Vocalizations
Dholes make a wide range of vocalizations that include whines, mews, yaps, squeaks, screams, growls, growl barks, and chatter calls; these are mainly used for short communications from dog to dog. They are also known to do a “huu-huu” type call, similar to the African painted dog (Fox 1974, Maisch, Ludwig). For long-distance communication, such as getting the pack together after a hunt or rising them from mid-day naps, they make a whistle. This whistle is how they got their nickname “the whistling dog” and where Fox got his title for his field book on dholes, The Whistling Hunters. The whistle call is great for the dense forest environment, as it travels well at ground level due to the double frequency and structure of the whistle (Sillero-Zubiri et al. 2004).
Pack Behavior
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Before a hunt, dhole’s will do a pep-rally greeting. This can look like play or aggressive play and includes mock chasing, jumping and running (Acharya 2007).
The pack will use one set latrine; here, the pack defecates communally. They are often located just off of paths, game trails, roads or intersections. Dhole latrines appear to functions as a home range marker, so not causing these sites to disappear is important to help maintain normalcy in the pack (Acharya 2007). |
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Pack Structure
The pack population tends to go on a boom-and-bust system. The pack size will often fluctuate greatly due to disease and dispersal (Karanth 1999). Other causes for this fluctuation include dholes immigrating into a new pack, individual migrations, pups being born or not being born for a year (Johnsingh 1983). This immigration and dispersal patterns does cause a skewing in the gender ration to have packs more male-heavy (Johnsingh 1982 and Venkataraman 1998). Even with fluctuating numbers, it’s been figured that the average pack of 14 dholes would still need 60 to 70 square kilometers to be sustainable (Acharya 2007).
Who is included in a pack tends to be a nuclear family and possibly non-reproductive siblings of the adult breeding pair (Acharya 2007). The largest pack that has been confirmed is 40 individual dholes; this did include pups (Davidar 1975). It’s thought that these larger packs are not a single pack but rather a group of smaller packs that ban together for one reason or another and disperse once breeding season comes around. These large gatherings are sometimes called clans, and this concentration of dholes and then dispersal has been dubbed a fusion-fission system (Johnsingh 1982). When the packs are separate, they may live close enough together that their territories overlap but not the internal core areas (Johnsingh 1983).
Who is included in a pack tends to be a nuclear family and possibly non-reproductive siblings of the adult breeding pair (Acharya 2007). The largest pack that has been confirmed is 40 individual dholes; this did include pups (Davidar 1975). It’s thought that these larger packs are not a single pack but rather a group of smaller packs that ban together for one reason or another and disperse once breeding season comes around. These large gatherings are sometimes called clans, and this concentration of dholes and then dispersal has been dubbed a fusion-fission system (Johnsingh 1982). When the packs are separate, they may live close enough together that their territories overlap but not the internal core areas (Johnsingh 1983).
Breeding and Pups
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Dholes are capable of reproducing at as young as one and a half of years, but three years of age is more typical of wild dholes (Venkataraman 1998), with females being reproductively viable up to ten years of age (Madhavan 2013). The breeding season varies widely depending on what latitude they are found and the dry and wet seasons. In European zoos, breeding season was observed to be December to February and pups were born March to May after a 63-day gestation. |
When it comes to breeding, it’s normally the alpha male and alpha female that breed. This has led to the development of a gender separate hierarchy where the alpha male is dominant over the males in the pack but not the females and vice versa (Ludwig 2004). Breeding time is really the only time that dhole packs get aggressive, with the alpha female suppressing the other females to prevent them from breeding (Volodina 2004). Most of the time, it’s only the alpha male that breeds, but sub males in the pack do show sexual interest in the alpha female and may contribute to the paternity of the litter. It’s interesting to note that once the pups are up and running with the rest of the pack, there is little behavior or aggression to indicate the hierarchy of the pack (Macisch 2006).
Dholes need the support of the pack to help raise pups. Subordinate females and larger packs had a better success rate than smaller packs (Pandey 2013). Lone females breeding outside the group have little or no success in raising the litter (Johnsingh 1982). The non-breeding females might also produce a litter, and it will be up to the alpha female if the litter survives and is raised with the rest of her litter (Fox 1974). There have also been notes of multiple lactating females in a pack; they all tend to nurse the pups (Davidar 1974). It’s also important for lower-ranking dholes to help raise pups, as it helps increase the likelihood of success when raising their own future litters (Masich 2003).
Dholes will either dig their own den or utilize and modify a hole that another animal has made. These dens can be simple single entrance and chamber dens to multi-chamber and entry networks. These dens are often used year after year and sometimes for generations (Fox 1974). These dens need to be close to water and hunting areas (Fox 1974). If at any time the pack feels the den is threatened or compromised, the whole pack will help in relocating the pups and move them to a safe new location (Johnsingh 1982).
The time after pups are born and restricted to the den is called the denning period. Denning causes the dhole pack to localize close to the den. A pack that might use 75 square kilometers during the regular part of the year the average being 30 to 60. When the breeding female is nursing a litter that range shrinks to just its core and smallest area at 15 to 20 square kilometers. Once the pups are older or post denning dholes from earlier litters will disperse, this causes the pack’s territory to expand to its largest sometimes swelling to larger than 100 square kilometers (Acharya 2007). During the denning time, other pack members will bring meat back to the den both for the pups and the nursing female. The meat is carried back in the stomach and is then regurgitated for the pups and mom (Sillero-Zubiri 2004). Once the mother is out hunting with the pack, she will often monopolize small kills for herself (Johnsing 1982).
The day after the pups are born is the first test; the alpha male will enter the den and pick up the pup and give a slight shake. If they pup whines or moves indicating it’s alive, it’s put back with the rest of the litter. For pups that are dead, they are removed from the den and consumed (Macisch 2006). Litter size varies from two up to twelve (Johnsingh 1982).
Pups weigh 200 to 350 grams when they are born and they are 25 to 27 centimeters long, but they don’t stay small for long; dhole pups can double in weight by ten days of age (Paulraj 1992). They are covered with a dark brown fluffy undercoat and a few long, dark guard hairs, no teeth and their eyes and ears are closed. By the time they are ten to twenty days old, their eyes start to open (Davidar 1975). Around this time, their upper and lower incisors come in (Shilo 1994). The next teeth to come in are the canines at around 20 days of age, molars around day 23 and last are the premolars around day 30 (Shilo, 1994, Sosnovkii 1967). For the first two months, the mother stays with the pups almost full-time, and she is fed by the pack (Johnsingh 1982). At five weeks, pups start to venture outside the den and are take meat from pack mates returning on the hunt (Davidar 1974); however, pups might take meat as young as 3 to 4 weeks of age (Sillero-Zubiri 2004). Dholes are fast to grow up and mature, and they have to be, because at 3 months of age, dhole pups are out from the den and following adults on hunts (Sillero-Zubiri 2004). It’s also around this time that the pups start to lose that brown fur color as it turns to the iconic red color (Paulraj 1992). Dhole pups are full-grown around nine to ten months of age (Shilo 2007), and around a year of age is when they get their mature coloring (Paulraj 1992).
Dholes need the support of the pack to help raise pups. Subordinate females and larger packs had a better success rate than smaller packs (Pandey 2013). Lone females breeding outside the group have little or no success in raising the litter (Johnsingh 1982). The non-breeding females might also produce a litter, and it will be up to the alpha female if the litter survives and is raised with the rest of her litter (Fox 1974). There have also been notes of multiple lactating females in a pack; they all tend to nurse the pups (Davidar 1974). It’s also important for lower-ranking dholes to help raise pups, as it helps increase the likelihood of success when raising their own future litters (Masich 2003).
Dholes will either dig their own den or utilize and modify a hole that another animal has made. These dens can be simple single entrance and chamber dens to multi-chamber and entry networks. These dens are often used year after year and sometimes for generations (Fox 1974). These dens need to be close to water and hunting areas (Fox 1974). If at any time the pack feels the den is threatened or compromised, the whole pack will help in relocating the pups and move them to a safe new location (Johnsingh 1982).
The time after pups are born and restricted to the den is called the denning period. Denning causes the dhole pack to localize close to the den. A pack that might use 75 square kilometers during the regular part of the year the average being 30 to 60. When the breeding female is nursing a litter that range shrinks to just its core and smallest area at 15 to 20 square kilometers. Once the pups are older or post denning dholes from earlier litters will disperse, this causes the pack’s territory to expand to its largest sometimes swelling to larger than 100 square kilometers (Acharya 2007). During the denning time, other pack members will bring meat back to the den both for the pups and the nursing female. The meat is carried back in the stomach and is then regurgitated for the pups and mom (Sillero-Zubiri 2004). Once the mother is out hunting with the pack, she will often monopolize small kills for herself (Johnsing 1982).
The day after the pups are born is the first test; the alpha male will enter the den and pick up the pup and give a slight shake. If they pup whines or moves indicating it’s alive, it’s put back with the rest of the litter. For pups that are dead, they are removed from the den and consumed (Macisch 2006). Litter size varies from two up to twelve (Johnsingh 1982).
Pups weigh 200 to 350 grams when they are born and they are 25 to 27 centimeters long, but they don’t stay small for long; dhole pups can double in weight by ten days of age (Paulraj 1992). They are covered with a dark brown fluffy undercoat and a few long, dark guard hairs, no teeth and their eyes and ears are closed. By the time they are ten to twenty days old, their eyes start to open (Davidar 1975). Around this time, their upper and lower incisors come in (Shilo 1994). The next teeth to come in are the canines at around 20 days of age, molars around day 23 and last are the premolars around day 30 (Shilo, 1994, Sosnovkii 1967). For the first two months, the mother stays with the pups almost full-time, and she is fed by the pack (Johnsingh 1982). At five weeks, pups start to venture outside the den and are take meat from pack mates returning on the hunt (Davidar 1974); however, pups might take meat as young as 3 to 4 weeks of age (Sillero-Zubiri 2004). Dholes are fast to grow up and mature, and they have to be, because at 3 months of age, dhole pups are out from the den and following adults on hunts (Sillero-Zubiri 2004). It’s also around this time that the pups start to lose that brown fur color as it turns to the iconic red color (Paulraj 1992). Dhole pups are full-grown around nine to ten months of age (Shilo 2007), and around a year of age is when they get their mature coloring (Paulraj 1992).
Hunting
Dholes are like other large pack canines and are hyper-carnivores, meaning their diet consists of over 70% meat. It’s thought that prey availability is one of the largest restriction for dholes in terms of habitat and pack-size restriction (Johnsingh 1985).
Prey Selection
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Prey choice in dholes has been well studied in many areas of its home range. It appears that two ungulate species in particular play an important role in dhole diets: sambar deer and chital deer. In many studies, these deer are the top two prey items taken by dholes. In most cases, it appears that dholes target male chital deer and fawns, and also targeted sambar sub-adults and fawns (Acharya 2007, Johnsingh 1908, Karanth 1999). The male deer might be targeted due the fact they have antlers when they are hunted. Male deer are known to range more wildly during the rut period; they are often alone and no longer have the benefit of the herd for early detection of predators (Patel 1992). It was also noted that the male deer that were taken with antlers tended to have a larger set of antlers. It’s been thought that the larger antler set might make it harder for the male deer to escape, considering the dense vegetation (Johnsingh 1983). In areas where chital and sambar are not available, dholes will turn to other medium- or even small-size game (Jenks 2012). Nepal gives a great example of an area that had dholes but no sambar or chital; the dholes there rely on a few different prey items with wild boar being the top taken food item followed by barking deer, musk deer, blue sheep and goral (Aryal 2015). In Malaysia, dholes again took wild pigs, but only occasionally; instead the main diet for dholes was mouse deer (Kawanishi 2008) In Loas, it appears that dholes didn’t favor a single prey item (Kamler 2012). Sadly, when their native prey is not available or is in low numbers, dholes will take livestock. This puts them into direct conflict with humans (Aryal 2015). In Nepal, it was noted that livestock makes up to 13% of a dhole’s diet (Aryal 2015).
Dholes appear to be opportunistic when it comes to prey choice. They do seem to have a preference for medium-size (26 – 100kg) ungulates. In areas where their preferred prey live, dholes take down prey averaging 55.3kg (Acharya 2007). The HuntHunting with dholes is a family affair, and almost all dholes in the pack help with the hunt. With each dhole estimated to eat 340kg of food a year (Karanth 1988), hunting seems to be a priority for these hyper-carnivore canines. Dholes are almost entirely diurnal and usually hunt during mornings and evenings. There are a few accounts of dholes hunting on full moons, but they tend to avoid the heat of mid-day (Johnsingh 1983, Acharya 2007). Unlike the wolf or the African painted dog, Dholes are more of an ambush predator chasing their prey down over a short distance, normally less than 500m (Johnsingh 1983).
Dholes have evolved a unique strategy to tackling larger prey than themselves. Some believe the pack formation is for hunting large game, but Venkataraman (1995) showed that there was no correlation between pack size and body weight of prey killed. In fact, he showed that the larger the pack meant there was less food per dhole when a kill was made. And in areas were dholes take down smaller prey items, like in Malaysia, dholes form smaller packs as it might be more energetically advantageous to do so (Kawanishi and Sunquist 2004). Instead of large numbers, dholes use the terrain and geographical features of their landscape to aid in the hunt. In India, it was noted that dholes often killed their prey in or just around waterways. At first, it was thought to be due to the fact that prey will run downhill and into valleys to out-run predators; however, the kills happened so often that it didn’t appear random. Feral dogs hunting in the same area as packs never made kills in waterways (Peacock 1928). Dholes are more water-loving than most dog species, so it could also be safer to take large prey in water compared to land. The water both tires the prey quicker but also makes it harder for the prey to kick out and injure a dhole. Once the prey is tired and is back on land, or even when the prey is still in the water, the pack will start feeding on the animal even if it’s still alive. Most prey appear to die of shock from blood loss, not from a single killing bite, like a tiger inflicts. In some instances, the prey drowns (Karanth 1999). Once the prey is down, all pack members feed at the same time. Pups and adults eat together and there is little in-fighting amongst the pack. In large packs and on large kills, it’s not uncommon for a lone dhole to stand watch. Tigers, leopards and humans have all been noted to take a dhole’s kill even with dholes were feeding on it (Karanth and Sunquist 2000). To help prevent a theft of their meal, dholes will eat their prey quickly. The change in dentation may allow for dholes to eat and tear off meat more efficiently, oftentimes finishing off a carcass in ten minutes. Even the small pups will gorge themselves at meal time; it’s been observed that even a 4kg pup can eat up to 1kg of meat (Maisch 2006). Once they have their fill, the dhole will leave the kill site. They do not cache their food and will often leave the kill out in the open (Acharya 2007). Dhole’s hunting success rate is about 20% (Keller 1973) or about 1 out of every 5 tries, which puts them at about average for carnivores. It hasn’t been studied much if a larger pack has a higher success rate than smaller packs. Due to the elusive nature and rugged environment dholes live in, it might be hard to figure this out (Venkataraman 1995). Dholes play a unique role in their ecosystem, both as an ecological indicator and as a regulator. Dholes, being a large predator, more due to their pack structure than individual size, require large numbers of prey to survive. They are highly sensitive to change in prey base and habitat quality (Acharya 2007). If an area has a healthy population of dholes, the area is most likely ecologically healthy. However, if an area’s dhole population declines, like they are across most of their range, it should be a warning there is something wrong in the ecosystem. It’s also thought that dholes play a role in prey regulation through trophic cascades that aren’t filled by other carnivores in the area (Woodroffe and Ginsberg 2005). |
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