Lion (Panthera leo)

[creature386]

Panthera leo

Classification:
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Family: Felidae
Genus: Panthera
Species: P. leo

Image source[1]
Size:
Panthera leo is a large feline, which can range in size from 83 kg (females) at minimum [2] and 225 kg (males)[3] at maximum, with males generally being 50% bigger[2]. The mean mass for males is 187 kg and for cubs 5 kg.[3] Females range in size from 83-168 kg and males from 145-225 kg. On average, females range from 119-140 kg and males from 175-193 kg depending on the location[3]. Some males can even weigh up tp 250 kg[4]. An extinct subspecies (Panthera leo atrox), may have weighed 210-297 kg [5]. It (the modern one, tough the extinct one likley aswell) has strong breasts and shoulders. Males can stand about 1,23m at the shoulder[2]. Head and body length is 170–250 cm in males and 140–175 cm in females [4]. The tail makes up 40-50% of the Lion's total length.[2].

Distribution:

Lions today live in most african contries below the sahara and in the Gir Forrests of India. They also lived in Marocco, but there they are now extinct[2]. Earlier, there was a subspecies in North America aswell.[5] Lions living in West- and Centralafrica are more related to the asian lions than to the lions in South- and Eastafrica.[6]

Predatory behavior and diet:
Lions are carnivores, who generally prefer medium size ungulates, but they also eat birds, hares, ostrich eggs, phytons, fish, crocodiles and even other lions. They have been recorded attacking pretty much everything ranging from 2-1000 kg, although ungultates make up 90% of the killed prey, but especially in the kalahari, they have to show flexibility for surviving. In India, they usually prey on 50 kg Chintial Deer.[2] But when available, the lions in India prefer larger prey.[1] Lions are generally ambush hunters, covering in the grass of the vegetation. Grass with a height of 40 cm is required for a succesful hunt, the success rate increases with 80 cm tall grass.[7]. Of course lions also sometimes attack at night, where their vision is superior to the vision of their prey. They usually try to get as close as 30m to their prey, before they start running towards it. In groups, they can kill their prey by seizing the muzzle, nape or throat. When they are together, lionesses are more succesful than alone, in the Etosha National Park, 27% of 486 cooperative hunts ended with catching and killing the victim. They use ambush strategies, because of their bad stamina. After a hundred meters, they can be exhausted yet. Tough, they can reach speeds of 48-59 km/h, they are also a bit more adapted for running than tigers[2].
For the killing, lions are equipped with 6 cm long canines[8] and a solidly built[2], 25.7 cm long [9] skull. For its bite force, the Lion has massive zygomatic arches and a large sagittal crest for the attachment of powerful jaw and neck muscles [2].
The prefered prey size of the Lion depends on it's habitat, but they usually prefer prey with a weight of 350 kg. The prefered weight range is 190-550 kg (in the serengeti 170-250 kg), animals out of that range are usually avoided[7]. Here a list of the prefered species:

©Matt W. Hayward and Graham I. H. Kerley[7]

Males prefer buffalo as prey item[1], tough the females attack bigger ones:

Image source[10]

Despite Black Rhinoceros being among the avoided prey, there is evidence of Lions attacking them. A subadult Black Rhinoceros was attacked by 3 male lions for 4 minutes. After 40 minutes (it was screaming a lot) it died.[11]
Regarding macropredatory, in Botswana, there are multiple accounts of Lions having taken Elephants. Of 74 elephants killed by lions (323 attempts were made by the lions), most attacked animals were 4-15 years old. The attacks on 4-9 year old elephants, yielded the most meals. One bull was killed, but he was injured by another bull before. Apart from this, the attacked adults were almost always females, tough the majority of the attacked juvenile to subadult elephants were males.[12]

Sources:
[1]Panthera leo Sarah K. Haas, Virginia Hayssen and Paul R. Krausman Mammalian Species No. 762, Panthera leo (Jul. 15, 2005), pp. 1-11 Published by: American Society of Mammalogists www.jstor.org/stable/3504539

[2]Sunquist, M. and Sunquist, F. (2002) Wild Cats of the World.
University of Chicago Press, Chicago. preview

[3] G .L. Smuts, G. A. Robinson, I. J. Whyte (1980). "Comparative growth of wild male and female lions (Panthera leo)"

[4] Nowak, Ronald M. (1999). Walker's Mammals of the World. Baltimore: Johns Hopkins University Press.

[5] Christiansen, Per, & Harris, John M. (2009). "Craniomandibular Morphology and Phylogenetic Affinities of Panthera atrox: Implications for the Evolution and Paleobiology of the Lion Lineage". Journal of Vertebrate Paleontology 29 (3): 934–945.

[6] L. D. Bertola, W. F. van Hooft, K. Vrieling, D. R. Uit de Weerd, D. S. York, H. Bauer, H. H. T. Prins, P. J. Funston, H. A. Udo de Haes, H. Leirs, W. A. van Haeringen, E. Sogbohossou, P. N. Tumenta, H. H. de Iongh. Genetic diversity, evolutionary history and implications for conservation of the lion (Panthera leo) in West and Central Africa. Journal of Biogeography, 2011 abstract

[7]Prey preferences of the lion (Panthera leo)
Matt W. Hayward, Graham I. H. Kerley
November 2005

pdf

[8]Bite forces, canine strength and skull allometry in carnivores
(Mammalia, Carnivora)
Per Christiansen* and Jan S. Adolfssen (2004)
pdf

[9]Body Size Overlap, Habitat Partitioning and Living Space Requirements of Terrestrial Vertebrate Predators: Implications for the Paleoecology of Large Theropod Dinosaurs
James O. Farlow, and Eric R. Pianka (2002) pdf

[10]Changes in home range size of African lions in relation to pride size and prey biomass in a semi-arid savanna Andrew J. Loveridge, Marion Valeix, Zeke Davidson, Felix Murindagomo, Hervé Fritz and David W. Macdonald Ecography Vol. 32, No. 6 (DECEMBER 2009), pp. 953-962 Published by: Wiley
www.jstor.org/stable/20696307

[11]C. Brian, O Forge and P. Erb (1999) "Lion predation on black rhinoceros (Diceros bicornis) in Etosha National Park." pdf

[12]Joubert D. 2006. Hunting behaviour of lions (Panthera leo) on elephants (Loxodonta africana) in the Chobe National Park, Botswana. Afr J Ecol 44:279-281. online

[creature386]

A paper about the distribution (the abstract):

Lions were the most widespread carnivores in the late Pleistocene, ranging from southern Africa to the southern USA, but little is known about the evolutionary relationships among these Pleistocene populations or the dynamics that led to their extinction. Using ancient DNA techniques, we obtained mitochondrial sequences from 52 individuals sampled across the present and former range of lions. Phylogenetic analysis revealed three distinct clusters: (i) modern lions, Panthera leo; (ii) extinct Pleistocene cave lions, which formed a homogeneous population extending from Europe across Beringia (Siberia, Alaska and western Canada); and (iii) extinct American lions, which formed a separate population south of the Pleistocene ice sheets. The American lion appears to have become genetically isolated around 340 000 years ago, despite the apparent lack of significant barriers to gene flow with Beringian populations through much of the late Pleistocene. We found potential evidence of a severe population bottleneck in the cave lion during the previous interstadial, sometime after 48 000 years, adding to evidence from bison, mammoths, horses and brown bears that megafaunal popu- lations underwent major genetic alterations throughout the last interstadial, potentially presaging the processes involved in the subsequent end-Pleistocene mass extinctions.

www.dur.ac.uk/greger.larson/DEADlab/Publications_files/Barnett_MEcavelions.pdf

[creature386]

Here some info about the prey preference of the lion in Addo (the information is based on abundance of the prey and how often they are encountered/hunted):

• Large herds of buffaloes were encountered far less frequently than their abundance would suggest, but when encountered, the buffaloes were hunted quite often.
• Zebras were encountered more often than their abundance would suggest and hunted as often as the encounter rate would suggest.
• Kudus were encountered as often as their abundance would suggested and hunted as often as the encounter rate would suggest.
• Warthogs were encountered more often than their abundance would suggest and hunted as often as the encounter rate would suggest.
• Elands were encountered as often as their abundance would suggest (as with the buffalo, it are large herds), but hunted more often than the encounter rate would suggest.
• Hartebeest were encountered more often than their abundance would suggest, but hunted less often than the encounter rate would suggest.
• Ostriches were encountered as often as their abundance would suggest, but hunted less often than the encounter rate would suggest.
• Elephants were encountered less frequently than their abundance would suggest and they were never hunted.

Figure 1. Variation in selectivity (Jacobs' index) for encounters with moving lions, hunts by lions, and kills by lions for the eight most abundant potential prey species in Addo Elephant National Park from December 2003 until November 2005.

Figure 2. Hunting success (number of kills observed divided by the total number of hunts observed) of lions in Addo Elephant National Park overall, and for the four species that were most frequently observed being hunted.

Conclusion:

Lions encounter preferred prey more frequently than expected (Table 1). Conversely, they hunt avoided prey less frequently than expected (Table 1).

Table 1:

Source: Hayward MW, Hayward GJ, Tambling CJ, Kerley GIH (2011) Do Lions Panthera leo Actively Select Prey or Do Prey Preferences Simply Reflect Chance Responses via Evolutionary Adaptations to Optimal Foraging? PLoS ONE 6(9): e23607. doi:10.1371/journal.pone.0023607

[creature386]

On the myth of lionesses doing all the work in a hunt:

Although it is often assumed that lionesses do most of the hunting, male lions are in fact considerably more effective and successful predators when it comes to targeting the largest prey—namely elephants and buffalo [23,25,26].

rspb.royalsocietypublishing.org/content/278/1722/3270.long

[creature386]

[creature386]

Aging in lions:
www.cbs.umn.edu/sites/default/files/public/downloads/Lion_Aging_Guide.pdf

[Vodmeister]

Size & weight of lions by population;

[creature386]

I only knew the table showing the size ranges, many thanks to @boldchamp!

[guategojira]

Hello to everyone. This is my first official post, and I will like to begin contributing with data. Here are my last tables about the body size and weight of the lions from East Africa:




The PDF document: www.scribd.com/doc/214211740/Valvert-2014-Body-Size-East-African-Lion

As we can see, the average weight overall is of c.174 kg for males and c.121 kg for females. This figures match perfectly with the previous figures from Smut et al. (1980) which stated an average weight of c.175 kg for males and c.120 kg for females, and those from Schaller (1972) which estimate an average male at 170 kg and an average female of 120 kg. This will rest the case that East African lions are the middle size population, been smaller than those of Southern Africa but larger than those in West Africa and India (Barbary lions would be of the same size).

Please take in count that the extreme weights of 180 kg for the females could include some stomach content. At this moment, I have not corroborated it yet, but I hope to do it soon.

Greetings to all.

[guategojira]

The size of the Barbary lion

Was the Barbary lion, the famous population in the north of Africa, a truly giant? Are those claims true? The next post show that the Barbary lion was of the same size than they genetic "brothers" of India and West Africa. None reliable weight has been reported from this population, but the few sizes and skull dimensions show that they are not exceptional in any particular characteristic.

1. Charles Guggisberg
It has been long debated which was the size of the Barbary lion. At this moment, the best source is the book “Simba, the life of the lion” of Charles Guggisberg (1961), check the image:


As we can see, the sizes reported are no larger than modern lions from East Africa, or a better comparison, of the same size than those from West Africa and India.

Some months ago, I buy the book “Wild Cats of the World”, also of Guggisberg (1975) and he put the same data about the Barbary lion that he puts in his previous book. Check the image:


So, I search the original source of the data which is this book:
* Brehm, A. E. 1925. Tierleben. Vol. 4. Leipzig.

After a few searches, I found it in the web here:
archive.org/details/brehmstierlebena12breh

The reference is different (year 1915, Vol. 12), but the data is exactly the same.

Now, check the measurements that Brehm states:


Brehm was probably one of the best Zoologists from Germany and the entire world. His data most be taken as mandatory and his statements are highly reliable, although now somewhat outdated.

Brehm states that the adult Barbary lions have a shoulder height of 80-100 cm, a head-body length of 160-190 cm and a tail length of 75-90 cm, which produce a total length of 235-280 cm. This size is similar to all the other lion populations and don’t present any exceptional dimension.

2. Nobuyuki Yamaguchi
Yamaguchi & Haddane (2002) made a good investigation about this lion, check it:

“How big was a Barbary lion? The famous French zoologist Cuvier measured a six-year-old captive-reared male Barbary lion, which had head and body length of 5 pieds 2 pouces (= c. 1.58 m), tail length c. 66.1 cm, height of forequarters c. 83.6 cm and of hindquarters c. 83.6 cm (Geoffroy-Saint-Hilaire and Cuvier, 1824). This lion was caught in eastern Algeria in 1795 at about one year old and died at ten years old in the Jardin des Plantes, Paris. Although the live lion may not have given Cuvier accurate measurements, the animal seems to have been very small for a male lion. It is, however, doubtful whether captive Barbary lions, usually captured as cubs and kept in menageries during the 18th or 19th centuries, attained the full body size. Cuvier himself referred to undesirable captive conditions at the menagerie (Geoffroy-Saint-Hilaire and Cuvier, 1824). Cornish (1899) reported that big cats only lived, on an average, for two years in London Zoo in the mid-1800s. Gérard (1856) also expressed his concern about the captive condition of lions at the Jardin des Plantes. On the other hand, he described a big wild Barbary lion he shot with the comment `This lion, compared to the finest of those which are exhibited in our menageries, or at the Jardin des Plantes, was what a horse is to a donkey. . .' There is, however, no credible record of body measurements of wild Barbary lions. Gérard (1856) described the size of wild male Barbary lions as c. 2.3 m from the tip of the nose to the root of the tail, which measured c. 90 cm, and their weight as c. 270–300 kg. If this had been true, Barbary lions would indeed have been big amongst lions. However, the methods of obtaining these measurements (e.g. straight or along the curve) were not specified, and the accuracy of the measurements themselves may be questionable, as Gérard made them in the field. Although Pease (1899, 1915) suggested that North African lions might have become very heavy because they fed on mutton so much, regarding the body length he seems not to have believed what he himself quoted – an Algerian lion whose head and body length was c. 2.5 m and the tail length 75 cm.”

“The largest Barbary lion skull so far measured, which is partly broken, has an estimated greatest length of c. 360 mm (Mazák, 1970; Yamaguchi, unpublished). Although 360 mm is not small, big skulls of sub-Saharan lions easily reach a maximum length of over 380 mm, and some even over 400 mm (Hemmer, 1974; Best, 1981; Yamaguchi, unpublished). Does this mean Barbary lions were not particularly big? Due to such a small sample size, we have to wait until more specimens may become available. The big lion Gérard shot in Algeria was presented to the Duchess of Orléans (Gérard, 1856), but the current whereabouts of this specimen and other wild-shot Barbary lions which decorated Gérard's Paris residence are not known.”
Source: www.izn.org.uk/Archive/321/Izn-321.htm#lion

Conclusion of Yamaguchi: there are no reliable sizes of wild Barbary lions, nor any weights. He presents a good case where he states that there is no evidence or at least enough specimens to say that this lion was larger than any other population. In fact, if we follow the new genetic evidence that prove that West African-Indian-Barbary lions form a single genetic subspecies (Dubach et al., 2013) and with the presented sizes, we can conclude that this lion was of the same size and weight than modern Indian and West African lions, two populations that incredible reach the same sizes despite the large geographical distance (average weight between 160-170 kg)

3. Skull size
About the skulls, Yamaguchi & Haddane (2002), quoting Mazák (1970) and they own data, states that the largest skull from this population was of c.360 mm, which is smaller than the maximum of other lion populations (South Africa, up to 419 mm according with Hemmer (1974)). However, a new investigation of J. H. Mazák (2010) provides new figures, check the image:


This table presents the following average for the population in the north of Africa:
Greatest skull length:
* North Africa:
Males – 372.3 mm (n=3)
Females – 318.3 mm (n=2)

The second sample from North East Africa (specifically Ethiopia, including those from Addis Ababa) is dubious on its classification, because some authors believe that they are descendants of the Barbary lion (Bruche et al., 2012), but the last genetic study show that they are, in fact, closer to the East African lions (Dubach et al., 2013), so its inclusion here will be incorrect.

Although small (n=3), the sample of Barbary lions show specimens that are clearly larger than the record stated by Yamaguchi, with a probable maximum between 410-420 mm. Even then, there are several records of lions with large skulls and relative small bodies, so this large skull size only suggest that Barbary lions were as large as other populations, but that they were no exceptional in any case.

Here are some images of a Barbary lion skull found in the tower of London:





Here is an image, with an available scale bar (5 cm), of the London Tower lion skulls:


The larger of the two skulls is the whitish one, and the scale shows that this one measured only 305 mm in length. Again, not an exceptional specimen; the darkest skull is even smaller.

The average for the Indian lions is of 338.9 mm (n=6) according with J. H. Mazák (2010) and of 344.0 mm using 4 specimens from Pocock (1939) and Prater (1921). The largest Indian lion skull available measured 365.8 mm (Prater, 1921). J. H. Mazák (2010) presents only one skull of 330 mm from Iran and one of 347.8 mm from West Africa. Obviously these samples are pretty small, but even then, it seems that the Barbary lions had larger skulls than those from Asia and West Africa. Could this mean that the largest specimens of the Barbary lions were larger than those from these regions? Probably they did, but we don’t know by how much. A study of the relation between skull length and body length of lions could help us with this problem, but we must take in count the possible inter-variations that present the different populations.

4. Conclusion
Using the little reliable information available, the Barbary lion (female and male) had a head-body length of 160-190 cm in straight line, a shoulder height of 80-100 cm and a greatest skull length with an average of 372.3 mm in males and 318.3 mm in females (c.420 mm probably maximum). These sizes are about the same than those of Indian and West Africa, with the exception of the relative longer skulls that resemble more those of the Sub-Saharan lions.

With the weight issue, judging by its body size and skull dimensions, Barbary lions probably weighed up to 200 kg (and much less in average, probably about 170 kg) although there is the possibility of some exceptional specimens of probably up to 230 kg like the East African lions, or even 250 kg like the Southern African lions, but this last figure will be probably just an exaggeration in the northern areas of Africa (poor prey base). There is only a single report of Gérard (1856) of males up to 270-300 kg, but these are simple estimations and are completely unreliable.

The idea of its large size came from the large mane, but there are captive Indian lions with heavy manes that weight less than 160 kg. There are several large lions with heavy manes in private facilities available in the web and the "Barbary-fans" proclaim without any evidence that they are “Barbary”, but the truth is that none of them is pure Barbary or from any other population (these captive lions are a mix of genes from several populations). Besides, any captive lion in cold climate can develop a large mane, so the mane is the worst factor to detect a Barbary lion (Patterson, 2007).

[Vodmeister]

Wow! Great info Guate!

Nice to have you here by the way.

[guategojira]

Mar 26, 2014 3:49:03 GMT 5 Vodmeister said:

Wow! Great info Guate!

Nice to have you here by the way.

Hello Vodmeister. I will go to every place that will welcome me.

Greetings my friend.

[Vodmeister]

All the data incidates that lions typically average 170-200 kg.

[guategojira]

In fact, the Sub-Saharan African lions have an average between 174-190 kg, depending of the population. For the West African lions the average is of c.170 kg and for those of India is of 160 kg.

Crater lions are good candidates for an average of over 200 kg, but sadly no weight is available (maximum chest girt reported is of 134 cm). On the other hand, Barbary lions probably were of the same weight than the West African population or a bit higher, like the East African ones.

The figure of 202 kg from Smithers & Wilson (1979) is reduced if we included all the other specimens from the same region (Zimbabwe).

[Vodmeister]

I remember how on AVA Boldchamp and MellivoraCapensis42 claimed that male lions in central Kenya averaged 203.3 kg, a sample which included 272 kg (599 lb!) specimen. Was this ever verified?