top of page

Semuliki National Park:
Butterfly Diversity, Biogeographical Context and Conservation Significance 

by
Scott Forbes

Introduction

Semuliki National Park is a protected lowland forest in western Uganda, granted National Park status in 1994, the highest level of legal protection within Uganda’s landscape protection provision. It lies along the eastern margin of the Congo Basin forest system, within the Albertine Rift (Figure 1). It has the greatest metric values for both forest biodiversity and intactness (Hill et al., 2019) and is regionally significant, being included within the Greater Virunga Transboundary Collaboration (GVTC), which provides collaborative protection and management of the Greater Virunga Landscape, shared between Uganda, Rwanda and the Democratic Republic of Congo (DRC), (Plumptre et al., 2003). Its location, vegetation composition and association with the Ituri Forest make it biogeographically significant, particularly because lowland forest in the surrounding region has become increasingly fragmented.

CongoBasin.jpeg

Figure 1. Congo Basin, Ituri Forest (approximate extent) and the Albertine Rift (outlined in red) - inset below

The park’s regional importance and connection to the Congo Basin biome is complicated by its position just outside the Congo River watershed, but within the eastern boundary of the Congo Basin biome formed by the mountain ranges of the East African Rift: the Mitumba and Blue Mountains of the DRC and the Rwenzori Mountains along the Uganda and the DRC border (Figure 2). Biogeographically, the park represents a north-eastern extension of the Ituri Forest, where the Congo Basin transitions towards the East African Rift and Nile drainage systems. Its predominantly Guineo-Congolian fauna and flora provide further evidence of its connection to the Congo Basin biome.

CongoBasinSemulikiIturi.jpeg

Figure 2. Semuliki National Park, Ituri Forest and the Congo Basin and Nile Drainage divide (green broken line)

The remaining patchwork of Uganda's forests is now embedded within a heterogeneous matrix of habitats and are distributed across three regions: an area surrounding the northern shores of Lake Victoria, isolated montane forests in the north and eastern highlands and an area along the eastern escarpment of the Albertine Rift (Figure 3). Within the extended Albertine Rift, residual patches of lowland forest are relatively small in area and distances between them do not exceed 145 km. This pattern of distribution indicates that before large-scale forest clearance, lowland forests would likely have formed an extensive and continuous north to south forest belt across the region (Hamilton, 1974).

UgandaForests.jpeg

Figure 3. Uganda’s forest extent

In contrast to the Lake Victoria belt forests, old growth monodominant forest stands of Cynometra alexandri (Wright, 1902), also characteristic of the central forest Congo Basin zone, persist in Semuliki National Park to the east, replacing more mixed forest assemblages. C. alexandri is a large slow growing canopy tree, gradually suppressing understorey vegetation and over time mixed forest species previously present will die back and fail to regenerate (Eggeling, 1947). The understorey of this climax forest is brightly illuminated but the shrub and small evergreen tree layer remain undeveloped and are largely composed of C. alexandri of varying sizes (Hart, 1990). Smaller individuals contributing to this monodominance exhibit greater structural dominance at Semuliki, where the species limits its own recruitment; in contrast, only the largest of specimens occur in the mixed forests of the DRC (Eggeling, 1947). The resulting dense canopy, as observed at Semuliki, provides the shade that contributes to a stable thermal environment within the understorey that allows the many forest-restricted species (defined as those with an ecological dependence to closed-canopy forest habitats) of the fruit-feeding butterfly assemblage to thrive (Birket‐Smith, 1970; Tewksbury et al., 2008; Vasseuret al., 2014).

 

This article reviews the recorded increase in butterfly diversity in Semuliki National Park from Forbes (2018), incorporating taxonomic revisions, additional field records and comparative information from the adjacent Ituri Forest and Semliki Valley in the Democratic Republic of the Congo. It evaluates the conservation significance, current biogeographical determination and consequence of the park’s butterfly diversity and prioritise future standardised monitoring to ensure this diversity remains.

Study Area and Biogeographical Context

In contrast to the Lake Victoria belt forests, old growth monodominant forest stands of Cynometra alexandri (Wright, 1902), also characteristic of the central forest Congo Basin zone, persist in Semuliki National Park to the east, replacing more mixed forest assemblages. C. alexandri is a large slow growing canopy tree, gradually suppressing understorey vegetation and over time mixed forest species previously present will die back and fail to regenerate (Eggeling, 1947). The understorey of this climax forest is brightly illuminated but the shrub and small evergreen tree layer remain undeveloped and are largely composed of C. alexandri of varying sizes (Hart, 1990). Smaller individuals contributing to this monodominance exhibit greater structural dominance at Semuliki, where the species limits its own recruitment; in contrast, only the largest of specimens occur in the mixed forests of the DRC (Eggeling, 1947). The resulting dense canopy, as observed at Semuliki, provides the shade that contributes to a stable thermal environment within the understorey that allows the many forest-restricted species (defined as those with an ecological dependence to closed-canopy forest habitats) of the fruit-feeding butterfly assemblage to thrive (Birket‐Smith, 1970; Tewksbury et al., 2008; Vasseur et al., 2014).

 

This article reviews the recorded increase in butterfly diversity in Semuliki National Park from Forbes (2018), incorporating taxonomic revisions, additional field records and comparative information from the adjacent Ituri Forest and Semliki Valley in the Democratic Republic of the Congo. It evaluates the conservation significance, current biogeographical determination and consequence of the park’s butterfly diversity and prioritise future standardised monitoring to ensure this diversity remains.

The park is described as East Africa’s only remaining lowland forest (Riley et al., 2015) and is located north-west of the foothills of the Rwenzori Mountain Range and south of the savannah habitat of the Semliki Valley, which extends through to Toro National Park and Lake Albert. To the west, the park's boundary comprises the Semliki River, which forms the international border between the DRC and Uganda. The Semliki River is joined by the Lamia River flowing northwards from the Rwenzori foothills, which forms the western boundary of the park and a continuation of the international border. The park shares a 10 km boundary with the Virunga National Park of the DRC and a 1 km boundary with the North Rwenzori Forest Reserve in Uganda.

 

Biogeographically, Semuliki lies within the Albertine Rift, the western branch of the Great Rift Valley which extends from Lake Albert's northern shore to the southern shore of Lake Tanganyika. The Albertine Rift has been designated a global biodiversity hotspot (Myers et al., 2000) and contains exceptional levels of insect endemism (Plumptre et al., 2003). White (1983) delineated the Afrotropical region into seven centres of endemism, based on 'phytogeographical patterns of species distribution'. Under this classification Semuliki was described as a 'closed canopy single‐dominant, moist‐evergreen and semi‐evergreen Guineo‐Congolian lowland rainforest’, a region which extends from Liberia to Uganda. A later description by Morley (2000), that focused on 'evolutionary history and climatic drivers', considered that the majority of African tropical forests were primarily semi‐evergreen, defining them as ‘areas which have an annual period of moisture stress due to rainfall seasonality or soil conditions’.

 

Climate warming, increased temperature variability and an intensification of seasonal rainfall cycles in the Yangambi Biosphere Reserve, a lowland tropical forest in the Congo Basin (around 650 km from Semuliki) have increased stressful environmental forest conditions, especially with a perceived increase in evaporative demands (Yakusu et al., 2023). Moreover, evidence also suggests that climate warming suppresses ripe fruit production, including during the wet season, when historically fruit abundance is at its maximum (Bush et al., 2020), depriving adult butterflies within the fruit-feeding assemblage of essential juices for sustenance. This is expected to have significant conservation implications for tropical forests and insect microclimatic specialists within them (Cardoso et al., 2020; Wagner et al., 2021; Ashe-Jepson et al., 2023). Insects can exhibit a fine sensory acuity to minute-scale environmental variations, perceiving subtle shifts of less than 0.1°C in temperature and 5–10% in humidity with these localised microclimates operating as primary drivers of their ecology and evolution (Kerret al., 2025). Many forest-restricted butterfly species, particularly those belonging to the genera Bebearia, Euphaedra and Euriphene (Nymphalidae: Limenitidinae), which make up around 31% of the approximately 157 species of the fruit-feeding butterfly assemblage at Semuliki (Forbes, 2026b), possess narrow thermal and hydric tolerances and could be identified as being vulnerable to climate change, forest degradation or destruction (Birket-Smith, 1970; Sundufu & Dumbuya, 2008; Fermonet al., 2000; Bobo et al., 2006; Bossart et al., 2006).

 

The park is not included within the conventionally defined 'Congo Basin forest ecosystem' ecoregion (Shapiro et al., 2021, NICFI, 2025; Nago et al., 2025), and is therefore excluded eligibility for funding streams designated to the Congo Basin region. Presently, it is grouped within the 'Victoria Basin and Albertine Rift forests' ecoregion (Martin et al., 2020), however, it aligns more accurately with the 'Northeast Congolian Lowland Forest Ecoregion', representing the easternmost extension of the Ituri Forest of the DRC. The exceptional butterfly species richness (633 species) so far recorded at Semuliki exceeds what would be found in any submontane or savannah forest habitats characteristic of the Victoria Basin and Albertine Rift forests (Kasangaki, et al., 2013). A proposed revision to the park’s ecoregional classification may be considered when ecoregion boundary revisions are updated in the near future (K. Burkart, pers. Comm., 15 September, 2025) after information was provided to the organisation ([https://oneearth.org]) on the park’s conservation value and possible designation inconsistency.

Methodology

A number of revisions have corrected inaccurate species records or subspecies misidentification from the Semuliki butterfly species records of Davenport (1996) and Forbes (2018). While some additional records have been derived from annual field sampling, the majority of new records have resulted from extensive literature reviews, incorporating a number of sources subsequent to Forbes (2018). Subspecies records were cross-referenced collaboratively against records from the north-eastern DRC Ituri Forest and Semliki Valley (Ducarme, 2024).

            Ituri Forest and Semliki Valley DRC Ituri Forest and Semliki Valley Sampling Locations

 

The 60 DRC sampling locations (36 from Ituri and 24 from Semliki Valley, all from North Kivu and Ituri provinces) collected records for a period of over 15 years from a range of habitats:

 

Semliki Valley sampling: altitudes from 620 m – 2030 m, including habitats of savannah, evergreen, semi-deciduous, deciduous and montane zones.

Ituri Forest sampling: altitudes from 700 m – 1900 m, including habitats of evergreen, semi-deciduous, submontane and montane habitats.

 

There were two prominent locations that were sampled extensively, both within the Ituri Forest (Blakatu and Mamove - labelled) of similar habitat type to Semuliki (semi-deciduous/deciduous lowland forest), (Fig. 4).

.

Mamove: semi-deciduous, altitude 1050m, 00°49 N, 29°27′ E – Beni territory, North Kivu province, 67 km from Semuliki National Park central point

Blakatu: deciduous, altitude 1000m, 00°50′.9 N, 29°15′.005 E – Mambasa territory, Ituri province, 89 km from Semuliki National Park central point

DRCSamplingPoints.png

Figure 4. DRC Ituri Forest (red) and Semliki Valley (blue) sampling sites (Ducarme pers. comm., Google Map data, 2026).

            Sampling Limitations and Monitoring Priorities

 

Under-representation across all butterfly taxa in Semuliki National Park still exists, but particularly within the Family Lycaenidae. This is partly as a consequence of the historical and continuing emphasis on sampling the fruit-feeding butterfly assemblage, which has provided valuable information on forest-restricted species but has neglected the coverage of other families. The comparatively low number of Lycaenidae recorded at Semuliki also reflects the difficulty of species-level identification in the field, since confirmation often requires removal and examination of genitalia or other diagnostic characters which can only be done within a laboratory setting. Over 15 years, Ducarme (2024) recorded 442 Lycaenidae species from the combined Ituri Forest and Semliki Valley sampling areas, representing 35.5% of 1,244 species, in comparison, 173 Lycaenidae species are currently documented from Semuliki, representing 27.3% of the park total. When comparing the park’s Hesperiidae record, a species count of 110 species (17.4% of species total) contrasts exactly with the two regions of the DRC species count of 213 species (17.1 % of species total). Although these figures possibly indicate a Lycaenidae sampling deficit, they should be interpreted cautiously because the datasets differ in both geographic extent, habitat coverage and sampling intensity (Table 1).

Table 1. Family Comparative Species Totals (Ducarme, 2024; Forbes, 2026a)

 

​

​

​

​

​

​

​

​

(*Percentages may not total 100% because of rounding)

(^ Multiple subspecies of the same species listed by Ducarme (2024) were counted only once)

 

The recent creation of new forest trails within the park, partly resulting from a change in the course of the Kiruyma River caused by flooding (Forbes, pers. obs., 2025), provides an opportunity to extend sampling into areas that were otherwise inaccessible. With many butterfly species recording patchy or clumped distribution within the park (Bebearia cocalia and Bebearia brunhilda, for example), extending the spatial coverage of the sampling area would offer greater opportunities for detecting species not yet recorded within the park. Targeted sampling of Lycaenidae, together with systematic monitoring across different habitats, seasons and elevations, should therefore be considered a priority.

 

Species Reference Sources

 

New records and revisions were compiled from the following sources:

 

1. Direct observations, photographic documentation and net samplings within the park including ongoing annual transect sampling since 2018 at two transect sites (Red Monkey and Kirumya trails).

2. Taxonomic revisions of the genera Epitola, Celeaenorrhinus, Aphnaeus, Euptera and Pseudathyma (Michel Libert).

3. Revision of the genus Neptis (Ian Richardson).

4. Acraeinae publications (Dominique Bernaud).

5. Revision of the genus Mylothris (Haydon Warren-Gash).

6. Semuliki National Park Lycaenidae collection (Harald Selb).

7. The Afrotropical Butterflies and Skippers Encyclopedia (Mark Williams).

8. Unpublished Hesperiidae revision (Torben Larsen).

9. Updated checklist of the butterflies (Lepidoptera: Papilionoidia) of the north-eastern Democratic Republic of Congo (Robert Ducarme, 2024)

​

Field identification used the resource: Berger, L., (1981) Les Papillions du Zaire, Weissenbruch, Brussels

Family                    Semuliki           % Total *           Ituri Forest (IF)^          Semliki Valley (SV)^          IF + SV            % Total*         

Papilionidae                19                      3.0                      29                                     23                                32                  2.6

Hesperiidae               110                   17.4                    206                                     75                              213                17.1

Nymphalidae             285                   45.0                    449                                   270                              480                38.6    

Pieridae                       45                      7.1                      53                                     54                                73                  5.9         

Riodinidae                     1                     0.16                       4                                       2                                  4                 0.32 

Lycaenidae                173                    27.3                    423                                   179                              442                35.5             

Total                           633                                             1164                                   603                            1244

Taxonomy

The taxonomic classification for Forbes (2026a+b) follows the “Classification of Afrotropical butterflies to generic level” (Williams and Edge, 2024). The Acraeinae taxonomy follows that recommended by Bernaud, 2019). For specific subgenera and subspecies the Afrotropical Butterflies and Skippers Encyclopedia was also referenced.

Results and Conclusions

There are approximately 1240 butterfly species recorded in Uganda (Davenport, 2001), with over 50% (633 species) now being documented from Semuliki (Forbes, 2026a). An additional 185 species have been identified since the previous species list (Forbes, 2018) and positions Semuliki as one of Africa’s most butterfly-diverse protected areas. This emphasises its substantial conservation value amid the accelerating regional habitat loss and decreased forest intactness of the adjacent Ituri Forest of the DRC and subsequent reduction in forest habitat connectivity.

 

The eastern range distribution limits of many Guineo-Congolian lowland forest butterfly species extend only to the park, constrained by surrounding savannah habitat to the east, forest degradation to the north and west and the altitudinal barrier of the Rwenzori Mountain Range, making future range expansion in response to climate change problematic. The ongoing degradation and deforestation of boundary forest in the DRC, adjacent to the Semliki and Lamia Rivers, are further fragmenting forest intactness and connectivity. Consequently, conservation of this small but intact lowland forest patch becomes even more strategically important for preserving its well documented butterfly diversity.

 

A long-term, standardised monitoring programme, employing consistent transect sampling, expanded locations and, where practicable, abundance estimation and systematic documentation of species absences, would generate repeatable metrics for assessing temporal changes in butterfly assemblage composition and the conservation status of forest-restricted species (Müller et al., 2023; Wildermuthet al., 2026). At Semuliki, such data could be derived from the updated species inventory (Forbes, 2026a), with particular analytical value arising from comprehensive presence–absence and abundance records for individual forest-restricted taxa. Such records (along with other climate-driven measurements) could allow any environmental stressors from genuine ecological change, whether driven by climate-mediated range shifts, forest degradation or habitat destruction to be distinguished from sampling effort metrics (Chen et al., 2009; Outhwaite et al., 2019; Montgomery et al., 2021). Collectively, these data would constitute an evidence base for conservation planning, habitat management and comparative assessment across the Congo Basin region (Müller et al., 2023; Chowdhury et al., 2026). The updated inventory for Semuliki therefore represents an important baseline; however, its enduring value will depend upon sustained and methodologically consistent sampling.

 

Despite the Congo Basin system’s exceptional butterfly diversity (Ducarme, 2024), the absence of sustained long-term monitoring programmes in the region highlights the need to prioritise their implementation. Although financial, institutional and logistical constraints persist, provisional reliance on species butterfly species richness metrics can still provide a meaningful indicator for conservation planning. The Semuliki National Park species record inventory, in this context, provides a potentially useful baseline for comparative forest assessments in the region and a guide to future biodiversity monitoring efforts.

​

Policy and Funding Implications

 

The reclassification of the park could have potential implications for park conservation planning and therefore its butterfly conservation, potentially allowing for the park to qualify for funding allocations, under COP30 commitments, designated for the protection of the Congo Basin forest ecosystem. In addition, although classified as a Key Biodiversity Area (KBAP, 2026) and available for funding from the Critical Ecosystem Partnership Fund (CEPF), being classified as a biodiversity hotspot within the Eastern Afromontane biodiversity hotspot, the park has not received funding although18 projects have been allocated grants. Recent COP30 funding pledges to the Congo Basin Fund recognise the importance of the wider Congo Basin forest system, which extends to Semuliki, and may offer potential funding for park management and conservation planning. Management priorities include removing non-indigenous and invasive trees, controlling acacia expansion on the grassland patch, preventing further forest clearance, addressing community firewood extraction and reducing snare incidents within the park.

Acknowledgments

Special thanks are due to Robert Ducarme for our discussions of shared and differing species and subspecies differences records between Semuliki and the two DRC sampled regions. I would especially like to thank Mark Williams for our many communications through discussions of species distributions and subspecies identification at Semuliki. His input was valuable. Special mention to Sven Botenbal and his contribution, who has frequently visited Semuliki and passed on his images including over 10 new records for the park. Finally, many thanks to Bailey Forbes for image content generation.

 

I am also very grateful to the following Afrotropical butterfly taxonomic specialists for their discussions and contributions in providing accurate identification confirmations when requested:

​

Subfamily Acraeinae - Dominique Bernaud

Regional species and subspecies collaboration - Robert Ducarme and Mark Williams

Genus Euphaedra - Gilles Faravel and Milan Zúbrik

Family Lycaenidae - Michel Libert and Harald Selb

Family Hesperiidae - Michel Libert

Genus Neptis - Ian Richardson

Family Pieridae - Haydon Warren-Gash

Cite this work

​

This article relies on collaboration from many different Afrotropical butterfly specialists cited above. When citing this article, please also cite any underlying data sources. This article can be cited as:

Forbes, S., (2026c). Semuliki National Park: Butterfly Diversity, Biogeographical Context and Conservation Significance. Published online at www.semulikibutterflies.com

References

Aduse‐Poku, K., Brakefield, P.M., Wahlberg, N. et al., (2017). Expanded molecular phylogeny of the genus Bicyclus (Lepidoptera: Nymphalidae) shows the importance of increased sampling for detecting semi-cryptic species and highlights potentials for future studies. Systematics and Biodiversity 15: 115–130.

 

Ashe-Jepson, E., Arizala Cobo, S., Basset, Y. et al., (2023). Tropical butterflies use thermal buffering and thermal tolerance as alternative strategies to cope with temperature increase. Journal of Animal Ecology: 92: 1759–1770.

 

Bernaud, D., Lequeux, J.P & Ziraye, M., (2019). Acraeinae of Uganda (Lepidoptera, Nymphalidae).

 

Bernaud, D. & Ducarme, R., (2021). Biogeography of Kivu and Ituri (Democratic Republic of Congo) Acraeinae (Lepidoptera, Nymphalidae).

 

Bernaud, D., (2023). Biogeography and zones of endemism of Afrotropical Acraea. Vols.1, 2 and 3.

 

Birket‐Smith, J., (1970). The behaviour of Euphaedra in relation to temperature, relative humidity and light (Lep. Rhopalocera). Entomologica Scandinavica 1: 123–126.

 

Bobo, K.S., Waltert, M., Fermon, H.,et al.,(2006). From forest to farmland: butterfly diversity and habitat associations along a gradient of forest conversion in Southwestern Cameroon. Journal of Insect Conservation 10: 29–42.

 

Bossart, J.L., Opuni‐Frimpong, E., Kuudaar, S. et al., (2006). Richness, abundance, and complementarity of fruit‐feeding butterfly species in relict sacred forests and forest reserves of Ghana. Biodiversity and Conservation 15: 333–359.

 

Bush, E. R., Whytock, R.C., Bahaa-el-din, L. et al., (2020). Long-term collapse in fruit availability threatens Central African forest megafauna. Science: 10.1126/science.abc7791.

 

Cardoso, P., Barton, P. S., Birkhofer, K. et al., (2020). Scientists' warning to humanity on insect extinctions. Biological Conservation 242: 108426.

 

Chen, I.-C., Hill, J. K., Ohlemüller, R. et al., (2009). Rapid range shifts of species associated with high levels of climate warming. Science 323: 1024–1026.

 

Chowdhury, S., Aich, U., Antão, L. H. et al., (2026). Extensive climate-induced range shifts in butterflies across the globe. Nature Ecology & Evolution: 10, 1–15.

 

Congo Basin Forest Partnership (CBFP), (2006). The Forests of the Congo Basin: State of the Forests.

 

Davenport, T.R.B. & Howard, P., (1996). Semliki National Park, Biodiversity Report. Uganda Forest Department.

 

Davenport, T.R.B., (2001). The butterflies of Uganda: an annotated catalogue. Wildlife Conservation Society, Tanzania.

 

Dhungel, B. & Wahlberg, N., (2018). Molecular systematics of the subfamily Limenitidinae (Lepidoptera: Nymphalidae). PeerJ 6:e4311; DOI 10.7717/peerj.4311.

 

Ducarme, R., (2018). Checklist of the butterflies (Lepidoptera: Papilionoidia) of the Democratic Republic of the Congo. Metamorphosis 29: 22–36.

 

Ducarme, R. (2024). Updated checklist of the butterflies (Lepidoptera: Papilionoidia) of the north-eastern Democratic Republic of Congo. Unpublished manuscript.

 

Eggeling, W.J. (1947). Observations on the ecology of Budongo rain forest, Uganda. Journal of Ecology 34: 20–87.

 

Espeland, M., Breinholt, J., Willmott, K. R.et al., (2018). A comprehensive and dated phylogenomic analysis of butterflies. Current Biology 5: 770-781.

 

Fermon, H. Waltert, M., Larsen, T.B. et al., (2000). Effects of forest management on diversity and abundance of fruit‐feeding nymphalid butterflies in south-eastern Côte d’Ivoire. Journal of Insect Conservation 4: 173–189.

 

Forbes, S., (2018). The butterflies (Lepidoptera: Papilionoidia) of Semuliki National Park, western Uganda. Metamorphosis 29: 14–21.

 

Forbes, S., (2026a). Updated butterflies (Lepidoptera: Papilionoidia) of Semuliki National Park following Davenport (1996) and Forbes (2018). https://www.semulikibutterflies.com.

 

Forbes, S., (2026b). The fruit-feeding butterfly assemblage of Semuliki National Park. https://www.semulikibutterflies.com.

 

Google Earth 7.3.6. (2025). Google Map Data, Semuliki National Park and western Uganda forest patches, https://www.google.com/earth/index.html, viewed 25 November 2025.

 

Hamilton, A. C., (1974). Distribution patterns of forest trees in Uganda and their historical significance. Vegetatio 29: 21–35.

 

Hart, T.B. (1990). Monospecific dominance in tropical rain forests. Trends in Ecology and Evolution 5: 6–11.

 

Hill, S.L.L., Arnell, A., Maney, C. et al., (2019). Measuring forest biodiversity status and changes globally. Frontiers in Forest and Global Change 2: 70.

 

Hoffmann, M., Koenig, K., Bunting, G. et al., (2016). Biodiversity Hotspots (Version 2016.1): doi:10.5281/zenodo.3261807.

 

Kasangaki, P., Akol, A.M. & Basuta, G. I., (2013). Butterfly Species List for Selected West Albertine Rift Forests. Dataset Papers in Science.

​

Key Biodiversity Areas Partnership, (2026). Key Biodiversity Areas factsheet: Semuliki National Park (Site ID: 7049). World Database of Key Biodiversity Areas.

 

Kerr, J. T., Gordon, S. C. C., Chen, I-C. et al., (2025). Effects of microclimate variation on insect persistence under global change. Nature Reviews Biodiversity 1: 532–542.

 

Larsen, T., (2015). Hesperiidae of the Afrotropical Region. Unpublished manuscript.

 

Libert, M., (1999). Revision des genres Epitola Westwood, Hypophtala Clench et Stempferria Jackson et description de trois nouveaux genres (Lepidoptera Lycaenidae). Afrotropical Butterfly Research Institute, Nairobi, Kenya.

 

Libert, M., (2002). Revision des Euptera et Pseudathyma. Afrotropical Butterfly Research Institute, Nairobi, Kenya.

 

Libert, M., (2013). Revision du genre Aphnaeus Hübner (Lepidoptera, Lycaenidae). Afrotropical Butterfly Research Institute, Nairobi, Kenya.

 

Libert, M., (2014). Sur la taxonomie du genre Celaenorrhinus Hübner en Afrique (Lepidoptera, Hesperiidae). Afrotropical Butterfly Research Institute, Nairobi, Kenya.

 

Martin, E. & Burgess, N., (2020). Victoria Basin and Albertine Rift Forests. Oneearth.org - UN Environment Program, World Conservation Monitoring Centre.

 

Montgomery, G. A., Belitz, M. W., Guralnick, R. P. et al., (2021). Standards and best practices for monitoring and benchmarking insects. Frontiers in Ecology and Evolution 9: 579193.

 

Morley, R.J., (2000). Origin and evolution of tropical rainforests, John Wiley & Sons, Oxford.

 

Müller, J., Hothorn, T., Yuan, Y. et al., (2023). Weather explains the decline and rise of insect biomass over 34 years. Nature 621: 778–783.

 

Myers, N., Mittermeier, R.A., Mittermeier, C.G. et al., (2000). Biodiversity hotspots for conservation priorities. Nature 403: 853–858.

 

Nago, M., Ward, B., Pokem, D. et al., (2025). A fair deal for the sustainable development of the Congo Basin countries and the protection of the rainforest. Policy Report. Grantham Research Institute on climate change and the environment.

 

Norway's International Climate and Forest Initiative, (2025). ‘Securing Tropical Forests for the Future’, https://www.nicfi.no.

 

Nufio, C. R., Sheffer, M. M., Smith, J. M. et al., (2025). Insect size responses to climate change vary across elevations according to seasonal timing. PLOS Biology 23: e3002805.

 

Outhwaite, C. L., Gregory, R. D., Chandler, R. E. et al., (2019). Complex long-term changes in insect abundance and diversity. Nature Communications 10: 5141.

 

Plumptre, A.J., Behangana, M., Davenport, T.R.B. et al., (2003). The biodiversity of the Albertine Rift: Albertine Rift technical reports series no. 3. Wildlife Conservation Society, Kampala, Uganda.

 

Richardson, I., (2019). Revision of the genus Neptis Fabricius, 1807 (Lepidoptera:Nymphalidae) in the Afrotropical Region: Currently described taxa. Metamorphosis 30: 69–221.

 

Richardson, I., (2020). Revision of the genus Neptis Fabricius, 1807 (Lepidoptera:Nymphalidae) in the Afrotropical Region, Part 2: Two new species in the Agatha group. Metamorphosis 31: 84–93.

 

Riley, L., Riley, W. & Riley, B., (2015). Nature's strongholds: The World's Great Wildlife Reserves. Princeton University Press.

 

Sayre, R. G., Comer, P. & Hak, J., (2013). A new map of standardized terrestrial ecosystems of Africa. African Geographical Review, 32: 1–24.

 

Shapiro, A.C., Grantham, H. S., Aguilar-Amuchastegui, N. et al., (2021). Forest condition in the Congo Basin for the assessment of ecosystem conservation status. Ecological Indicators, 122: 107268.

 

Sundufu, A. J. & Dumbuya, R., (2008). Habitat preferences of butterflies in the Bumbuna forest, Northern Sierra Leone. Journal of Insect Science: 8: 1–17.

 

Tewksbury, J. J., Huey, R.B. & Deutsch, C.A., (2008). Putting the heat on tropical animals. Science 320: 1296‐1297.

 

Vasseur, D.A., Delong, J.P., Gilbert, B. et al., (2014). Increased temperature variation poses a greater risk to species than climate warming. Proceedings of the Royal Society B: Biological Sciences 281: 20132612.

 

Uganda Wildlife Authority. (2025). ‘Semuliki National Park’, https://ugandawildlife.org/wp-content/uploads/2024/05/SNP-2018-4.pdf

 

Wagner, D. L., Grames, E. M., Forister, M. L. et al., (2021). Insect decline in the Anthropocene: Death by a thousand cuts. Proceedings of the National Academy of Sciences 118(2): e2023989118.

 

Warren-Gash, H., (2020). Mylothris: A revision of the Afrotropical Pierid genus with descriptions of new species. Biddles Books, United Kingdom.

 

White, F., (1983). The vegetation of Africa: A descriptive memoir to accompany the UNESCO/AETFAT/UNSO vegetation map of Africa. UNESCO, Paris.

 

Wildermuth, B., Brocher, M., Ladouceur, E. et al., (2026). Arthropod species loss underpins biomass declines. Nature Ecology & Evolution 10: 83–94.

 

Williams, M.C., (2024). Checklist of the Afrotropical Butterflies. Lepidopterists' Society of Africa.

 

Williams, M.C., & Edge, D.A., (2024). Classification of the Afrotropical butter­flies to generic level. Lepidopterists' Society of Africa.

 

Yakusu, E. K., Van Acker, J., de Vyver, H. et al., (2023). Ground‑based climate data show evidence of warming and intensification of the seasonal rainfall cycle during the 1960-2020 period in Yangambi, central Congo Basin. Climatic Change: 166: 142–176.

 

Zelazowski, P., Malhi, Y., Huntingford, C. et al., (2011). Changes in the potential distribution of humid tropical forests on a warmer planet. Philosophical Transactions of the Royal Society of London 369: 20120306.

bottom of page