From HofWiki

Ecology

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"Whatever may be in store for old Gandalf, I'll wager it isn't a wolf's belly."
-Sam Gamgee, The Fellowship of the Ring

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Habitat

If all species and sub-species are considered, canids are found throughout Middle-earth. They can adapt to many environments and some have even been found to thrive in areas of Middle-earth, despite significant persecution by humans.

Wolves

True wolves are geographically located throughout most of Middle-earth, and in a wide range of habitat types (although less often in the south). They are found in dense lowland forests, mountainous forests, boreal forests, tundra, temperate grasslands, and prairies. They usually require areas that are less populated by humans, as humans are the most significant source of mortality facing wolves (aside from the occasional territorial battle with hybrids). Although much smaller than hybrids, true wolves operate in much larger packs and can efficiently defend their territories from their larger cousins.

Hybrids

Despite the fact that hybrids are large and ferocious in their behavior and hunting tactics, this sub-specie is relatively rare in Middle-earth. They generally avoid humans (although have been known to exist near urbanized areas in heavily wooded northern habitats), and avoid true wolves when possible. They seem to prefer dense old-growth forests, with many trees and little under-story vegetation (i.e., plants growing at the base of trees, such as grasses, shrubs and raspberries). Because of this, they are more frequently found in Northern Middle-earth and most of Mirkwood, outside of the Wood- elf's realm (Note: While Elves do not spare hybrids when encountered, they feel great pity for them. This is because hybrids are creatures that have been indirectly perverted by the Dark Lord and, although dangerous, act only on the survival instincts instilled in them by Morgoth and Sauron.). They also exist in the northern Misty Mountains, the Grey Mountains, and the boreal forests north of there.

Wargs

Wargs are a captive animal and are not found existing on their own in the wild. They are normally found near or within the dwellings of orcs, goblins and other servants of evil. Therefore, you are more likely to find Wargs in the realms dominated by the minions of the Dark Lord (such as Angmar, the northern Misty Mountains, and near Mordor).

Preferred Food and Hunting Tactics

As said previously, as in most carnivores, canids consume only animal material. Furthermore, because they are endothermic, most must hunt and eat constantly to maintain their internal physiology. Therefore, not only are many aspects of their anatomy highly evolved specifically for hunting prey, even their behavior has evolved to help in the efficiency of prey capture, while still reducing the amount of energy spent during hunting. To insure that the energy acquired from food exceeds that expended during hunting, all canids will readily stalk prey that is larger than themselves (e.g. taking the time to stalk and kill an animal the size of a deer, which results in more calories consumed as food, is more efficient than wasting the same amount of energy and time hunting an animal the size of a mouse, although smaller animals are eaten in leaner times).

True Wolves

The preferred food items of wolves are mostly warm-blooded prey. This includes deer, harts, elk, moose, caribou and other smaller mammal species. True wolves hunt in large packs and work in a coordinated fashion to bring down large prey. Usually, this includes carefully selecting the sick, weak, or old members of a prey animal's herd and making a coordinated effort to separate this individual from the pack for killing. Because the wolf is an endotherm, it must consume large amounts of meat to fuel its internal metabolism. Therefore, a wolf can consume over 9 lbs of food in a single sitting.

Wolves are very shy and usually run from humans and dwarves. Furthermore, and despite popular belief, they also very rarely attack humans. Despite the fact that some farmers believe wolves to have a negative effect on their livestock, this is largely untrue. While wolves have been reported to attack and kill domestic animals on occasion, this usually only occurs in areas where humans have significantly encroached upon the wolf's territory or reduced the wolf's food source through the over-hunting of deer, and the like. In these situations, hungry wolf packs may resort to attacking and consuming livestock as a necessity for survival. However, most canid attacks on livestock are actually due to hybrids and not true wolves. Telling the difference between whether a true wolf or a hybrid has attacked domesticated animals is very easy: wolves work silently when hunting and sneak up on their prey. Hybrids, on the other hand, use surprise and brute force to catch prey. Therefore, a hybrid attack on domestic animals can be identified based on the fact that any pen or fence surrounding the herd of livestock being attacked will be completely obliterated in several places (because hybrids generally do not bother jumping fences, and simply crash through them into the trapped herd of livestock; see below for more information on hybrid hunting tactics). In addition, a wolf attack will usually result in one or two individual livestock missing. A hybrid attack will result in the death of most, or all, of a farmer's herd, as these animals require much more food than do true wolves.

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"The Warg that one hears is worse than the orc that one fears."
-Boromir, The Fellowship of the Ring

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Wargs

While Wargs are definitely equipped with the tools necessary to bring down considerably large prey, all Wargs are captive bred and have been raised by the minions of the Dark Lord since the First Age and have no need to hunt. The Dark Lord specifically selected their natural weapons and prowess when he began attempting to breed them for his own devices. Because of this, the ferocious nature of Wargs, which has been strengthened by selective breeding, is solely for combat and not survival. They have no instinctual drive to hunt for food because it has been provided for them since they were brought into existence, and their only desire is to be allowed the opportunity to kill on the battlefield. Therefore, Wargs are an unnatural creation; a perversion of true wolves essentially created by the Dark Lord.

Under normal conditions, Wargs are fed by their captors and eat whatever is offered them. Favorite foods include the flesh of the free peoples (Elves, in particular, are said to be relished). In fact, a Warg handler who is trying to persuade a sick or fastidious animal to eat will use Elf flesh (if available), as no Warg can resist it and this food item often stimulates the finicky animal's appetite. In situations where this flesh is unavailable, Wargs will eat nearly anything tossed at them; this includes rotten meat, the meat of other Wargs, and even the flesh of orcs/goblins. In fact, it has been reported that in certain orc clans where the Warg is revered, old or weak family members will readily be sacrificed to feed growing Wargs.

Hybrids

Hybrids will eat any kind of warm-blooded animal that they are presented with and feel that they can catch. They will hunt any and all of the animals that true wolves will, and to supplement this they will hunt humans, dwarves, elves, and livestock. Hybrids are just as dangerous (if not more so) than Wargs. These beasts originally came into existence as a result of copulation between a true wolf and an escaped Warg, which basically means that they retain the best qualities of both: the true wolf's instinctual ability to hunt and survive outside of captivity, and the Warg's incredible size and prowess in combat. While they hunt in packs, as do true wolves, there are several fundamental differences in the tactics that they employ. Unlike wolves, which hunt in much larger groups, hybrids only hunt in packs of three (which is also the average size of a family group without pups). Furthermore, while wolves generally single out the sick or old members of a prey item's herd by carefully isolating and taking down those that lag behind, hybrids do no such thing. When hunting, two individuals (usually females) move to one side of the herd being stalked, while a single individual (usually the male) moves to the opposite side of the herd in question and waits. When the time is right, the two females rush into the group of prey at great speed. Because of their amazing ability to stalk prey and mask their scents, the herd is often taken completely unawares. Once into the herd, the females quickly lash out with fang and claw attacking any animal in their path. At this point, Hybrids can employ one of several very effective tactics for quickly taking down large prey. Most often, they grab the prey animal's throat in their teeth and shake violently, breaking the victim's neck, severing major arteries/veins and dropping the animal instantly. However, they have also been reported to employ their massive forepaws and swat prey animals with enough force to break the spinal cord, legs, or hips, rendering the animal immobile. Often, the animals that are killed during the surprise attack are initially left where they lay as the females continue to drive the rest of the herd head-long into the waiting male, who employs similar tactics to kill as many more of the unsuspecting herd as he can. Then, the pack can feed go back and feed on the previously slain animals at their leisure. It is a savage display of hunting ability combined with deadly force, and it's amazingly effective. The amount of prey killed during an episode such as this is also vastly different from what is reported in true wolf attacks, largely because hybrids are significantly bigger and require more food to survive. In fact, because of their size and fast paced metabolism, it is very difficult for hybrids to acquire enough food, thus the presence of adequate prey populations is a limiting factor in the geographic distribution of hybrids. Furthermore, many efforts have been made by the free peoples in the mid latitudes to eradicate them, thus they are more often found in less populated areas of northern Middle-earth. While a wolf can consume more than 9 lbs of meat at a single meal, a Hybrid must consume nearly 30 lbs.

Competition

While there is no direct competition between Wargs and other canids, wolves and hybrids feed on many of the same prey items, and are therefore constantly at odds. Because they are larger and eat more, the presence of hybrids within a geographical area may result in other carnivores living nearby to be pushed out due to lack of food. This leads in bitter territorial bouts between the two species. While in one on one combat a true wolf has no chance of defeating a hybrid, true wolves usually face intruders as a pack. Therefore, a pack of wolves (consisting of up to 30 or more individuals) can easily force a pack of hybrids out of an area that they are defending.

"At a gap in the circle, a great wolf shape could be seen halted, gazing at them. A shuddering howl broke from him, as if he were a captain summoning his pack to the assault."
- The Fellowship of the Ring

Pack behavior and dynamics

Most wild canid species are highly social, having very specialized and extensive means of communicating with one another. This includes vocalization, production of scent, and posturing. All of these allow most pack-living canids the ability to be very organized in their behavior.

Wolves

Wolves are highly social animals, often living in packs of two to thirty-six individuals. Wolf packs are typically composed of an alpha mating pair and their offspring, including young of previous years. Unrelated immigrants may also be allowed into the pack if immediately submissive to the current alpha pair upon approach. The territory of an average wolf pack ranges from 50 to 5,000 square miles, and is vehemently defended against intruders.

There is a strong dominance hierarchy within each pack. The pack leader, usually the alpha male, is dominant over all other individuals. The next dominant individual is the alpha female, who is subordinate only to the alpha male. In the event that the alpha male becomes injured or is otherwise unable to maintain his dominance, the beta male will take his place in the hierarchy. Rank within the pack hierarchy determines which animals have breeding rights and which eat first after a kill. Rank is demonstrated by postural cues and facial expressions, such as crouching, chin touching, and rolling over to show the stomach.

Vocalizations, such as howling, allow pack members to communicate with each other during hunting and to advertise territories to other packs. Scent marking is ordinarily only done by the alpha male, and is used for communication with other packs.

Each year, wolf packs have both a stationary and nomadic phase. Stationary phases occur during the spring and summer, while pups are being reared. Nomadic phases occur during the fall and winter, when adequate food supplies can only be found by roaming over large areas. Wolves are mostly nocturnal in their habits and may cover long distances on a daily basis (i.e. up to 120 miles, with a pace of 5 mph). Wolves can run at speeds up to 34 to 43 mph.

Wolves may live thirteen years in the wild, though average lifespan is 5 to 6 years. In captivity they may live to be fifteen years of age.

Wargs

Because they are essentially domesticated animals, Wargs have few social interactions of note. Unlike true wolves, they do not have the instinct to communicate whether they are dominant or sub- ordinate and have no way of forming a social structure. Thus, they display no instinctive pack behavior and no understanding of family dynamics and do not pay any heed to relatives during their daily life (unless one jumps its turn during feeding time). However, this should not be confused with having no means of communicating with conspecifics or handlers. For example, Wargs are able to use vocalizations to denote location, temper, or pain to one another. Furthermore, Wargs have the ability to use vocalizations to express their general mood to their handler, allowing a bond (albeit weak) between master and mount. Despite popular belief, Wargs do not serve their handler out of love and this bond is not one of friendship (as is found in most domestic canids), rather Wargs are slaves who have been bred into servitude. Furthermore, the Dark Lord was clever in breeding out their instinct for survival, but not their instinctive aggressiveness. In doing this, he bred a species that is formidable in combat and loyal, not out of desire, but out of necessity.

Because of all these points, true Wargs have very little intraspecific social interaction. For example, in the case of feeding order, the largest male simply attacks those that do not wait their turn. The others do not see him as dominant, only as the one who eats first until he becomes too old and isn't strong enough to defend that right. Breeding is accomplished by separating the larger, stronger males and placing them in breeding pens with larger, stronger females until copulation is complete. At all other times (except during combat), sexes are kept separate to insure that no indiscriminant breeding occurs and that males do not harass pregnant females by attempting to breed when the females are not receptive (which can lead to conflict). Furthermore, females vehemently defend their cubs as males are known to kill juveniles outright (the reasoning for this is unknown) and the two sexes generally do not intermingle well.

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"(Wargriders) were very swift and skilled in avoiding ordered men in close array....but at speed they would pass with reckless ferocity through any gaps in companies of horsemen, slashing at the bellies of the horses."
-Unfinished Tales

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There are several myths surrounding Wargs that should here be corrected: Wargs are not nocturnal by necessity. In fact, they can readily exist in full daylight with no inhibition. However, it is their goblin handlers that have an innate fear of the sun and, thus, Wargs are rarely seen about unless it is nighttime. Furthermore, Wargs are not evil spirits or phantasms that take the shape of wolves. They are real, corporeal creatures. Finally, Wargs are cannot outrun horses. Yet, most horses (aside from Mearas and Elven Horses) fear them and are easily struck dumb by the presence of the beasts, allowing Wargs to readily catch most horses that they are confronted with. In fact, a Warg is much slower than even a true wolf and can only attain speeds of 25-30 mph at the most. Their longevity is normally 10-12 years.

Hybrids

Hybrids are much more social animals than Wargs, but exist in harem-like packs of no more than 3 or 4 individuals (1 male and 2 or 3 females). The male is always considered dominant in the pack, although females do most of the strenuous work during group hunts (see above). Most alpha males attain their position by killing or defeating an existing alpha male in combat. Therefore, because rogue males (which are usually younger individuals) will almost always attempt to take over the alpha position within packs, they are chased away immediately by current alpha males if encountered.

Rogue females are also chased off (usually by current females within the pack). This is presumably done to give the current females confidence in the fact that they will have breeding opportunities each year. The result of too many females existing within a pack is less breeding opportunities. Packs also occasionally form when a rogue male comes into contact with several rogue females. Because parents chase off all offspring as soon as they can fend for themselves, juveniles are usually still relatively young when on their own and not able to defeat a current alpha male. Therefore, the only rogue males that pose an actual threat to current alpha males are over 5 years old.

Interestingly, hybrids have only dominant visual and vocal cues, and no submissive ones. Therefore, when two unacquainted hybrids meet, the individual that displays the more impressive dominate cues will win the bout. These cues include the bearing of teeth, holding the tail erect, pointing the ears forward, and growling. Furthermore, hybrids rarely howl to communicate. Instead, they use deep guttural barks to communicate with one another.

Although hybrids have a stationary and nomadic phase (similar to wolves), the stationary phase is much more abbreviated (usually only in the spring). The rest of the year, these animals roam about in territories averaging 30-50 square miles in size. A hybrid can run at speeds of 27-35 mph, adopting an average speed of 5 mph when searching for food within its territory. Longevity is normally 5-10 years.

Reproduction and Development

The breeding cycles of wolves hybrids begin in early to mid spring, with northern populations breeding later in the season than southern populations. Female wolves and hybrids choose their mates and often form a life-long pair bond (Note: unlike wolves, hybrid males will often have a harem of two or three females). Wolf and hybrid pairs or trios spend much of their time together. Females come into estrus once each year, which lasts 5 to 14 days, mating also occurs during this time.

After the mating season has finished, the female digs a den in which to raise her young. The den often includes an entrance that first slopes down and then up again before reaching a higher area, which helps prevent den-flooding during periods of heavy rain. Pups are born in the den and will remain there for several weeks after birth. Aside from digging their own dens, wolves have also been reported to den under cliffs, under fallen trees, and in caves. Although they have been known to dig their own dens, hybrids more often take over and modify the dens of other mammals or den in caves. The gestation period lasts between 60 and 63 days in wolves and 90 to 100 days in hybrids, litter size ranges from one to fourteen in wolves and one to three in hybrids.

Pups remain in the den until they are 8 to 10 weeks old in wolves and 20 weeks in hybrids. Females stay with their pups almost exclusively for the first 3 weeks and then are cared for by all members of the pack. Until they are about 45 days old, the pups are fed regurgitated food by all pack members. They are fed meat provided by pack members after that age. Wolf and hybrid pups are born blind and deaf. They weigh approximately 1 lb in wolves and 3 lbs in hybrids. All pups are relatively helpless and must depend on the mother even for warmth. At ten to fifteen days of age, the pups' blue eyes open, but they only have control over their front legs, thus crawling is their only mode of mobility. Five to ten days later, the young are able to stand, walk, and vocalize. Weaning occurs at about 9 weeks of age. Once pups leave the den they begin to learn to hunt and begin play fighting. Interactions at this time, as well as the dominance status of the mother, ultimately determine their position in the pack hierarchy. Wolf pups develop rapidly they must be large and accomplished enough to hunt with the pack with the onset of winter. At approximately ten months old, the young begin to hunt with the pack. For reasons unknown, hybrids develop much more slowly and pups often do not survive their first winter. Furthermore, as soon as pups are large enough to hunt, the adults force them out of the pack.

Female wolf pups reach maturity at two years of age, while males will not reach full maturity until three years of age. Most young wolves disperse from their natal pack when they are between 1 and 3 years old. Female hybrids reach maturity at 3 to 4 years of age, while males reach maturity at 5 years. Because food availability is a limiting factor in the size of hybrid packs, young hybrids represent an extra mouth to feed are forced out of the pack as soon as they can fend for themselves (usually at three years regardless of sex).

Wargs and domestic dogs generally are more receptive to breeding in the spring, although breeding can be induced year round. Developmental rates for domestic dogs are similar to wolves, while developmental rates of Wargs are similar to hybrids.

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Tune in Next Month when we will discuss Canids in game-terms. This will include the statistics for the different species of wild canids mentioned already, as well as descriptions and statistics for the common breeds of domestic dog found throughout Middle-earth!