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ON THE EDGE OF EXTINCTION - A - THE TIGER (Panthera tigris)
ON THE EDGE OF EXTINCTION - A - THE TIGER (Panthera tigris)
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ON THE EDGE OF EXTINCTION - A - THE TIGER (Panthera tigris)
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B - LANGS TIJGERPADEN (L. Denninghoff Stelling, The Hague, 1966, 233 pages)
 *This image is copyright of its original author 1 - INTRODUCTION While we're at it (Sumatran tiger and sun bear), this one should be discussed as well. It is one of of the best books about tigers I read. Although Denninghoff Stelling wasn't as able with his pencil as Corbett, the book is vividly written and very interesting. It also has some good photographs. In the days Indonesia was a Dutch colony, many left Holland to work in the 'Empire of Insulinde', as Indonesia was known. Many never returned. The great grandparents of Denninghoff Stelling moved to Indonesia a long time ago. His grandparents probably were born there and so was his father. I don't know of Denninghoff Stelling was, but it seems very likely. In the early fifties of the last century, Indonesia became independant. Many thousands were shipped back to Holland. I remember the big passenger ships in the Amsterdam harbour. Although many ended up in The Hague, Amsterdam also was popular. I know quite many who were born in Indonesia. Some of my friends were and so was Dr. P. van Bree. Some were white, some were Asian and others were of mixed origin. They thought and acted as we did and they did not. They interacted with their new neighbours and they did not. They adjusted and they did not. In many cases, they kept to themselves. Friendly and silent, they often were. When I got older, I came to the conclusion that many just never found their way over here. They never really left Indonesia. Their children, although as Dutch as it gets, often visit Indonesia. At times, they meet a partner. Some return and some don't. As a child, Indonesia seemed like a far away paradise to me. My guess is many of those who had to return to Holland felt the same way. They seemed to long for Indonesia and never got over it. Denninghoff Stelling probably was one of them. He dedicated his book to his father and his friends in Indonesia. The ones with whom he walked through the steaming jungles of Sumatra. I don't know when he left Indonesia, but my guess was he returned somewhere in the early fifties. In Holland, a country he probably had never visited before, he wrote a book about his experiences in Indonesia. It was published 15 years after he had left the country in which he had lived for so long. The introduction, written by Tjalie Robinson (probably also born in Indonesia), is interesting. He said there's only one kind of hunting (on foot), only one kind of motivation (to be one with the natural world) and only one real hunt. The reason tiger hunting was considered as the ultimate hunt was the tiger himself. In Indonesia, many saw him as a magician. He could mimic all other creatures, he could select and kill the most able of men and yet spare the playing child, he could be a human if he wanted to and he could outwit anyone trying to hunt him. Hunting tigers wasn't considered a sport. It was much more than that. To Robinson, the book isn't about hunting tigers. It is a study about them. Denninghoff Stelling's great grandparents hunted tigers. Same for his grandparents and his father. Every generation trained the new one and in the end the ultimate hunter was born. The young Denninghoff Stelling was trained to have a career. He had one. But one day, he left and never returned. He sailed to Sumatra. Like all true hunters, he couldn't stay away from the forest and the animals. He caught, shipped and killed many and yet he respected them, more so than his friends. They were his peers and talked to him. It is clear Denninghoff felt strongly connected. In his day, it was considered as out quite something to leave a career and enter the forest for a living. Not only because he gave up what he had (a lot), but also because he entered a world feared by many. In the twenties and thirties of the last century, Sumatra still had very large tracts of forest. It also had dangerous animals. The newspapers often had weird stories about large unknown animals (like the lion-tiger), giant snakes and crocs, killer elephants, ruined estates, deserted villages and countless man-eaters. The result was Sumatra became known an alien place. One you should stay away from. 2 - LIFE AS A CATCHER AND HUNTER Denninghoff settled somewhere in the southeast of Sumatra. Initially, he hunted deer and sold the meat. Later, he caught, sold and shipped animals. Sun bears, tigers, birds, small cats, monkeys, tapirs, crocs and snakes. Pythons in particular were easy to sell. The longer, the better. Quite many reached a length of 17-20 feet and their heads were large. Denninghoff and his friends often travelled to unknown places. He saw villages where a hunter had never set foot. When travelling, they used the rivers. And they walked, at times up to thirty miles a day. He noticed the people he met were more developed and aware than those on Java. Different culture. Men and women has equal rights. Catching, selling and shipping animals took much more time than hunting. When he hunted, he often went for deer and elephants. Meat and ivory were easy to sell. Not many were interested in tigers and tiger skins. Apart from that, they were considered as dangerous everywhere. Denninghood, however, hunted them whenever he could, sometimes for months at an end. In Sumatra, hunters used a 'belor' (a long stick with a light at the end) when they were sitting over a bait. Most animals didn't seem to be bothered, but tigers and, in particular, bears didn't like it. When he got a reputation, people in remote villages asked him to hunt the animals who raided their fields. He also hunted man-killers (very often roque elephants) and worked for estate-owners troubled by wild boars, sun bears and tigers. 3 - SUN BEARS In general, estates in the southeast of Sumatra (the first region to be developed) were attractive for sun bears. The reason was food. Sun bears had to be hunted in daylight. And not on foot if it could be prevented. The father of one of the trackers of Denninghoff Stelling, when walking beneath an overhanging branch, startled a bear. The bear immediately attacked, shredded his belly with the claws of his hind legs and bit his skull into small pieces. After the poor man had died, the bear decided to become afraid. He left the man screaming like a madman, running for his life. According to Denninghoff, bears were more courageous than tigers. They also were more loyal to their partner. If one was in trouble, the other would come to his assistence, no matter what. The front paws of a sun bear compared to those of the tiger and his hindlegs were even more terrible. When shot at a vital place, they often managed to get away. One, after his heart had been shattered by a bullit, managed to walk 15 yards and hide. Denninghoff was amazed at their strength and power and considered them way more dangerous than tigers. When in a bad mood or at the wrong place, they would attack on sight. 4 - GOLIATH AND THE SUN BEAR When working in a remote part of an estate, Denninghoff Stelling saw the largest tiger he had ever seen. He described his head as monstrous. He shot the tiger, but never saw him again. Others found his remains. It wasn't the only large tiger he saw. On an estate known as 'Bekri', a very large male tiger he called 'Goliath' was one of the tigers he was after. During a number of days, they followed his trail. His pug marks were the largest he had seen. One day, when following the tiger, he came to a place where something had happened:  *This image is copyright of its original author  *This image is copyright of its original author Here's the rough translation: " ... We followed the trail for about a hundred yards, when it suddenly stopped. I returned to the last prints ... and saw that the tiger had left the trail abruptly and entered a garden. The distance between the prints become shorter, as if he had advanced slowly. Then they suddenly became wide apart, as if he had taken very long jumps. They stopped at a place where large claws had left many gashes in the ground. This was the place where a fight had taken place. There was blood everywhere. Goliath had won the fight. From the battlefield, something heavy had been dragged to an alang-alang field a hundred yards wide. Directly behind the field, the forest started. We expected to find the remains of the animal killed by Goliath, but only found bear skin, bear claws and gnawed bones. I became enthousiastic. Tigers hesitate to attack bears. The reason is these innocent creatures, very solidly built and armed with strong teeth and dangerous claws, are more than capable to defend themselves ... " (pp. 187-188). The bear had been completely eaten. In order to get to the giant tiger, Denninghoff Stelling shot a large wild boar. He was to be his next meal. The next day, they found that Goliath had broken the rope and taken the heavyweight in his jaws. He did it in such a way, there was hardly a drag visible. It took Denninghoff and his tracker Oesin some time to get to the place where the giant had decided to start his meal. The wild boar had been completely eaten, but Goliath had not been the only one who had feasted. They found the pug marks of a small tigress close to the boar. They decided to sit up over the left-overs, lighted the 'belor' and thought it would be wasted energy. To their surprise, Goliath returned to his kill. But he didn't show himself. He made a detour, discovered the machan (called a 'prangong' in Sumatra) and protested for a long time before leaving. But minutes later, still in the prangong, Denninghoff saw him out in the open in a nearby field. The tiger wasn't going to give in and waited for them to leave. When he approached his kill, Denninghoff Stelling, aiming at the only thing he could see, the eye, didn't miss. It proved to be a giant tiger:  *This image is copyright of its original author Here's a rough translation of what he saw: " ... I had Goliath! What a giant and how perfect he was. His skin was beautiful and dark, a clear sign he still was in his prime. There was not a scar visible on his skin, which was unusual for animals of his size. My shot hadn't damaged anything. What a trophy! I couldn't stop looking at him. His head was as large as mine and my torso together and his canines were thick and undamaged. He measured 295 cm. Not as large as the other, second, male I had shot at the estate, but he was close. He was, however, more beautiful and as heavy as a result of his robust built. I estimated he was about twice as heavy as the wild boar ... " (pp. 199-200). The same day, a little later in the evening, he went to see a friend who had sat up in a different corner of the estate. His tigress had not showed. When he saw Goliath, he said he didn't know tigers could grow to such a size in Sumatra. Neither did I:  *This image is copyright of its original author Have a good look at this tiger and remember Denninghoff Stelling was as experienced as they come. Also remember he wasn't out for records. The only measurement in the book is this one. Like many of you, I think he could have been measured 'over curves'. Maybe he was measured in a generous way, although Denninghoff Stelling wasn't involved in exaggerations anywhere in the book. He just said it as it was. In northern India, tigers were also measured 'over curves' as a rule. It was done in a very careful way, as measurements were sacred over there. In all cases, the difference between both methods used ('between pegs' and 'over curves') was 2-5 inches. But in the largest tiger shot by Col. Stewart Capper (an 'immense animal'), the difference was no less than 12 inches. Let's assume the difference between both methods was somewhere between 6-12 inches in Goliath. That would still leave a total length of 8.8-9.2 (264,16-279,40 cm.) 'between pegs'. Unheard of for a Sumatran, but he wasn't the only one who reached that size. The tiger he had shot on the same estate (the one he had not been able to find after he had shot him) could have been even larger. Also remember a renowned hunter in Indonesia (a man called Pieters) stated he had shot males of up to 300 cm. in total length and 185 kg. (Hoogerwerf, 1970). V. Mazak (1983) wrote males ranged between 100-140 kg. I don't doubt he was right. But in the days there were ten tigers to every one today, things could have been different. In the ZMA (the former Zoological Museum of Amsterdam), I measured a skull of 350,00 mm. in greatest total length. Not saying the owner would have been as large as an average male Indian tiger (Sumatran tigers have relatively larger skulls), but there's too much information about very large animals to ignore. Also remember I talked to men who had hunted tigers in Indonesia. They had seen a lot and were convinced some males were only marginally smaller than an average male Indian tiger. And yes, they had been in India. I'm not going to speculate about the weight, but it is clear the tiger was a very robust male, well over average in many respects. Even without the wild boar, he would have been quite a bit heavier than 'Slamet', who was 148,2 kg. I compared both photographs and there's no doubt Goliath was a larger animal. Denninghoff Stelling thought he was about twice the weight of a large male wild boar. In order to show you wild boars in Sumatra were anything but small, two more pictures to finish the post:  *This image is copyright of its original author  *This image is copyright of its original author I have no reliable records of Sumatran wild boars, but the photographs I have suggest large males could have been 80-100 kg. (177-221 lbs.). Some no doubt exceeded 100 kg. 
Massive Sumatran Male. What a tiger and great info Peter.
Sumatra is such a unique area, before we destroyed it, it was so vast with its wild life. Very cool to here the stories from old hunters. Sad that such a beautiful tiger was taken in his prime, but such was the times. TFS
SLAMET AND GOLIATH COMPARED
I said I had compared of Slamet and Goliath and concluded Goliath was a larger animal, but you decide for yourself. Here's the two photographs. 1 - SLAMET I have no information about his length, but Slamet was 148,2 kg. A bit over Mazak's maximum for Sumatrans:  *This image is copyright of its original author 2 - GOLIATH At 295 cm. in total length (very probably measured 'over curves'), Denninghoff Stelling estimated him to be twice the weight of a large male wild boar. But what is the weight of a large male wild boar? Remember Slamet was empty when they darted him, whereas Goliath and a small tigress had consumed most of a large male wild boar hours before he was shot. His belly is clearly extended. In spite of that, it is clear Goliath was a more robust animal:  *This image is copyright of its original author Goliath was closer to the camera than Slamet, but the angles in both photographs is not that different. I don't know how tall Denninghoff Stelling was, but the Dutch were not short. Maybe he was similar to the second man on the left in the first photograph or slightly shorter. 3 - WEIGHT When I first saw the photograph of Goliath, my estimate was 160-175 kg. empty. After comparing both photographs, I would get to the upper end of it (170-175 kg.). As to the reasons of my estimate. Reliable records say Sumatran males are a bit longer in head and body than many expect (160-176 cm. in a straight line). Tails usually are short (not over 80 cm.). If Goliath really was 264-279 cm. in total length in a straight line and his tail would have been about normal in length, it could, at his size, have measured 75-85 cm. This means his head and body length would have been 180-200 cm. A bit over the top, I concluded. Let's assume I overestimated his length. Maybe he was 10 cm. shorter 'between pegs' than I thought (254-269 cm.). Let's assume his tail was 75-85 cm. If so, he would have been 170-195 cm. in head and body. Assuming he would have had about 0,8-1,0 kg. per cm. in length, this would have resulted in 136-195 kg. I have no information about Sumatran tigers exceeding 185 kg., but we know Slamet was 148,2. As we also know Goliath was a larger animal, this would narrow it down to 148-185. If Goliath really had about 50 lbs. on Slamet, I would get to 170 kg., maybe a bit more. Let's say 170-180 kg. empty. As we also know photographs and overestimates usually get along just fine, I propose to deduct 10 kg., which would result in 160-170 kg. As Denninghoff Stelling estimated him to be twice as heavy as a large male wild boar, I added 5-10 kg., resulting in 165-180 kg. I propose 380 pounds (172,37 kg.) empty as the best guess for now. But. Just suppose the measurement of Denninghoff Stelling was an accurate measurement taken 'over curves'. Also suppose my initial estimate regarding his total length 'between pegs' (264-279 cm.) was right and the tiger was as robust as I think he was. If so, it can't be excluded the tiger was at least 180 cm. in head and body length and 400 pounds or slightly over in weight. The photograph of Denninghoff Stelling compares to the photograph of the Sungari River tiger in that both (to an extent) prove that records of large tigers in the past can't be dismissed out of hand. Some Amur tigers might have approached or exceeded 650 pounds and some very large Sumatran tigers might have topped 400 pounds. 4 - UNKNOWN JAVA TIGER Now that we're at it, what about this tiger? Skullwise, Java tigers had slightly better averages than Sumatrans, but they, like today's Amur tigers, also showed less individual variation. Based on the photographs I saw, I would say Sumatrans were a bit more compact, especially in the body, whereas Javan tigers, apart from longer skulls, had longer and more muscular limbs. Javan tigers could have been closer to mainland tigers (longer, taller and more athletic). What would be your estimate for this male?  *This image is copyright of its original author It would help when the three photographs, angle and distancewise, could be compared. If someone would succeed, we could use Slamet's weight as a start. Was he heavier than Slamet? If so, by how much? Why? 
A brand new study on tiger taxonomy is attached, "Planning tiger recovery: Understanding intraspecific variation for effective conservation". Kitchener is involved in this one as well. The conclusion is that only to subspecies exist, the mainland and the island forms. I think Kitchener has already proposed something like this in another paper, I cannot remember.
Excellent information Tigerluver, great paper, I need to read it right now.
About Dr Kitchener, in fact he was the FIRST in propose this new point of view (only two or three subspecies). The first time was published in the book "Riding the Tiger" of 1999, in chapter 2 "Tiger distribution, phenotypic variation and conservation issues". This chapter concluded that the taxonomy of the tiger, popularized by Mazák (1981) was based in too few specimens that it was probably invalid. He claimed that only two or three subspecies existed: mainland, island and Caspian. Latter in 2010, in the book "Tigers of the World" in chapter 3, he repeated exactly the same, although he improved some parts with new studies and with the help of Yamaguchi. This time, although he claimed again that "2 or 3" subspecies existed, he inclined more to the idea of two, as the genetic analysis showed that the Caspian tigers are the same than the Amur ones. This hypothesis was summarized by Valmik Thapar in his book "Tiger: the ultimate guide" of 2005. Well, I am going to read the paper between today and tomorrow and I will put my appreciations in Sunday. Greetings.
The issue with subspecies classification is that there is no unanimously accepted definition as to what a subspecies is. Until the taxonomic community can set a concrete definition, the conclusions of such genetic studies are more based on author interpretation than anything else. The Kitpipit et al. (2011) study (which Kitchener coauthored and I've posted here a while back) states that the genetic differences give reason to "subspeciate" eleven forms.
Wow, imagine that, eleven forms!
I read some time ago, from a study of Dr Stephen O'Brien, that the genetic differences between tiger population and lion populations (intra-specifically, of course), are smaller than that between human populations. Using also skulls only, don't provide all the image, check how different are the skulls from a human in Europe from one from India, Namibia or Australia (native ones). Still, we are all humans, despite our external differences and colors. I think that, at some point, tigers are about the same, with the only difference that the Island population is different from the mainland one, just like the Homo Floreciensis was different from the Homo Sapiens or the Neanderthal cousin. Dr Kitchener proposed a difference of 75% for been a "subspecies", but if we follow this, many accepted "subspecies" will be only distant populations of a same population, just like jaguars for example (there is only one single species with no subspecies, just clinal variations).
The sequence(s) being tested also matters. For example, Xue et al. (2015) couldn't subspeciate P.t. jacksoni based on certain types of genes, but was able to on others. Kitchener has limited himself to certain genes only, and this could cause some error. Also, the citation for my last post was supposed ot be Xue et al. (2015) and the number of forms is 9.
Clinal variation is dangerous to conservation in my opinion. There are at least distinct ecotypes between the Bengal, Southeast Asian, and Amur forms just based on the body characteristics (which are coded by genes). Drop off an Amur tiger population in southeast Asia and you likely won't be able to replicate the Southeast Asian tiger form within enough evolutionary time. In that manner, I feel these ecotypes are unique, even if they at this point are man-made.
This is also interesting to know, thank you Tigerluver.
Now, I have not read the full document yet, but from the news articles and a few lines that I have read, I think that although they recognize only two "subspecies", they also propose three conservation units, separating the Amur-Caspian tigers from the southern Asian population, based in the fact that they are more than 1,000 km from the nearest population. I think this is correct, as the population of the Amur-Caspian tigers separated from the other mainland tigers about 12,000 - 15,000 years ago, about the same tiger than Bengal tigers, but at difference than this last one, they don't sustain any genetic interchange until about 2,000 years ago, when the Amur tigers invaded Manchuria and they contacted again with the South China tiger population. I insist, if you see the skulls of the Amur tigers, they look very different from other mainland populations, but if you see those of the Bengal, which sustained constant interchange with all the other southern tigers, they look very similar with not much differences. In this case, Amur tigers will be managed separately, Sumatran tigers also follow the same way. while all southern Asian tigers (Bengal, Indochina, South China and Malaysia) were managed as a single population. Here are the images from the document itself:  *This image is copyright of its original author  *This image is copyright of its original author The problem are the South China and Indochina tigers, which captive populations are very small with less than 100 specimens between the two, while the wild population of Indochinese tiger is probably less than 1,200 specimens (South China tigers are extinct in the wild). This new document suggest reintroduction of genes, in this case, Indochinese genes to the captive South China tigers (specially those from South Africa, as they are already wild and are a viable population, but they will surely need more genetic variation to they pool), or Bengal genes to the Indochinese captive population. Previously, all south Asian tigers, from India, Indochina, South China and Malaysia were considered P. t. tigris, and this paper suggest the same (I think). By the way, check this new paper on the genetic of Indian tigers, from July 2015 (pretty new too): Demographic loss, genetic structure and the conservation implications for Indian tigers. Full paper: http://rspb.royalsocietypublishing.org/c...6.full.pdf I think we have enough material to read and to discuss, which is excellent. [img]images/smilies/tongue.gif[/img]
TIGERLUVER AND GUATE
Very interesting articles! Much obliged. I quickly read the article of Balkenhof, Fickel, Kramer-Schadt and Kitchener when it was printed. An interesting evaluation of a number of recent articles on tiger evolution and taxonomy. They are no doubt right. In spite of that, I think a number of issues should be discussed. One of these is definitions. 
So two intraspecies clades for Panthera tigris tigris; the northern clade (Amur-Caspian) and the southern clade (Chinese-Bengal-Indochinese-Malay).
And Bengal tiger and other southern Asian tigers are now just considered as the regional variations, maybe some Bengal tigers like the Sunderban tigers are no more closer to the mainland Bengal tigers than to the Chinese/Indochinese tigers.
I could not say it better, two subspecies and one of them divided in two clades:
1. Mainland tiger - Panthera tigris tigris: Northern clade - virgata. Southern clade - tigris. 2. Sunda tiger - Panthera tigris sondaica Good description GrizzlyClaws.
- I bet that if we have enough sample for the Chinese/Indochinese specimens, then the genetic distance might show that the Sunderban tiger could just sit in between the mainland Bengal and the Chinese/Indochinese.
- The Ngandong tiger could be in fact an extinct clade of Panthera tigris sondaica. - Panthera tigris acutidens could be the progenitor for both Panthera tigris tigris and Panthera tigris sondaica.
In fact, it is, the Wanhsien tiger southern population gives origin to modern tigers overall, even the Ngandong tiger came from it. The South China tiger original population (described by Luo et al. (2004) by a single nucleotide) was probably the closer thing of a Wanhsien tiger specimen.
The Ngandong tiger was just the beginning of the Sunda subspecies, because it had the large size from mainland but also the characteristics of the island tiger skulls.
Wanhsien tiger/Panthera youngi/Manchurian mandible: almost identical
Wanhsien tiger/South Chinese tiger: extremely similar Wanhsien tiger/Ngandong tiger: very similar but with some minor differences Wanhsien tiger/Modern tiger: quite different Ngandong tiger/Modern tiger: quite different The diversification of the tiger species is simply intriguing, but as you said, Homo sapiens just shows the same kind of diversification as Panthera tigris. |
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