Barbary Lion Research Articles

The natural history of lions in North Africa and the relevance of their depiction in the “Lions Attacking a Dromedary” diorama

AbstractThe diorama “Lions Attacking a Dromedary” is notorious, not just as a remarkable 19th century example of taxidermy, but also its controversial representation of human culture and animals, its questionable accuracy and the murky ethics of the materials sourced for its construction. This study examines whether the diorama is a reasonable representation of the Barbary lion in North Africa. We review the history of lions in North Africa and their interaction with humans in the 19th and 20th centuries. The ecology and biology of North African lions, supported by scientific knowledge of the species, enables assessment of the value of diorama depiction, whether it is realistic and informative, or an artistic interpretation, or mere fiction. Furthermore, if the depiction is fictional, whether the representation is entirely unrepresentative, or at worst a stereotypic fabrication based on Western colonial perspectives of the 1800s. The paper does not explore cultural aspects, although refers to the experiences of people in local communities as well as colonial visitors through 150 documented accounts of lions in the region (modern day Morocco, Algeria, and Tunisia) between 1830 and 1960. The analysis explores whether real experiences are reflected in the depiction presented in the diorama and identifies that some aspects of the exhibit can be considered authentic, while others appear to emphasize drama rather than lived reality. Recommendations are offered for well‐informed future presentation of “Lions Attacking a Dromedary.”

Fire human-climate interaction in Atlas cedar forests of Aurès, Northern Algeria

Forty-one fire-scarred samples from two new Cedrus atlantica sites in the Ouled-Yaâgoub Forest were collected and combined with the two previous sites from the Chelia Forest reported in Slimani et al. (2014) to develop a regional analysis of fire history in the Aurès Mountains, northern Algeria. Prior to 1850, cedar forests were characterized by high frequency, low-intensity surface fire regimes. Fire frequency declined after 1850 after the elimination of the Barbary lion and the enforcement of the first laws governing land and forest use in Algeria. Most of the fire events in both forests occurred in the summer season, the latter part of the growing season, or in the dormant season. We compared past fires occurrences and an October-June precipitation reconstruction. On a regional scale, precipitation was significantly reduced in the months prior to the fire occurrence. In addition, precipitation during the first year preceding the fire years was significantly above average. This study provides a baseline for planning and justifying improved ecosystem management programs.

Genetic diversity of Barbary lion based on genealogic analysis

Genetic diversity of Barbary lion based on genealogic analysis

Are extinction opinions extinct?

Extinction models vary in the information they require, the simplest considering the rate of certain sightings only. More complicated methods include uncertain sightings and allow for variation in the reliability of uncertain sightings. Generally extinction models require expert opinion, either as a prior belief that a species is extinct, or to establish the quality of a sighting record, or both. Is this subjectivity necessary? We present two models to explore whether the individual quality of sightings, judged by experts, is strongly informative of the probability of extinction: the ‘quality breakpoint method’ and the ‘quality as variance method’. For the first method we use the Barbary lion as an exemplar. For the second method we use the Barbary lion, Alaotra grebe, Jamaican petrel and Pohnpei starling as exemplars. The ‘quality breakpoint method’ uses certain and uncertain sighting records, and the quality of uncertain records, to establish whether a change point in the rate of sightings can be established using a simultaneous Bayesian optimisation with a non-informative prior. For the Barbary lion, there is a change in subjective quality of sightings around 1930. Unexpectedly sighting quality increases after this date. This suggests that including quality scores from experts can lead to irregular effects and may not offer reliable results. As an alternative, we use quality as a measure of variance around the sightings, not a change in quality. This leads to predictions with larger standard deviations, however the results remain consistent across any prior belief of extinction. Nonetheless, replacing actual quality scores with random quality scores showed little difference, inferring that the quality scores from experts are superfluous. Therefore, we deem the expensive process of obtaining pooled expert estimates as unnecessary, and even when used we recommend that sighting data should have minimal input from experts in terms of assessing the sighting quality at a fine scale. Rather, sightings should be classed as certain or uncertain, using a framework that is as independent of human bias as possible.

The Challenges and Relevance of Exploring the Genetics of North Africa's "Barbary Lion" and the Conservation of Putative Descendants in Captivity.

The lions of North Africa were unique in ecological terms as well as from a human cultural perspective and were the definitive lions of Roman and Medieval Europe. Labelled “Barbary” lions, they were once numerous in North Africa but were exterminated by the mid-20th century. Despite subsequent degeneration of the Atlas Mountain ecosystem through human pressures, the feasibility of lion reintroduction has been debated since the 1970s. Research on the long-established captive lion collection traditionally kept by the sultans and kings of Morocco has enabled selective breeding coordinated across Moroccan and European zoos involving a significant number of animals. Molecular genetic research has recently provided insights into lion phylogeny which, despite previous suggestions that all lions share recent common ancestry, now indicates clear distinctions between lions in North, West, and Central Africa, the Middle East, and India versus those in Southern and Eastern Africa. A review of the evolutionary relevance of North African lions highlights the important challenges and opportunities in understanding relationships between Moroccan lions, extinct North African lions, and extant lion populations in India and West and Central Africa and the potential role for lions in ecosystem recovery in those regions.

Assessing uncertainty in sighting records: an example of the Barbary lion.

As species become rare and approach extinction, purported sightings can be controversial, especially when scarce management resources are at stake. We consider the probability that each individual sighting of a series is valid. Obtaining these probabilities requires a strict framework to ensure that they are as accurately representative as possible. We used a process, which has proven to provide accurate estimates from a group of experts, to obtain probabilities for the validation of 32 sightings of the Barbary lion. We consider the scenario where experts are simply asked whether a sighting was valid, as well as asking them to score the sighting based on distinguishablity, observer competence, and verifiability. We find that asking experts to provide scores for these three aspects resulted in each sighting being considered more individually, meaning that this new questioning method provides very different estimated probabilities that a sighting is valid, which greatly affects the outcome from an extinction model. We consider linear opinion pooling and logarithm opinion pooling to combine the three scores, and also to combine opinions on each sighting. We find the two methods produce similar outcomes, allowing the user to focus on chosen features of each method, such as satisfying the marginalisation property or being externally Bayesian.

From dinosaurs to dodos: who could and should we de-extinct?

Reviving extinct species with new synthetic biology tools is as exciting an idea as it is controversial. Genomic manipulation of extinct species’ close relatives and/or cloning suitably preserved cells are the two main ways synthetic biology could be used to revive species. Discussions of where to target initial revival efforts have focused on species’ charisma (e.g. Woolly mammoth, Passenger pigeon) with less emphasis on feasibility or the ecological, ethical and legal considerations. Here I discuss who we could and should de-extinct, focussing on these latter criteria. Given the current devastating anthropogenic pressures on biodiversity, I suggest that a better use of de-extinction technologies would be to focus them on preventing species extinctions by restoring populations of critically endangered species. For example, this could be through increasing population numbers through cloning or genomic manipulation to better enable susceptible species to adapt to global change or by restoring genetic diversity by reviving extinct sub-species (e.g. Quagga, Barbary lion). This idea circumvents many of the criticisms of de-extinction from conservationists, whilst retaining public interest in de-extinction.

Revealing the maternal demographic history of Panthera leo using ancient DNA and a spatially explicit genealogical analysis

BackgroundUnderstanding the demographic history of a population is critical to conservation and to our broader understanding of evolutionary processes. For many tropical large mammals, however, this aim is confounded by the absence of fossil material and by the misleading signal obtained from genetic data of recently fragmented and isolated populations. This is particularly true for the lion which as a consequence of millennia of human persecution, has large gaps in its natural distribution and several recently extinct populations.ResultsWe sequenced mitochondrial DNA from museum-preserved individuals, including the extinct Barbary lion (Panthera leo leo) and Iranian lion (P. l. persica), as well as lions from West and Central Africa. We added these to a broader sample of lion sequences, resulting in a data set spanning the historical range of lions. Our Bayesian phylogeographical analyses provide evidence for highly supported, reciprocally monophyletic lion clades. Using a molecular clock, we estimated that recent lion lineages began to diverge in the Late Pleistocene. Expanding equatorial rainforest probably separated lions in South and East Africa from other populations. West African lions then expanded into Central Africa during periods of rainforest contraction. Lastly, we found evidence of two separate incursions into Asia from North Africa, first into India and later into the Middle East.ConclusionsWe have identified deep, well-supported splits within the mitochondrial phylogeny of African lions, arguing for recognition of some regional populations as worthy of independent conservation. More morphological and nuclear DNA data are now needed to test these subdivisions.

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.

Embryo Retrieval after Hormonal Treatment to Control Ovarian Function and Non‐surgical Artificial Insemination in African Lions (Panthera leo)

Assisted reproduction technologies are essential for propagating endangered wild felids. Artificial insemination (AI) has been reported in several wild feline species, but pregnancy rates are low, partially owing to failures of current hormonal stimulation protocols. Therefore, this study describes the application of reliable methods to monitor ovarian activity and the development of an effective hormonal protocol to induce oestrus and ovulation in African lions. Application of porcine FSH and porcine LH was shown to be effective for inducing follicular growth and ovulation, and this regimen appeared to be superior to protocols described earlier in terms of ovulation and fertilization rates. Furthermore, non-surgical AI was performed successfully in lions, and uterine-stage embryos were collected and cryopreserved. African lions may serve as a valuable model to develop assisted reproduction for propagation of relic zoo populations in the critically endangered Asian lion or Barbary lion.

LEISHMANIA INFANTUMINFECTION IN TWO CAPTIVE BARBARY LIONS (PANTHERA LEO LEO)

A female barbary lion (Panthera leo leo) from the Montpellier Zoological Park (France) showing colitis, epistaxis, and lameness with pad ulcers was positive by polymerase chain reaction (PCR) for Leishmania infantum. Further indirect immunofluorescence (IFAT) tests on the banked sera from all lions of the park detected another infected but asymptomatic female, which was confirmed by PCR on ethylenediaminetetraacetic acid (EDTA) blood sample. Leishmania infantum zymodeme MON-1 was cultured from EDTA bone marrow samples sampled from this second animal. The first female was successfully treated with marbofloxacine at 2 mg/kg s.i.d. for 28 days (Marbocyl, Vetoquinol 70204 Lure, France) and allopurinol at 30 mg/kg s.i.d. for 3 mo (Allopurinol Mylan, Mylan SAS, 69800 Saint-Priest, France) and then 1 wk/mo. Both positive animals were born at the Rabat Zoological Park, Morocco, and arrived together at Montpellier in 2003. The chronicity and source of this current infection are unknown since Morocco and southern France are well-known to be enzootic for leishmaniasis.

Past and present distribution of the North African–Asian lion subgroup: a review

ABSTRACT The North African–Asian lion subgroup, which is composed of two subspecies, the Barbary lion, Panthera leo leo, and the Asian lion, P. l. persica, was nearly exterminated during the last centuries. The remaining free‐ranging population of Asian lions consists of c. 350 animals in India. The Barbary lion subsists as captive animals in zoos or circuses, all of which originate from the Moroccan Royal Collection. There have been multiple genetic hybridizations with sub‐Saharan lions. Several incomplete distribution maps of the past occupation of lions were proposed in the literature. I examine the distribution of these lions with the help of a larger database, extracted from zoological, archaeological and historical reports. Some records are however not completely reliable. Data were collected over a long period of time (from the Epipaleolithic to the modern times) in Eurasia, North Africa and the Arabian Peninsula. Dispersal took place in latitudes as far south as 15°N (Yemen) to 18°N (Mali; Chad) and as far north as 45–48°N (Bulgaria); and to longitudes as far west as 5.57°W (Morocco) and as far east as 84°E (India). Expansion was probably constrained by natural ecological factors northwards (higher seasonality, harsher winters) and southwards (extension of aridity after 3000 BP). At latitudes between 40 and 43°N, lions seem to have become a permanent part of the fauna during 6–8 millennia. Lions were brought to extinction by humans through hunting, captures for exhibition in zoos and by the associated natural fragmentation of wild populations. The dates at which extinction took place varied greatly according to the geographical sites: 3000 BC for temperate Europe; 1000 BC in the south of Greece; 12–13th century in the Near East, Arabian Peninsula, Trans‐Caucasia or north of Afghanistan; and 19–20th century for North Africa, the Middle East and India.

Maintaining the genetic health of putative Barbary lions in captivity: an analysis of Moroccan Royal Lions

The last representatives of the Barbary lion (Panthera leo leo), once numerous in North Africa but exterminated from the wild by the 1940s, are believed to be the captive lions descended from the Moroccan Royal Collection, numbering less than 90 animals in zoos worldwide. The genetic fitness of these captive “Royal Lions” may now be under threat since, although most zoos have avoided hybridisation with animals of other origin, no formal breeding programme currently exists and several institutions have halted breeding activities. This situation has arisen since the distinctiveness of Barbary lions and the representative status of Royal Lions remain inconclusive and definitive molecular studies have yet to be completed. Previously, in the 1970s, morphological and phenotypic traits were used to match Royal Lions and the historic Barbary lion and an ex situ breeding programme was initiated involving a number of selected “founder” animals. This paper outlines the status of the descendent population within zoos in Morocco and Europe, including all known pure-bred descendents from the Royal Palace collection. Founder representation is shown to be greater across European collections than the Moroccan collection. Breeding exchanges are recommended between institutions in order to improve genetic diversity and maintain the genetic health of the population and a studbook for European zoo animals has been developed to support this action. This analysis serves as a benchmark for guiding effective maintenance of the captive population, thereby allowing time to clarify the conservation value of Royal Lions and their relevance to North African ecology.

A lion’s share of attention: Archaeozoology and the historical record

This is a review worth starting with Aesop (?620–564 BC). Eight classical Greek texts written in the Age of Homer (8–7th c. BC) reveal that lion was most frequently mentioned among wild animals, surpassed only by the six common domesticates that have directly surrounded people.1 References to this beast far outnumbered those to comparably large, dangerous and highly valued carnivores (Fig. 1). This is in sharp contrast with the relative underrepresentation of lion bones in archaeozoological assemblages. As A. Meyer noted with immense foresight over a century ago, “From the philological side there is evidently no basis to doubt... that lions indeed lived in Europe... around 500 BC... [Although] until recently no lion bones have been found in Greece, one should consider how few excavations took place in those lands”.2 Only a quarter of a century later, ecavations at the 5th–2nd century BC site of Ol’bia, a Greek colony on the Black Sea coast in modern-day Ukraine indeed yielded the fi rst bones of large Felids,3 attributed at the time to animals kept in a menagerie or the import of precious pelts.4 Although these possibilities must always be taken into consideration, reconfi rming Meyer’s optimistic view large faunal assemblages recovered in the Balkans during the last three decades have produced increasing numbers of lion remains from prehistoric as well as early historical times. This study seeks reconciling the remianing discrepancies between the historical record and archaeozoological evidence. As for the zoological aspect of this work, there are fundamental diffi culties in distinguishing between the bones of Late-Glacial and Holocene lions in Europe. Morphometric differences between the Pleistocene cave lion (Panthera leo spelaea Goldfuss, 1810) and the recent lion (Panthera leo Linnaeus, 1758) are negligible. While the bones of cave lion tend to be larger, precise identifi cation is often impossible using only sporadically occurring remains. Ancient DNA sequences indicate that cave lion from Austria and Southern Germany represent lineages that diverged from lions in Africa and Asia about 600,000 BP, and became extinct without mitochondrial descendants in those two continents. Depending on convention, even this ancient, distant form, however, may fi t within a single species nomenclature.5 Modern lions are closely related. Craniological traits of the now extinct North African Barbary lion (Panthera leo leo Linnaeus, 1758) and Asiatic lions (Panthera leo persica Meyer, 1826) are very similar. There must have been a contiguous population inhabiting North Africa and Asia. Moreover, the population in South-East Europe possibly once linked lions in the Middle East and North Africa, although the European form

Lost populations and preserving genetic diversity in the lion Panthera leo: Implications for its ex situ conservation

Two of the eight recognized lion subspecies, North African Barbary lion (Panthera leo leo) and South African Cape lion (Panthera leo melanochaita), have become extinct in the wild in the last 150 years. Based on sequences of mitochondrial DNA (mtDNA) control region (HVR1) extracted from museum specimens of four Barbary and one Cape lion, the former was probably a distinct population characterized by an invariable, unique mtDNA haplotype, whilst the latter was likely a part of the extant southern African lion population. Extinction of the Barbary line, which may still be found in ‘‘generic’’ zoo lions, would further erode lion genetic diversity. Therefore, appropriate management of such animals is important for maintaining the overall genetic diversity of the species. The mtDNA haplotype unique to the Barbary lion, in combination with the small size of the HVR1 analyzed (c. 130 bp), makes it possible and cost-effective to identify unlabelled Barbary specimens kept in museums and ‘‘generic’’ captive lions that may carry the Barbary line. An initial study of five samples from the lion collection of the King of Morocco, tested using this method, shows that they are not maternally Barbary.

Urgent call for further breeding of the relic zoo population of the critically endangered Barbary lion (Panthera leo leo Linnaeus 1758)

The Barbary lion became extinct in the wild around 1942. Since the end of the 19th century, a last purebred captive breeding stock existed at the court of Morocco. The rest of these animals became the core exhibition of the new Rabat Zoo after passing through repeated bottlenecks and possibly some introgression events by foreign lions. This study uses mitochondrial DNA sequencing data to clarify the relationship among these lions and their sub-Saharan and Asian relatives. We analysed mitochondrial cytochrome b sequences obtained from a sample from a Barbary lion descended from a young female of the Barbary lion breeding group at the Rabat Zoo and various other members of the genus Panthera. In our cytochrome-b-based phylogenetic tree, the North African Barbary lion, represented by a biopsy sample from the Neuwied Zoo, joins the Asian lion clade, although it is slightly different from its Asian sister group. However, it is clearly distinct from sub-Saharan lions and can be considered as a genetically defined phylogeographic group of its own. Molecular dating of the extant sub-Saharan and Asian lion groups shows that the split between North African Barbary lions and Asian lions must be considerably more recent than 74–203 kilo years ago.