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Conservation status, distribution and richness of small in

Emmanuel DO LINH SAN1, Adam W. FERGUSON2 3 4,5, Michael HOFFMANN4,6 7, Francesco M. ANGELICI8 9,10 , Jerrold L. BELANT , Jan SCHIPPER Abstract , Philippe GAUBERT and Michael J. SOMERS We assessed the global conservation status of small carnivores in Africa based on the IUCN Red List of Threatened Species. African small carnivores represent about 34% of extant small carnivores worldwide. Familial diversity is intermediate, with four of the world’s nine families represented (Herpestidae: 47% of African species; : 20%; Nandiniidae: 2%; and :

(with a maximum of 26 in any one country). Of the 55 small species found in Africa, 51 (93%) are predominantly dis- 31%).tributed Greatest in Africa species and 48 richness (87%) areis recorded endemic. in In equatorial terms of IUCNAfrica, Red although List conservation most sub-Saharan status, 43 countries species arehost Least at least Concern 15 species (LC), smallthree arecarnivore Near Threatened species (i.e. (NT), non-DD), four are8% Vulnerable were considered (VU) and threatened five are Data (all VU),Deficient primarily (DD). a No result African of population small carnivore declines species and smallis currently distribution listed asranges Endangered (encompassing (EN), Critically only 2–6 Endangered countries). (CR),The exact Extinct percentage in the Wild of threatened (EW) or Extinct species (EX). is unknown, For data-sufficient but is be- while 27% are thought stable and 27% are believed decreasing. Compared with worldwide, the overall conservation statustween of7% small (if no carnivores DD species in is Africa threatened) appears and relatively 16% (if favourable.all are). Population However, trends declining are adjudged populations unknown of many for species 46% of and the existingspecies, (habitat loss, degradation and fragmentation; exploitation for meat) and new threats (rapid economic development expanding the wild meat market, possibly to ) hint that additional small carnivore species may become threatened unless effective - nivore species are endemic to Africa. Actions to remove or mitigate factors threatening Vulnerable and Near Threatened species constituteconservation the strategies short-term are priority implemented. for small This carnivore is of prime conservation importance in Africa. considering that over a quarter of the world’s small car

Keywords IUCN Red List, Least Concern, Mustelidae, Nandiniidae, Near Threat- ened, population trends, species richness, Viverridae, Vulnerable : conservation status, Data Deficient, Herpestidae, Statut de conservation, répartition et richesse spécifique des petits carnivores en Afrique Résumé Liste Rouge des Espèces Menacées de l’UICN. Les petits carnivores africains représentent environ 34% des petits carnivores existant à travers le monde. La diversité Nous avons évalué l’état de conservation global des petits carnivores en Afrique en utilisant la - familiale est intermédiaire, avec quatre des neufs familles de la planète représentées (Herpestidae: 47% des espèces africaines; Mustelidae: 20%; Nandiniidae: 2%, et Viverridae: 31%). La plus grande richesse en espèces est enregistrée en Afrique équato- riale, bien que la plupart des pays d’Afrique subsaharienne hébergent au moins 15 espèces (avec un maximumListe Rouge de 26 de dans l’UICN un, même pays). Sur les 55 espèces de petits carnivores qui se trouvent sur le continent africain, 51 (93%) sont essentiellement dis tribuées en Afrique et 48 (87%) sont endémiques. En ce qui concerne leurs statuts de conservation sur la 43 espèces sont dans la catégorie « Préoccupation mineure » (LC), trois sont « Quasi menacé » (NT), quatre sont « Vulnérable » (VU) et cinq sont classées dans la catégorie « Données insuffisantes » (DD). Aucune espèce de petits carnivore n’est actuellement considéréecomme menacées « En danger (toutes » (EN), VU), principalement« En danger critique en raison » (CR), du « déclinÉteint desà l’état populations sauvage » concernées (EW) ou « Éteintet de leurs» (EX). aires Parmi de lesrépartition espèces de petits carnivores pour lesquelles il existe des données suffisantes afin de leur attribuer un statut, 8% ont été considérées géographique réduites (incluant seulement de 2–6 pays). Le pourcentage exact d’espèces menacées est inconnu, mais il est compris entre 7% (si aucune espèce DD n’est menacée) et 16% (si toutes le sont). Les tendances démographiques sont adjugées favorable.inconnues Toutefois, pour 46% le des déclin espèces, des populations tandis que de27% nombreuses sont considérées espèces stableset les menaces et 27% existantessont estimées (perte, être dégradation en baisse. Paret fragmen rapport- aux mammifères à travers le monde, le statut général de conservation des petits carnivores en Afrique semble relativement tation de l’habitat; exploitation pour la viande) et les nouvelles menaces (développement économique rapide entraînant une- expansion du marché de la viande sauvage, peut-être jusqu’en Asie) laissent entendre que d’autres espèces de petits carnivores pourraient devenir menacées à moins que des stratégies de conservation efficaces soient mises en œuvre. Ceci est d’une impor tance primordiale étant donné que plus d’un quart des espèces de petits carnivores du monde sont endémiques à l’Afrique. Des actions pour éliminer ou atténuer les facteurs menaçant les espèces « Vulnérable » et « Quasi menacé » constituent la priorité à Motscourt clésterme pour la conservation des petits carnivoresListe en Rouge Afrique. de l’UICN, Mustelidae, Nandiniidae, « Préoccupation mineure »,

: « Données insuffisantes », Herpestidae, « Quasi menacé », richesse spécifique, statut de conservation, tendances démographiques, Viverridae, « Vulnérable »

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Introduction (1.05 US billion) of the estimated global human population Encompassing nearly 30.3 million km² or 20% of the Earth’s (6.97 US billion) live in Africa (UN 2011). People exert local, land surface, Africa represents the second largest continent regional and national pressure on wildlife populations, espe- and one of the oldest and most geologically stable land masses cially mammals of medium to large size (i.e. >3 kg; Cardillo et on Earth, existing in continental form for at least 3,800 million al. 2005), often enhancing their risk of extinction. - tor, Africa is the only continent to occupy both northern and value (e.g. food and tourism, including hunting), many large years of Earth’s history (Schlüter 2008). Spanning the Equa Because of their size, often diurnal habits and economic- ranges from 37°20′N in Tunisia to 34°50′S along ’s damental (see Kingdon et al. 2013) and applied research, southernCape region, temperate longitude zones of the (O’Brien mainland & Peters from 17°311999).′ WLatitude in Da- primarilyAfrican mammals serving thehave purposes been subjected of the wildlife to considerable industry (seefun kar, , to 51°25′E in coastal . With the exception of the Atlas Mountains, running from southwestern carnivores have drawn considerable attention from research- along the Mediterranean coastline to the eastern edge of Tuni- ersBothma (and & conservation du Toit 2010). organisations), Among those initially species, through large African their sia, northern Africa is dominated by the world’s largest desert: the Desert covers approximately 9 million km², nearly resulting threats to several species and populations (Gittle- 30% of continental Africa. In the east, the Great Rift Valley, a mancharisma et al. but2001). also Large for theircarnivores potential are important for conflicts in andregulat the- massive depression bordered by numerous mountain chains, et al. 2011) through cascading interactions across trophic levels (Steneck 2005, ing land and aquatic ecosystems (Estes intersectingruns from northern 10 African Syria countries to central and . running over The 6,695 world’s km although more species-rich and generally more common, are fromlongest its river,origin the in Rwanda Nile, flows (Liu northward et al. 2009) through to its mouth the Sahara, along mistakenlyTerborgh & thoughtEstes 2010). to have Small a lower carnivores, impact on at the the other ecosystem hand, ’s Mediterranean coastline (Reader 1997). The Sahel, a level (Roemer et al. 2009). Indeed, although their impacts are broad expanse of semi-arid spanning the southern not on the same guild of prey as large carnivores, small carni- edge of the Sahara Desert, separates the dry deserts of North vores are similarly important ecosystem regulators through Africa from the tropical Sudanian Savannah of north-central structuring small and/or invertebrate communities - (Virgós et al. 1999), which in turn might affect higher trophic pian Highlands, a contiguous region where altitude rarely falls levels. They may also be important in seed dispersal, affect- Africa. Bordering the eastern limits of the Sahel are the Ethio et al. the Congo in Central Africa, help to shape the tropical forests et al. 2010). Possi- ofbelow west-central 1,500 m. TwoAfrica. major Spanning rivers, morethe Niger than in 4,000 the west km, andthe bleing rolesplant ofgene small flow carnivores or ecology in shaping(Herrera ecosystems 1989, Jordano have also Congo forms the second largest river basin in the world, cov- been2007, shownNakashima accidently & Sukor through 2009, Mudappaintroductions. For example, ering nearly 3.7 million km² across seven countries (Reader American Neovison vison introduced to can 1997). South of the Central African lie the miombo woodlands, a broad belt of wooded savannah running west - from into eastern (Le Houérou 2009). The aldcause 2008). a shift There in are similarbreeding examples sites (Nordström of important & Korpimäki ecological Southern African Subregion, south of the Kunene and Zam- impacts2004) and from compete introduced with local species species in several (Harrington families & Macdon of land bezi Rivers, is dominated by the Kalahari and Namib Deserts that contain small to mid-sized species (Roemer et al. 2009). Finally, where larger carnivores are exterminated and northern miombo woodlands) and Namibia. Additional by humans (directly or through habitat change), small carni- ecoregionswhich cover include most of the Botswana Karoo of (excepting South Africa; the Okavangothe bushveld Delta of vores have or may become de facto apex predators in these - et al. 2009), po- bezian and mopane woodlands of southeast Africa. These di- tentially altering their ecological roles and importance in verseeastern ecoregions, Botswana, delimited South Africa by distinct and ; bioclimatic and parameters the Zam suchecosystems systems. (Crooks & Soulé 1999, Roemer such as soil types, climate and vegetation (Le Houérou 2009), In Africa, only some of the diurnal, social small carnivore combined with historical geomorphological changes, climatic Suricata suricatta (see back cov- oscillations, colonisation patterns and in situ evolution have mungo, Common Dwarf Mon- all helped shape Africa’s modern mammal community (Sand- species,goose specifically parvula and Yellow Cynictis peni- cillataer), Banded (see cover),Mongoose have been extensively studied. All others, 16 orders (Kingdon et al - including the widely distributed Common ingers &mammals, Werdelin Africa 2010). is Supportingsecond only 1,161 to the mammal Neotropics species (with in sanguineus (Fig. 1), have received limited attention. 1,282 species) in overall .mammal 2013), nearlyspecies a richness quarter of(Mace all liv et Hence, with few exceptions, the behaviour and ecology, and al. 2005). However, local African mammal communities tend therefore the ecological role, of most African small carnivore to hold more species than their Neotropical counterparts, species remain unknown. The conservation status of all mam- despite similarities in area, latitudinal position, landscapes mals worldwide was assessed for the 2008 IUCN Red List of Threatened Species (Schipper et al. 2008b) and results were et al. 2005), often attributable to differences in abundance summarised for small carnivores globally (Schipper et al. and regional species pools (Vivo & Carmignotto 2004, Nieto et al. 2009). Here, Carmignotto 2004, Nieto et al. 2005). Unfortunately, Africa’s we report on the conservation status, distribution and species mammalianof medium to communities large species currently (Cristoffer face & extreme Peres 2003, threats Vivo from & richness2008a) as of well small as carnivores in the Americas (Herpestidae, (Belant Mustelidae, Nandi- one species in particular, Homo sapiens. Approximately 15% niidae and Viverridae) in Africa.

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Western M. putorius are speculated to be native (Gip- politi 2011, Ahmim 2013). Several mainland species occur on African islands, either naturally or through human transport (Appendix 2). Several non-African species have also been in- troduced to various African islands. These species are not in- cluded in analyses and discussion. As examples, Small Indian Viverricula indica -

Mongoose Herpestes javanicusis found on Unguja, Pemba and Mafia Is 2009).lands (Pakenham There seem 1984, to be Kock no non-native & Stanley small 2009) carnivores and Small estab Asian- lished in mainland Africa. Analyses on Mafia of distributional Island (Kock & and Stanley spe- cies-richness patterns cover only the countries of mainland Africa. Thus, the Macaronesian islands (various countries) and the island nation of Sao Tome and Principe were excluded. None of these archipelagos is believed to support native small Fig. 1. Common Slender Mongoose Herpestes sanguineus (here two carnivores (Appendix 2). These country-based analyses also juveniles in Kruger National Park, South Africa) is one of the most ignore the two European exclaves in mainland Africa, Ceuta widespread and commonly seen in Africa. Yet, little is known and Melilla (Spain; total 30.8 km²); we traced no information about its behavioural ecology (Photo: E. Do Linh San). on which small carnivore species these support. IUCN Red List (see Appendix 1) currently largely follows Wozencraft (2005),The although classification Sokoke for AfricanDog Mongoose small carnivores on omnivora the Methods and Congo Clawless congicus were considered con- Methods to assess the conservation status of the world’s mam- B. crassicauda and Afri- mals through the Global Mammal Assessment in 2008 were can Clawless Otter Aonyx capensis, respectively, by Wozencraft reported by Schipper et al. (2008a, 2008b). Contrary to pre- (2005).specific Thewith many Bushy-tailed points of Mongoose taxonomic uncertainty with Afri- vious mammal IUCN Red List assessments, that in 2008 used can small carnivores mean that the species count presented an expert review process. General information was gathered in this paper (n = 55) is sure to change, perhaps substantially, on distribution, population size and trends, habitat use, ecol- ogy, threats and conservation actions for each species. A digi- morphological analyses. For example, South African Small- tal map of the geographic range of each species was also de- spottedwith further research Genetta involving felina (Fig. both 2) molecularis now widely techniques recognised and veloped in a Geographic Information System. Supporting in- as distinct (e.g. Gaubert et al. formation for most African species was reviewed during the Old World Small Carnivore Red List Assessment workshop in 2013, IUCN 2013) and, following 2004, the latter, 2005, is Jennings here considered & Veron 2009) although not universally G. so genetta (e.g. Delibes. IUCN Red & GaubertList sta- and a preliminary assessment of the IUCN Red List status of tus and population trend information for each species refers to theseCuc Phuong species National was made Park, using Viet theNam, IUCN from Red 3 toList 7 CategoriesJuly 2006, conspecific with - and Criteria version 3.1. The remaining species were assessed cies also occur outside Africa; Appendix 1). Some data used in and reviewed through email correspondence with experts. Fi- thisits global paper status, are freely not availableto its status online specific (IUCN to 2013). Africa (some spe nally, the Red List Authority Coordinators of the IUCN Species Survival Commission (SCC) Small Carnivore Specialist Group (SCSG) and the IUCN SCC Otter Specialist Group (OSG) re- viewed the assessments. The former covers and allies except (Mustelidae except Lutrinae), (Nandiniidae), and allies (Viverridae) and mongooses (Herpestidae); the latter treats otters (Mustelidae: Lutrinae). of the SCSG and OSG, and took the Suez Canal as the eastern boundaryWe defined of Africa. small We carnivores therefore as do species not consider within the Marbled remits Polecat Vormela peregusna, a Eurasian species recently dis- - uony 1998), part of the African fauna. Madagascar was simi- larlycovered excluded; to inhabit no species,the Sinai or Peninsula even , of Egypt of carnivores (Saleh & is Bas na- tive to both Africa and Madagascar (Goodman 2012). Analysis and discussion treats only species native to Africa, including mainland Africa to the north, for which origin in Africa is not clear.(pending Contemporary further clarification) research points two species, to an anthropogenic both confined in in- Fig. 2. South African Small-spotted Genet Genetta (genetta) felina (here troduction for Least Mustela nivalis (Dobson 1998, one radio-collared individual from Great Fish River Reserve, South Africa) Lebarbenchon et al. 2010), but taxonomically close to is now often treated as a species (Photo: E. Do Linh San).

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- man rank correlation; variables considered highly correlated ber of small carnivore species here assessed as occurring nat- (rS > 0.3, P < 0.05) were not included in the same models. As urallyIn in this each study, of the species 48 mainland richness African was defined countries, as 1) and the 2) num spe- suggested by Norušis (2008), in all GzLMs the scale param- cies density, the number of species per 100,000 km² of coun- eter was estimated by dividing the deviance by its degrees of try (see Appendix ES1). Species lists for each country were freedom. The possible effects of independent variables were compiled from several sources (Appendix 1), including exten- evaluated with a Type III test, which does not depend on the sions of known range described in this special issue of Small Carnivore Conservation. Sudan and are treated as a single unit, because most references did not differentiate be- entry order of variables (Norušis 2008). The significance level tween the former Southern Sudan autonomous region (which Resultsfor all analyses was set at α < 0.05. became a country in 2011) and the residual Sudan in terms Under the IUCN Red List’s , four families of small car- of species presence or absence. We used Generalized Linear Models (GzLMs) to test whether species threat level is associ- in Africa, encompassing 23 genera and 55 species. The most ated with the extent of species distribution, expressed as the speciosenivores occur family partly in Africa or entirely is Herpestidae (Nandiniidae; (26 species, monospecific) 47% of number of countries in which each species is indicated to oc- African small carnivore species), followed by Viverridae (17 cur by these sources, or its range area as in the IUCN Red List. species, 31%) and Mustelidae (11 species, 20%). On average, For this purpose, IUCN Red List categories were converted to mainland African countries contain 15 (SD = 5.5) small carni- an ordinal scale according to an increasing threat level (Least vore species (Table 1). Countries of greatest species richness proportionConcern = 1,of Near their Threatened world range = 2,in VulnerableAfrica (see = Appendix 3). The five 1) 15occur species in equatorial (Fig. 3; Appendix Africa (countries ES1). Lowest roughly per-country within 15° species of the wereData Deficientexcluded speciesfrom analyses. and the Athree multinomial species with distribution only a small and richnessEquator) isalthough in North most Africa. sub-Saharan The pattern countries is somewhat hold more different than a cumulative logit function were used to generate GzLMs, and when using each country’s species density as an index. On aver- age, each country contains 13.5 (SD = 24) species per 100,000 (AICc) was used to compare the models (Norušis 2008). Po- km² of its area (Table 1). Figure 3 (right map) shows that the finite sample corrected Aikaike’s Information Criterion were further tested with Mann-Whitney U tests. Similarly, we testedtential whethersignificant species differences richness between is affected threat by level four categories possible several small African countries (The Gambia, Bissau, - moreSierra speciesLeone, ,per unit Togo, area Benin,of country Equatorial than do Guinea, larger, Eritrea, neigh- sity and societal development: country area (km²), number of bouringDjibouti, countries. Rwanda, Burundi,This is because Malawi, the Swaziland, calculated Lesotho) country-spe host- inhabitants/km²,predictors (or covariates) gross national reflecting product country (GNP) size, per human inhabitant den (US$) and gross domestic product (GDP) per inhabitant (US$) per 100,000 km² (i.e. the number of species per 100,000 km² (raw data in Appendix ES1). Negative binomial distributions block,cific species irrespective density of iscountry affected boundaries) by both true and species by country richness size and log link functions were used for the count variable (ab- solute species number), whereas gamma distributions and species density). However, species density is again lower in power functions were used to model the continuous variable North(i.e. the Africa. size ofMost the countriesblock used fall to on derive or very the close country-specific to the indi- (average number of species per 100,000 km² of country). For cated power regression curve (Fig. 4), suggesting that most of the negative binomial distributions, the dispersion param- the variation in this character stems from a basic relationship eter k was set at 0.1. The ratio of the deviance to its degree of species richness increasing with country area, but not in lin- of freedom was close to 1 for the response variable in all the ear proportion. However, two countries (Western Sahara and models, indicating that the variability in observed data was Tunisia) lie noticeably below the curve, indicating that these similar to that predicted by the underlying distributions used countries (both dominated by poorly-vegetated arid habitats) support anomalously few species for their area. Perhaps sur- assessed multicollinearity of predictor variables using Spear- prisingly, no country supports an unexpectedly large number for the models (Norušis 2008). Before conducting GzLMs, we

Table 1. Small carnivore species richness and density in mainland African countries and size, demographic and socio-economic characteristics of those countries. n Average Standard deviation Minimum Maximum Species per country1 48 14.92 5.61 3 26 Species density2 48 13.56 24.06 0.17 134.87 Country area (km²) 48 579,996 584,636 10,380 2,381,740 Number of inhabitants/km² 48 74.22 95.36 2.04 448.26 GNP3/inhabitant (US$) 473 969 1,308 66 5,398 GDP3/inhabitant (US$) 473 2,729 4,906 231 29,332 1The two countries Sudan and South Sudan are treated as one unit; countries with only island and/or exclave in Africa are omitted (see text). 2Number of species per 100,000 km² of a country. 3GNP = Gross National Product, GDP = Gross Domestic Product; no data were available for Western Sahara. Data country by country are provided in Appendix ES1.

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the covariates (P > 0.13) affected country species richness. In

in country species density than did the intercept-only model (Omnibuscontrast, several Test, P fitted models explained better the variation that ‘Country area’ (B = –2.071 × 10–7 df = 1, P < 0.001) and ≤ 0.014; ‘GNP/inhabitant’ Table 2). Univariate (B models showed 23.048, df = 1, P < 0.001) were negatively,, Wald and χ² the = 141.578,‘Number of inhabitants/km², positively (B = –0.003, Walddf χ² = 1,= P = 0.013), associated with country species richness. However, the best model incorporated ‘Country = 5.025, area’ Wald ( Bχ² = = –4.339 6.112, × 10–7, df = 1, P < 0.001) and ‘GNP/inhabitant’ (B = –5.956 × 10–5 df = 1, P = 0.024) (Table 2). WaldOf χ² the= 128.434, 55 small carnivore species native to Africa, 51 (93%) are predominantly, Wald χ² = 5.114,African and 48 (87%) are endemic (Appendix 1). Ratel Mellivora capensis is the only species with large proportions of its range both in and outside Africa. In terms of conservation status, the IUCN Red List indicates that 43 species (78%) are Least Concern (LC), three (6%) are Near

listedThreatened as Endangered (NT), four (EN), (7%) Critically are Vulnerable Endangered (VU) and (CR), five Extinct (9%) inare the Data Wild Deficient (EW) or (DD). Extinct No African(EX). small carnivore species are All four species considered globally threatened (all VU; Fig. 3. Country-based species richness of small carnivores in continental Sokoke Mongoose, Liberiictis kuhni, Africa based on the IUCN Red List and overview and location of mainland Crested Genet Genetta cristata G. john- African countries (countries with only island territory in Africa were not stoni) were listed under the A Criterion (population decline) included in the comparison; and Sudan and South Sudan were treated as and Johnston’s Genet - one unit). Left: species richness per country; right: species density, i.e. graphic range size), C Criterion (population size and decline), number of species per 100,000 km² of country. D(Appendix Criterion (very 1). None small was or restricted listed using population) the B Criterion or E Criterion (geo - son’s Mongoose Bdeogale jacksoni G. bourloni of species for its area. The results of the GzLM procedure in- and(quantitative Eurasian analysis).Otter Similarly, lutra) were the three listed NT using species the A(Jack Cri- dicated that the intercept-only model had a greater explana- terion (Appendix 1). All threatened, Bourlon’s and two Genet of the three NT tory power (Omnibus Test, P small carnivore species are endemic to Africa, while Eurasian including different combinations of predictors; hence, none of Otter is widespread across Eurasia (Appendix 1). While for > 0.13) than the fitted models

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Table 2. Results of the GzLM procedures (Omnibus tests) testing the potential effects of country size (km²) and demographic and socio- economic characteristics on country-specific small carnivore species density, i.e. species richness per 100,000 km² of a country’s area. Variables in the alternative GzLMs Power Scale LR χ² df P AICc Country area, GNP/inhabitant 0.20 0.988 55.356 2 <0.001 292.714 Country area, GDP/inhabitant 0.14 1.000 54.178 2 <0.001 293.235 Country area 0.10 0.975 57.937 1 <0.001 294.046 Number of inhabitants/km² 2.20 1.549 19.426 1 <0.001 320.440 GNP/inhabitant 1.00 1.922 6.074 1 0.014 333.455 GDP/inhabitant 0.20 2.176 0.131 1 0.717 344.845 GzLM = Generalized Linear Model, LR = Likelihood Ratio, AICc = finite sample corrected Akaike’s Information Criterion, GNP = Gross National Product (US$), GDP = Gross Domestic Product (US$). Only models incorporating uncorrelated predictors were considered.

Fig. 4. The relation of country-based species density of small carnivores Fig. 5. in continental Africa with country area. Line represents a fitted power Threat levels of small carnivore species by family in Africa based IUCN Red List function regression curve. on the . The family Nandiniidae, monospecific (LC), is not displayed but has been taken into account in the overall evaluation (category ‘Combined’). Note that Near Threatened is a category of ‘not threatened’, not of ‘globally threatened’. threatened (Fig. 5), the exact threat level is between 7% (if no DDdata-sufficient species is threatened) small carnivore and 16% species, (if all 8% are). were considered As could be expected, the range area and the number of countries in which each African small carnivore species oc- cur are highly positively correlated (Spearman rank correla- being so classified (Fig. 5). The geographic distribution of DD species includes equatorial African countries (Côte d’Ivoire, tion, rS = 0.915, P < 0.001). The GzLM procedure indicated that African Republic, Uganda and Democratic Republic of Congo), ‘Range area’ is negatively associated with an increase in small Liberia, Ghana, Equatorial Guinea, Republic of Congo, Central- carnivore threat level (B = –9.36 × 10–6 df = cies, the percentage of threatened species increases slightly to 1, P < 0.001, AICc = 26.708). A model integrating ‘Number of 14%as well for as Viverridae, Angola and while Sudan. remaining Treating 8% only for data-sufficient Herpestidae. spe countries’ even had a slightly better explanatory, Wald χ² = power,17.727, with Overall, population trends for 46% (n = 25) of small car- an increase in the number of countries being linked to a de- nivore species in Africa are assessed as unknown globally, crease in species threat level (B including 27% of species of Mustelidae, 35% of Herpestidae df = 1, P < 0.001, AICc = 23.721). Vulnerable and Near Threat- and 71% of Viverridae (Fig. 6). Of the 30 species with assessed ened small carnivores have typically = –0.383, more Wald restricted χ² = 17.258,ranges population trends, 50% (27% of species overall) are believed than Least Concern species (Table 3). Overall, Least Concern to be stable and 50% (27%) to be decreasing; none is thought species occur in more countries (Mann-Whitney U test, n1 = to be increasing.

43, n2 = 7, U = 59, P = 0.009) and have larger geographic ranges

(n1 = 41, n2 = 6, U = 7, P < 0.001) than do Vulnerable and Near Discussion Small carnivores in Africa represent 34% of the extant small even more restricted distribution ranges than do threatened carnivores worldwide (n = 163 species; Schipper et al. 2008a). speciesThreatened (Table species 3). combined. Data Deficient species possess Familial richness is intermediate, with four of the world’s The percentage of species considered globally threatened nine families represented (, , , varies across families, from 0% (Mustelidae and Nandiniidae) and Prionodontidae are all extralimital). Coun- to about 8% in Herpestidae and 12% in Viverridae (Fig. 5). try species richness was not affected by country size, human There are no DD species in the family Mustelidae, in contrast population density or by coarse-scale socio-economic indices. to about 8% of species of Herpestidae and 18% of Viverridae

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Table 3. Distribution of small carnivores in Africa, compared based on their IUCN Red List category. Red List category Number of Number of mainland countries per species species1 in Mean Standard deviation Minimum Maximum Africa LC 43 15.72 14.40 1 44 NT 3 3.33 0.58 3 4 VU 4 3.75 1.71 2 6 DD 5 2.80 1.64 1 5 Total 55 13.02 13.76 1 44 Red List category Number of Geographic range area per species (km²) species2 in Mean Standard deviation Minimum Maximum Africa LC 41 5,785,851 6,614,627 43,777 28,885,834 NT 2 105,408 58,626 63,953 146,863 VU 4 129,949 121,033 34,426 306,732 DD 5 298,473 236,848 49,336 602,126 Total 52 4,604,671 6,294,881 34,426 28,885,834 LC = Least Concern, NT = Near Threatened, VU = Vulnerable, DD = Data Deficient. 1Comparison considers all species of small carnivores occurring in Africa. 2Comparison omits the three species in Africa (Mustela nivalis, ‘M. putorius’ and Lutra lutra) with ranges predominantly in other continents.

ised’ small carnivore species does not increase in proportion. These results support the notions that although some habi- tat size threshold might be essential to ensure population vi-

that the larger an area the more species generally it will hold (Ruggieroability (e.g. et Crawley al. 1994), & areaHaral per 2001, se is Brito not the & Grelle only variable 2006) and for explaining local patterns of mammalian species richness. For conservation purposes, other factors such as habitat diversity and species interactions should be taken into account when assessing species richness and diversity at the landscape scale

associated with country species richness. This could suggest that( an& increaseFox 2000). in potential Human population human population density pressurewas positively might not necessarily be detrimental to small carnivores, at least for generalist species that are probably less sensitive to habitat Fig. 6. Global population trends of small carnivore species by family in change. However, it could as well indicate that areas produc- Africa based on the IUCN Red List. The family Nandiniidae, monospecific tive for people, and thus supporting higher human population (unknown population trend), is not displayed but has been taken into densities, are also inherently rich in small carnivore species. If account in the overall evaluation (category ‘Combined’). indeed so, this might mean that species-rich small carnivore

with, humans. This situation would add to conservation chal- lenges.communities Precise are data more on likely small to carnivore be threatened species by, compositions or in conflict in Africa seems to follow the general pattern of overall land and densities in human-populated areas would be needed for mammal species richness (Schipper et al. 2008b) and of et al. IUCN Red 2009), peaking in the tropics and therefore in areas of high Listfirm, smallconclusions. carnivore species in Africa appear more secure than ecologicalsmall carnivore and possibly species richnesstopographic in the complexity. Americas (BelantSmall carni - smallBased carnivores on extinction or mammals risk globally. as measured However, by thethis compari- vore species richness generally declines with increasing lati- et al. 2009). In Africa, small carnivore richness contain some degree of uncertainty and the particularly low is indeed lower in the north, but not in the south, suggesting levelsson requires of knowledge a caveat. for Assessments many African of species species mean status that always they thattude the(Belant low productivity of the vast and arid Sahara might are at elevated risk of incorrect assessment. That said, ‘only’ explain the observed difference. That country species density was markedly greater in smaller countries throughout Africa as globally threatened, compared with 20% in the Ameri- indicates that as country size increases, the number of ‘new’ 8% of data-sufficientet al. 2009) African and 22% small worldwide carnivores (Schipper were assessed et al. habitats favourable to host additional, possibly more ‘special- 2008a). Overall, 25% of all mammals worldwide are consid- cas (Belant

Small Carnivore Conservation 10

, Vol. 48, July 2013 Status of small carnivores in Africa ered globally threatened (Schipper et al. 2008b). In Africa, the 20% over the 20 years prior to 2008 based on estimates of four globally threatened (in this case Vulnerable) and two of ongoing forest loss (although not as severe in Liberia, the core the Near Threatened species were listed as such by the IUCN Red List based on an estimated population decline. Most Afri- can small carnivores have distribution ranges that well exceed (IUCNof the 2013).species’s All range,Vulnerable as elsewhere and Near inThreatened Upper Guinea; small carniPapeş- vore& Gaubert species 2007), are in coupled need of furtherwith the survey likely work impacts to clarify of hunting their increase in threat levels is associated with a decrease in range conservation status (distribution and population density/ the thresholds for listing under the B criterion. However, an trends) in the wild, and also, when relevant, to determine sus- ranges and considering the likely threats to such species (see tainable levels of offtake from the wild and general manage- below),areas (see the above).comparatively All five low DD speciesthreat levels have to extremely African small ment and conservation measures. carnivores provided above should be interpreted with cau- The percentage of small carnivores in Africa with inad-

9%) is similar to that of small carnivores in the Americas tion. Threats In addition, to the even globally some threatened LC species African would benefitsmall carni from- equate data toet assess al. 2009) conservation and worldwide status (9%;(i.e. DataSchipper Deficient; et al. voresclarification vary between of conservation species. Sokokestatus. Dog Mongoose, restricted 2008a), and slightly lower than the percentage of such land to coastal forests of and Tanzania, is believed to have mammals(11%; Belant overall (15%; Schipper et al. 2008b). Categorising declined substantially through impacts of extensive, ongo- - ing habitat loss related to illegal logging (Taylor 2013). In the tion is available to evaluate ongoing threats and/or there is Shimba Hills National Reserve (Kenya), the resident popula- seriousa species doubt as Data that Deficientspecies rank means is taxonomically that insufficient appropriate. informa tion was, and might still be, under potential threat from af- Taxonomic uncertainty does not drive the DD listing for any of forestation with non-native Pinus together with regular burning of the undergrowth to favour Antelope Hippotra- Mongoose Dologale dybowskii is perhaps the least known Afri- gus niger grazing canthe fivesmall African carnivore: small known carnivores from soonly categorised. 31 specimens, Pousargues’s it has not agriculture, logging(Engel and mining& Van Rompaey within their 1995). Upper In West Guinea Africa, for- 2013; but see Aebischer et al Genetta estsboth ranges,Liberian and Mongoose are hunted and (mostly Johnston’s for Genetmeat andlose skin)habitat with to piscivorabeen conclusively from Democratic recorded inRepublic several decadesof Congo (Stuart is also & poorlyStuart known, rarely observed, and .taken 2013). as Aquatic Genet (Van Rom- 2013). The lack of den sites in secondary forests might restrict Liberiandogs, shotguns Mongoose and distribution, snares (Dunham while & in Gaubert forest 2013,plantations IUCN criteria A or C. The taxonomically recently resurrected King this species might also suffer from pesticide use, because the Genetpaey & G. Colyn poensis 2013b); it may possibly warrant listing under- worms it forages on accumulate toxins to levels threatening to ria to Republic of Congo and although not reliably recorded for has a disjunct distribution in forest from Libe Genet, endemic to Nigeria and Cameroon, and perhaps the Re- so it may well be more common than it seems (Gaubert 2003, mammalian predators (Taylor & Dunham 2013). For Crested 2013).over 50 West years, African this likely Oyan reflects confusion leightoni haswith a othervery narrowgenets, et al. 2006), be- - causepublic the of Congonon-protected and Cross (Hunter River State & Barrett forests 2011), (Nigeria) habitat are lyn 2013c), and may have a status akin to that of other co-oc- graduallyloss is probably being alsoconverted a major into threat farms (Gaubert or wastelands and the distribution in the upper Guinean forests (Van Rompaey & Co Genet (VU) and Liberian Mongoose (VU). Finally, Ansorge’s Luiselli 2005). It probably also suffers from high hunting pres- Cusimansecurring species with similar ansorgei narrow from ranges, Central such Africa as Johnston’s is poorly Niger Delta is exploited as an oil-production area (Angelici & Among the Near Threatened species, little is known as Least Concern with further information (IUCN 2013). Only aboutsure (Van threats Rompaey to Eurasian & Colyn Otter 2013a). in its limited African range (Al- furtherknown (Vanresearch Rompaey and survey & Colyn work 2013e), can clarifybut likely the topopulation be listed geria, Morocco and Tunisia). These populations have shown - little sign of recovery, unlike those in parts of Europe. In Mo- onestatus, region trends where and investigation threats of these and researchspecies. Theseefforts Data should Defi be especially in the north, where otters have apparently disap- concentrated.cient species occur primarily in equatorial Africa, making this pearedrocco, pollution from rivers has in increased the lowland dramatically plains, and in dam the major building rivers, has Finally, populations of mainland species on islands might also reduced habitat and fragmented populations (Delibes et be worth investigating in further detail, especially endemic al is thought to have declined by 20–25% over the 15 years preceding the IUCN Winther-Hansen (2003) mentioned three such endemic sub- Red. 2012, List assessment Kruuk 2013). (IUCN Jackson’s 2013). Mongoose Its probable dependence on species:subspecies Servaline perhaps Genet under Genetta threat. servalina In , archeri Goldman, Common & forest means its main threat is likely to be ongoing clearance Slender Mongoose Herpestes sanguineus rufescens at the restricted number of sites in Uganda, Kenya and Tanza- tailed Mongoose Bdeogale crassicauda tenuis. The overall assessed conservation status of African and Bushy- small such forests is crucial, and other East African groundwater- carnivores is relatively favourable compared with mammals nia it occupies (Van Rompaey & Kingdon 2013). Protection of worldwide. However, the four Vulnerable and three Near - stricteddependent to foreststhe Upper should Guinean be surveyed rainforests for Jackson’s (see countries Mongoose in their persistence, yet for none does there seem to be a con- Appendix(De Luca &ES1) Rovero and is2006). believed Bourlon’s to have Genet declined is essentially by more than re servationThreatened programme species warrant in place specific to remove interventions and/or mitigate to ensure the

11 Small Carnivore Conservation

, Vol. 48, July 2013 Do Linh San et al. factors threatening it. Effective action for these species is the M. (eds) Mammals of Africa, V. Carnivores, pangolins, equids and short-term priority for small carnivore conservation in Africa. rhinoceroses In future the threat levels of African small carnivores are likely - lationships between. Bloomsbury, two genets, , Genetta U.K. maculata and Genetta Angelici,cristata F. M.. Revue & Luiselli, d’Écologie L. 2005. (Terre Habitat et Vie) associations 60: 341–354. and dietary re withto worsen. Africa Athaving least thea quarter highest are human assessed population by IUCN growth (2013) rate as ofalready any continent in decline. (UN All 2011),major threats existing originate threats willfrom surely people, inten and- sify and probably diversify. The high demand for wildlife meat Bahaa-el-din,Rabanal, L., Starkey,Henschel, M., P., Taylor, Aba’a, G.,R., Vanthomme,Abernethy, K., A., Bohm, Nakashima, T., Bout, Y. and other products in East Asia is not abating and already ani- N., Coad, L., Head, J., Inoue, E., Lahm, S., Lee, M. E., Maisels, F., mals much declined in Asia are sourced from Africa to meet carnivores in Gabon. Small Carnivore Conservation 48: 19–29. & Hunter, L. 2013. Notes on the distribution and status of small et al. 2004) and, as their South- small carnivores in the Americas. Small Carnivore Conservation this demand (Bennett 2011). Small carnivores are a large part Belant,41: J. 3–8. L., Schipper, J. & Conroy, J. 2009. The conservation status of fromof Asian Africa wildlife to East trade Asia. (e.g. Simultaneously, Bell continued high lev- - elseast of Asian evergreen populations forest declineconversion they and are likelyfragmentation to join the are trade re- navirus: possible links with the international trade in small car- ducing habitat block size (Newmark 1998), meaning that the Bell, nivores.D., Roberton, Philosophical S. & Hunter, Transactions P. R. 2004. Animalof the Royal origins Society, of SARS B coro359: species of those habitats will become increasingly susceptible 1107–1114. to hunting even if levels remain constant. Altogether, this sug- - gests that additional African small carnivore species will meet forcement systems to save charismatic species. Oryx 45: 476– globally threatened criteria if no effective conservation strate- Bennett,479. E. L. 2011. Another inconvenient truth: the failure of en gies are implemented, in particular to combat wildlife meat Game ranch management, trade and illegal logging. Considering the paucity of informa- 5th edn. Van Schaik, Pretoria, South Africa. tion available on this fascinating group of species and the high Bothma, J. du P. & du Toit, J. G. (eds) 2010. level of endemism of small carnivores in Africa, both research and conservation will be of prime importance in the future. Brito,( BrachytelesD. & Grelle, hypoxanthusC. E. 2006. Estimating). minimum and Conservation area of suitable 15: 4197–4210.habitat and viable population size for the Northern Muriqui Acknowledgements The 2008 IUCN Red List assessment was made possible by the efforts - of more than 1,700 experts from 130 countries who assessed the Bronner,list of G. theN., Hoffmann,extant mammals M., Taylor, of the P. J.,southern Chimimba, African C. T., subregion.Best, P. B., conservation status of all mammal species worldwide (see Schipper DurbanMatthee, Museum C. A. & Robinson, Novitates T.28: J. 56–106.2003. A revised systematic check et al. 2008b); many thanks to these individuals and the organisations - Cardillo,of high M., Mace,extinction G. M., risk Jones, in K.large E., Bielby,mammal J., Bininda-Emonds, species. Science 309:O. R. who supported their efforts. Specifically, we thank Keith and Col 1239–1241.P., Sechrest, W., Orme, C. D. L. & Purvis, A. 2005. Multiple causes Kingdon,leen Begg, David Paolo Macdonald, Cavallini, João Robbie Crawford-Cabral, McDonald, Francesco Scott Creel, Palomares, Fabrice - Cuzin, Koenraad De Smet, Amy Dunham, Corey Goldman, Jonathan sity. Science 291: 864–868. Stuart, Mark Taylor, Peter Taylor and the late Harry Van Rompaey for Crawley, M. J. & Harral, J. E. 2001. Scale dependence in plant biodiver- providingGalen Rathbun, valuable Justina information Ray, Claudio to compile Sillero-Zubiri, the accounts Chris ofand the Mathilde African logical role of large herbivores in the evolutionary history of two small carnivores, and assisting with assessments. We also acknowl- Cristoffer,tropical C. & worlds. Peres, C.Journal A. 2003. of Biogeography Elephants versus 30: 1357–1380.butterflies: the eco

African carnivore assessments. We are grateful to Federica Chiozza for extinctions in a fragmented system. 400: 563–566. generouslyedge the role providing of J. W. Duckworth us the information and S. A. aboutHussain the as geographic reviewers ofrange the Crooks, K. R. & Soulé, M. E. 1999. Meso-predator release and avifaunal- Bdeogale jacksoni (Herpestidae). kindly provided last-minute information on the small carnivore spe- De Luca,Oryx D. 40: W. 468–471. & Rovero, F. 2006. First records in Tanzania of the Vul ciessizes present of the world’s in Lesotho small and carnivores. Swaziland, Nico respectively. Avenant and Frank Ara Monadjem Hawkins, nerable Jackson’s Mongoose Genetta genetta. Pp. - Mammals of Africa, knowledged for their comments on an earlier draft of this paper. Delibes,V. Carnivores, M. & Gaubert, pangolins, P. 2013. equids Common and Genetrhinoceroses Arno Gutleb, Nicole Duplaix and J. W. Duckworth are gratefully ac London,224–229 U.K. in Kingdon, J. & Hoffmann, M. (eds) References . Bloomsbury,

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Africa, V. Carnivores, pangolins, equids and rhinoceroses - Conservation International, 2011 Crystal Drive, bury, London, U.K. Arlington, VA 22202, U.S.A. . Blooms Email: [email protected] genet (Genetta genetta L.) diet: a literature review. Mammal Re- 6IUCN Species Survival Commission, c/o United Nations Virgós,view E., 29: Llorente, 119–128. M. & Cortés, Y. 1999. Geographical variation in Environment Programme - World Conservation Monitoring Centre, 219 Huntingdon Rd, Cambridge, CB3 the mammal faunas of South America and Africa. Journal of Bio- 0DL, U.K. Vivo,geography M. & Carmignotto, 31: 943–957. A. P. 2004. Holocene vegetation change and Email: [email protected] Wozencraft, W. C. 2005. Order Carnivora. Pp. 532–628 in Wilson, 7Muséum National d’Histoire Naturelle, UMR BOREA IRD Mammalian species of the world, 207, 43 rue Cuvier, 75005 Paris, . Email: [email protected] U.S.A.D. E. & Reeder, D. M. (eds) 8Italian Foundation of Vertebrate Zoology (FIZV), Via 3rd edn. John Hopkins University Press, Baltimore, Maryland, Cleonia 30, Scala C, I-00152 Roma, . 1Department of Zoology and Entomology, Email: [email protected] University of Fort Hare, Private Bag X1314, Alice, 5700, 9Centre for Wildlife Management, University of Pretoria, South Africa. Pretoria, 0002, South Africa. Email: [email protected] Email: [email protected] 2Department of Biological Sciences, 10Centre for Invasion Biology, University of Pretoria, Texas Tech University, Box 43131, Lubbock, Pretoria, 0002, South Africa. Texas, U.S.A. Email: [email protected] Additional material related to this paper* can be found as a 3Carnivore Ecology Laboratory, Forest and Wildlife Research Center, Mississippi State University, Box 9690, following link: http://www.smallcarnivoreconservation.org/ Mississippi State, Mississippi, U.S.A. PDF file (SCC48_Appendix-ES1.pdf) on the SCC website in the Email: [email protected] 4IUCN Species Programme, IUCN, 28 Rue Mauverney, home/journal/SCC48_Appendix-ES1. 1196 Gland, Switzerland. *Appendix ES1. Raw data for each African country used in 5IUCN/SSC-CI/CABS Biodiversity Assessment Unit, analyses including individual lists of species reported in each c/o Center for Applied Biodiversity Science, country.

Appendix 1. Taxonomic affiliation, conservation status, population trends and distribution of small carnivores in Africa. Taxon 1 English name IUCN Red List Population Number of Distribution ­categorisation2 trend countries3 Family Mustelidae Mustela nivalis4 Least (Common) Weasel LC Stable 2 Eurasia, North Ameri- ca, North Africa Mustela subpalmata (M. Egyptian Weasel LC Stable 1 Africa nivalis subpalmata) Mustela putorius4 Western (European) Polecat (Ferret?) LC Decreasing 2 Europe, North Africa libycus5 (Poecilictis Libyan (Libyan Striped, Saharan LC Unknown 15 Africa libyca) Striped, North A­frican Striped) Wea- sel (Saharan ) Ictonyx striatus Zorilla (Striped Polecat, African Pole- LC Stable 39 Africa ) Poecilogale albinucha African Striped (Striped, African, LC Unknown 17 Africa White-naped, ) Weasel Aonyx capensis African (Cape) Clawless Otter LC Stable 36 Africa Aonyx congicus (A. capensis Congo (Cameroon, Small-toothed) LC Unknown 9 Africa congicus) Clawless (Swamp) Otter Lutra lutra Eurasian (Common) Otter NT (A2cd) Decreasing 3 Eurasia, North Africa Lutra maculicollis (Hydrictis Spotted-necked (Spot-necked, Speck- LC Decreasing 34 Africa maculicollis) le-throated) Otter Mellivora capensis Honey (Ratel) LC Decreasing 44 Africa, Arabia, South Asia

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Taxon 1 English name IUCN Red List Population Number of Distribution ­categorisation2 trend countries3 Family Nandiniidae Nandinia binotata African (Two-spotted) Palm (Tree) LC Unknown 28 Africa Civet

Family Viverridae Genetta abyssinica Abyssinian (Ethiopian) Genet LC Unknown 5 Africa Genetta angolensis Angolan (Miombo) Genet LC Unknown 6 Africa Genetta bourloni Bourlon’s Genet NT (A2cd) Decreasing 4 Africa Genetta cristata Crested (Crested Servaline) Genet VU (A2cd) Decreasing 4 Africa Genetta genetta6 Common (Small-spotted) Genet LC Stable 37 Africa, South-west Europe, Arabia Genetta johnstoni Johnston’s Genet VU (A2cd) Decreasing 6 Africa Genetta maculata Rusty-spotted (Blotched, Central LC Unknown 31 Africa African Large-spotted, Large-spotted) Genet Genetta pardina Pardine (West African Large-spotted) LC Unknown 11 Africa Genet Genetta piscivora DD Unknown 1 Africa Genetta poensis King Genet DD Unknown 5 Africa Genetta servalina Servaline Genet LC Unknown 11 Africa Genetta thierryi Hausa (Haussa) Genet LC Unknown 13 Africa Genetta tigrina Cape (South African Large-spotted) LC Unknown 2 Africa Genet Genetta victoriae Giant (Giant Forest) Genet LC Unknown 3 Africa Poiana leightoni Leighton’s (West African) Oyan DD Decreasing 2 Africa (Leighton’s Linsang, West African Linsang) Poiana richardsonii Central African Oyan (African, Central LC Unknown 6 Africa African Linsang) Civettictis civetta LC Unknown 37 Africa

Family Herpestidae Atilax paludinosus Marsh (Water) Mongoose LC Decreasing 37 Africa Herpestes naso (Xenogale Long-nosed (Long-snouted) LC Decreasing 10 Africa naso) ­Mongoose Herpestes flavescens Kaokoveld Slender (Angolan Slender, LC Stable 2 Africa ( flavescens) , Larger Red) Mongoose Herpestes ichneumon Egyptian (Large Grey) Mongoose LC Stable 44 Africa, South-west ­(Ichneumon) Europe, Middle East Herpestes ochraceus Somali (Somalian) Slender Mongoose LC Unknown 3 Africa (Galerella ochracea) Herpestes pulverulentus Cape (Small) Grey Mongoose LC Stable 3 Africa (Galerella pulverulenta) Herpestes sanguineus Common Slender (Slender) LC Stable 39 Africa (Galerella sanguinea) ­Mongoose Bdeogale crassicauda Bushy-tailed Mongoose LC Unknown 7 Africa Bdeogale jacksoni Jackson’s Mongoose NT (A2cd) Decreasing 3 Africa Bdeogale nigripes Black-footed (Black-legged) Mon- LC Decreasing 6 Africa goose Bdeogale omnivora (B. Sokoke Dog (Sokoke Bushy-tailed) VU (A2c) Decreasing 2 Africa crassicauda omnivora) Mongoose Rhynchogale melleri Meller’s Mongoose LC Unknown 8 Africa Cynictis penicillata LC Stable 6 Africa Paracynictis selousi Selous’s Mongoose LC Unknown 8 Africa Ichneumia albicauda White-tailed Mongoose LC Stable 33 Africa, Arabia

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Taxon 1 English name IUCN Red List Population Number of Distribution ­categorisation2 trend countries3 Suricata suricatta Meerkat (Suricate, Slender-tailed LC Unknown 4 Africa Meerkat, Grey Meerkat) Mungos gambianus LC Stable 10 Africa Mungos mungo LC Stable 33 Africa Liberiictis kuhni Liberian Mongoose VU (A2cd) Decreasing 3 Africa Dologale dybowskii Pousargues’s (Savannah) Mongoose DD Unknown 4 Africa Helogale hirtula Somali (Ethiopian, Desert) Dwarf LC Stable 3 Africa Mongoose Helogale parvula Common Dwarf (Dwarf) Mongoose LC Stable 15 Africa Crossarchus alexandri Alexander’s Cusimanse LC Decreasing 4 Africa Crossarchus ansorgei Ansorge’s (Angolan) Cusimanse (An- DD Unknown 2 Africa golan Mongoose) Crossarchus obscurus Common (Long-nosed) Cusimanse LC Unknown 5 Africa Crossarchus platycephalus Cameroon (Flat-headed) Cusimanse LC Unknown 7 Africa (C. obscurus platycephalus) 1Genus and species limits and spellings follow IUCN (2013), itself based on Wozencraft (2005), selected to be a readily available, widely used, source. Some of the more widely-used alternative taxonomic treatments and English names are given, but listings are far from comprehensive. Notably, genet taxonomy has been particularly unstable recently. 2DD = Data Deficient, LC = Least Concern, NT = Near Threatened, VU = Vulnerable; A2 = Population reduction observed, estimated, inferred, or suspected in the past where the causes of reduction may not have ceased or may not be understood or may not be reversible, c = assessment for category ‘A2’ based on a decline in area of occupancy, extent of occurrence and/or habitat quality, d = assessment for category ‘A2’ based on actual or potential levels of exploitation (IUCN 2012). 3Refers to the number of mainland African countries in which each species is here taken to occur, based on Bronner et al. (2003), Saleh & Basuony (2005), Wozencraft (2005), Dinets (2011), Ahmim (2013), N. Avenant (verbally 2013), Bahaa-el-din et al. (2013), IUCN (2013), Kingdon et al. (2013), A. Monadjem (verbally 2013) and Pacheco et al. (2013). To some extent these sources provide generalised distributions; they are not restricted to verifiable records. So the pattern analyses undertaken here are based upon the plausible inferred distribution of each species, rather than specific validated records of each species in each country. 4The origins of Mustela nivalis and animals identified as M. putorius in North Africa remain unresolved (see text). 5Ictonyx is a masculine genus so this species’s name is thus correctly I. libycus, not I. libyca. 6South African Small-spotted Genet is sometimes given species rank (see text).

Appendix 2. Endemic African small carnivores present on African islands1. Island name Country Taxon 2 English name2 Remark(s) References Bioko Equatorial Guinea Lutra maculicollis Spotted-necked Otter Former resident; now Harrington et al. (2002), extirpated; contro- D’Inzillo Carranza & versial3 ­Rowe-Rowe (2013) Nandinia binotata African Palm Civet Possibly present; his- Harrington et al. (2002), Van torically rare Rompaey & Ray (2013) Genetta maculata Rusty-spotted Genet Possibly present Harrington et al. (2002), ­Angelici & Gaubert (2013) Genetta poensis King Genet Possibly present Harrington et al. (2002), Gaubert (2003, 2013) Poiana richardsonii Central African Oyan Present Harrington et al. (2002), Van Rompaey & Colyn (2013d) Sao Tome Island4 Sao Tome and Civettictis civetta African Civet Purportedly intro- Dutton (1994) Principe duced Pemba Tanzania Atilax paludinosus Absent from Unguja Pakenham (1984) Unguja (Zanzibar) Tanzania Nandinia binotata African Palm Civet Present Perkin (2004, 2005) Genetta servalina Servaline Genet Present Van Rompaey & Colyn (1998), archeri Goldman & Winther-Hansen (2003) Civettictis civetta African Civet Present Pakenham (1984), Stuart & Stuart (1998) Herpestes sanguineus Common Slender Present Pakenham (1984), Stuart & rufescens ­Mongoose Stuart (1998), Goldman & Winther-Hansen (2003)

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Island name Country Taxon 2 English name2 Remark(s) References Bdeogale crassicauda Bushy-tailed Mongoose Present Pakenham (1984), Stuart & tenuis Stuart (1998), Goldman & Winther-Hansen (2003) Mungos mungo Banded Mongoose Purportedly intro- Pakenham (1984), Stuart & duced; no recent Stuart (1998), Goldman & records Winther-Hansen (2003) 1The Macaronesian islands (the Azores, Madeira, Savage, Canary and Cape Verde archipelagos) contain no native species of small carnivore nor any introduced African endemics, although Mustela nivalis and Western Polecat M. putorius and/or Domestic Ferret M. furo have been introduced; past reports of Common Slender Mongoose Herpestes sanguineus on the Cape Verde archipelago are in error (Masseti 2010, Hazevoet & Masseti 2011). 2Alternative taxonomic treatments and names are given in Appendix 1. 3Extensively treated in the past as a subspecies of Aonyx congicus, A. c. poensis; now considered synonymous with Lutra maculicollis (d’Inzillo Car- ranza & Rowe-Rowe 2013). 4Also supports Mustela nivalis, purportedly introduced (Dutton 1994, Sheffield & King 1994).

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