| Волк Временной диапазон: Средний плейстоцен - настоящее время (810 000–0 лет назад ) [1] | |
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Евразийский волк ( Canis lupus lupus ) в парке дикой природы Колморден , Швеция | |
Волчья стая воет | |
Научная классификация  | |
| Королевство: | Animalia |
| Тип: | Хордовые |
| Класс: | Млекопитающие |
| Заказ: | Хищник |
| Семья: | Canidae |
| Род: | Canis |
| Разновидность: | C. волчанка |
| Биномиальное имя | |
| Canis lupus | |
| Подвиды | |
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| Мировой ареал волков на основе оценки МСОП за 2018 год. [2] | |
Волк ( Canis волчанка [а] ), также известный как серый волк или серый волк , большой собачьей произрастающих в Евразии и Северной Америке . Выявлено более тридцати подвидов Canis lupus , и серые волки, как принято понимать в разговорной речи, составляют не домашние / дикие подвиды. Волк - самый крупный из ныне живущих членов семейства Canidae , самцы в среднем весят 40 кг (88 фунтов), а самки - 37 кг (82 фунта). Волки достигают 105–160 см (41–63 дюйма) в длину и 80–85 см (31–33 дюйма) на высоте плеч. Волк также отличается от других Canis.разновидности менее заостренными ушами и мордой, а также более коротким туловищем и более длинным хвостом. Тем не менее, волк достаточно тесно связан с более мелкими видами Canis , такими как койот и золотой шакал , чтобы производить с ними плодородные гибриды . Пластинчатый мех волка обычно пестрый белый, коричневый, серый и черный, хотя подвидов в арктической области может быть почти все белые.
Из всех представителей рода Canis волк наиболее специализирован для совместной охоты, что демонстрируется его физической адаптацией к схватке с крупной добычей, его более социальным характером и высокоразвитым экспрессивным поведением . Он путешествует в нуклеарных семьях, состоящих из пары, сопровождаемой их потомством. Потомство может уйти, чтобы сформировать свои собственные стаи с наступлением половой зрелости и в ответ на конкуренцию за пищу внутри стаи. Волки также территориальны, и драки за территорию являются одной из основных причин смертности волков. Волк в основном хищники питается крупными дикими копытными млекопитающими, а также более мелкими животными, домашним скотом, падалью и мусором. Одиночные волки или спариваемые пары обычно более успешны на охоте, чем большие стаи. Патогены и паразиты, особенно вирус бешенства , могут заразить волков.
The global wild wolf population was estimated to be 300,000 in 2003 and is considered to be of Least Concern by the International Union for Conservation of Nature (IUCN). Wolves have a long history of interactions with humans, having been despised and hunted in most pastoral communities because of their attacks on livestock, while conversely being respected in some agrarian and hunter-gatherer societies. The wolf is also considered the ancestor of most domestic dog breeds. Although the fear of wolves exists in many human societies, the majority of recorded attacks on people have been attributed to animals suffering from rabies. Wolf attacks на людях встречаются редко, потому что волков относительно мало, они живут вдали от людей и у них развился страх перед людьми из-за своего опыта общения с охотниками, владельцами ранчо и пастухами.
Этимология
Английское «волк» происходит от древнеанглийского wulf , которое, как считается, происходит от протогерманского * wulfaz . Прото-индо-европейский корень * wĺ̥kʷos может также быть источником латинского слова для животных волчанки (* lúkʷos ). [4] [5] Название «серый волк» относится к сероватому цвету этого вида. [6]
С дохристианских времен, германские народы , такие как англо-саксы взяли на Вульфа в качестве префикса или суффикса в их именах. Примеры включают Wulfhere («Армия волков»), Cynewulf («Королевский волк»), Cēnwulf («Смелый волк»), Wulfheard («Жесткий волк»), Earnwulf («Орлиный волк»), Wulfstān («Волчий камень»). Челвульф («Благородный волк»), Вольфхрок («Волк-Фрок»), Вольфетан («Волчья шкура»), Исангрим («Серая маска»), Скратольф («Одеяние волка»), Вольфганг («Волчья походка») и Вольфдрегил («Бегущий по волку»). [7]
Таксономия
| Филогения собак с возрастом дивергенции | |||||||||||||||||||||||||||||||||||||||||||||||||||
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| Кладограмма и расхождение серого волка (включая домашнюю собаку) среди его ближайших дошедших до нас родственников [8] |
В 1758 году шведский ботаник и зоолог Карл Линней опубликовал в своем Systema Naturae биномиального номенклатуры . [3] Canis - это латинское слово, означающее « собака », [9] и под этим родом он перечислил собакоподобных хищников, включая домашних собак, волков и шакалов . Он классифицировал домашнюю собаку как Canis familis , а волка как Canis lupus . [3] Линней считал собаку отдельным видом от волка из-за ее « каучукового рекурсивного хвоста» ( вздернутого кверху хвоста), которого нет ни у одного другого псового . [10]
Подвиды
В третьем издании « Виды млекопитающих мира», опубликованном в 2005 г., маммолог У. Кристофер Возенкрафт перечислил в C. lupus 36 диких подвидов и предложил два дополнительных подвида: фамильяр (Linnaeus, 1758) и динго (Meyer, 1793). Wozencraft включены hallstromi -The Новой Гвинеи пение собака -Как таксономического синонима для динго . Возенкрафт сослался на исследование митохондриальной ДНК 1999 года как на одно из руководств при принятии своего решения и перечислил 38 подвидов C. lupus под общим биологическим названием.слова «волк», номинальным подвидом которого является евразийский волк ( C. l. lupus ) на основе типового образца, который Линней изучал в Швеции. [11] Исследования с использованием палеогеномных методов показывают, что современные волк и собака являются сестринскими таксонами , поскольку современные волки не имеют близкого родства с популяцией волков, которая была впервые приручена . [12] В 2019 году на семинаре, организованном Группой специалистов по собакам Комиссии по выживанию видов МСОП, поющая собака Новой Гвинеи и динго были признаны одичавшими собаками Canis knownis и поэтому не должны оцениваться на предметКрасный список МСОП . [13]
Эволюция
Филогенетическое спуск сохранившегося волка С. волчанка из C. Этруска через C. mosbachensis получила широкое признание. [14] Самые ранние окаменелости C. lupus были найдены на территории бывшей восточной Берингии в Олд Кроу, Юкон , Канада, и в Криппл Крик Самп, Фэрбенкс , Аляска. Возраст не согласован, но может быть около миллиона лет назад. Значительное морфологическое разнообразие волков существовало к позднему плейстоцену . У них были более крепкие черепа и зубы, чем у современных волков, часто с укороченной мордой , что являлось выраженным развитиемвисочная мышца и крепкие премоляры . Предполагается, что эти особенности были специализированными приспособлениями для обработки туш и костей, связанными с охотой и выловом падальщиков мегафауны плейстоцена . По сравнению с современными волками, у некоторых плейстоценовых волков наблюдалось увеличение числа поломок зубов, аналогичное тому, которое наблюдается у вымерших ужасных волков . Это говорит о том, что они либо часто перерабатывали туши, либо конкурировали с другими хищниками и им нужно было быстро съесть свою добычу. По сравнению с теми, что встречаются у современных пятнистых гиен , частота и расположение переломов зубов у этих волков указывает на то, что они были обычными костными трещинами. [15] В июне 2019 года отрубленная, но сохранившаяся головаПлейстоценовый волк , возраст которого насчитывает более 40 000 лет назад, был найден недалеко от реки Тирехтях в Якутии , Россия , недалеко от полярного круга . Голова была около 16 дюймов (41 см) в длину, что намного больше, чем голова современного волка. [16] [17] [18]
Геномные исследования показывают, что современные волки и собаки произошли от общей предковой популяции волков [19] [20] [21] , существовавшей 20 000 лет назад. [19] Исследования, проведенные в 2017 и 2018 годах, показали, что гималайский волк является частью линии, которая является базовой по отношению к другим волкам и отделилась от них 691 000–740 000 лет назад. [22] [23] Другие волки, по-видимому, произошли в Берингии в результате расширения, которое было вызвано огромными экологическими изменениями в конце позднего плейстоцена. [23] Исследование, проведенное в 2016 году, показывает, что за « узким местом» населения последовало быстроерадиация от древнего населения во время или сразу после последнего ледникового максимума . Это означает, что первоначальные морфологически разнообразные популяции волков были вытеснены и заменены более современными волками. [24]
Геномное исследование 2016 года предполагает, что волки Старого и Нового Света разделились примерно 12 500 лет назад, после чего произошло расхождение в родословной, которое привело к появлению собак от других волков Старого Света примерно 11 100–12 300 лет назад. [21] Вымерший волк позднего плейстоцена, возможно, был предком собаки, [25] [15] при этом сходство собаки с существующим волком является результатом генетической смеси между ними. [15] Динго, басенджи , тибетский мастиф.а местные породы Китая являются основными членами клады домашних собак. Время расхождения волков в Европе, на Ближнем Востоке и в Азии, по оценкам, наступило сравнительно недавно - около 1600 лет назад. Среди волков Нового Света мексиканские волки разошлись около 5400 лет назад. [21]
Смешивание с другими псовыми
В далеком прошлом существовал обмен генами между африканскими золотыми волками , золотыми шакалами и серыми волками. Африканский золотой волк является потомком генетически смешанных псовых, 72% волка и 28% эфиопского волка. Один африканский золотой волк с египетского Синайского полуострова демонстрирует смешение с ближневосточными волками и собаками. [26] Есть свидетельства обмена генами между золотыми шакалами и ближневосточными волками, в меньшей степени у европейских и азиатских волков и меньше всего у североамериканских волков. Это указывает на то, что происхождение золотого шакала, обнаруженное у североамериканских волков, могло произойти до расхождения евразийских и североамериканских волков. [27]
Общий предок койота и волка смешался с популяцией призраков вымерших неопознанных псовых. Этот псовый генетически близок к волку и эволюционировал после расхождения африканской охотничьей собаки от других видов псовых. Предполагается, что базальное положение койота по сравнению с волком связано с тем, что койот сохраняет большую часть митохондриального генома этого неопознанного псового. [26] Точно так же музейный образец волка из южного Китая, собранный в 1963 году, показал геном, на 12–14% состоящий из смеси этой неизвестной собаки. [28] В Северной Америке большинство койотов и волков показывают различную степень прошлой генетической примеси. Красный волкна юго-востоке Соединенных Штатов - это гибридное животное с 40%: 60% родословной волка и койота. Кроме того, было обнаружено, что 60%: 40% генетики волков и койотов у восточных лесных волков и 75%: 25% у волков в районе Великих озер . [27]
В последнее время некоторые самцы итальянских волков произошли от собачьих предков, что указывает на то, что самки волков будут размножаться с самцами собак в дикой природе. [29] В Кавказских горах десять процентов собак, включая собак-охранников , являются гибридами первого поколения. [30] Хотя спаривание золотых шакалов и волков никогда не наблюдалось, доказательства гибридизации шакала и волка были обнаружены посредством анализа митохондриальной ДНК шакалов, обитающих в горах Кавказа [30] и в Болгарии. [31]
В 2021 году генетическое исследование показало, что сходство собаки с существующим серым волком было результатом значительного потока генов от собаки к волку , с почти незначительным потоком генов от волка к собаке с момента приручения собаки. Некоторые серые волки были родственниками всех древних и современных собак. [32]
Описание
Волк - самый крупный из ныне живущих представителей семейства Canidae , [33] и отличается от койотов и шакалов более широкой мордой, более короткими ушами, более коротким туловищем и более длинным хвостом. [34] [33] Он стройный и мощно сложенный, с большой глубоко опускающейся грудной клеткой , покатой спиной и сильно мускулистой шеей. [35] Ноги волка умеренно длиннее, чем у других псовых, что позволяет животному быстро двигаться и преодолевать глубокий снег, который покрывает большую часть его ареала зимой. [36] Уши относительно маленькие и треугольные. [35]Голова волка большая и тяжелая, с широким лбом, сильными челюстями и длинной тупой мордой. [37] Череп составляет 230–280 мм (9–11 дюймов) в длину и 130–150 мм (5–6 дюймов) в ширину. [38] Зубы тяжелые и большие, поэтому они лучше подходят для дробления костей, чем у других псовых. Однако они не так специализированы, как гиены . [39] [40] Его коренные зубы имеют плоскую жевательную поверхность, но не в такой степени, как у койота, чья диета содержит больше растительных веществ. [41] У самок более узкая морда и лоб, тонкая шея, немного короче ноги и менее массивные плечи, чем у самцов. [42]
Взрослые волки достигают 105–160 см (41–63 дюйма) в длину и 80–85 см (31–33 дюйма) на высоте плеч. [37] Длина хвоста 29–50 см (11–20 дюймов), уши 90–110 мм ( 3+1 ⁄ 2 - 4+3 ⁄ 8 дюйма) в высоту, а задние лапы -220–250 мм ( 8+5 ⁄ 8 - 9+7 ⁄ 8 дюйма). [43] Размер и вес современного волка увеличивается пропорционально широте в соответствии с правилом Бергманна . [44] Средняя масса тела волка составляет 40 кг (88 фунтов), самый маленький зарегистрированный образец - 12 кг (26 фунтов), а самый большой - 79,4 кг (175 фунтов). [45] [37] В среднем европейские волки весят 38,5 кг (85 фунтов), североамериканские волки - 36 кг (79 фунтов), а индийские и арабские волки - 25 кг (55 фунтов). [46]Самки в любой популяции волков обычно весят на 2,3–4,5 кг (5–10 фунтов) меньше, чем самцы. Волки весом более 54 кг (119 фунтов) - редкость, хотя исключительно крупные особи были зарегистрированы на Аляске и в Канаде. [47] В средней полосе России исключительно крупным самцам дают максимальный вес 69–79 кг (152–174 фунта). [43]
Пелаж
У волка очень густой и пушистый зимний мех, с коротким подшерстком и длинными жесткими остовыми волосами . [37] Большая часть подшерстка и некоторые остевые волосы сбрасываются весной и снова отрастают осенью. [46] Самые длинные волосы встречаются на спине, особенно на передней части и шее. Особенно длинные волосы растут на плечах и почти образуют гребень на верхней части шеи. Волосы на щеках удлиненные, образуют пучки. Уши покрыты короткими волосками и выступают из меха. На конечностях от локтей до пяточных сухожилий имеются короткие, эластичные и плотно прилегающие волоски . [37]Зимний мех очень устойчив к холоду. Волки в северном климате могут комфортно отдыхать на открытых площадках при -40 ° C (-40 ° F), помещая морду между задними лапами и прикрывая морду хвостом. Мех волка обеспечивает лучшую изоляцию, чем мех собаки, и не собирает лед, когда на него конденсируется теплое дыхание. [46]
В холодном климате волк может уменьшить приток крови к своей коже, чтобы сохранить тепло тела. Тепло подушечек для ног регулируется независимо от остальной части тела и поддерживается на уровне чуть выше точки замерзания тканей, когда подушечки соприкасаются со льдом и снегом. [48] В теплом климате мех грубее и реже, чем у северных волков. [37] Волки-самки, как правило, имеют более гладкие конечности, чем самцы, и, как правило, с возрастом у них становится более гладкая шерсть. У старых волков обычно больше белых волосков на кончике хвоста, вдоль носа и на лбу. Зимний мех дольше всего сохраняется у кормящих самок, хотя и с некоторой потерей шерсти вокруг сосков. [42] Длина волос посередине спины составляет60–70 мм ( 2+3 ⁄ 8 - 2+3 ⁄ 4 дюйма), а остевые волосы на плечах обычно не превышают90 мм ( 3+1 ⁄ 2 дюйма), но может достигать110–130 мм ( 4+3 ⁄ 8 - 5+1 ⁄ 8 дюйма). [37]
Цвет шерсти волка определяется его остевым волосом. Волки обычно имеют белые, коричневые, серые и черные волосы. [49] Шерсть евразийского волка представляет собой смесь охристого (от желтого до оранжевого) и ржавого.охристые (оранжевый / красный / коричневый) цвета со светло-серым. Морда бледно-охристо-серая, а область губ, щек, подбородка и горла белые. Верх головы, лоб, под глазами и между ними, а также между глазами и ушами серые с красноватым налетом. Шея охристая. Длинные черные кончики волос вдоль спины образуют широкую полосу с черными кончиками волос на плечах, верхней части груди и задней части тела. Бока туловища, хвоста и внешних конечностей бледно-грязно-охристого цвета, а внутренние стороны конечностей, живота и паха - белого цвета. Если не считать чисто белых или черных волков, эти тона мало различаются по географическим регионам, хотя образцы этих цветов различаются у разных людей. [50]
В Северной Америке окраска шерсти волков соответствует правилу Глогера: волки в канадской Арктике - белые, а в южной Канаде, США и Мексике - преимущественно серые. В некоторых районах Скалистых гор Альберты и Британской Колумбии окрас шерсти преимущественно черный, некоторые - сине-серые, а некоторые - серебристо-черные. [49] Различия в цвете шерсти между полами отсутствуют в Евразии; [51] у женщин в Северной Америке обычно более красный оттенок. [52] Волки черного окраса в Северной Америке приобрели свой окрас из-за примеси собак и волков после того, как собаки впервые пересекли Берингов пролив 12–14 тысяч лет назад. [53]Исследования по наследству белого окраса от собак волкам еще не проводились. [54]
Экология
Распространение и среда обитания
Первоначально волки обитали в Евразии и Северной Америке. Преднамеренное преследование людей из-за хищничества домашнего скота и страха перед нападениями на людей сократило ареал волка примерно до одной трети от того, что было раньше. В настоящее время волк искоренен (локально вымерли) в большей части Западной Европы, США и Мексики, а также в Японии. В наше время волк встречается в основном в дикой природе и отдаленных районах. Волка обитают на высоте от уровня моря до 3000 м (9800 футов). Волки живут в лесах, внутренних водно-болотных угодьях , кустарниках , лугах (включая арктическую тундру ), пастбищах , пустынях и скалистых вершинах гор. [2]Использование волками среды обитания зависит от обилия добычи, снежных условий, плотности скота, плотности дорог, присутствия человека и топографии . [41]
Диета
Как и все наземные млекопитающие, которые являются стайными охотниками , волк питается преимущественно дикими травоядными копытными млекопитающими , которых можно разделить на крупных 240–650 кг (530–1 430 фунтов) и средних 23–130 кг (51–287 фунтов), а также иметь массу тела, аналогичную совокупной массе членов стаи. [55] [56] Волк специализируется на охоте на уязвимых особей крупной добычи, [41] со стаей из 15 человек, способной сбить взрослого лося . [57] Различия в рационе волков, живущих на разных континентах, основаны на разнообразии копытных млекопитающих и доступной более мелкой и домашней добычи. [58]
В Северной Америке в рационе волка преобладают дикие крупные копытные (копытные) и млекопитающие среднего размера. В Азии и Европе в их рационе преобладают дикие копытные млекопитающие среднего размера и домашние виды. Волк зависит от диких видов, и если они недоступны, как в Азии, волк больше полагается на домашних. [58] По всей Евразии волки охотятся в основном на лосей, благородных оленей , косуль и кабанов . [59] В Северной Америке важной добычей для широкого ареала являются лоси , лоси, карибу , белохвостые олени и олени-мулы . [60]Волки переваривают пищу за несколько часов и могут кормиться несколько раз в день, быстро потребляя большое количество мяса. [61] Сытый волк откладывает жир под кожей, вокруг сердца, кишечника, почек и костного мозга, особенно осенью и зимой. [62]
Тем не менее, волки не привередливы в еде. Животные меньшего размера, которые могут дополнять их рацион, включают грызунов , зайцев , насекомоядных и более мелких плотоядных. Они часто едят водоплавающих птиц и их яйца. Когда такой пищи недостаточно, они охотятся на ящериц , змей , лягушек и крупных насекомых, когда они доступны. [63] Волки в северной Миннесоте охотятся на северную щуку в пресноводных ручьях. [64] Рацион прибрежных волков на Аляске включает 20% лосося , [65] while those of coastal wolves in British Columbia includes 25% marine sources, and those on the nearby islands 75%.[66]
In Europe, wolves eat apples, pears, figs, melons, berries and cherries. In North America, wolves eat blueberries and raspberries. Wolves also eat grass, which may provide some vitamins.[67] They are known to eat the berries of mountain-ash, lily of the valley, bilberries, cowberries, European black nightshade, grain crops, and the shoots of reeds.[63]
In times of scarcity, wolves will readily eat carrion.[63] In Eurasian areas with dense human activity, many wolf populations are forced to subsist largely on livestock and garbage.[59] Prey in North America continue to occupy suitable habitats with low human density, the wolves eating livestock and garbage only in dire circumstances.[68] Cannibalism is not uncommon in wolves during harsh winters, when packs often attack weak or injured wolves and may eat the bodies of dead pack members.[63][69][70]
Interactions with other predators
Wolves typically dominate other canid species in areas where they both occur. In North America, incidents of wolves killing coyotes are common, particularly in winter, when coyotes feed on wolf kills. Wolves may attack coyote den sites, digging out and killing their pups, though rarely eating them. There are no records of coyotes killing wolves, though coyotes may chase wolves if they outnumber them.[71] According to a press release by the U.S. Department of Agriculture in 1921, the infamous Custer Wolf relied on coyotes to accompany him and warn him of danger. Though they fed from his kills, he never allowed them to approach him.[72] Interactions have been observed in Eurasia between wolves and golden jackals, the latter's numbers being comparatively small in areas with high wolf densities.[37][71][73] Wolves also kill red, Arctic and corsac foxes, usually in disputes over carcasses, sometimes eating them.[37][74]
Brown bears typically dominate wolf packs in disputes over carcasses, while wolf packs mostly prevail against bears when defending their den sites. Both species kill each other's young. Wolves eat the brown bears they kill, while brown bears seem to eat only young wolves.[75] Wolf interactions with American black bears are much rarer because of differences in habitat preferences. Wolves have been recorded on numerous occasions actively seeking out American black bears in their dens and killing them without eating them. Unlike brown bears, American black bears frequently lose against wolves in disputes over kills.[76] Wolves also dominate and sometimes kill wolverines, and will chase off those that attempt to scavenge from their kills. Wolverines escape from wolves in caves or up trees.[77]
Wolves may interact and compete with felids, such as the Eurasian lynx, which may feed on smaller prey where wolves are present[78] and may be suppressed by large wolf populations.[79] Wolves encounter cougars along portions of the Rocky Mountains and adjacent mountain ranges. Wolves and cougars typically avoid encountering each other by hunting at different elevations for different prey (niche partitioning). In winter, when snow accumulation forces their prey into valleys, interactions between the two species become more likely. Wolves in packs usually dominate cougars and can steal their kills or even kill them,[80] while one-to-one encounters tend to be dominated by the cat. There are several documented cases of cougars killing wolves.[81] Wolves more broadly affect cougar population dynamics and distribution by dominating territory and prey opportunities and disrupting the feline's behaviour.[82] Wolf and Siberian tiger interactions are well-documented in the Russian Far East, where tigers significantly depress wolf numbers, sometimes to the point of localized extinction. Only human depletion of tiger numbers appears to protect wolves from competitive exclusion from them. With perhaps only four proven records of tigers killing wolves, these cases are rare; attacks appear to be competitive rather than predatory in nature.[83][78]
In Israel, Central Asia and India wolves may encounter striped hyenas, usually in disputes over carcasses. Striped hyenas feed extensively on wolf-killed carcasses in areas where the two species interact. One-to-one, hyenas dominate wolves, and may prey on them,[84] but wolf packs can drive off single or outnumbered hyenas.[85][86] There is at least one case in Israel of a hyena associating and cooperating with a wolf pack. It is proposed that the hyena could benefit from the wolves' superior ability to hunt large, agile prey. The wolves could benefit from the hyena's superior sense of smell, to locate and dig out tortoises, to crack open large bones, and to tear open discarded food containers like tin cans.[87]
Behaviour
Social structure
The wolf is a social animal.[37] Its populations consist of packs and lone wolves, most lone wolves being temporarily alone while they disperse from packs to form their own or join another one.[88] The wolf's basic social unit is the nuclear family consisting of a mated pair accompanied by their offspring.[37] The average pack size in North America is eight wolves and in Europe 5.5 wolves.[44] The average pack across Eurasia consists of a family of eight wolves (two adults, juveniles, and yearlings),[37] or sometimes two or three such families,[41] with examples of exceptionally large packs consisting of up to 42 wolves being known.[89] Cortisol levels in wolves rise significantly when a pack member dies, indicating the presence of stress.[90] During times of prey abundance caused by calving or migration, different wolf packs may join together temporarily.[37]
Offspring typically stay in the pack for 10–54 months before dispersing.[91] Triggers for dispersal include the onset of sexual maturity and competition within the pack for food.[92] The distance travelled by dispersing wolves varies widely; some stay in the vicinity of the parental group, while other individuals may travel great distances of upwards of 206 km (128 mi), 390 km (240 mi), and 670 km (420 mi) from their natal (birth) packs.[93] A new pack is usually founded by an unrelated dispersing male and female, travelling together in search of an area devoid of other hostile packs.[94] Wolf packs rarely adopt other wolves into their fold and typically kill them. In the rare cases where other wolves are adopted, the adoptee is almost invariably an immature animal of one to three years old, and unlikely to compete for breeding rights with the mated pair. This usually occurs between the months of February and May. Adoptee males may mate with an available pack female and then form their own pack. In some cases, a lone wolf is adopted into a pack to replace a deceased breeder.[89]
Wolves are territorial and generally establish territories far larger than they require to survive assuring a steady supply of prey. Territory size depends largely on the amount of prey available and the age of the pack's pups. They tend to increase in size in areas with low prey populations,[95] or when the pups reach the age of six months when they have the same nutritional needs as adults.[96] Wolf packs travel constantly in search of prey, covering roughly 9% of their territory per day, on average 25 km/d (16 mi/d). The core of their territory is on average 35 km2 (14 sq mi) where they spend 50% of their time.[95] Prey density tends to be much higher on the territory's periphery. Except out of desperation, wolves tend to avoid hunting on the fringes of their range to avoid fatal confrontations with neighbouring packs.[97] The smallest territory on record was held by a pack of six wolves in northeastern Minnesota, which occupied an estimated 33 km2 (13 sq mi), while the largest was held by an Alaskan pack of ten wolves encompassing 6,272 km2 (2,422 sq mi).[96] Wolf packs are typically settled, and usually leave their accustomed ranges only during severe food shortages.[37]
Wolves advertise their territories to other packs through howling and scent marking. Scent marking involves urine, feces, and anal gland scents. This is more effective at advertising territory than howling and is often used in combination with scratch marks. Wolves increase their rate of scent marking when they encounter the marks of wolves from other packs. Lone wolves will rarely mark, but newly bonded pairs will scent mark the most.[41] These marks are generally left every 240 m (260 yd) throughout the territory on regular travelways and junctions. Such markers can last for two to three weeks,[96] and are typically placed near rocks, boulders, trees, or the skeletons of large animals.[37] Territorial fights are among the principal causes of wolf mortality, one study concluding that 14–65% of wolf deaths in Minnesota and the Denali National Park and Preserve were due to other wolves.[98]
Wolves communicate to anticipate what their pack mates or other wolves might do next.[99] This includes the use of vocalization, body posture, scent, touch, and taste.[100] The phases of the moon have no effect on wolf vocalization, and despite popular belief, wolves do not howl at the moon.[101] Wolves howl to assemble the pack usually before and after hunts, to pass on an alarm particularly at a den site, to locate each other during a storm, while crossing unfamiliar territory, and to communicate across great distances.[102] Wolf howls can under certain conditions be heard over areas of up to 130 km2 (50 sq mi).[41] Other vocalizations include growls, barks and whines. Wolves do not bark as loudly or continuously as dogs do in confrontations, rather barking a few times and then retreating from a perceived danger.[103] Aggressive or self-assertive wolves are characterized by their slow and deliberate movements, high body posture and raised hackles, while submissive ones carry their bodies low, flatten their fur, and lower their ears and tail.[104] Raised leg urination is considered to be one of the most important forms of scent communication in the wolf, making up 60–80% of all scent marks observed.[105]
Reproduction
Wolves are monogamous, mated pairs usually remaining together for life. Should one of the pair die, another mate is found quickly.[106] With wolves in the wild, inbreeding does not occur where outbreeding is possible.[107] Wolves become mature at the age of two years and sexually mature from the age of three years.[106] The age of first breeding in wolves depends largely on environmental factors: when food is plentiful, or when wolf populations are heavily managed, wolves can rear pups at younger ages to better exploit abundant resources. Females are capable of producing pups every year, one litter annually being the average.[108] Oestrus and rut begin in the second half of winter and lasts for two weeks.[106]
Dens are usually constructed for pups during the summer period. When building dens, females make use of natural shelters like fissures in rocks, cliffs overhanging riverbanks and holes thickly covered by vegetation. Sometimes, the den is the appropriated burrow of smaller animals such as foxes, badgers or marmots. An appropriated den is often widened and partly remade. On rare occasions, female wolves dig burrows themselves, which are usually small and short with one to three openings. The den is usually constructed not more than 500 m (550 yd) away from a water source. It typically faces southwards where it can be better warmed by sunlight exposure, and the snow can thaw more quickly. Resting places, play areas for the pups, and food remains are commonly found around wolf dens. The odor of urine and rotting food emanating from the denning area often attracts scavenging birds like magpies and ravens. Though they mostly avoid areas within human sight, wolves have been known to nest near domiciles, paved roads and railways.[109] During pregnancy, female wolves remain in a den located away from the peripheral zone of their territories, where violent encounters with other packs are less likely to occur.[110]
The gestation period lasts 62–75 days with pups usually being born in the spring months or early summer in very cold places such as on the tundra. Young females give birth to four to five young, and older females from six to eight young and up to 14. Their mortality rate is 60–80%.[111] Newborn wolf pups look similar to German Shepherd Dog pups.[112] They are born blind and deaf and are covered in short soft grayish-brown fur. They weigh 300–500 g (10+1⁄2–17+3⁄4 oz) at birth and begin to see after nine to 12 days. The milk canines erupt after one month. Pups first leave the den after three weeks. At one-and-a-half months of age, they are agile enough to flee from danger. Mother wolves do not leave the den for the first few weeks, relying on the fathers to provide food for them and their young. Pups begin to eat solid food at the age of three to four weeks. They have a fast growth rate during their first four months of life: during this period, a pup's weight can increase nearly 30 times.[111][113] Wolf pups begin play-fighting at the age of three weeks, though unlike young coyotes and foxes, their bites are gentle and controlled. Actual fights to establish hierarchy usually occur at five to eight weeks of age. This is in contrast to young coyotes and foxes, which may begin fighting even before the onset of play behaviour.[114] By autumn, the pups are mature enough to accompany the adults on hunts for large prey.[110]
Hunting
Single wolves or mated pairs typically have higher success rates in hunting than do large packs; single wolves have occasionally been observed to kill large prey such as moose, bison and muskoxen unaided.[115][116] This contrasts with the commonly held belief that larger packs benefit from cooperative hunting to bring down large game.[116] The size of a wolf hunting pack is related to the number of pups that survived the previous winter, adult survival, and the rate of dispersing wolves leaving the pack. The optimal pack size for hunting elk is four wolves, and for bison a large pack size is more successful.[117]
As well as their physical adaptations for hunting hoofed mammals, wolves possess certain behavioural, cognitive, and psychological adaptations to assist with their hunting lifestyle. Wolves are excellent learners that match or outperform domestic dogs. They can use gaze to focus attention on where other wolves are looking. This is important because wolves do not use vocalization when hunting. In laboratory tests, they appear to exhibit insight, foresight, understanding, and the ability to plan. To survive, wolves must be able to solve two problems—finding a prey animal, then confronting it.[118]
Tracking
Wolves move around their territory when hunting, using the same trails for extended periods. After snowfalls, wolves find their old trails and continue using them. These follow the banks of rivers, the shorelines of lakes, ravines overgrown with shrubs, plantations, or roads and human paths.[119] Wolves are nocturnal predators. During the winter, a pack will commence hunting in the twilight of early evening and will hunt all night, traveling tens of kilometres. Sometimes hunting large prey occurs during the day. During the summer, wolves generally tend to hunt individually, ambushing their prey and rarely giving pursuit.[120]
The wolf usually travels at a loping pace, placing one of its paws directly in front of the other. This gait can be maintained for hours at a rate of 8–9 km/h (5.0–5.6 mph).[121] On bare paths, a wolf can quickly achieve speeds of 50–60 km/h (31–37 mph). The wolf has a running gait of 55–70 km/h (34–43 mph), can leap 5 m (16 ft) horizontally in a single bound, and can maintain rapid pursuit for at least 20 minutes.[93] A wolf's foot is large and flexible, which allows it to tread on a wide variety of terrain. A wolf's legs are long compared to their body size allowing them to travel up to 76 km (47 mi) in 12 hours. This adaptation allows wolves to locate prey within hours, but it can take days to find prey that can be killed without great risk. Moose and deer live singly in the summer. Caribou live in herds of thousands which presents dangers for wolves. Elk live in small herds and these are a safer target.[118]
A wolf carries its head at the same level as its back, lifting it only when alert.[37] In one study, wolves detected moose using scent ten times, vision six times, and once by following tracks in the snow. Their vision is as good as a human's, and they can smell prey at least 2.4 km (1+1⁄2 mi) away. One wolf travelled to a herd 103 km (64 mi) away. A human can detect the smell of a forest fire over the same distance from downwind. The wolf's sense of smell is at least comparable to that of the domestic dog, which is at least ten thousand times more sensitive than a human's.[118]
Pursuit
When hunting large gregarious prey, wolves will try to isolate an individual from its group.[122] If successful, a wolf pack can bring down game that will feed it for days, but one error in judgement can lead to serious injury or death. Most large prey have developed defensive adaptations and behaviours. Wolves have been killed while attempting to bring down bison, elk, moose, muskoxen, and even by one of their smallest hoofed prey, the white-tailed deer. With smaller prey like beaver, geese, and hares, there is no risk to the wolf.[118] Although people often believe wolves can easily overcome any of their prey, their success rate in hunting hoofed prey is usually low.[123]
Generally, bison, elk, and moose will stand their ground, then the wolves must struggle with them to bring them down. Often caribou and deer will flee, but sometimes deer also make a stand.[118] If the targeted animal stands its ground, wolves either ignore it, or try to intimidate it into running.[115] Wolves, or even a wolf on its own, will attempt to frighten a herd into panicking and dispersing.[124]
When wolves encounter prey that flees, they give chase. The speed of sprinting prey is closely related to the speed of their main predators. Wolves can run at 56–64 km/h (35–40 mph) across several kilometres and will often pursue prey for at least 1 km (0.62 mi). One wolf chased a caribou for 8 km (5 mi), another chased and tracked a deer for 20 km (12 mi), and one 11-year-old wolf chased and caught an Arctic hare after seven minutes. Most wolf prey will try to run to water, where they will either escape or be better placed to attempt to ward off the wolves.[118]
Disablement
The wolf must give chase and gain on its fleeing prey, slow it down by biting through thick hair and hide, and then disable it enough to begin feeding.[118] After chasing and then confronting a large prey animal, the wolf makes use of its 6 cm (2+1⁄2 in) fangs and its powerful masseter muscles to deliver a bite force of 28 kg/cm2 (400 lbf/in2), which is capable of breaking open the skulls of many of its prey animals. The wolf leaps at its quarry and tears at it. One wolf was observed being dragged for dozens of metres attached to the hind leg of a moose; another was seen being dragged over a fallen log while attached to a bull elk's nose.[117]
The most common point of wolf attacks on moose is the upper hind legs.[125][126][127] Hind leg wounds are inflicted from the rear, midway up the hock with the canine teeth. These leave gaping skin perforations over 4 cm (1+1⁄2 in) in diameter. Although blood loss, muscle damage, and tendon exposure may occur, there is no evidence of hamstringing. Attacks also occur on the fleshy nose, the back and sides of the neck, the ears, and the perineum.[127] Wolves may wound large prey and then lie around resting for hours before killing it when it is weaker due to blood loss, thereby lessening the risk of injury to themselves.[128]
With medium-sized prey, such as roe deer or sheep, wolves kill by biting the throat, severing nerve tracks and the carotid artery, thus causing the animal to die within a few seconds to a minute. With small, mouselike prey, wolves leap in a high arc and immobilize it with their forepaws.[129] When prey is vulnerable and abundant, wolves may occasionally surplus kill. Such instances are common with domestic animals, but rare with wild prey. In the wild, surplus killing occurs primarily during late winter or spring, when snow is unusually deep (thus impeding the movements of prey)[130] or during the denning period, when den bound wolves require a ready supply of meat.[131] Medium-sized prey are especially vulnerable to surplus killing, as the swift throat-biting method allows wolves to kill one animal quickly and move on to another.[129]
Feeding
Once prey is brought down, wolves begin to feed excitedly, ripping and tugging at the carcass in all directions, and bolting down large chunks of it.[132] The breeding pair typically monopolizes food to continue producing pups. When food is scarce, this is done at the expense of other family members, especially non-pups.[133] The breeding pair typically eats first. They usually work the hardest at killing prey, and may rest after a long hunt and allow the rest of the family to eat undisturbed. Once the breeding pair has finished eating, the rest of the family tears off pieces of the carcass and transports them to secluded areas where they can eat in peace. Wolves typically commence feeding by consuming the larger internal organs, like the heart, liver, lungs, and stomach lining. The kidneys and spleen are eaten once they are exposed, followed by the muscles.[134] A wolf can eat 15–19% of its body weight in a single feeding.[62]
Infections
Viral and bacterial
Viral diseases carried by wolves include: rabies, canine distemper, canine parvovirus, infectious canine hepatitis, papillomatosis, and canine coronavirus.[135] Wolves are a major host for rabies in Russia, Iran, Afghanistan, Iraq and India.[136] In wolves, the incubation period is eight to 21 days, and results in the host becoming agitated, deserting its pack, and travelling up to 80 km (50 mi) a day, thus increasing the risk of infecting other wolves. Infected wolves do not show any fear of humans, most documented wolf attacks on people being attributed to rabid animals. Although canine distemper is lethal in dogs, it has not been recorded to kill wolves, except in Canada and Alaska. The canine parvovirus, which causes death by dehydration, electrolyte imbalance, and endotoxic shock or sepsis, is largely survivable in wolves, but can be lethal to pups. Wolves may catch infectious canine hepatitis from dogs, though there are no records of wolves dying from it. Papillomatosis has been recorded only once in wolves, and likely does not cause serious illness or death, though it may alter feeding behaviours. The canine coronavirus has been recorded in Alaskan wolves, infections being most prevalent in winter months.[135]
Bacterial diseases carried by wolves include: brucellosis, Lyme disease, leptospirosis, tularemia, bovine tuberculosis,[137] listeriosis and anthrax.[136] Wolves can catch Brucella suis from wild and domestic reindeer. While adult wolves tend not to show any clinical signs, it can severely weaken the pups of infected females. Although lyme disease can debilitate individual wolves, it does not appear to significantly affect wolf populations. Leptospirosis can be contracted through contact with infected prey or urine, and can cause fever, anorexia, vomiting, anemia, hematuria, icterus, and death. Wolves living near farms are more vulnerable to the disease than those living in the wilderness, probably because of prolonged contact with infected domestic animal waste. Wolves may catch tularemia from lagomorph prey, though its effect on wolves is unknown. Although bovine tuberculosis is not considered a major threat to wolves, it has been recorded to have killed two wolf pups in Canada.[137]
Parasitic
Wolves carry ectoparasites and endoparasites; those in the former Soviet Union have been recorded to carry at least 50 species.[136] Most of these parasites infect wolves without adverse effects, though the effects may become more serious in sick or malnourished specimens.[138] Parasitic infection in wolves is of particular concern to people. Wolves can spread them to dogs, which in turn can carry the parasites to humans. In areas where wolves inhabit pastoral areas, the parasites can be spread to livestock.[136]
Wolves are often infested with a variety of arthropod exoparasites, including fleas, ticks, lice, and mites. The most harmful to wolves, particularly pups, is the mange mite (Sarcoptes scabiei),[138] though they rarely develop full-blown mange, unlike foxes.[37] Lice, such as Trichodectes canis, may cause sickness in wolves, but rarely death. Ticks of the genus Ixodes can infect wolves with Lyme disease and Rocky Mountain spotted fever.[138] The tick Dermacentor pictus also infests wolves. Other ectoparasites include chewing lice, sucking lice and the fleas Pulex irritans and Ctenocephalides canis.[37]
Endoparasites known to infect wolves include: protozoans and helminths (flukes, tapeworms, roundworms and thorny-headed worms). Of 30,000 protozoan species, only a few have been recorded to infect wolves: Isospora, Toxoplasma, Sarcocystis, Babesia, and Giardia.[138] Some wolves carry Neospora caninum, which can be spread to cattle and is correlated with bovine miscarriages.[139] Among flukes, the most common in North American wolves is Alaria, which infects small rodents and amphibians which are eaten by wolves. Upon reaching maturity, Alaria migrates to the wolf's intestine, but does little harm. Metorchis conjunctus, which enters wolves through eating fish, infects the wolf's liver or gall bladder, causing liver disease, inflammation of the pancreas, and emaciation. Most other fluke species reside in the wolf's intestine, though Paragonimus westermani lives in the lungs. Tapeworms are commonly found in wolves, as their primary hosts are ungulates, small mammals, and fish, which wolves feed upon. Tapeworms generally cause little harm in wolves, though this depends on the number and size of the parasites, and the sensitivity of the host. Symptoms often include constipation, toxic and allergic reactions, irritation of the intestinal mucosa, and malnutrition. Infections by the tapeworm Echinococcus granulosus in ungulate populations tend to increase in areas with high wolf densities, as wolves can shed Echinoccocus eggs in their feces onto grazing areas.[138]
Wolves can carry over 30 roundworm species, though most roundworm infections appear benign, depending on the number of worms and the age of the host. Ancylostoma caninum attaches itself on the intestinal wall to feed on the host's blood, and can cause hyperchromic anemia, emaciation, diarrhea, and possibly death. Toxocara canis, a hookworm known to infect wolf pups in the uterus, can cause intestinal irritation, bloating, vomiting, and diarrhea. Wolves may catch Dioctophyma renale from minks, which infects the kidneys, and can grow to lengths of 100 cm (40 in). D. renale causes the complete destruction of the kidney's functional tissue and can be fatal if both kidneys are infected. Wolves can tolerate low levels of Dirofilaria immitis for many years without showing any ill effects, though high levels can kill wolves through cardiac enlargement and congestive hepatopathy. Wolves probably become infected with Trichinella spiralis by eating infected ungulates. Although T. spiralis is not known to produce clinical signs in wolves, it can cause emaciation, salivation, and crippling muscle pains in dogs. Thorny-headed worms rarely infect wolves, though three species have been identified in Russian wolves: Nicolla skrjabini, Macracanthorhynchus catulinus, and Moniliformis moniliformis.[138]
Status and conservation
The global wild wolf population in 2003 was estimated at 300,000.[140] Wolf population declines have been arrested since the 1970s. This has fostered recolonization and reintroduction in parts of its former range as a result of legal protection, changes in land use, and rural human population shifts to cities. Competition with humans for livestock and game species, concerns over the danger posed by wolves to people, and habitat fragmentation pose a continued threat to the wolf. Despite these threats, the IUCN classifies the wolf as Least Concern on its Red List due to its relatively widespread range and stable population. The species is listed by the Convention on International Trade in Endangered Species of Wild Fauna and Flora in its Appendix II, indicating that it is not threatened with extinction. However, those wolf populations living in Bhutan, India, Nepal and Pakistan are listed in its Appendix I, indicating that these may become extinct without restrictions on their trade.[2]
North America
In Canada, 50,000–60,000 wolves live in 80% of their historical range, making Canada an important stronghold for the species.[41] Under Canadian law, First Nations people can hunt wolves without restrictions, but others must acquire licenses for the hunting and trapping seasons. As many as 4,000 wolves may be harvested in Canada each year.[141] The wolf is a protected species in national parks under the Canada National Parks Act.[142] In Alaska, 7,000–11,000 wolves are found on 85% of the state's 1,517,733 km2 (586,000 sq mi). Wolves may be hunted or trapped with a license; around 1,200 wolves are harvested annually.[143]
In the contiguous United States, wolf declines were caused by the expansion of agriculture, the decimation of the wolf's main prey species like the American bison, and extermination campaigns.[41] Wolves were given protection under the Endangered Species Act (ESA) of 1973, and have since returned to parts of their former range thanks to both natural recolonizations and reintroductions.[144] Wolf populations in the Great Lakes states of Minnesota, Wisconsin and Michigan number over 4,000 as of 2018.[145] Wolves also occupy much of the northern Rocky Mountains region, with at least 1,704 wolves in Montana, Idaho and Wyoming as of 2015. They have also established populations in Washington and Oregon.[146] On October 29, 2020, it was announced that the wolf would be delisted from the ESA.[147][148] In Mexico and parts of the southwestern United States, the Mexican and U.S. governments collaborated from 1977 to 1980 in capturing all Mexican wolves remaining in the wild to prevent their extinction and established captive breeding programs for reintroduction.[149]
Eurasia
Europe, excluding Russia, Belarus and Ukraine, has 17,000 wolves in more than 28 countries.[150] In many countries of the European Union, the wolf is strictly protected under the 1979 Berne Convention on the Conservation of European Wildlife and Natural Habitats (Appendix II) and the 1992 Council Directive 92/43/EEC on the Conservation of Natural Habitats and of Wild Fauna and Flora (Annex II and IV). There is extensive legal protection in many European countries, although there are national exceptions.[2][151]
Wolves have been persecuted in Europe for centuries, having been exterminated in Great Britain by 1684, in Ireland by 1770, in Central Europe by 1899, in France by the 1930s, and in much of Scandinavia by the early 1970s. They continued to survive in parts of Finland, Eastern Europe and Southern Europe.[152] Since 1980, European wolves have rebounded and expanded into parts of their former range. The decline of the traditional pastoral and rural economies seems to have ended the need to exterminate the wolf in parts of Europe.[141] As of 2016, estimates of wolf numbers include: 4,000 in the Balkans, 3,460–3,849 in the Carpathian Mountains, 1,700–2,240 in the Baltic states, 1,100–2,400 in the Italian peninsula, and around 2,500 in the northwest Iberian peninsula as of 2007.[150]
In the former Soviet Union, wolf populations have retained much of their historical range despite Soviet-era large scale extermination campaigns. Their numbers range from 1,500 in Georgia, to 20,000 in Kazakhstan and up to 45,000 in Russia.[153] In Russia, the wolf is regarded as a pest because of its attacks on livestock, and wolf management means controlling their numbers by destroying them throughout the year. Russian history over the past century shows that reduced hunting leads to an abundance of wolves.[154] The Russian government has continued to pay bounties for wolves and annual harvests of 20–30% do not appear to significantly affect their numbers.[155]
During the 19th century, wolves were widespread in many parts of the Holy Land east and west of the Jordan River, but decreased considerably in number between 1964 and 1980, largely due to persecution by farmers.[156] In the Middle East, only Israel and Oman give wolves explicit legal protection.[157] Israel has protected its wolves since 1954 and has maintained a moderately sized population of 150 through effective enforcement of conservation policies. These wolves have moved into neighboring countries. Approximately 300–600 wolves inhabit the Arabian Peninsula.[158] The wolf also appears to be widespread in Iran.[159] Turkey has an estimated population of about 7,000 wolves.[160] Outside of Turkey, wolf populations in the Middle East may total 1,000–2,000.[157]
In southern Asia, the northern regions of Afghanistan and Pakistan are important strongholds for wolves. The wolf has been protected in India since 1972.[161] Hindus traditionally considered the hunting of wolves, even dangerous ones, as taboo, for fear of causing a bad harvest. The Santals considered them fair game, as they did every other forest-dwelling animal.[162] During British rule in India, wolves were not considered game species, and were killed primarily in response to them attacking game herds, livestock, and people.[163] The Indian wolf is distributed across the states of Gujarat, Rajasthan, Haryana, Uttar Pradesh, Madhya Pradesh, Maharashtra, Karnataka and Andhra Pradesh.[164] As of 2019, it is estimated that there are around 2,000–3,000 Indian wolves in the country.[165] In East Asia, Mongolia's population numbers 10,000–20,000. In China, Heilongjiang has roughly 650 wolves, Xinjiang has 10,000 and Tibet has 2,000.[166] 2017 evidence suggests that wolves range across all of mainland China.[167] Wolves have been historically persecuted in China[168] but have been legally protected since 1998.[169] The last Japanese wolf was captured and killed in 1905.[170]
Relationships with humans
In culture
In folklore, religion and mythology
The wolf is a common motif in the mythologies and cosmologies of peoples throughout its historical range. The Ancient Greeks associated wolves with Apollo, the god of light and order.[171] The Ancient Romans connected the wolf with their god of war and agriculture Mars,[172] and believed their city's founders, Romulus and Remus, were suckled by a she-wolf.[173] Norse mythology includes the feared giant wolf Fenrir,[174] and Geri and Freki, Odin's faithful pets.[175]
In Chinese astronomy, the wolf represents Sirius and guards the heavenly gate. In China, the wolf was traditionally associated with greed and cruelty and wolf epithets were used to describe negative behaviours such as cruelty ("wolf's heart"), mistrust ("wolf's look") and lechery ("wolf-sex"). In both Hinduism and Buddhism, the wolf is ridden by gods of protection. In Vedic Hinduism, the wolf is a symbol of the night and the daytime quail must escape from its jaws. In Tantric Buddhism, wolves are depicted as inhabitants of graveyards and destroyers of corpses.[174]
In the Pawnee creation myth, the wolf was the first animal brought to Earth. When humans killed it, they were punished with death, destruction and the loss of immortality.[176] For the Pawnee, Sirius is the "wolf star" and its disappearance and reappearance signified the wolf moving to and from the spirit world. Both Pawnee and Blackfoot call the Milky Way the "wolf trail".[177] The wolf is also an important crest symbol for clans of the Pacific Northwest like the Kwakwakaʼwakw.[174]
The concept of people turning into wolves, and the inverse, has been present in many cultures. One Greek myth tells of Lycaon of Arcadia being transformed into a wolf by Zeus as punishment for his evil deeds.[178] The legend of the werewolf has been widespread in European folklore and involves people willingly turning into wolves to attack and kill others.[179] The Navajo have traditionally believed that witches would turn into wolves by donning wolf skins and would kill people and raid graveyards.[180] The Dena'ina believed wolves were once men and viewed them as brothers.[171]
In fable and literature
Aesop featured wolves in several of his fables, playing on the concerns of Ancient Greece's settled, sheep-herding world. His most famous is the fable of "The Boy Who Cried Wolf", which is directed at those who knowingly raise false alarms, and from which the idiomatic phrase "to cry wolf" is derived. Some of his other fables concentrate on maintaining the trust between shepherds and guard dogs in their vigilance against wolves, as well as anxieties over the close relationship between wolves and dogs. Although Aesop used wolves to warn, criticize and moralize about human behaviour, his portrayals added to the wolf's image as a deceitful and dangerous animal. The Bible uses an image of a wolf lying with a lamb in a utopian vision of the future. In the New Testament, Jesus is said to have used wolves as illustrations of the dangers his followers, whom he represents as sheep, would face should they follow him.[181]
Isengrim the wolf, a character first appearing in the 12th-century Latin poem Ysengrimus, is a major character in the Reynard Cycle, where he stands for the low nobility, whilst his adversary, Reynard the fox, represents the peasant hero. Isengrim is forever the victim of Reynard's wit and cruelty, often dying at the end of each story.[182] The tale of "Little Red Riding Hood", first written in 1697 by Charles Perrault, is considered to have further contributed to the wolf's negative reputation in the Western world. The Big Bad Wolf is portrayed as a villain capable of imitating human speech and disguising itself with human clothing. The character has been interpreted as an allegorical sexual predator.[183] Villainous wolf characters also appear in The Three Little Pigs and "The Wolf and the Seven Young Goats".[184] The hunting of wolves, and their attacks on humans and livestock, feature prominently in Russian literature, and are included in the works of Leo Tolstoy, Anton Chekhov, Nikolay Nekrasov, Ivan Bunin, Leonid Pavlovich Sabaneyev, and others. Tolstoy's War and Peace and Chekhov's Peasants both feature scenes in which wolves are hunted with hounds and Borzois.[185] The musical Peter and the Wolf involves a wolf being captured for eating a farm duck, but is spared and sent to a zoo.[186]
Wolves are among the central characters of Rudyard Kipling's The Jungle Book. His portrayal of wolves has been praised posthumously by wolf biologists for his depiction of them: rather than being villainous or gluttonous, as was common in wolf portrayals at the time of the book's publication, they are shown as living in amiable family groups and drawing on the experience of infirm but experienced elder pack members.[187] Farley Mowat's largely fictional 1963 memoir Never Cry Wolf is widely considered to be the most popular book on wolves, having been adapted into a Hollywood film and taught in several schools decades after its publication. Although credited with having changed popular perceptions on wolves by portraying them as loving, cooperative and noble, it has been criticized for its idealization of wolves and its factual inaccuracies.[188][189][190]
In heraldry and emblems
The wolf is a frequent charge in English heraldry. It is illustrated as a supporter on the shields of Lord Welby, Rendel, and Viscount Wolseley, and can be found on the coat of arms of Lovett and the vast majority of the Wilsons and Lows. The demi-wolf is a common crest, appearing in the arms and crests of members of many families, including that of the Wolfes, whose crest depicts a demi-wolf holding a crown in its paws, in reference to the assistance the family gave to Charles II during the Battle of Worcester. Wolf heads are common in Scottish heraldry, particularly in the coats of Clan Robertson and Skene. The wolf is the most common animal in Spanish heraldry and is often depicted as carrying a lamb in its mouth, or across its back.[191]
The wolf is featured on the flags of the Confederated Tribes of the Colville Reservation, the Oneida Nation of Wisconsin and the Pawnee.[192] The Chechen wolf has been a symbol of the Chechen Nation.[193] In modern times, the wolf is widely used as an emblem for military and paramilitary groups. It is the unofficial symbol of the spetsnaz, and serves as the logo of the Turkish Gray Wolves. During the Yugoslav Wars, several Serb paramilitary units adopted the wolf as their symbol, including the White Wolves and the Wolves of Vučjak.[194]
Conflicts
Human presence appears to stress wolves, as seen by increased cortisol levels in instances such as snowmobiling near their territory.[195]
Predation on livestock
Livestock depredation has been one of the primary reasons for hunting wolves and can pose a severe problem for wolf conservation. As well as causing economic losses, the threat of wolf predation causes great stress on livestock producers, and no foolproof solution of preventing such attacks short of exterminating wolves has been found.[196] Some nations help offset economic losses to wolves through compensation programs or state insurance.[197] Domesticated animals are easy prey for wolves, as they have been bred under constant human protection, and are thus unable to defend themselves very well.[198] Wolves typically resort to attacking livestock when wild prey is depleted.[199] In Eurasia, a large part of the diet of some wolf populations consists of livestock, while such incidents are rare in North America, where healthy populations of wild prey have been largely restored.[196]
The majority of losses occur during the summer grazing period, untended livestock in remote pastures being the most vulnerable to wolf predation.[200] The most frequently targeted livestock species are sheep (Europe), domestic reindeer (northern Scandinavia), goats (India), horses (Mongolia), cattle and turkeys (North America).[196] The number of animals killed in single attacks varies according to species: most attacks on cattle and horses result in one death, while turkeys, sheep and domestic reindeer may be killed in surplus.[201] Wolves mainly attack livestock when the animals are grazing, though they occasionally break into fenced enclosures.[202]
Competition with dogs
A review of the studies on the competitive effects of dogs on sympatric carnivores did not mention any research on competition between dogs and wolves.[203][204] Competition would favour the wolf, which is known to kill dogs; however wolves usually live in pairs or in small packs in areas with high human persecution, giving them a disadvantage when facing large groups of dogs.[204][205]
Wolves kill dogs on occasion, and some wolf populations rely on dogs as an important food source. In Croatia, wolves kill more dogs than sheep, and wolves in Russia appear to limit stray dog populations. Wolves may display unusually bold behaviour when attacking dogs accompanied by people, sometimes ignoring nearby humans. Wolf attacks on dogs may occur both in house yards and in forests. Wolf attacks on hunting dogs are considered a major problem in Scandinavia and Wisconsin.[196][206] The most frequently killed hunting breeds in Scandinavia are Harriers, older animals being most at risk, likely because they are less timid than younger animals, and react to the presence of wolves differently. Large hunting dogs such as Swedish Elkhounds are more likely to survive wolf attacks because of their better ability to defend themselves.[206]
Although the number of dogs killed each year by wolves is relatively low, it induces a fear of wolves' entering villages and farmyards to prey on them. In many cultures, dogs are seen as family members, or at least working team members, and losing one can lead to strong emotional responses such as demanding more liberal hunting regulations.[204]
Dogs that are employed to guard sheep help to mitigate human–wolf conflicts, and are often proposed as one of the non-lethal tools in the conservation of wolves.[204][207] Shepherd dogs are not particularly aggressive, but they can disrupt potential wolf predation by displaying what is to the wolf ambiguous behaviours, such as barking, social greeting, invitation to play or aggression. The historical use of shepherd dogs across Eurasia has been effective against wolf predation,[204][208] especially when confining sheep in the presence of several livestock guardian dogs.[204][209] Shepherd dogs are sometimes killed by wolves.[204]
Attacks on humans
The fear of wolves has been pervasive in many societies, though humans are not part of the wolf's natural prey.[210] How wolves react to humans depends largely on their prior experience with people: wolves lacking any negative experience of humans, or which are food-conditioned, may show little fear of people.[211] Although wolves may react aggressively when provoked, such attacks are mostly limited to quick bites on extremities, and the attacks are not pressed.[210]
Predatory attacks may be preceded by a long period of habituation, in which wolves gradually lose their fear of humans. The victims are repeatedly bitten on the head and face, and are then dragged off and consumed unless the wolves are driven off. Such attacks typically occur only locally and do not stop until the wolves involved are eliminated. Predatory attacks can occur at any time of the year, with a peak in the June–August period, when the chances of people entering forested areas (for livestock grazing or berry and mushroom picking) increase.[210] Cases of non-rabid wolf attacks in winter have been recorded in Belarus, Kirov and Irkutsk oblasts, Karelia and Ukraine. Also, wolves with pups experience greater food stresses during this period.[37] The majority of victims of predatory wolf attacks are children under the age of 18 and, in the rare cases where adults are killed, the victims are almost always women.[210] Indian wolves have a history of preying on children, a phenomenon called "child-lifting". They may be taken primarily in the summer period in the evening hours, and often within human settlements.[212]
Cases of rabid wolves are low when compared to other species, as wolves do not serve as primary reservoirs of the disease, but can be infected by animals such as dogs, jackals and foxes. Incidents of rabies in wolves are very rare in North America, though numerous in the eastern Mediterranean, the Middle East and Central Asia. Wolves apparently develop the "furious" phase of rabies to a very high degree. This, coupled with their size and strength, makes rabid wolves perhaps the most dangerous of rabid animals.[210] Bites from rabid wolves are 15 times more dangerous than those of rabid dogs.[213] Rabid wolves usually act alone, travelling large distances and often biting large numbers of people and domestic animals. Most rabid wolf attacks occur in the spring and autumn periods. Unlike with predatory attacks, the victims of rabid wolves are not eaten, and the attacks generally occur only on a single day. The victims are chosen at random, though most cases involve adult men. During the fifty years up to 2002, there were eight fatal attacks in Europe and Russia, and more than two hundred in southern Asia.[210]
Human hunting of wolves
Theodore Roosevelt said wolves are difficult to hunt because of their elusiveness, sharp senses, high endurance, and ability to quickly incapacitate and kill a dog.[214] Historic methods included killing of spring-born litters in their dens, coursing with dogs (usually combinations of sighthounds, Bloodhounds and Fox Terriers), poisoning with strychnine, and trapping.[215][216]
A popular method of wolf hunting in Russia involves trapping a pack within a small area by encircling it with fladry poles carrying a human scent. This method relies heavily on the wolf's fear of human scents, though it can lose its effectiveness when wolves become accustomed to the odor. Some hunters can lure wolves by imitating their calls. In Kazakhstan and Mongolia, wolves are traditionally hunted with eagles and falcons, though this practice is declining, as experienced falconers are becoming few in number. Shooting wolves from aircraft is highly effective, due to increased visibility and direct lines of fire.[216] Several types of dog, including the Borzoi and Kyrgyz Tajgan, have been specifically bred for wolf hunting.[204]
As pets and working animals
Wolves and wolf-dog hybrids are sometimes kept as exotic pets. Although closely related to domestic dogs, wolves do not show the same tractability as dogs in living alongside humans, being generally less responsive to human commands and more likely to act aggressively. A person is more likely to be fatally mauled by a pet wolf or wolf-dog hybrid than by a dog.[217]
Notes
- ^ The domesticated dog (C.l. familiaris) and the dingo/New Guinea singing dog (C.l. dingo) are included in the phylogenetic but not colloquial definition of 'wolf', and thus not in the scope of this article.
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Bibliography
- Busch, R. H. (2007). Wolf Almanac, New and Revised: A Celebration Of Wolves And Their World (3 ed.). Rowman & Littlefield. ISBN 978-1-59921-069-8.
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- Lopez, Barry H. (1978). Of Wolves and Men. J. M. Dent and Sons Limited. ISBN 978-0-7432-4936-2.
- Marvin, Garry (2012). Wolf. Reaktion Books Ltd. ISBN 978-1-86189-879-1.
- Mech, L. David (1981). The Wolf: The Ecology and Behaviour of an Endangered Species. University of Minnesota Press. ISBN 978-0-8166-1026-6.
- Mech, L. David; Boitani, Luigi, eds. (2003). Wolves: Behaviour, Ecology and Conservation. University of Chicago Press. ISBN 978-0-226-51696-7.
- Mech, David L.; Smith, Douglas W.; MacNulty, Daniel R. (2015). Wolves on the Hunt: The Behavior of Wolves Hunting Wild Prey. University of Chicago Press. ISBN 978-0-226-25514-9.
External links
 | Wikimedia Commons has media related to Canis lupus. |
 | Wikispecies has information related to Canis lupus. |
 | Wikiquote has quotations related to: Wolf |
- Gray wolf (mammal) at the Encyclopædia Britannica
- The International Wolf Center
- Staying Safe in Wolf Country, ADFG (January 2009)
- Watch Death of a Legend and Cry of the Wild by Bill Mason
