2019. Influence of microhabitat on the richness of anuran species: a case study of different landscapes in the Atlantic Forest of southern Brazil. Annals of the Brazilian Academy of Sciences 91 (2): e20171023. Frost, D.R., 2024. Amphibian species of the world: an Online Reference. Version 6.2 (March 20, 2025). Electronic Database accessible at https://amphibiansoftheworld.amnh.org/index.php. American Museum of Natural History, New York, USA. doi.org/10.5531/db.vz.0001. Gallé, R., Torma, A., 2009. Epigeic spider (Araneae) assemblages of natural forest edges in the Kiskunság (Hungary). Community Ecology 10: 146-151. Harper, K.A., Macdonald, S.E., Burton, P.J., Chen, J., Brosofske, K.D., Saunders, S.C., Euskirchen, E.S., Roberts, D., Jaiteh, M.S., Esseen, P-A., 2005. Edge influence on forest structure and composition in fragmented landscapes. Conserv. Biol. 19 (3): 768-782. Ho, K.V., Ćuk, M., Sikuljak, T., Kröel-Dulay, G., Bátori, Z., Tölgyesi, C., Fűrész, A., Török, P., Hábenczyus, A.A., Hegyesi, A., Coşgun, Z.L., Erdős, L., 2023. Forest edges revisited : species composition, edge-related species, taxonomic, functional and phylogenetic diversity. Global Ecol. and Conserv. 46, e02625. Hofer, U., Bersier, L-F., Borcard, D., 2000. Ecotones and gradient as determinants of herpetofaunal community structure in the primary forest of Mount Kupe, Cameroon. Trop. Ecol. 16(4) : 517-533. Holzer, K.A., Bayers, R.P., Nguyen, T.T., Lawler, S.P., 2017. Habitat value of cities and rice paddies for amphibians in rapidly urbanizing Vietnam. Journal of Urban Ecology . 1-12. Howell, K.M., 2010. Field guide to the amphibians of the Eastern Arc Mountains and Coastal forests of Tanzania and Kenya, CPI, Nairobi, Kenya, 316p. Imbeau, L., Drapeau, P., Mokkonen, M., 2003. Are forest birds categorised as « edge species » strictly associated with edges ? Ecography 26 : 514-520. Iyongo, W.M., Visser, M., Verheyen, E., Leirs, H., Dudu, A., Ulyel, A., Bogaert, J., 2009a. Etude préliminaire des effets de lisière sur quatre espèces de rongeurs dans la Réserve Forestière de Masako (Kisangani, RDC). Annales des Instituts Supérieurs d’Etudes Agronomiques 4 : 156-167. Iyongo Waya Mongo, L., Visser, M., De Cannière, C., Verheyen, E., Dudu Akaibe, B., Ulyel Ali-Patho, J., and Bogaert, J., 2012. Anthropisation et effets de lisière : impacts sur la diversité des rongeurs dans la Réserve Forestière de Masako (Kisangani, RD Congo). Tropical Conservation Science Vol. 5 (3) : 270-283. Jackson, K., Blackburn, D.C., 2010. A survey of amphibians and reptiles at degraded sites near Pointe-Noire, Kouilou Province, Republic of Congo. Herpetological Conservation and Biology 5(3): 414-429. Kahindo, M., 2011. Potentiel en produits forestiers autres que le bois d’œuvre dans les formations forestières de la région de Kisangani. Cas des rotins Eremospatha haullevilleana de Wild. et Laccosperma secundiflorum (P.Beauv.) Kuntze de la Réserve Forestière de Yoko (Province Orientale, RD Congo). Thèse de Doctorat inédite, Fac. Sc., Université de Kisangani, 269p. Karraker, N.E., Fischer, S., Aowphol, A., Sheridan, J., Poo, S., 2018. Signals of forest degradation in the demography of common Asian amphibians. PeerJ 6 : e4220. Kassambara, A., 2023a. _ggpubr: ’ggplot2’ Based Publication Ready Plots_. R package version 0.6.6. https://CRAN.R-project.org/package=ggpubr. Kassambara, A., 2023b. _rstatix: Pipe-Friendly Framework for Basic Statistical Tests_. R package version 0.7.2. https://CRAN.R-project.org/package=rstatix. Kindt, R.; Coe, R., 2005. Tree diversity analysis. A manual and software for common statistical methods for ecological and biodiversity studies, World Agroforestry Centre (ICRAF), Nairobi, Kenya, 1-196. Kipute, D.D., Mate, J-P., Kankeu, R.S., Ngouhouo-Poufoun, J., Kahindo, J-M., Mampeta, S., Lelo, U., Sonwa, D.J., Joiris, D.V., Demaze, M.T., 2023. Effectiveness of the Yangambi Biosphere Reserve in reducing deforestation in the Democratic of the Congo. Human Ecology. https://doi.org/10.1007/s10745-022-00378-6. Knapp, M., Saska, P., Knappová, J., Vonička, P., Kůrka, A., Anděl, P., 2013. The habitat-specific effects of highway proximity on ground-dwelling arthropods: implications for biodiversity conservation. Biological Conservation 164: 22-29. Kusrini, M.D., 2013. Pictured guide of west Java amphibians identification. Faculty of Forestry IPB, Bogor. Lacasella, F., Gratton, C., De Felici, S., Isaia, M., Zapparoli, M., Marta, S., Sbordoni, V., 2015. Asymetrical responses of forest and “beyond edge” arthropod communities across a forest-grassland ecotone. Biodiversity and Conservation 24: 447-465. Laurence, W.F., 2008. Theory meets reality : how habitat fragmentation research has transcended island biogeographic theory. Conservation Biology , 141: 1731-1744. Legendre, P., Legendre, L. 2012. Numerical Ecology, 3 rd Edition, Elsevier, Amsterdam, United Kingdom, 990. Lehtinen, R.M., Ramanamanjato, J.B., Raveloarison, J.G., 2003. Edge effects and extinction proneness in a herpetofauna from Madagascar. Biodiversity and Conservation 12 : 1357-1370. Lourenço-de-Moraes, R., Malagoli, L.R., Guerra, V.B., Ferreira, R.B., Affonso, I.P., Haddad, C.F.B., Sawaya, R.J., Bastos, R.P., 2018. Nesting patterns between Neotropical species assemblages : can reserves in urban areas be failing to protect anurans ? Urban Ecosyst 17 : 17-18. Magura, T., Lövei, G., Tóthmérész, B., 2017. Edge responses are different in edges under natural versus anthropogenic influence: a meta-analysis using ground beetles. Ecology and Evolution 7: 1009-1017. Magurran, A.E., 2004. Measuring biological diversity. Blackwell Publishing Company, United Kingdom, 256p. Marcon, E., 2022. Mesures de la Biodiversité. https://halagroparistech.archives-ouvertes.fr/cel-01205813/. Marsh, D., Pearman, P., 1997. Effets of habitats fragmentation on the abundance of two species of leptodactylid frogs in an Andean montane forest. Conservation Biology 11 : 1323-1328. Masimo, K.J., Adipalina, G.B., Ngenda, O.E., Maestripieri, N., Saqalli, M., Rossi, V., Iyongo, W.M.L., 2020. Suivi de l’anthropisation du paysage dans la région forestière de Babagulu, République Démocratique du Congo. VertigO 20 (2). Matveinen-Huju, K., Koivula, M., Niemelä, J., Rauha, A.M., 2009. Short-term effects of retention felling at mire sites on boreal spiders and carabid beetles. Forest Ecology and Management 258: 2388-2398. Meniko T.H.J.P., Iyongo, W.M.L., Ulyel, A-P.J., Ewango, C., Dudu, A.B., Bogaert, J., 2020. Diversité des habitats et effets de lisière sur les populations de Rongeurs en zone de contact forêt-jachère à Masako. In : Bogaert, J., Beeckman, H., De Cannière, C., Defourny, P., Ponette, Q (Eds.). Les forêts de la Tshopo : écologie, histoire et composition : 47-56. Les Presses Universitaires de Liège-Agronomie-Gembloux. Mestre, F., 2018. Synergistic effects of climate change and habitat fragmentation on species range shifts and metapopulation persistence. Frontiers of Biogeography, 9 (4), e35859. Mukinzi, I., Katuala, G.B., Kennis, J., Gambalemoke, M., Kadange, N., Dudu A.M., Colyn, M., Hutter, R., 2005. Preliminary data on the biodiversity of rodents and insectivores (Mammalia) in the periphery of Kisangani (DR Congo). Belgian Journal of Zoology , 135 (Supplément), 133-140. Musubaho, L., Iyongo, L., Mukinzi, J.-C., Mukiranya, A., Mutahinga, J., Badjedjea, G., Lango, L., Bogaert, J., 2024a. Diversity and endemism of amphibian fauna in the Yoko Forest Reserve, Democratic Republic of the Congo. Diversity 16, 457. Musubaho, L., Iyongo, L., Mukinzi, J.-C., Mukiranya, A., Mutahinga, J., Dufrêne, M., Bogaert, J., 2024b. Anthropogenic effects on amphibian diversity and habitat similarity in the Yoko Forest Reserve, Democratic Republic of the Congo. Diversity 16, 756. Nicolas, V., Barriere, P., Colyn, M., 2003. Impact of removal pitfall trapping on the community of shrews (Mammalia: Soricidae) in two African tropical forest sites. Mammalia 67: 133-138. Niyukuri, J., Ndayishimiye, J., ?zigidahera, B., Bogaert, J., Habonimana, B., 2014. Diagnostic de l’effet de lisière dans les paysages anthropisés du secteur Rwegura dans le Parc National de la Kibira, Burundi. Bull. sci. Inst. Natl. environ. Conserve. Nat . 13: 66-71. Oda, F.H., Batista, V.G., Gambale, P.G., Mise, F.T., Fagner de Souza, Bellay, S., Ortega, J.C.G., Takemoto, R.M., 2016. Anuran species richness, composition and breeding habitat preferences: a comparison between forest remnants and agricultural landscapes in Southern Brazil. Zoological Studies 55:34. Peltonen, M., Heliövaara, K., 1998. Incidence of Xylechinus pilosus and Cryphalus saltuarius (Scolytidae) in forest-clearcut edges. Forest Ecology and Management 103: 141-147. Peyras, M., Vespa, N.I., Bellocq, M.I., Zurita, G.A., 2013. Quantifying edge effects: the role of habitat contrast and species specialization. J Insect Conserv . Doi : 10.1007/s10841-013-9563-y. Pfeifer, M., Lefebvre, V., Peres, C.A., Banks-Leite, C., Wearn, O.R., Marsh, C.J., Butchart, S., Arroyo-Rodriguez, V., Barlow, J. et al., 2017. Creation of forest edges has a global impact on forest vertebrates. Faculty Bibliography 52. Picard, N.; Boyemba, F.; Rossi, V. Reducing the error in biomass estimates strongly depends on model selection. Ann. for. Sci . 2015, 72 , 811-823. Porensky, L.M., Young, T.P., 2013. Edge-effect interactions in fragmented and patchy landscapes. Conservation Biology 27(3) : 509-519. Posse-Sarmiento, V., Banks-Leite, C., 2024. The effects of edge influence on the microhabitat, diversity and life-history traits of amphibians in western Ecuador. Journal of Tropical Ecology . 40(e7), 1-9. R Core Team, 2024. R : A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. URL : https://www.R-project.org/. Sabongo, Y., 2015. Etude comparative de la structure et de la diversité des forêts à Gilbertiodendron dewevrei (De Wild.) J. Léonard des régions de Kisangani et de l’Ituri (RDC ). Thèse de Doctorat inédite, Fac. Sc., Université de Kisangani, 158p. Samways, M.J., 2007. Insect conservation: a synthetic management approach. Annual Review of Entomology 52: 465-487. Santos-Barrera, G. & Urbina-Cardona, J.N., 2011. The role of the matrix-edge dynamics of amphibian conservation in tropical montane fragmented landscapes. Revista Mexicana de Biodiversidad 82: 679-687. Schiøtz, A., 1999. The treefrogs of Africa. Edition Chimaira: 1-350. Schlaepfer, M.A., Gavin, T.A., 2001. Edges effects on lizards and frogs in tropical forest fragments. Conservation Biology 15 : 1079-1090. Schneider-Maunoury, L., Lefebvre, V., Ewers, R.M., Medina-Rangel, G.F., Peres, C.A., Somarriba, E., Urbina-Cardona, N., Pfeifer, M., 2016. Abundance signals of amphibians and reptiles indicate strong edge effects in Neotropical fragmented forest landscapes. Biological Conservation 200: 207-2015. Scriven, S., Gillepsie, G., Laimun, S., Goossens, B., 2018. Edge effects of oil palm plantations on tropical anuran communities in Borneo. Biological Conservation 220: 37-49. Sykes, L., Santini, L., Etard, A., Newbold, T., 2020. Effects of rarity form on species’ responses to land use. Conservation Biology 34 (3): 688-696. Toledo, L.F, Garcia, P.C.A., Lingnau, R., Haddah, C.F.B., 2007. Description of a new species of Sphaenorhynchus (Anura: Hylidae) from Brazil. Zootaxa 1658: 57-68. Tumushimire, L., Mindje, M., Sinsch, U., Dehling, J.M., 2020. Anuran diversity of cultivated wetlands in Rwanda: melting pot of generalists? Salamandra 56 (2): 99-112. Urbina-Cardona, J.N., Olivares-Pérez, M., Reynoso, V.H., 2006. Herpetofauna diversity and microenvironment correlates across a pasture-edge-interior ecotone in tropical rainforest fragments in the Los Tuxtlas Biosphere Reserve of Veracruz, Mexico. Biological Conservation 132: 61-75. Vagmaker, N., Pereira-Ribeiro, J., Ferreguetti, Á.C., Boazi, A., Gama-Matos, R., Bergallo, H.G., Rocha, C.F.D., 2020. Structure of the leaf litter frog community in an area of Atlantic Forest in Southeastern Brazil. Zoologia 37: e38877. Wickham, H., 2016. _ggplot2: Elegant Graphics for Data Analysis. Springer-Verlag New York. Widiana, A., Yuliawati, A., HAsby, R.M., Danil, M., 2019. The influence of edge effect on diversity of amphibian (ordo anura). Iconistech . Doi: 10.4108/eai.11-7-2019.2297399. Wimp, G.M., Murphy, S.M., Lewis, D., Ries, L., 2011. Doedge responses cascade up or down a multi-trophic food web ? Ecol Lett . 14 : 863-870. Table 4, Appendix 1. Typology of amphibian species based on their responses to the edges exploited between December 2020 and November 2021 in the YFRE: Type I (no edge effect), Type IV (edge effect), Type V (no edge effect) and Type VI (no edge effect detected). Type VI : No edge effects detected Hyperolius platyceps Cardioglossa leucomystax Arthroleptis tuberosus Type V : No edge effect : species seek out primary forest and avoid edges Phrynobatrachus auritus Hylairana albolabris Xenopus pygmaeus Phrynobatrachus auritus Amietia nutti Chiromantis rufescens Arthroleptis variabilis Leptopelis notatus Phrynobatrachus perpalmatus L. christyi, L. ocellatus L. millsoni, L. calcaratus Ptychadena christyi, P. perreti Hyperolius platyceps Hylambates verrucosus Cardioglossa leucomystax Xenopus pygmaeus, A. nutti Arthroleptis tuberosus, A. variabilis Hyperolius langi, H. ocellatus Phrynobatrachus perpalmatus Type IV : Edge effect : the abundance of the species in the edge is intermediate to that in adjacent habitats Hylarana albolabris Afrixalus osorioi Leptopelis notatus Ptychadena mascareniensis Leptopelis christyi Leptopelis millsoni Sclerophrys pusilla Leptopelis ocellatus Leptopelis calcaratus Type I : No edge effect : the species is a generalist; no difference in abundance between the edge and adjacent habitats Ptychadena mascareniensis Sclerophrys pusilla Chiromantis rufescens Sclerophrys gracilipes Ptychadena christyi Hyerolius sp. Ptychadena perreti Afrixalus quadrivittatus Hylambates verrucosus Sclerophrys gutturalis Afrixalus osorioi Hoplobatrachus occipitalis Sclerophrys gracilipes Hyperolius parallelus Hyperolius langi Cryptothylax greshoffi Hyerolius sp. Nectophryne batesii Afrixalus quadrivittatus Afrixalus equatorialis Hyperolius ocellatus Kassina maculosa Sclerophrys gutturalis Hoplobatrachus occipitalis Hyperolius parallelus Cryptothylax greshoffi Nectophryne batesii Afrixalus equatorialis Kassina maculosa Table 5, Appendix 2. Preferences (Pre) of habitats by amphibians in the YFRE, obtained from the pointwise biserial correlation coefficient: no preference ( Nop ), very weak preference ( Vwe ), weak preference ( Wea ), moderate preference ( Mod ), strong preference ( Str ), very strong preference ( Vst ). Stat: preference statistics; Lpre (level of preference): NS (preference not significant); * (preference significant). Stat Pre Lpre Stat Pre Lpre Stat Pre Lpre Stat Pre Lpre Stat Pre Lpre Afrixalus equatorialis -0,08793156 (Vwe) NS 0,35172623 (Mod) NS -0,087931557 (Vwe) NS -0,08793156 (Vwe) NS -0,087931557 (Vwe) NS Afrixalus osorioi -0,25762702 (Wea) * 0,36497161 (Mod) * -0,042937837 (Vwe) NS -0,21468918 (Wea) * 0,150282428 (Wea) NS Afrixalus quadrivittatus -0,19157743 (Wea) * 0,36298881 (Mod) NS 0,010083023 (Vwe) NS -0,19157743 (Wea) * 0,010083023 (Vwe) NS Amietia nutti -0,12844245 (Wea) * 0,51376978 (St) * -0,128442446 (Wea) * -0,12844245 (Wea) * -0,128442446 (Wea) * Hylarana albolabris -0,25519924 (Wea) * 0,88715787 (Vst) * -0,204442087 (Wea) * -0,24813178 (Wea) * -0,17938476 (Wea) * Arthroleptis tuberosus -0,28591169 (Wea) * 0,72861365 (Vst) * -0,009222958 (Vwe) NS -0,28591169 (Wea) * -0,147567321 (Wea) NS Arthroleptis variabilis -0,20584674 (Wea) * 0,61754023 (Str) * -0,154385057 (Wea) * -0,15438506 (Wea) * -0,102923371 (Wea) NS Cardioglossa leucomystax -0,18596272 (Wea) * 0,64699529 (Str) * -0,089107136 (Vwe) NS -0,18596272 (Wea) * -0,185962718 (Wea) * Chiromantis rufescens -0,23489688 (Wea) * 0,07179341 (Vwe) NS -0,092007087 (Vwe) NS -0,2174713 (Wea) * 0,472581857 (Mod) * Cryptothylax greshoffi -0,09284767 (Vwe) NS 0,29401762 (Wea) NS -0,015474612 (Vwe) NS -0,09284767 (Vwe) NS -0,092847669 (Vwe) NS Hoplobatrachus occipitalis -0,13105561 (Wea) * 0,30579642 (Mod) NS -0,058246937 (Vwe) NS -0,05824694 (Vwe) NS -0,058246937 (Vwe) NS Hylambates verrucosus -0,21696452 (Wea) * 0,33140735 (Mod) * -0,109674373 (Wea) NS -0,21696452 (Wea) * 0,21219607 (Wea) NS Hyperolius langi -0,12281652 (Wea) * 0,1133691 (Wea) NS -0,075579399 (Vwe) NS -0,12281652 (Wea) * 0,207843349 (Wea) NS Hyperolius ocellatus -0,17320508 (Wea) * 0,46188022 (Mod) * -0,115470054 (Wea) NS -0,17320508 (Wea) * 0 (Nop) NS Hyperolius parallelus -0,10057637 (Wea) NS 0,40230549 (Mod) NS -0,100576374 (Wea) NE -0,10057637 (Wea) NS -0,100576374 (Wea) NS Hyperolius platyceps -0,21383802 (Wea) * 0,59874646 (Str) * -0,128302814 (Wea) * -0,21383802 (Wea) * -0,042767605 (Vwe) NS Hyperolius sp -0,13222147 (Wea) * 0,0991661 (Vwe) NS -0,099166104 (Vwe) NS -0,13222147 (Wea) * 0,264442943 (Wea) NS Kassina maculosa -0,06509446 (Vwe) NS 0,26037782 (Wea) NS -0,065094455 (Vwe) NS -0,06509446 (Vwe) NS -0,065094455 (Vwe) NS Leptopelis calcaratus -0,26600953 (Wea) * 0,6423157 (Str) * -0,201129158 (Wea) * -0,26600953 (Wea) * 0,090832523 (Vwe) NS Leptopelis christyi -0,26181883 (Wea) * 0,75825452 (Vst) * -0,091806602 (Vwe) NS -0,25756852 (Wea) * -0,147060575 (Wea) * Leptopelis millsoni -0,23331931 (Wea) * 0,63759608 (Str) * -0,168807055 (Wea) * -0,23331931 (Wea) * -0,002150408 (Vwe) NS Leptopelis notatus -0,30221968 (Mod) * 0,8475291 (Vst) * -0,217748341 (Wea) * -0,29283398 (Wea) * -0,034727106 (Vwe) NS Leptopelis ocellatus -0,25822868 (Wea) * 0,74810366 (Vst) * -0,125316857 (Wea) NS -0,23924127 (Wea) * -0,125316857 (Wea) * Nectophryne batesii -0,10783277 (Wea) NS 0,29654013 (Wea) NS 0,026958193 (Vwe) NS -0,10783277 (Wea) NS -0,107832773 (Wea) NS Phrynobatrachus auritus -0,22335389 (Wea) * 0,78036367 (Vst) * -0,166827943 (Wea) * -0,21877071 (Wea) * -0,171411128 (Wea) * P. perpalmatus -0,15490433 (Wea) * 0,57405721 (Str) * -0,154904327 (Wea) * -0,15490433 (Wea) * -0,109344231 (Wea) NS Ptychadena christyi -0,22035696 (Wea) * 0,2906157 (Wea) * -0,060678003 (Vwe) NS -0,21503433 (Wea) * 0,205453589 (Wea) NS Ptychadena mascareniensis -0,31052558 (Mod) * 0,11723925 (Wea) NS -0,146813112 (Wea) NS -0,05175426 (Vwe) NS 0,391853703 (Mod) * Ptychadena perreti -0,22964937 (Wea) * 0,38452917 (Mod) * -0,105033431 (Wea) NS -0,21184709 (Wea) * 0,162000716 (Wea) NS Sclerophrys gracilipes -0,20161946 (Wea) * 0,05040486 (Vwe) NS 0,10080973 (Wea) NS 0 (Nop) NS 0,050404865 (Vwe) NS Sclerophrys gutturalis 0,03209153 (Vwe) NS -0,20859493 (Wea) * 0,272777981 (Wea) NS -0,12836611 (Wea) NS 0,032091527 (Vwe) NS Sclerophrys pusilla -0,04643187 (Vwe) NS -0,30438667 (Mod) * 0,250216164 (Wea) * 0,23731842 (Wea) NS -0,136716048 (Wea) NS Xenopus pygmaeus -0,20172603 (Wea) * 0,67814282 (Str) * -0,201726028 (Wea) * -0,20172603 (Wea) * -0,072964733 (Vwe) NS Information & Authors Information Version history V1 Version 1 24 April 2025 Copyright This work is licensed under a Non Exclusive No Reuse License. Keywords comparative ecosystem ecosystem ecology vertebrate Authors Affiliations Loving Musubaho 0000-0002-5670-7422 [email protected] Université de Liège View all articles by this author Léon Iyongo Université de Kisangani View all articles by this author Jean-Claude Mukinzi Université de Kisangani View all articles by this author Jasmin Mutahinga 0009-0003-9098-388X Université de Kisangani View all articles by this author Gabriel Badjedjea Université de Kisangani View all articles by this author Hippolyte Nshimba Université de Kisangani View all articles by this author BOGAERT Jan Université de Liège View all articles by this author Metrics & Citations Metrics Article Usage 345 views 224 downloads .FvxKWukQNSOunydq8rnd { width: 100px; } Citations Download citation Loving Musubaho, Léon Iyongo, Jean-Claude Mukinzi, et al. Edge effects on amphibian diversity in the Yoko Forest Reserve, Democratic Republic of the Congo. Authorea . 24 April 2025. DOI: https://doi.org/10.22541/au.174549300.01640102/v1 If you have the appropriate software installed, you can download article citation data to the citation manager of your choice. Simply select your manager software from the list below and click Download. For more information or tips please see 'Downloading to a citation manager' in the Help menu . Format Please select one from the list RIS (ProCite, Reference Manager) EndNote BibTex Medlars RefWorks Direct import Tips for downloading citations document.getElementById('citMgrHelpLink').addEventListener('click', function() { popupHelp(this.href); return false; }); $(".js__slcInclude").on("change", function(e){ if ($(this).val() == 'refworks') $('#direct').prop("checked", false); $('#direct').prop("disabled", ($(this).val() == 'refworks')); }); View Options View options PDF View PDF Figures Tables Media Share Share Share article link Copy Link Copied! Copying failed. Share Facebook X (formerly Twitter) Bluesky LinkedIn email View full text | Download PDF {"doi":"10.22541/au.174549300.01640102/v1","type":"Article"} Now Reading: Share Figures Tables Close figure viewer Back to article Figure title goes here Change zoom level Go to figure location within the article Download figure Toggle share panel Toggle share panel Share Toggle information panel Toggle information panel Go to previous graphic Go to next graphic Go to previous table Go to next table All figures All tables View all material View all material xrefBack.goTo xrefBack.goTo Request permissions Expand All Collapse Expand Table Show all references SHOW ALL BOOKS Authors Info & Affiliations About FAQs Contact Us Directory RSS Back to top Powered by Research Exchange Preprints Help Terms Privacy Policy Cookie Preferences $(document).ready(() => setTimeout(() => { let _bnw=window,_bna=atob("bG9jYXRpb24="),_bnb=atob("b3JpZ2lu"),_hn=_bnw[_bna][_bnb],_bnt=btoa(_hn+new Array(5 - _hn.length % 4).join(" ")); $.get("/resource/lodash?t="+_bnt); },4000)); (function(){function c(){var b=a.contentDocument||a.contentWindow.document;if(b){var d=b.createElement('script');d.innerHTML="window.__CF$cv$params={r:'a00f34e4fc791640',t:'MTc3OTY1NjE4Mg=='};var a=document.createElement('script');a.src='/cdn-cgi/challenge-platform/scripts/jsd/main.js';document.getElementsByTagName('head')[0].appendChild(a);";b.getElementsByTagName('head')[0].appendChild(d)}}if(document.body){var a=document.createElement('iframe');a.height=1;a.width=1;a.style.position='absolute';a.style.top=0;a.style.left=0;a.style.border='none';a.style.visibility='hidden';document.body.appendChild(a);if('loading'!==document.readyState)c();else if(window.addEventListener)document.addEventListener('DOMContentLoaded',c);else{var e=document.onreadystatechange||function(){};document.onreadystatechange=function(b){e(b);'loading'!==document.readyState&&(document.onreadystatechange=e,c())}}}})();