Individual Identification and Genetic Variation of Lions (Panthera leo) from Two Protected Areas in Nigeria

This survey was conducted in two protected areas in Nigeria to genetically identify individual lions and to determine the genetic variation within and between the populations. We used faecal sample DNA, a non-invasive alternative to the risky and laborious task of taking samples directly from the animals, often preceded by catching and immobilization. Data collection in Yankari Game Reserve (YGR) spanned through a period of five years (2008 –2012), whereas data in Kainji Lake National Park (KLNP) was gathered for a period of three years (2009, 2010 and 2012). We identified a minimum of eight individuals (2 males, 3 females, 3 unknown) from YGR and a minimum of ten individuals (7 males, 3 females) from KLNP. The two populations were found to be genetically distinct as shown by the relatively high fixation index (F_ST  = 0.17) with each population exhibiting signs of inbreeding (YGR F_IS  = 0.49, KLNP F_IS  = 0.38). The genetic differentiation between the Yankari and Kainji lions is assumed to result from large spatial geographic distance and physical barriers reducing gene flow between these two remaining wild lion populations in Nigeria. To mitigate the probable inbreeding depression in the lion populations within Nigeria it might be important to transfer lions between parks or reserves or to reintroduce lions from the zoos back to the wild.     

Threat of rapid extermination of the lion (Panthera leo leo) in Waza National Park,Northern Cameroon

Lion populations in West and Central Africa are small and fragmented. In areas where park management is weak, threats will likely facilitate the extinction of the lion. Wildlife management requires knowledge of the population estimate. The population of lions in Waza National Park (Waza NP) was assessed by individual identification of members in the population. The population was assessed to comprise of 14–21 adult individual lions. The age structure was skewed towards adults; cubs comprised 22% of all lions identified while the sex ratio was 1 : 3. Two out of four collared lions were lost to illegal, retaliatory killings within 1 year; and probably two more males and one more female were also killed during this period. The lion population appears to have declined during the last 5 years with six lions dying per year, which is at a much higher rate than observed in the previous decades. Human‐livestock pressure has increased tremendously in this period, resulting to frequent human‐lion conflicts. To ensure the survival of the lion in Waza NP and in the entire region, management needs to intensify efforts to mitigate the pressure from humans and their livestock.     

A lion lentivirus related to feline immunodeficiency virus: epidemiologic and phylogenetic aspects.

Feline immunodeficiency virus (FIV) is a novel lentivirus that is genetically homologous and functionally analogous to the human AIDS viruses, human immunodeficiency virus types 1 and 2. FIV causes immunosuppression in domestic cats by destroying the CD4 T-lymphocyte subsets in infected hosts. A serological survey of over 400 free-ranging African and Asian lions (Panthera leo) for antibodies to FIV revealed endemic lentivirus prevalence with an incidence of seropositivity as high as 90%. A lion lentivirus (FIV-Ple) was isolated by infection of lion lymphocytes in vitro. Seroconversion was documented in two Serengeti lions, and discordance of mother-cub serological status argues against maternal transmission (in favor of horizontal spread) as a major route of infection among lions. A phylogenetic analysis of cloned FIV-Ple pol gene sequences from 27 lions from four African populations (from the Serengeti reserve, Ngorongoro Crater, Lake Manyara, and Kruger Park) revealed remarkably high intra- and interindividual genetic diversity at the sequence level. Three FIV-Ple phylogenetic clusters or clades were resolved with phenetic, parsimony, and likelihood analytical procedures. The three clades, which occurred not only together in the same population but throughout Africa, were as divergent from each other as were homologous pol sequences of lentivirus isolated from distinct feline species, i.e., puma and domestic cat. The FIV-Ple clades, however, were more closely related to each other than to other feline lentiviruses (monophyletic for lion species), suggesting that the ancestors of FIV-Ple evolved in allopatric (geographically isolated) lion populations that converged recently. To date, there is no clear evidence of FIV-Ple-associated pathology, raising the possibility of a historic genetic accommodation of the lion lentivirus and its host leading to a coevolved host-parasite symbiosis (or commensalism) in the population similar to that hypothesized for endemic simian immunodeficiency virus without pathology in free-ranging African monkey species.     

Social Structure of Lions (Panthera leo) Is Affected by Management in Pendjari Biosphere Reserve,Benin

Lion populations have undergone a severe decline in West Africa. As baseline for conservation management, we assessed the group structure of lions in the Pendjari Biosphere Reserve in Benin. This reserve, composed of one National Park and two Hunting Zones, is part of the WAP transboundary complex of protected areas. Overall mean group size was 2.6±1.7 individuals (n = 296), it was significantly higher in the National Park (2.7±1.7, n = 168) than in the Hunting Zones (2.2±1.5, n = 128). Overall adult sex ratio was even, but significantly biased towards females (0.67) in the National Park and towards males (1.67) in the Hunting Zones. Our results suggest that the Pendjari lion population is affected by perturbations, such as trophy hunting.     

Diversity and phylogenetic relationships of <Emphasis Type="Italic">Glossina</Emphasis> populations in Nigeria and the Cameroonian border region

Background

Tsetse flies are vectors of trypanosomes, parasites that cause devastating disease in humans and livestock. In the course of vector control programmes it is necessary to know about the Glossina species present in the study area, the population dynamics and the genetic exchange between tsetse fly populations.

Results

To achieve an overview of the tsetse fly diversity in Nigeria and at the Nigeria-Cameroon border, tsetse flies were trapped and collected between February and March 2014 and December 2016. Species diversity was determined morphologically and by analysis of Cytochrome C Oxidase SU1 (COI) gene sequences. Internal transcribed spacer-1 (ITS-1) sequences were compared to analyse variations within populations. The most dominant species were G. m. submorsitans, G. tachinoides and G. p. palpalis. In Yankari Game Reserve and Kainji Lake National Park, G. submorsitans and G. tachinoides were most frequent, whereas in Old Oyo National Park and Ijah Gwari G. p. palpalis was the dominant species. Interestingly, four unidentified species were recorded during the survey, for which no information on COI or ITS-1 sequences exists. G. p. palpalis populations showed a segregation in two clusters along the Cameroon-Nigerian border.

Conclusions

The improved understanding of the tsetse populations in Nigeria will support decisions on the scale in which vector control is likely to be more effective. In order to understand in more detail how isolated these populations are, it is recommended that further studies on gene flow be carried out using other markers, including microsatellites.

     

The Lion in West Africa Is Critically Endangered

The African lion has declined to <35,000 individuals occupying 25% of its historic range. The situation is most critical for the geographically isolated populations in West Africa, where the species is considered regionally endangered. Elevating their conservation significance, recent molecular studies establish the genetic distinctiveness of West and Central African lions from other extant African populations. Interventions to save West African lions are urgently required. However formulating effective conservation strategies has been hampered by a lack of data on the species'' current distribution, status, and potential management deficiencies of protected areas (PAs) harboring lions. Our study synthesized available expert opinion and field data to close this knowledge gap, and formulate recommendations for the conservation of West African lions. We undertook lion surveys in 13 large (>500 km²) PAs and compiled evidence of lion presence/absence for a further eight PAs. All PAs were situated within Lion Conservation Units, geographical units designated as priority lion areas by wildlife experts at a regional lion conservation workshop in 2005. Lions were confirmed in only 4 PAs, and our results suggest that only 406 (273–605) lions remain in West Africa, representing <250 mature individuals. Confirmed lion range is estimated at 49,000 km², or 1.1% of historical range in West Africa. PAs retaining lions were larger than PAs without lions and had significantly higher management budgets. We encourage revision of lion taxonomy, to recognize the genetic distinctiveness of West African lions and highlight their potentially unique conservation value. Further, we call for listing of the lion as critically endangered in West Africa, under criterion C2a(ii) for populations with <250 mature individuals. Finally, considering the relative poverty of lion range states in West Africa, we call for urgent mobilization of investment from the international community to assist range states to increase management effectiveness of PAs retaining lions.     

Implications of closed ecosystem conservation management: the decline of Rothschild’s giraffe (Giraffa camelopardalis rothschildi) in Lake Nakuru National Park,Kenya

Giraffe were historically free‐ranging across most of sub‐Saharan Africa but are now most often confined to national parks, conservation areas, or private ranches. Five viable populations of Rothschild’s giraffe (Giraffa camelopardalis rothschildi) remain in protected areas in Kenya and Uganda. The viable population in Uganda is Murchison Falls National Park and the four populations in Kenya are Lake Nakuru National Park (LNNP), Ruma National Park, Giraffe Manor, and Mwea Natural Reserve. The Kenya Wildlife Service queried a rapid decline in LNNP giraffe numbers falling from 153 individuals (1995) to 62 individuals (2002) and the failure of that population to recruit young in those years. Significantly reduced browse options, inbreeding depression and preferential lion predation were considered as potential reasons for this trend. Population genetic parameter estimates derived from multilocus genotype analyses suggest that the LNNP population was in good genetic health with respect to the likelihood of inbreeding depression. The population decline coincided with the drought attributed to the 1994 El Niño. Possible dietary complications from highly concentrated tannin levels because of forced over consumption of the park’s declining acacia trees may have compromised young giraffe, making them easy and opportunistic prey for the park’s lion population.     

Population size of lions in Yankari Game Reserve as revealed by faecal DNA sampling

Studies have shown that lion (Panthera leo) populations in West Africa are small, isolated and fragmented. In Nigeria, lions have disappeared from unprotected areas and are nowadays found only in parks and reserves where these populations may still decline. It is therefore urgent to obtain reliable estimates of population sizes at different localities. Direct observational surveys may either fail to count all individuals or count some individuals repeatedly and are therefore associated with unknown levels of estimation errors. More accurate estimates can be obtained if direct counting is combined with DNA‐based individual identification. As lions are difficult to identify individually, presented here is a method that can be a valuable addition to the existing census methods.     

Molecular phylogeny of the extinct cave lion Panthera leo spelaea

To reconstruct the phylogenetic position of the extinct cave lion (Panthera leo spelaea), we sequenced 1 kb of the mitochondrial cytochrome b gene from two Pleistocene cave lion DNA samples (47 and 32 ky B.P.). Phylogenetic analysis shows that the ancient sequences form a clade that is most closely related to the extant lions from Africa and Asia; at the same time, cave lions appear to be highly distinct from their living relatives. Our data show that these cave lion sequences represent lineages that were isolated from lions in Africa and Asia since their dispersal over Europe about 600 ky B.P., as they are not found among our sample of extant populations. The cave lion lineages presented here went extinct without mitochondrial descendants on other continents. The high sequence divergence in the cytochrome b gene between cave and modern lions is notable.     

Genetic diversity,evolutionary history and implications for conservation of the lion (Panthera leo) in West and Central Africa

Aim In recent decades there has been a marked decline in the numbers of African lions (Panthera leo), especially in West Africa where the species is regionally endangered. Based on the climatological history of western Africa, we hypothesize that West and Central African lions have a unique evolutionary history, which is reflected by their genetic makeup. Location Sub‐Saharan Africa and India, with special focus on West and Central Africa. Method In this study 126 samples, throughout the lion’s complete geographic range, were subjected to phylogenetic analyses. DNA sequences of a mitochondrial region, containing cytochrome b, tRNAPro, tRNAThr and the left part of the control region, were analysed. Results Bayesian, maximum likelihood and maximum parsimony analyses consistently showed a distinction between lions from West and Central Africa and lions from southern and East Africa. West and Central African lions are more closely related to Asiatic lions than to the southern and East African lions. This can be explained by a Pleistocene extinction and subsequent recolonization of West Africa from refugia in the Middle East. This is further supported by the fact that the West and Central African clade shows relatively little genetic diversity and is therefore thought to be an evolutionarily young clade. Main conclusions The taxonomic division between an African and an Asian subspecies does not fully reflect the overall genetic diversity within lions. In order to conserve genetic diversity within the species, genetically distinct lineages should be prioritized. Understanding the geographic pattern of genetic diversity is key to developing conservation strategies, both for in situ management and for breeding of captive stocks.     

Molecular genetic variation across the southern and eastern geographic ranges of the African lion, <Emphasis Type="Italic">Panthera leo</Emphasis>

We examined sequence variation in the mitochondrial cytochrome b and NADH dehydrogenase subunit 5 genes (2,360 bp total) for 26 lions from eleven locations throughout sub-Saharan Africa. Six distinct haplotypes were observed in the combined sequences, forming two clades: the eastern and the western savannas. The Uganda-Western Kenya haplotype grouped at a basal position with the eastern clade of lions from Tsavo south to the Transvaal and Natal regions. The phylogenetic position of the haplotype from Sabi Sands in the southern part of Kruger National Park remained poorly resolved. The haplotypes found in Namibia and Botswana formed the western clade. The modest genetic variation documented here argues against taxonomic distinctions among living African lions.     

Autosomal and mtDNA Markers Affirm the Distinctiveness of Lions in West and Central Africa

The evolutionary history of a species is key for understanding the taxonomy and for the design of effective management strategies for species conservation. The knowledge about the phylogenetic position of the lion (Panthera leo) in West/Central Africa is largely based on mitochondrial markers. Previous studies using mtDNA only have shown this region to hold a distinct evolutionary lineage. In addition, anthropogenic factors have led to a strong decline in West/Central African lion numbers, thus, the conservation value of these populations is particularly high. Here, we investigate whether autosomal markers are concordant with previously described phylogeographic patterns, and confirm the unique position of the West/Central African lion. Analysis of 20 microsatellites and 1,454 bp of the mitochondrial DNA in 16 lion populations representing the entire geographic range of the species found congruence in both types of markers, identifying four clusters: 1) West/Central Africa, 2) East Africa, 3) Southern Africa and 4) India. This is not in line with the current taxonomy, as defined by the IUCN, which only recognizes an African and an Asiatic subspecies. There are no indications that genetic diversity in West/Central Africa lions is lower than in either East or Southern Africa, however, given this genetic distinction and the recent declines of lion numbers in this region, we strongly recommend prioritization of conservation projects in West/Central Africa. As the current taxonomic nomenclature does not reflect the evolutionary history of the lion, we suggest that a taxonomic revision of the lion is warranted.     

Research needs for lion conservation in West and Central Africa

The lion has historically probably been widespread at low densities in West and Central Africa, nowadays they are largely restricted to small isolated populations inside protected areas. The total number is probably between 1200 and 2700, the best possible guesstimate would be 1700. Mankind is the main cause for the suspected decline of lion populations, both inside and outside protected areas. Very little research has been done on West and Central African lions a few examples are summarized here. The international community is slowly becoming aware of threats to lions in the region and some initiatives for lion conservation have started.     

Examining the Extinction of the Barbary Lion and Its Implications for Felid Conservation

Estimations of species extinction dates are rarely definitive, yet declarations of extinction or extirpation are important as they define when conservation efforts may cease. Erroneous declarations of extinctions not only destabilize conservation efforts but also corrode local community support. Mismatches in perceptions by the scientific and local communities risk undermining sensitive, but important partnerships. We examine observations relating to the decline and extinction of Barbary lions in North Africa. Whilst the extinction predates the era of the scientific conservation movement, the decline is relatively well documented in historical records. Recently unearthed accounts suggest Barbary lions survived later than previously assumed. We use probabilistic methods to estimate a more recent extinction date for the subspecies. The evidence presented for a much later persistence of lions in North Africa, including generations when sightings were nil, suggests caution when considering felid populations as extinct in the wild. The case raises the possibility that captive animals descended from the Moroccan royal collection are closer contemporaries to wild Barbary lions. Furthermore, our results highlight the vulnerability of very small lion populations and the significance of continued conservation of remnant lion populations in Central and West Africa.     

A possible new piroplasm in lions from the Republic of South Africa

A small piroplasm was detected in blood smears from lions (Panthera leo) in the Kruger National Park (KNP; Republic of South Africa) during 1991/1992. The parasite was identified provisionally as Babesia felis, but sera from these lions tested negative to B. felis antigen in the indirect immunofluorescent antibody test (IFAT). Blood from an infected lion was subsequently subinoculated into a domestic cat and two leopards in an attempt to identify the parasite. A lion also was infected with B. felis (from a cat). Serum samples collected from these animals were tested against B. felis, the KNP small piroplasm, and Cytauxzoon felis antigen in the IFAT. The serological results indicate that the KNP small piroplasm isolated from the lion is probably a distinct species from B. felis and C. felis.     

Translocating lions into an inbred lion population in the Hluhluwe-iMfolozi Park, South Africa

A fundamental problem in conservation biology is the risk of inbreeding in fragmented and declining populations. In the Hluhluwe-iMfolozi Park (HiP), a small, enclosed reserve in South Africa, a large lion Panthera leo population arose from a founder group of five individuals in the 1960s. The HiP lion population went through a persistent decline and showed indications of inbreeding depression. To restore the genetic variation of the inbred HiP lion population, new lions were translocated into the existing population. Translocated females formed stable associations and established enduring pride areas with other translocated lionesses, but did not bond into native female prides. The translocated male coalition was more successful in gaining and maintaining residence in a pride than the translocated lone male that split off on his own from the male coalition. Litter size and cub survival was about twice as high for pairings involving at least one translocated parent than for pairings of two native lions. It is therefore possible to infuse new genes rapidly and successfully into a small, isolated lion population. Such translocations may become an important adaptive management tool as lion populations become increasingly fragmented.     

Inbreeding depression increases susceptibility to bovine tuberculosis in lions: an experimental test using an inbred-outbred contrast through translocation

Disease can dramatically influence the dynamics of endangered wildlife populations, especially when they are small and isolated, with increased risk of inbreeding. In Hluhluwe-iMfolozi Park (HiP), a small, enclosed reserve in South Africa, a large lion (Panthera leo) population arose from a small founder group in the 1960s and started showing conspicuous signs of inbreeding. To restore the health status of the HiP lion population, outbred lions were translocated into the existing population. In this study, we determined the susceptibility to bovine tuberculosis (bTB), and the prevalence of antibody to feline viruses of native lions, and compared the findings with those from translocated outbred lions and their offspring. Antibodies to feline herpesvirus, feline calicivirus, feline parvovirus, and feline coronavirus were present in the lion population, but there was no significant difference in antibody prevalence between native and translocated lions and their offspring, and these feline viruses did not appear to have an effect on the clinical health of HiP lions. However, feline immunodeficiency virus (FIV), which was previously absent from HiP, appears to have been introduced into the lion population through translocation. Within 7 yr, the prevalence of antibody to FIV increased up to 42%. Bovine tuberculosis posed a major threat to the inbred native lion population, but not to translocated lions and their offspring. More than 30% of the native lion population died from bTB or malnutrition compared with <2% of the translocated lions and their offspring. We have demonstrated that management of population genetics through supplementation can successfully combat a disease that threatens population persistence. However, great care must be taken not to introduce new diseases into populations through translocation.     

The size of savannah Africa: a lion’s (Panthera leo) view

We define African savannahs as being those areas that receive between 300 and 1,500 mm of rain annually. This broad definition encompasses a variety of habitats. Thus defined, savannahs comprise 13.5 million km² and encompass most of the present range of the African lion (Panthera leo). Dense human populations and extensive conversion of land to human use preclude use by lions. Using high-resolution satellite imagery and human population density data we define lion areas, places that likely have resident lion populations. In 1960, 11.9 million km² of these savannahs had fewer than 25 people per km². The comparable area shrank to 9.7 million km² by 2000. Areas of savannah Africa with few people have shrunk considerably in the last 50 years and human population projections suggest they will likely shrink significantly in the next 40. The current extent of free-ranging lion populations is 3.4 million km² or about 25 % of savannah area. Habitats across this area are fragmented; all available data indicate that between 32,000 and 35,000 free-ranging lions live in 67 lion areas. Although these numbers are similar to previous estimates, they are geographically more comprehensive. There is abundant evidence of widespread declines and local extinctions. Under the criteria we outline, ten lion areas qualify as lion strongholds: four in East Africa and six in Southern Africa. Approximately 24,000 lions are in strongholds, with an additional 4,000 in potential ones. However, over 6,000 lions are in populations of doubtful long-term viability. Lion populations in West and Central Africa are acutely threatened with many recent, local extinctions even in nominally protected areas.     

Change in percentage ground cover of perennial grasses under different burning regimes

Summary Differences in the ground and perennial grass covers under different burning and grazing regimes are presented for three vegetation types in Kainji Lake National Park. It is shown from the results that perennial grasses established higher basal cover under late burning treatments than under early burning and no burning treatments. Late fires destroy litter which hinders grass growth while no burning treatments cause accumulation of litter. The highest number of grass tillers are produced under late burning treatments, while the lowest occurred under no burning treatments. The early burning plots have intermediate results. Occasional late burning in controlled small blocks of land is recommended for optimum grass production in the study area.Nomenclature follows Hutchinson & Dalziel 1954–1972. Flora of West Tropical Africa 2nd ed., edited by R.W.J. Keay & F.N. Hepper, Crown Agents, London.I am most grateful to Kainji Lake Research Institute for giving me great moral and financial support during the course of field work for this study. I am also very grateful to Dr. S.S. Ajayi of the Department of Forest Resources Management for allowing me to use part of his Senate Research Grant for the study.     

Seasonal changes in the nutrient content of East African grassland vegetation

Significant seasonal changes in the concentrations of N, P, K, and Ca were observed in the herb layer vegetation at Nairobi National Park and Masai Mara Game Reserve in Kenya. These changes were related to the alternation of wet and dry seasons. Live forbs typically had the highest concentrations of all nutrients, especially N and Ca, at both locations. Standing dead and litter compartments showed less seasonal variation and had the lowest concentrations of all nutrients except Ca when compared with those of other compartments. Standing stocks of all nutrients were higher at Nairobi National Park than at Masai Mara due to larger litter and standing dead compartments resulting from lower grazing intensity and a lower frequency of fire. Most of the aboveground nutrients at Nairobi National Park appeared to be recycled through the decomposer pathway, while a large proportion of the standing stock at Masai Mara appeared to be recycled through the excreta of large mammalian herbivores and through fires. The crude protein content of the live grass compartment fell below maintenance levels required for ruminants (5% crude protein) during September 1980 and February 1981 at Nairobi National Park and during June and July 1980 at Masai Mara Game Reserve. Thus, herbivore populations in these preserves may be limited by shortages of nutritionally adequate food during dry seasons, as previously reported for other regions in East Africa.