Der Wolf ( Canis lupus ), auch als grau bekannt ] / grauer Wolf oder Timberwolf [4][5] ist ein in der Wildnis und in abgelegenen Gebieten von Eurasien und Nordamerika beheimateter Hund. Es ist das größte verbliebene Mitglied seiner Familie, mit Männern von durchschnittlich 43 bis 45 kg und Frauen mit 36 bis 38,5 kg. [6] Es unterscheidet sich von anderen Canis durch ihre größere Größe und weniger spitze Züge, besonders an den Ohren und am Maulkorb. [7] Sein Winterfell ist lang und buschig und überwiegend fleckig grau, obwohl auch fast reines Weiß, Rot und Braun bis Schwarz vorkommen. [5] Säugetierarten der Welt (3. Auflage, 2005), ein Standard-Nachschlagewerk in der Zoologie, erkennt 38 Unterarten von C an. Lupus . [8]
Der graue Wolf ist das zweithäufigste Mitglied der Gattung Canis nach dem äthiopischen Wolf, wie seine morphologischen Anpassungen an die Jagd auf große Beute, seine eher gesellige Natur zeigen. 19659009] und sein hochentwickeltes Ausdrucksverhalten. [10][11] Trotzdem ist es eng verwandt mit kleineren Canis -Sorten wie dem Kojoten [12] und dem Goldschakal [13][14] um fruchtbare Hybriden zu erzeugen. Es ist die einzige Spezies von Canis die eine Reihe umfasst, die sowohl die Alte als auch die Neue Welt umfasst [15] und ihren Ursprung in Eurasien während des Pleistozäns hatte und Nordamerika mindestens dreimal während des Rancholabrean besiedelte. 19659014] Es ist ein soziales Tier, das in Kernfamilien unterwegs ist und aus einem Paar besteht, das von einem erwachsenen Nachwuchs des Paares begleitet wird. [16] Der graue Wolf ist in seiner gesamten Reichweite typischerweise ein Raubtier, wobei nur Menschen und Tiger [9][17][18][19] ein Tier zeigen ernsthafte Bedrohung Es ernährt sich hauptsächlich von großen Huftieren, frisst aber auch kleinere Tiere, Vieh, Aas und Müll. [5] Ein siebenjähriger Wolf gilt als relativ alt und die maximale Lebensdauer beträgt etwa 16 Jahre. [20]
Die Weltbevölkerung für graue Wölfe wird auf 300.000 geschätzt. [21] Der graue Wolf ist eines der bekanntesten und am meisten erforschten Tiere der Welt, mit wahrscheinlich mehr Büchern darüber geschrieben als jede andere Tierart. [22] Es hat eine Eine lange Geschichte der Verbindung mit Menschen, die in den meisten Hirtengemeinschaften wegen ihrer Angriffe auf Vieh verachtet und gejagt wurde, während sie in einigen Agrar- und Jäger-Sammler-Gesellschaften umgekehrt respektiert wurde. [23] Obwohl die Angst vor Wölfen in vielen menschlichen Gesellschaften allgegenwärtig ist Die Mehrzahl der aufgezeichneten Angriffe auf Menschen wurde auf Tollwut erkrankte Tiere zurückgeführt. Nicht tollwütige Wölfe haben Menschen angegriffen und getötet, hauptsächlich Kinder. Dies ist jedoch selten, da Wölfe relativ wenige Menschen sind und von Menschen leben und eine Angst vor Menschen durch Jäger und Hirten entwickelt haben. [24]
Etymology
The English "Wolf" stammt aus dem Altenglisch Wulf von dem angenommen wird, dass er aus dem Protogermanischen * Wulfaz stammt. Der lateinische Lupus ist ein Sabine-Lehnwort. [23] Beide stammen von der Proto-Indo-Europäischen Wurzel * wlq w os oder * luk w os . [25]
Taxonomie und Evolution
Taxonomie
Die Art Canis Lupus wurde erstmals von Carl Linnaeus in seiner Publikation Systema Naturae aufgezeichnet ] im Jahre 1758, [3] mit der lateinischen Klassifikation übersetzt in die englischen Wörter "Hundewolf". Die siebenunddreißig Unterarten von Canis lupus sind unter der Bezeichnung "Wolf" in "Säugetierarten der dritten Ausgabe" von 2005 aufgeführt. [8] Die nominierte Unterart ist der Eurasische Wolf ( Canis lupus lupus ), auch bekannt als der gewöhnliche Wolf. [26] Die Unterart umfasst den Haushund, den Dingo, den östlichen Wolf und den roten Wolf, listet jedoch auf C. l. italicus als Synonym für C. l. Lupus . [8] Allerdings wurde die Einstufung von mehreren als Arten oder Unterarten in Frage gestellt.
Origin
Die Entwicklung des Wolfes fand über eine geologische Zeitspanne von mindestens 300.000 Jahren statt.
Der graue Wolf Canis lupus ist eine sehr anpassungsfähige Spezies, die in einer Reihe von Umgebungen existieren kann und eine weite Verbreitung in der Holarktis besitzt. Studien über moderne graue Wölfe haben unterschiedliche Bevölkerungsgruppen identifiziert, die nahe beieinander leben. [27] [28] Diese Variation in den Unterpopulationen ist eng mit den Unterschieden in den Habitaten - Niederschlag, Temperatur, Vegetation und Beutespezialisierung - verbunden, die die cranio-dentale Plastizität beeinflussen. [29] [30] [1965000] [19650008] [31] [19659035] [32]
Die archäologischen und paläontologischen Aufzeichnungen zeigen, dass der graue Wolf seit mindestens 300.000 Jahren anwesend ist. [33] Diese ständige Präsenz steht im Gegensatz zu genomischen Analysen, die darauf schließen lassen, dass alle modernen Wölfe und Hunde von einer gemeinsamen Wolfspopulation abstammen [34] [35] [36] die erst vor 20.000 existierte Jahren [34] Diese Analysen deuten auf einen Engpass in der Bevölkerung hin, gefolgt von einer schnellen Bestrahlung einer Ahnenpopulation zu einem Zeitpunkt während oder kurz nach dem Letzten Gletschermaximum. Der geographische Ursprung dieser Strahlung ist jedoch nicht bekannt.
Im Jahr 2018 wurde die gesamte Genomsequenzierung verwendet, um Mitglieder der Gattung Canis sowie den Dhole ( Cuon alpinus ) und den afrikanischen Jagdhund ( Lycaon pictus [19]) zu vergleichen. ). Es gibt Hinweise auf Genfluss zwischen afrikanischen Goldwölfen, Goldschakalen und grauen Wölfen. Ein afrikanischer Goldwolf von der ägyptischen Sinai-Halbinsel zeigte eine starke Vermischung mit den Wölfen und Hunden des Nahen Ostens, was die Rolle der Landbrücke zwischen dem afrikanischen und dem eurasischen Kontinent in der Canid-Evolution unterstreicht. Es gab Anzeichen für einen Genfluss zwischen Goldschakalen und Wölfen des Nahen Ostens, weniger bei europäischen und asiatischen Wölfen und am wenigsten bei nordamerikanischen Wölfen. Die Studie schlägt vor, dass die Vorfahren der Goldschakale, die bei nordamerikanischen Wölfen gefunden wurden, möglicherweise vor der Divergenz der eurasischen und nordamerikanischen Grauwölfe aufgetreten sind. Die Studie zeigt, dass der gemeinsame Vorfahr des Kojoten und des Grauen Wolfs genetisch mit einer Geisterpopulation eines ausgestorbenen, nicht identifizierten Canids vermischt wurde. Die Canid ist genetisch nahe an der Dhole und hat sich nach der Abweichung des afrikanischen Jagdhundes von den anderen Canid-Arten entwickelt. Die Grundposition des Kojoten im Vergleich zum Wolf wird vermutet, weil der Kojote mehr Mitochondrien-Genoms dieses unbekannten Canids beibehielt. [37]
Bevölkerungsstruktur
Im Jahr 2013 wurde eine genetische Studie gefunden dass die Wolfspopulation in Europa entlang einer Nord-Süd-Achse aufgeteilt wurde und fünf große Cluster bildete. Es wurden drei Cluster identifiziert, die Süd- und Mitteleuropa in Italien besetzten, der Dinaric-Balkan und die Karpaten. Es wurden zwei weitere Cluster identifiziert, die Nordmitteleuropa und die ukrainische Steppe besetzten. Der italienische Wolf bestand aus einer isolierten Population mit geringer genetischer Vielfalt. Wölfe aus Kroatien, Bulgarien und Griechenland bildeten den Dinaric-Balkan-Cluster. Wölfe aus Finnland, Lettland, Weißrussland, Polen und Russland bildeten den Cluster Nord-Mitteleuropa. Wölfe aus den Karpaten stammen aus einer Mischung von Wölfen aus dem Nord-Zentral-Cluster und dem Dinaric-Balkan-Cluster. Die Wölfe aus den Karpaten waren den Wölfen aus der Pontisch-Kaspischen Steppe ähnlicher als Wölfen aus Nordmitteleuropa. Diese Cluster könnten das Ergebnis der Expansion von Gletscherrefugien gewesen sein, eine Anpassung an die örtlichen Gegebenheiten, die Fragmentierung der Landschaft und das Töten von Wölfen in einigen Gebieten durch den Menschen. [38]
Im Jahr 2016 zwei genetische Studien von North Amerikanische graue Wölfe stellten fest, dass sie sechs Ökotypen bildeten - genetisch und ökologisch unterscheidbare Populationen, die aufgrund ihres unterschiedlichen Lebensraums von anderen Populationen getrennt waren. Diese sechs Wolf-Ökotypen wurden West Forest, Boreal Forest, Arctic, High Arctic, Baffin und British Columbia genannt. Die Studien fanden heraus, dass der Niederschlag und der mittlere Tagestemperaturbereich die einflussreichsten Variablen waren. [28] [39] Diese Ergebnisse stimmten mit früheren Studien überein, dass Niederschlag die Morphologie beeinflusste und dass Vegetation und Lebensraumtyp die Wolfsunterschiede beeinflussten. [39] Die lokale Anpassung eines Wolf-Ökotyps spiegelt höchstwahrscheinlich die Präferenz eines Wolfs wider, in dem Lebensraum zu bleiben, in dem er geboren wurde. [28]
Hybridisierung mit anderen Canis
Man dachte, dies sei einmal der Fall Hunde und graue Wölfe haben sich nicht freiwillig in freier Wildbahn miteinander gekreuzt, obwohl sie fruchtbare Wolfshundkinder hervorbringen können. [40] Im Jahr 2010 ergab eine Studie von 74 italienischen männlichen Wolfslinien, dass fünf von ihnen von der Herkunft der Hunde stammten, was darauf hinweist, dass weibliche Wölfe mit streunenden männlichen Hunden in freier Wildbahn brüten werden. [41] In Nordamerika erhielten schwarze Wölfe ihre Färbung durch die vor 10.000 bis 15.000 Jahren erfolgte Wolf-Hund-Hybridisierung. [42] Wie reine Wölfe brüten Hybriden einmal im Jahr, obwohl ihre Paarungszeit drei Monate früher erfolgt, wobei die Welpen meistens im Winter geboren werden, wodurch ihre Überlebenschancen verringert werden. [40] Eine im Kaukasus durchgeführte genetische Studie zeigte jedoch, dass bis zu 10% der Hunde in der Umgebung, einschließlich Viehhüterhunde, Hybriden der ersten Generation sind. [43] Die Zucht von Wolf-Hund-Hybriden in Gefangenschaft hat sich in den Vereinigten Staaten verbreitet, wo schätzungsweise 300.000 Einwohner leben. [44]
Der graue Wolf mischt sich nicht ohne weiteres mit Kojoten. [15] Dennoch im Jahr 2018 In einer Studie wurde die genomische Struktur und Beimischung nordamerikanischer Wölfe, Kojoten und anderer wolfähnlicher Caniden untersucht, wobei Proben aus ihrem gesamten Bereich verwendet wurden, die den größten Datensatz von Kerngenomsequenzen kartierten und mit dem Wolfsreferenzgenom verglichen hatten. Die Studie unterstützt die Ergebnisse früherer Studien, dass nordamerikanische Grauwölfe und andere wolfartige Caniden das Ergebnis einer komplexen Mischung von Grauwolf und Kojoten waren. Ein polnischer Wolf aus Grönland und ein Kojote aus Mexiko waren die reinsten Exemplare. Die Kojoten aus Alaska, Kalifornien, Alabama und Quebec zeigen fast keine Wolfsvorfahren. Kojoten aus Missouri, Illinois und Florida weisen 5–10% Wolfsvorfahren auf. Es gab 40%: 60% Wolf-Kojote-Abstammung bei roten Wölfen, 60%: 40% bei östlichen Wölfen aus Ontario und 75%: 25% bei Wölfen in der Region Great Lakes. Es gab 10% Coyote-Abstammung bei mexikanischen Wölfen, 5% bei Pazifikküste und Yellowstone-Wölfen und weniger als 3% bei kanadischen Archipelwölfen. [45] Die Studie zeigt, dass die genomische Abstammung der Rot-, Ost- und Great Lakes-Wölfe das Ergebnis der Beimischung moderner Grauwölfe zu modernen Kojoten war. Es folgte die Entwicklung zur lokalen Bevölkerung. Individuen in jeder Gruppe wiesen ein einheitliches Verhältnis von Kojote zu Wolf auf, was darauf hindeutet, dass dies das Ergebnis einer relativ alten Vermischung war. Der östliche Wolf (Angonquin Provincial Park) ist genetisch eng mit dem Wolf der Großen Seen (Minnisota, Isle Royale National Park) verwandt. Wenn eine dritte Canid an der Beimischung der nordamerikanischen wolfähnlichen Caniden beteiligt gewesen wäre, wäre ihre genetische Signatur bei Kojoten und Wölfen gefunden worden, was sie nicht gefunden hat. [45]
Außerdem genetische Marker für die Kojote wurden in einigen wilden, isolierten Grauwolfpopulationen im Süden der Vereinigten Staaten gefunden. Y-Chromosomen von grauen Wölfen wurden auch in texanischen Kojotenhaplotypen gefunden. [46] In Tests, die an einem texanischen Canid aus mehrdeutigen Spezies durchgeführt wurden, zeigte die mtDNA-Analyse, dass es sich um einen Kojoten handelt, während nachfolgende Tests zeigten, dass es sich um einen Kojotengrau-Wolf-Hybriden handelte, der von einem männlichen mexikanischen Grauwolf vererbt wurde. [47] Im Jahr 2013 brachte ein Zuchtversuch in Gefangenschaft in Utah zwischen grauen Wölfen und westlichen Kojoten sechs Hybriden durch künstliche Befruchtung hervor. Damit war dies der erste Hybridisierungsfall zwischen reinen Kojoten und nordwestlichen Wölfen. Im Alter von sechs Monaten wurden die Hybriden engmaschig überwacht und zeigten bei beiden Spezies sowohl physikalische als auch Verhaltensmerkmale. [12]
Obwohl Hybridisierung zwischen Wölfen und Goldschakalen niemals beobachtet wurde, sind Hinweise auf solche Vorkommnisse zu beobachten wurde durch mtDNA-Analyse an Schakalen in Bulgarien entdeckt. [14] Obwohl es im Kaukasus keine genetischen Beweise für eine Grau-Wolf-Schakal-Hybridisierung gibt, gab es Fälle, in denen ansonsten genetisch reine Goldschakale bemerkenswert graue Wolf-ähnliche Phänotypen aufwiesen der Punkt, von ausgebildeten Biologen für Wölfe gehalten zu werden [43]
Körperliche Beschreibung
Anatomie und Abmessungen
Der graue Wolf ist das größte noch existierende Mitglied der Canidae mit Ausnahme einiger großer Hausrassen. [48] Das Gewicht und die Größe von Grauwölfen können sich weltweit stark unterscheiden und neigen dazu, proportional mit dem Breitengrad zuzunehmen, wie von Bergmanns Regel [9] vorhergesagt. Die großen Wölfe Alaskas und Kanadas wiegen manchmal 3–6-mal mehr als ihre Verwandten aus dem Nahen Osten und Südasien. [49] Erwachsene Wölfe sind im Durchschnitt 105–160 cm (41–63 in) lang und 80–85 cm (31–33 in) in Schulterhöhe. [9] Der Schwanz misst 29–50 cm (11–20 in) Länge. Die Ohren sind 90–110 mm hoch und die Hinterfüße sind 220–250 mm. [9] Die mittlere Körpermasse des vorhandenen grauen Wolfes beträgt 40 kg ), wobei das kleinste Exemplar bei 12 kg (26 lb) und das größte bei 79,4 kg (175 lb) gemessen wurde. [50] [51] [52] [19650061] [53] 19659062] [54] Das Gewicht des grauen Wolfs variiert geografisch. Im Durchschnitt können europäische Wölfe 38,5 kg wiegen, nordamerikanische Wölfe 36 kg und indische und arabische Wölfe 25 kg. [55] Frauen in einer bestimmten Wolfspopulation wiegen normalerweise 2,3 bis 4,5 kg weniger als Männer. [56] Wölfe mit einem Gewicht von über 54 kg sind ungewöhnlich, obwohl in Alaska (Kanada), [56] und den Wäldern Westrusslands außergewöhnlich große Individuen registriert wurden. [9] Der schwerste aufgezeichnete graue Wolf im Norden Amerika wurde am 12. Juli 1939 am 70 Mile River in Ost-Zentral-Alaska getötet und wog 79,4 kg. [57]
Im Vergleich zu seinen nächsten wilden Cousins (dem Kojoten und Goldschakal) Der graue Wolf ist größer und schwerer mit breiterer Schnauze, kürzeren Ohren, kurzem Rumpf und längerem Schwanz. [9][13] [48] Es ist ein schlankes, kräftig gebautes Tier mit einem großen, tief abfallenden Brustkorb, einem abfallenden Rücken und einem stark muskulösen Hals. [9] Die Wolfsbeine sind mäßig länger als die der anderen Caniden, wodurch sich das Tier schnell bewegen kann und ermöglicht um den Tiefschnee zu überwinden, der den größten Teil seines geografischen Verbreitungsgebietes bedeckt. [58] Die Ohren sind relativ klein und dreieckig. [9] Frauen neigen zu schmaleren Mündungen und Stirnen, dünneren Hälsen, etwas kürzeren Beinen und weniger massiven Schultern als Männer. [59]
Der graue Wolf trägt gewöhnlich seinen Kopf auf der gleichen Ebene wie der Rücken, nur wenn er wach ist. [9] Er bewegt sich normalerweise in einem langsamen Tempo und setzt seine Pfoten direkt vor die andere. Dieser Gang kann stundenlang mit einer Geschwindigkeit von 8–9 km / h gehalten werden, [60] und erlaubt dem Wolf, große Entfernungen zu überwinden. Auf nackten Wegen kann ein Wolf schnell Geschwindigkeiten von 50 bis 60 km / h erreichen. Der graue Wolf hat einen Laufschritt von 55–70 km / h, kann in einer einzelnen Schräglage horizontal um 5 m springen und kann für mindestens 20 Minuten eine schnelle Verfolgung aufrechterhalten. 19460059]
Im Allgemeinen haben Wölfe ein hohes Herzgewicht von 0,93% bis 1,07% Gesamtkörpermasse im Vergleich zum durchschnittlichen Säugetier bei 0,59% Gesamtkörpermasse. [62] Wölfe haben eine verminderte Herzfrequenz, was auf eine Herzvergrößerung und Hypertrophie schließen lässt. Die tibetischen Grauwölfe, die Gebiete bis zu 3.000 Meter über dem Meeresspiegel besetzen, haben Herzen entwickelt, die den niedrigen Sauerstoffwerten widerstehen. [63] Insbesondere haben diese Wölfe eine starke Selektion für RYR2, ein Gen, das die Herzanregung initiiert.
Schädel und Gebiss
Der graue Wolfskopf ist groß und schwer, hat eine breite Stirn, kräftige Kiefer und eine lange, stumpfe Schnauze. [9] Der Schädel hat eine durchschnittliche Länge von 230–280 mm und 130–150 mm (5,1–5,9 in) breit. [64] Die Zähne sind schwer und groß und eignen sich besser zum Zerquetschen von Knochen als die von anderen vorhandenen Caniden, wenn auch nicht so spezialisiert wie in Hyänen. [65] [66] Seine Backenzähne haben eine flache Kaufläche, jedoch nicht in dem Maße wie der Kojote, dessen Ernährung mehr pflanzliche Stoffe enthält. [4] Die Kiefer des grauen Wolfes können einen Druck von vielleicht 10.340 kPa (1.500 psi) ausüben, verglichen mit 5.200 kPa ( 750 psi) für einen deutschen Schäferhund. Diese Kraft reicht aus, um die meisten Knochen aufzubrechen. [67] Eine Untersuchung der geschätzten Bisskraft an den Eckzähnen einer großen Stichprobe lebender und fossiler Säugetier-Raubtiere, wenn sie für die Körpermasse eingestellt wurde, ergab, dass bei Plazentasäugern die Bisskraft an den Eckzähnen (in Newton / Kilogramm Körpergewicht) am größten war in dem ausgestorbenen Wolf (163), dann unter den noch vorhandenen Caniden die vier Hyperkarnivore, die oft Tiere jagen, die größer sind als sie selbst: der afrikanische Jagdhund (142), der graue Wolf (136), der Dhole (112) und der Dingo (108). Ein ähnlicher Trend wurde bei der karnassischen Zahnbisskraft festgestellt, wobei jedoch der ausgestorbene Wolf und der graue Wolf beide gemessen wurden (141), gefolgt von dem afrikanischen Jagdhund (136), dem Dhole (114) und dem Dingo (113). [68]
Pelz
Der graue Wolf hat ein sehr dichtes und flauschiges Winterfell mit kurzem Unterpelz und langen, groben Wachhaaren. [9] Die meisten Unterpelzhaare und einige Wachhaare sind abgelegt im Frühling und im Herbst nachwachsen. [55] Die längsten Haare treten am Rücken auf, insbesondere an der vorderen Seite und am Hals. Besonders lange Haare befinden sich auf den Schultern und bilden im oberen Teil des Halses fast einen Kamm. Die Haare auf den Wangen sind länglich und bilden Büschel. Die Ohren sind mit kurzen Haaren bedeckt, die stark aus dem Fell herausragen. An den Gliedmaßen sind kurze, elastische und eng anliegende Haare von den Ellbogen bis zu den Sehnen des Kalzanealachs vorhanden. [9] Das Winterfell ist sehr widerstandsfähig gegen Kälte; Wölfe in nördlichen Klimazonen können sich in offenen Bereichen bei -40 ° bequem ausruhen, indem sie ihre Schnauze zwischen den Hinterbeinen platzieren und ihre Gesichter mit ihrem Schwanz bedecken. Wolfspelz bietet eine bessere Isolierung als Hundefell und sammelt kein Eis, wenn der warme Atem dagegen verdichtet wird. [55] In warmen Klimazonen ist das Fell gröber und knapper als in Nordwölfen. [9] Weibliche Wölfe neigen zu glatteren Gliedmaßen als Männer und entwickeln im Allgemeinen mit dem Alter die glattesten Mäntel. Ältere Wölfe haben in der Regel mehr weiße Haare in der Schwanzspitze, entlang der Nase und an der Stirn. Das Winterfell bleibt bei säugenden Weibchen am längsten erhalten, allerdings mit etwas Haarausfall um die Brustwarzen. [59] Die Haarlänge in der Mitte des Rückens beträgt 60–70 mm (2,4–2,8 Zoll). Die Haarlänge der Schutzhaare auf den Schultern überschreitet im Allgemeinen nicht 90 mm (3,5 Zoll), kann jedoch 110–130 mm (4,3–5,1 Zoll) betragen. [9]
Die Fellfarbe reicht von fast reinem Weiß über verschiedene Schattierungen von Blond, Creme und Ocker bis hin zu Grau, Braun und Schwarz [69] wobei die Fellfarbe in höheren Breiten variiert. 19460077] Unterschiede in der Fellfarbe zwischen den Geschlechtern sind weitgehend nicht vorhanden, [9] obwohl Frauen rötere Töne haben können. [71] Schwarze Wölfe in Nordamerika erbten das für den Melanismus verantwortliche K b aus früheren Kreuzungen mit Hunden [42] während die Mutation bei Wölfen aus dem Iran als natürlich vorkommend befunden wurde. [72] Schwarze Exemplare sind in Nordamerika häufiger als in Eurasien, wobei etwa die Hälfte der Wölfe im Yellowstone-Nationalpark schwarz ist. [42]
Verhalten
Soziales und territoriales Verhalten
Der graue Wolf ist ein sozialer Tier, dessen soziale Grundeinheit aus einem Paar besteht, begleitet von den erwachsenen Nachkommen des Paares. [b] Das durchschnittliche Rudel besteht aus einer Familie von 5–11 Tieren (1–2 Erwachsene, 3–6 Jugendliche und 1–3 Jährige), [9] oder manchmal zwei oder drei solcher Familien, [4] mit außergewöhnlich großen Rudeln, die aus bis zu drei Tieren bestehen 42 Wölfe sind bekannt. [74] Unter idealen Bedingungen bringt das Paar jedes Jahr Welpen hervor, die normalerweise 10 bis 54 Monate im Rudel bleiben, bevor sie sich zerstreuen. [16] Zu den Auslösern für die Verbreitung gehören der Ausbruch der Geschlechtsreife und der Wettbewerb innerhalb des Packs um Nahrung. [75] Die Entfernung, die die zerstreuten Wölfe zurücklegen, variiert stark. einige bleiben in der Nähe der elterlichen Gruppe, während andere Personen große Entfernungen von 206 km (390 Meilen), 390 km (240 Meilen) und 670 km (420 Meilen) von ihren Geburtspaketen zurücklegen können. [61] Ein neues Rudel wird gewöhnlich von einem nicht verwandten Mann und einer Frau gegründet, die zusammen auf der Suche nach einem Gebiet unterwegs sind, das frei von anderen feindlichen Rudeln ist. [76] Wolfsrudel nehmen selten andere Wölfe in ihre Herde auf und töten sie normalerweise. In den seltenen Fällen, in denen andere Wölfe adoptiert werden, handelt es sich bei dem Adoptierten fast immer um ein unreifes Tier (1–3 Jahre), bei dem es unwahrscheinlich ist, dass es um das Zuchtrecht mit dem Paar steht. In einigen Fällen wird ein einzelner Wolf in einen Rudel aufgenommen, um einen verstorbenen Züchter zu ersetzen. [74] In Zeiten ungleichmäßiger Huftiere (Migration, Kalbung usw.) können sich vorübergehend verschiedene Wolfsrudel zusammenschließen. [9] Untersuchungen des Cortisolspiegels von Wölfen zeigen, dass sie erheblich ansteigen, wenn ein Rudelmitglied stirbt, was auf Stress hindeutet. [77]
Wölfe sind höchst territoriale Tiere und errichten im Allgemeinen weit größere Gebiete, als sie zum Überleben benötigen, um eine konstante Beute zu gewährleisten. Die Größe des Territoriums hängt weitgehend von der Menge der verfügbaren Beute und dem Alter der Jungtiere ab, die in Gebieten mit niedrigen Beutepopulationen tendenziell an Größe zunehmen [78] oder wenn die Jungen das Alter von 6 Monaten erreichen mit den gleichen Ernährungsbedürfnissen wie Erwachsene. [79] Wolfsrudel sind ständig auf der Suche nach Beute und decken ungefähr 9% ihres Territoriums pro Tag ab (durchschnittlich 25 km / d). Der Kern ihres Territoriums beträgt im Durchschnitt 35 km 2 (14 sq mi), in dem sie 50% ihrer Zeit verbringen. [78] Die Beutendichte ist in den umliegenden Gebieten des Territoriums tendenziell viel höher, obwohl Wölfe die Jagd in den Randbereichen ihres Verbreitungsgebiets vermeiden, es sei denn, sie sind verzweifelt, da möglicherweise tödliche Begegnungen mit benachbarten Rudeln auftreten. [80] Das kleinste verzeichnete Gebiet wurde von einem Rudel von sechs Wölfen im Nordosten von Minnesota gehalten, das geschätzte 33 km einnahm 2 (13 sq mi), während das größte von einem alaskischen Rudel von zehn Wölfen gehalten wurde, die ein Wale umfassten 6.272 km 2 (2,422 km²) Fläche. [79] Wolfsrudel sind in der Regel besiedelt und verlassen ihre gewohnten Weiten meist nur bei schwerem Nahrungsmittelknappheit. [19459171[9]
Wölfe verteidigen ihr Territorium durch eine Kombination aus Duftmarkierungen, direkten Angriffen und Heulen (siehe Mitteilung). Duftmarkierung wird für territoriale Werbung verwendet und umfasst Urinieren, Defäkation und Bodenkratzer. [81] [82] [83] [84] [85] [85] [194560104] Duftmarken werden in der Regel alle 240 m auf regelmäßigen Wegen und Kreuzungen im gesamten Gebiet hinterlassen. Solche Marker können 2–3 Wochen bestehen [79] und werden typischerweise in der Nähe von Felsen, Felsbrocken, Bäumen oder den Skeletten großer Tiere platziert. [9] Territoriale Kämpfe gehören zu den Hauptursachen der Wolfsterblichkeit Eine Studie schlussfolgerte, dass 14–65% der Todesfälle durch Wolf in Minnesota und im Denali National Park and Preserve auf Raubtiere anderer Wölfe zurückzuführen waren. [86]
Fortpflanzung und Entwicklung
Der graue Wolf ist im Allgemeinen monogam, [87] mit zusammengehörenden Paaren, die normalerweise lebenslang zusammen bleiben. Nach dem Tod eines verpaarten Wolfes werden Paare schnell wieder hergestellt. Da Männchen in einer bestimmten Wolfspopulation häufig dominieren, sind ungepaarte Weibchen eine Seltenheit. [9] Wenn ein sich ausbreitender grauer Wolf kein Territorium errichten oder einen Partner finden kann, paart er sich mit den Töchtern bereits etablierter Brutpaare aus anderen Rudeln. Solche grauen Wölfe werden "Casanova-Wölfe" genannt und bilden im Gegensatz zu Männern aus etablierten Rudeln keine Paarbeziehungen mit den Weibchen, mit denen sie sich paaren. Einige graue Wolfsrudel können auf diese Weise mehrere Zuchtweibchen haben, wie dies im Yellowstone National Park der Fall ist. [88] Graue Wölfe üben auch eine Alloparental-Pflege aus, in der ein Wolfspaar die Welpen oder Welpen einer anderen aufnehmen kann. Dies kann geschehen, wenn die ursprünglichen Eltern sterben oder aus irgendeinem Grund von ihnen getrennt werden. [89] Neben dem heterosexuellen Verhalten wurde homosexuelles Verhalten bei grauen Wölfen beobachtet. [90] Männliche graue Wölfe reihen sich oft zusammen, wenn die ranghöchste Frau im Rudel in Hitze gerät. [91]
Das Alter der ersten Zucht bei Grauwölfen hängt stark von den Umweltfaktoren ab: Wenn reichlich Futter vorhanden ist oder wenn Wolfspopulationen stark bewirtschaftet werden, können Wölfe Welpen im jüngeren Alter aufziehen, um reichlich vorhandene Ressourcen besser zu nutzen. Dies zeigt sich auch durch die Tatsache, dass bekannt ist, dass in Gefangenschaft gehaltene Wölfe nach Erreichen des Zeitraums von 9–10 Monaten brüten, während die jüngsten registrierten Brutwölfe in freier Wildbahn 2 Jahre alt waren. Weibchen können jedes Jahr Welpen hervorbringen, wobei ein Wurf jährlich der Durchschnitt ist. Im Gegensatz zum Kojoten erreicht der graue Wolf niemals die reproduktive Alterung. [92] Estrus tritt typischerweise im späten Winter auf, wobei ältere, multiparer Weibchen 2–3 Wochen früher als jüngere Frauen in den Östrus gelangen. [9] weibliche Wölfe bleiben in einer Höhle, die von der Randzone ihres Territoriums entfernt liegt, wo gewalttätige Begegnungen mit anderen Rudeln wahrscheinlicher sind. [93] Alte Weibchen jagen normalerweise in der Höhle ihres vorherigen Wurfes, während jüngere Weibchen normalerweise in der Nähe ihres Geburtsortes leben. Die Trächtigkeit dauert 62–75 Tage, wobei die Welpen normalerweise im Sommer geboren werden. [9]
Obwohl soziale Tiere, einzelne Wölfe oder Paarpaare bei der Jagd typischerweise höhere Erfolgsraten aufweisen als große Rudel, gelegentlich wurde beobachtet, dass einzelne Wölfe große Beutetiere wie Elche, Bisons und Moschusochsen ohne Hilfe töten. [97] Der Geruchssinn des grauen Wolfs ist im Vergleich zu einigen Jagdhunderassen relativ schwach entwickelt, da er Aas vor dem Wind nicht weiter als 2–3 Kilometer (1,2–1,9 mi) erkennen kann. Aus diesem Grund gelingt es selten, versteckte Hasen oder Vögel einzufangen, obwohl sie leicht neuen Spuren folgen kann. Its auditory perception is acute enough to be able to hear up to a frequency of 26 kHz,[98] which is sufficient to register the fall of leaves in the autumn period.[9] A gray wolf hunt can be divided into five stages: The actual killing method varies according to prey species. With large prey, mature wolves usually avoid attacking frontally, instead focusing on the rear and sides of the animal. Large prey, such as moose, is killed by biting large chunks of flesh from the soft perineum area, causing massive blood loss. Such bites can cause wounds 10–15 cm (3.9–5.9 in) in length, with three such bites to the perineum usually being sufficient to bring down a large deer in optimum health.[105] 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, mouse-like prey, wolves leap in a high arc and immobilize it with their forepaws.[106] When prey is vulnerable and abundant, wolves may occasionally surplus kill. Such instances are common in domestic animals, but rare in the wild. In the wild, surplus killing primarily occurs during late winter or spring, when snow is unusually deep (thus impeding the movements of prey)[107] or during the denning period, when wolves require a ready supply of meat when denbound.[108] Medium-sized prey are especially vulnerable to surplus killing, as the swift throat-biting method by which they are killed allows wolves to quickly kill one animal and move on to another.[106] 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.[109] The breeding pair typically monopolizes food in order to continue producing pups. When food is scarce, this is done at the expense of other family members, especially non-pups.[110] The breeding pair typically eats first, though as it is they who usually work the hardest in killing prey, they may rest after a long hunt and allow the rest of the family to eat unmolested. Once the breeding pair has finished eating, the rest of the family tears off pieces of the carcass and transport them to secluded areas where they can eat in peace. Wolves typically commence feeding by consuming the larger internal organs of their prey, such as the heart, liver, lungs and stomach lining. The kidneys and spleen are eaten once they are exposed, followed by the muscles.[111] A single wolf can eat 15–19% of its body weight in a single feeding.[112] The gray wolf's expressive behavior is more complex than that of the coyote and golden jackal, as necessitated by its group living and hunting habits. While less gregarious canids generally possess simple repertoires of visual signals, wolves have more varied signals that subtly inter grade in intensity.[10][11] When neutral, the legs are not stiffened, the tail hangs down loosely, the face is smooth, the lips untensed, and the ears point in no particular direction.[113] Postural communication in wolves consists of a variety of facial expressions, tail positions and piloerection.[98] 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, sleeken their fur and lower their ears and tail.[114] When a breeding male encounters a subordinate family member, it may stare at it, standing erect and still with the tails horizontal to its spine.[115] Two forms of submissive behavior are recognized: passive and active. Passive submission usually occurs as a reaction to the approach of a dominant animal, and consists of the submissive wolf lying partly on its back and allowing the dominant wolf to sniff its anogenital area. Active submission occurs often as a form of greeting, and involves the submissive wolf approaching another in a low posture, and licking the other wolf's face.[116] When wolves are together, they commonly indulge in behaviors such as nose pushing, jaw wrestling, cheek rubbing and facial licking. The mouthing of each other's muzzles is a friendly gesture, while clamping on the muzzle with bared teeth is a dominance display.[117] Similar to humans, gray wolves have facial color patterns in which the gaze direction can be easily identified, although this is often not the case in other canid species. In 2014, a study compared the facial color pattern across 25 canid species. The results suggested that the facial color pattern of canid species is related to their gaze communication, and that especially gray wolves use the gaze signal in conspecific communication.[118] Gray 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 or unfamiliar territory and to communicate across great distances.[119] Wolf howls can under certain conditions be heard over areas of up to 130 km2 (50 sq mi).[4] Wolf howls are generally indistinguishable from those of large dogs.[120] Male wolves give voice through an octave, passing to a deep bass with a stress on "O", while females produce a modulated nasal baritone with stress on "U". Pups almost never howl, while yearling wolves produce howls ending in a series of dog-like yelps.[9] Howling consists of a fundamental frequency that may lie between 150 and 780 Hz, and consists of up to 12 harmonically related overtones. The pitch usually remains constant or varies smoothly, and may change direction as many as four or five times.[48] Howls used for calling pack mates to a kill are long, smooth sounds similar to the beginning of the cry of a horned owl. When pursuing prey, they emit a higher pitched howl, vibrating on two notes. When closing in on their prey, they emit a combination of a short bark and a howl.[120] When howling together, wolves harmonize rather than chorus on the same note, thus creating the illusion of there being more wolves than there actually are.[119] Lone wolves typically avoid howling in areas where other packs are present.[121] Wolves from different geographic locations may howl in different fashions: the howls of European wolves are much more protracted and melodious than those of North American wolves, whose howls are louder and have a stronger emphasis on the first syllable. The two are however mutually intelligible, as North American wolves have been recorded to respond to European-style howls made by biologists.[122] Other vocalisations of wolves are usually divided into three categories: growls, barks and whines.[123] Barking has a fundamental frequency between 320–904 Hz,[48] and is usually emitted by startled wolves. Wolves do not bark as loudly or continuously as dogs do, but bark a few times and retreat from perceived danger.[123] Growling has a fundamental frequency of 380–450 Hz,[48] and is usually emitted during food challenges. Pups commonly growl when playing. One variation of the howl is accompanied by a high pitched whine, which precedes a lunging attack.[119] Whining is associated with situations of anxiety, curiosity, inquiry and intimacy such as greeting, feeding pups and playing.[123] Olfaction is probably the wolf's most acute sense, and plays a fundamental role in communication. The wolf has a large number of apocrine sweat glands on the face, lips, back, and between the toes. The odor produced by these glands varies according to the individual wolf's microflora and diet, giving each a distinct "odor fingerprint". A combination of apocrine and eccrine sweat glands on the feet allows the wolf to deposit its scent whilst scratching the ground, which usually occurs after urine marking and defecation during the breeding season. The follicles present on the guard hairs from the wolf's back have clusters of apocrine and sebaceous glands at their bases. As the skin on the back is usually folded, this provides a microclimate for bacterial propagation around the glands. During piloerection, the guard hairs on the back are raised and the skin folds spread, thus releasing scent.[124] The pre-caudal scent glands may play a role in expressing aggression, as combative wolves raise the base of their tails whilst drooping the tip, thus positioning the scent glands at the highest point.[120] The wolf possesses a pair of anal sacs beneath the rectum, which contain both apocrine and sebaceous glands. The components of anal sac secretions vary according to season and gender, thus indicating that the secretions provide information related to gender and reproductive state. The secretions of the preputial glands may advertise hormonal condition or social position, as dominant wolves have been observed to stand over subordinates, apparently presenting the genital area for investigation,[124] which may include genital licking.[125] During the breeding season, female wolves secrete substances from the vagina, which communicate the females' reproductive state, and can be detected by males from long distances. Urine marking is the best-studied means of olfactory communication in wolves. Its exact function is debated, though most researchers agree that its primary purpose is to establish boundaries. Wolves urine mark more frequently and vigorously in unfamiliar areas, or areas of intrusion, where the scent of other wolves or canids is present. So-called raised leg urination (RLU) is more common in male wolves than in females, and may serve the purpose of maximizing the possibility of detection by conspecifics, as well as reflect the height of the marking wolf. Only dominant wolves typically use RLU, with subordinate males continuing to use the juvenile standing posture throughout adulthood.[124] RLU 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.[126] The gray wolf is a habitat generalist, and can occur in deserts, grasslands, forests and arctic tundras. Habitat use by gray wolves is strongly correlated with the abundance of prey, snow conditions, absence or low livestock densities, road densities, human presence and topography.[4] In cold climates, the gray wolf can reduce the flow of blood near its skin to conserve body heat. The warmth of the footpads is regulated independently of the rest of the body, and is maintained at just above tissue-freezing point where the pads come in contact with ice and snow.[127] Gray wolves use different places for their diurnal rest: places with cover are preferred during cold, damp and windy weather, while wolves in dry, calm and warm weather readily rest in the open. During the autumn-spring period, when wolves are more active, they willingly lie out in the open, whatever their location. Actual dens are usually constructed for pups during the summer period. When building dens, females make use of natural shelters such as 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 1–3 openings. The den is usually constructed not more than 500 m (550 yd) away from a water source,[9] and typically faces southwards, thus ensuring enough sunlight exposure, keeping the denning area relatively snow free.[4] Resting places, play areas for the pups and food remains are commonly found around wolf dens. The odour of urine and rotting food emanating from the denning area often attracts scavenging birds such as magpies and ravens. As there are few convenient places for burrows, wolf dens are usually occupied by animals of the same family. Though they mostly avoid areas within human sight, wolves have been known to nest near domiciles, paved roads and railways.[9] Globally, gray wolf diet is predominantly composed of large (240–650 kg (530–1,430 lb)) and medium-sized (23–130 kg (51–287 lb)) wild ungulates, with local population variations due to the mix of wild ungulates, smaller prey and domestic species consumed.[128] All terrestrial mammalian social predators feed predominantly on terrestrial herbivorous mammals that have a body mass similar to that of the combined mass of the social group members.[129][130] The gray wolf generally specializes in preying on the vulnerable individuals of large prey,[4] with pack of timber wolves capable of bringing down a 500 kg (1,100 lb) moose.[53][131]Digestion only takes a few hours, thus wolves can feed several times in one day, making quick use of large quantities of meat.[132] Although wolves primarily feed on medium to large sized ungulates, they are not fussy eaters. Smaller sized animals that may supplement the diet of wolves include marmots, hares, badgers, foxes, weasels, ground squirrels, mice, hamsters, voles and other rodents, as well as insectivores. They frequently eat waterfowl and their eggs. When such foods are insufficient, they prey on lizards, snakes, frogs, rarely toads and large insects as available. In times of scarcity, wolves readily eat carrion, visiting cattle burial grounds and slaughter houses.[9]Cannibalism is not uncommon in wolves: during harsh winters, packs often attack weak or injured wolves, and may eat the bodies of dead pack members.[9][133][134] Wolf packs in Astrakhan hunt Caspian seals on the Caspian Sea coastline[135] and some wolf packs in Alaska and Western Canada have been observed to feed on salmon.[136] Wolves in northern Minnesota have also been observed hunting northern pike in freshwater streams.[137]Humans are rarely, but occasionally preyed upon.[9][24] Other primates occasionally taken by wolves include grey langurs in Nepal[138] and hamadryas baboons in Saudi Arabia.[139] In Eurasia, many gray wolf populations are forced to subsist largely on livestock and garbage in areas with dense human activity, though wild ungulates such as moose, red deer, roe deer and wild boar are still the most important food sources in Russia and the more mountainous regions of Eastern Europe. Other prey species include reindeer, argali, mouflon, wisent, saiga, ibex, chamois, wild goats, fallow deer and musk deer.[140] The prey animals of North American wolves have largely continued to occupy suitable habitats with low human density, and cases of wolves subsisting largely on garbage or livestock are exceptional. Animals preferred as prey by North American wolves include moose, white-tailed deer, elk, mule deer, bighorn sheep, Dall's sheep, American bison, muskox and caribou.[141] Gray wolves typically dominate other canid species in areas where they both occur. In North America, incidents of gray 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.[142][143] Near identical interactions have been observed in Eurasia between gray wolves and golden jackals, with the latter's numbers being comparatively small in areas with high wolf densities.[9][142][144] Gray wolves are the most important predator of raccoon dogs, killing large numbers of them in the spring and summer periods.[9] Wolves also kill red, arctic and corsac foxes, usually in disputes over carcasses, sometimes eating them.[9][145] In Asia, they may compete with dholes,[9] though there is at least one record of a lone wolf associating with a pair of dholes in Debrigarh Wildlife Sanctuary.[146] 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 only eat young wolves.[147] Wolf interactions with American black bears are much rarer than with brown bears, because of differences in habitat preferences. The majority of black bear encounters with wolves occur in the species' northern range, with no interactions being recorded in Mexico. Wolves have been recorded on numerous occasions to actively seek out black bears in their dens and kill them without eating them. Unlike brown bears, black bears frequently lose against wolves in disputes over kills.[148] While encounters with brown and black bears appear to be common, polar bears are rarely encountered by wolves, though there are two records of wolf packs killing polar bear cubs.[149] Wolves also kill the cubs of Asian black bears.[9] Wolves may encounter striped hyenas in Israel, Central Asia and India, 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,[150] but wolf packs can drive off single or outnumbered hyenas.[151][152] However, there was a case of a female striped hyena dominating 12 Arabian wolves.[153] Two cases are known however from southern Israel, where wolves and striped hyenas associated closely with each other in an apparently amicable way.[154] Large wolf populations limit the numbers of small to medium-sized felines. Wolves encounter cougars along portions of the Rocky Mountains and adjacent mountain ranges. Wolves and cougars typically avoid encountering each other by hunting on different elevations. In winter, however, 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 kills. They have been reported killing mothers and their kittens.[155][156] Wolves hunt steppe cats, and may scavenge from snow leopard kills.[157][158] Wolves may also reduce Eurasian lynx populations. Wolves may kill lynxes by running them down, or killing them before they can escape into trees.[159] Similar reports of encounters between wolves and bobcats have been documented.[160] Leftovers of wolf kills are sometimes scavenged by wolverines. Wolverines usually wait until the wolves are done feeding, but have been known to drive away wolves from kills. However, there have been confirmed reports of wolf packs killing wolverines.[161] Other than humans, tigers appear to be the only serious predators of wolves.[9][17][18][19][157] Wolf and tiger interactions are well documented in Sikhote-Alin, where tigers depress wolf numbers, either to the point of localized extinction or to such low numbers as to make them a functionally insignificant component of the ecosystem. Wolves appear capable of escaping competitive exclusion from tigers only when human persecution decreases tiger numbers. Proven cases of tigers killing wolves are rare and attacks appear to be competitive rather than predatory in nature, with at least four proven records of tigers killing wolves without consuming them.[162] The gray wolf was once one of the world's most widely distributed mammals, living throughout the northern hemisphere north of 15°N latitude in North America and 12°N in India. However, deliberate human persecution has reduced the species' range to about one third, because of livestock predation and fear of attacks on humans. The species is now extinct in much of Western Europe, in Mexico and much of the United States. In modern times, the gray wolf occurs mostly in wilderness and remote areas, particularly in Canada, Alaska and northern USA, Europe, and Asia from about 75°N to 12°N. Wolf population declines have been arrested since the 1970s, and have fostered recolonization and reintroduction in parts of its former range, due to 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 species. Despite these threats, because of the gray wolf's relatively widespread range and stable population, it is classified as Least Concern on the IUCN Red List.[2] In Europe, the oldest gray wolf remains were found in France and date to 400,000–350,000 years ago.[163] Wolf populations strongly declined across Europe during the 18th and 19th centuries largely due to human persecution, and by the end of the Second World War they had been eradicated from all of Central Europe and almost all of Northern Europe.[164] The extermination of Northern Europe's wolves first became an organized effort during the Middle Ages, and continued until the late 19th century. In England, wolf eradication was enforced by legislation, and the last wolf was killed in the early sixteenth century during the reign of Henry VII. Wolves lasted longer in Scotland, where they sheltered in vast tracts of forest, which were subsequently burned down. Wolves managed to survive in the forests of Braemar and Sutherland until 1684. The extirpation of wolves in Ireland followed a similar course, with the last wolf believed to have been killed in 1786.[165] A wolf bounty was introduced in Sweden in 1647, after the extermination of moose and reindeer forced wolves to feed on livestock. The Sami extirpated wolves in northern Sweden in organized drives. By 1960, few wolves remained in Sweden, because of the use of snowmobiles in hunting them, with the last specimen being killed in 1966. The gray wolf was exterminated in Denmark in 1772 and Norway's last wolf was killed in 1973. The species was decimated in 20th century Finland, despite regular dispersals from Russia. The gray wolf was only present in the eastern and northern parts of Finland by 1900, though its numbers increased after World War II.[166] In Central Europe, wolves were dramatically reduced in number during the early nineteenth century, because of organized hunts and reductions in ungulate populations. In Bavaria, the last wolf was killed in 1847, and had disappeared from the Rhine regions by 1899.[166] In Switzerland, wolves were extinct in the twentieth century; they are naturally coming back from Italy since the 1990s.[167] In 1934, Nazi Germany became the first state in modern history to place the wolf under protection, though the species was already extirpated in Germany at this point.[168] The last free-living wolf to be killed on the soil of present-day Germany before 1945 was the so-called "Tiger of Sabrodt", which was shot near Hoyerswerda, Lusatia (then Lower Silesia) in 1904. Today, wolves have returned to the area.[169] Wolf hunting in France was first institutionalized by Charlemagne between 800–813, when he established the louveteriea special corps of wolf hunters. The louveterie was abolished after the French Revolution in 1789, but was re-established in 1814. In 1883, up to 1,386 wolves were killed, with many more by poison.[166] In Eastern Europe, wolves were never fully exterminated, because of the area's contiguity with Asia and its large forested areas. However, Eastern European wolf populations were reduced to very low numbers by the late nineteenth century. Wolves were extirpated in Slovakia during the first decade of the twentieth century and, by the mid-twentieth century, could only be found in a few forested areas in eastern Poland. Wolves in the eastern Balkans benefitted from the region's contiguity with the former Soviet Union and large areas of plains, mountains and farmlands. Wolves in Hungary occurred in only half the country around the start of the 20th century, and were largely restricted to the Carpathian Basin. Wolf populations in Romania remained largely substantial, with an average of 2,800 wolves being killed annually out of a population of 4,600 from 1955–1965. An all-time low was reached in 1967, when the population was reduced to 1,550 animals. The extermination of wolves in Bulgaria was relatively recent, as a previous population of about 1,000 animals in 1955 was reduced to about 100–200 in 1964. In Greece, the species disappeared from the southern Peloponnese in 1930. Despite periods of intense hunting during the eighteenth century, wolves never disappeared in the western Balkans, from Albania to the former Yugoslavia. Organized persecution of wolves began in Yugoslavia in 1923, with the setting up of the Wolf Extermination Committee (WEC) in Kocevje, Slovenia. The WEC was successful in reducing wolf numbers in the Dinaric Alps.[166] In Southern Europe, wolf extermination was not as complete as in Northern Europe, because of greater cultural tolerance of the species. Wolf populations only began declining in the Iberian Peninsula in the early 19th-century, and was reduced by a half of its original size by 1900. Wolf bounties were regularly paid in Italy as late as 1950. Wolves were exterminated in the Alps by 1800, and numbered only 100 by 1973, inhabiting only 3–5% of their former Italian range.[166] The recovery of European wolf populations began after the 1950s, when traditional pastoral and rural economies declined and thus removed the need to heavily persecute wolves. By the 1980s, small and isolated wolf populations expanded in the wake of decreased human density in rural areas and the recovery of wild prey populations.[170] The gray wolf has been fully protected in Italy since 1976, and now holds a population of over 1,269–1,800.[171] Italian wolves entered France's Mercantour National Park in 1993, and at least fifty wolves were discovered in the western Alps in 2000. By 2013 the 250 wolves in the Western Alps imposed a significant burden on traditional sheep and goat husbandry with a loss of over 5,000 animals in 2012.[172] There are approximately 2,000 wolves inhabiting the Iberian Peninsula, of which 150 reside in northeastern Portugal. In Spain, the species occurs in Galicia, Leon, and Asturias. Although hundreds of Iberian wolves are illegally killed annually, the population has expanded south across the river Duero and east to the Asturias and Pyrenees Mountains.[170] In 1978, wolves began recolonising central Sweden after a twelve-year absence, and have since expanded into southern Norway. As of 2005, the total number of Swedish and Norwegian wolves is estimated to be at least one hundred, including eleven breeding pairs. The gray wolf is fully protected in Sweden and partially controlled in Norway. The Scandinavian wolf populations owe their continued existence to neighbouring Finland's contiguity with the Republic of Karelia, which houses a large population of wolves. Wolves in Finland are protected only in the southern third of the country, and can be hunted in other areas during specific seasons,[170] though poaching remains common, with 90% of young wolf deaths being due to human predation, and the number of wolves killed exceeds the number of hunting licenses, in some areas by a factor of two. Furthermore, the decline in the moose populations has reduced the wolf's food supply.[173][174] Since 2011, the Netherlands, Belgium and Denmark have also reported wolf sightings presumably by natural migration from adjacent countries.[175][176] In 2016, a female wolf tracked 550 kilometers from a region southwest of Berlin to settle in Jutland, Denmark where male wolves had been reported in 2012 for the first time in 200 years.[177] Wolves have also commenced breeding in Lower Austria's Waldviertel region for the first time in over 130 years.[178] Wolf populations in Poland have increased to about 800–900 individuals since being classified as a game species in 1976. Poland plays a fundamental role in providing routes of expansion into neighbouring Central European countries. In the east, its range overlaps with populations in Lithuania, Belarus, Ukraine, and Slovakia. A population in western Poland expanded into eastern Germany and in 2000 the first pups were born on German territory.[179] In 2012, an estimated 14 wolf packs were living in Germany (mostly in the east and north) and a pack with pups has been sighted within 15 miles of Berlin;[180] the number increased to 46 packs in 2016.[181] The gray wolf is protected in Slovakia, though an exception is made for wolves killing livestock. A few Slovakian wolves disperse into the Czech Republic, where they are afforded full protection. Wolves in Slovakia, Ukraine and Croatia may disperse into Hungary, where the lack of cover hinders the buildup of an autonomous population. Although wolves have special status in Hungary, they may be hunted with a year-round permit if they cause problems.[170] Romania has a large population of wolves, numbering 2,500 animals. The wolf has been a protected animal in Romania since 1996, although the law is not enforced. The number of wolves in Albania and Macedonia is largely unknown, despite the importance the two countries have in linking wolf populations from Greece to those of Bosnia and Herzegovina and Croatia. Although protected, sometimes wolves are still illegally killed in Greece, and their future is uncertain. Wolf numbers have declined in Bosnia and Herzegovina since 1986, while the species is fully protected in neighbouring Croatia and Slovenia.[170] Although wolf-dog hybridization in Europe has raised concern among conservation groups fearing for the gray wolf's purity, genetic tests show that introgression of dog genes into European gray wolf populations does not pose a significant threat. Also, as wolf and dog mating seasons do not fully coincide, the likelihood of wild wolves and dogs mating and producing surviving offspring is small.[182] During the 19th century, gray wolves were widespread in many parts of the Holy Land east and west of the Jordan River. However, they decreased considerably in number between 1964 and 1980, largely because of persecution by farmers.[183] The species was not considered common in northern and central Saudi Arabia during the 19th century, with most early publications involving animals either from southwestern Asir, northern rocky areas bordering Jordan, or areas surrounding Riyadh.[184] The gray wolf's range in the Soviet Union encompassed nearly the entire territory of the country, being absent only on the Solovetsky Islands, Franz-Josef Land, Severnaya Zemlya, and the Karagin, Commander and Shantar Islands. The species was exterminated twice in Crimea, once after the Russian Civil War, and again after World War II.[9] Following the two world wars, Soviet wolf populations peaked twice. 30,000 wolves were harvested annually out of a population of 200,000 during the 1940s, with 40,000–50,000 harvested during peak years. Soviet wolf populations reached a low around 1970, disappearing over much of European Russia. The population increased again by 1980 to about 75,000, with 32,000 being killed in 1979.[185] Wolf populations in northern inner Mongolia declined during the 1940s, primarily because of poaching of gazelles, the wolf's main prey.[186] In British-ruled India, wolves were heavily persecuted because of their attacks on sheep, goats and children. In 1876, 2,825 wolves were bountied in the North-Western Provinces (NWP) and Bihar. By the 1920s, wolf extermination remained a priority in the NWP and Awadh. Overall, over 100,000 wolves were killed for bounties in British India between 1871 and 1916.[187] Wolves in Japan became extinct during the Meiji restoration period, an extermination known as ōkami no kujo. The wolf was deemed a threat to ranching, which the Meiji government promoted at the time, and targeted via a bounty system and a direct chemical extermination campaign inspired by the similar contemporary American campaign. The last Japanese wolf was a male killed on January 23, 1905 near Washikaguchi (now called Higashi Yoshiro).[188] The now extinct Japanese wolves were descended from large Siberian wolves, which colonized the Korean Peninsula and Japan, before it separated from mainland Asia, 20,000 years ago during the Pleistocene. During the Holocene, the Tsugaru Strait widened and isolated Honshu from Hokkaidō, thus causing climatic changes leading to the extinction of most large bodied ungulates inhabiting the archipelago. Japanese wolves likely underwent a process of island dwarfism 7,000–13,000 years ago in response to these climatological and ecological pressures. C. l. hattai (formerly native to Hokkaidō) was significantly larger than its southern cousin C. l. hodophilaxas it inhabited higher elevations and had access to larger prey, as well as a continuing genetic interaction with dispersing wolves from Siberia.[189] There is little reliable data on the status of wolves in the Middle East, save for those in Israel and Saudi Arabia, though their numbers appear to be stable, and are likely to remain so. Israel's conservation policies and effective law enforcement maintain a moderately sized wolf population, which radiates into neighbouring countries, while Saudi Arabia has vast tracts of desert, where about 300–600 wolves live undisturbed.[191] The wolf survives throughout most of its historical range in Saudi Arabia, probably because of a lack of pastoralism and abundant human waste.[184] Turkey may play an important role in maintaining wolves in the region, because of its contiguity with Central Asia. The mountains of Turkey have served as a refuge for the few wolves remaining in Syria. A small wolf population occurs in the Golan Heights, and is well protected by the military activities there. Wolves living in the southern Negev desert are contiguous with populations living in the Egyptian Sinai and Jordan. Throughout the Middle East, the species is only protected in Israel. Elsewhere, it can be hunted year-round by Bedouins.[191] Little is known of current wolf populations in Iran, which once occurred throughout the country in low densities during the mid-1970s. The northern regions of Afghanistan and Pakistan are important strongholds for the wolf. It has been estimated that there are about 300 wolves in approximately 60,000 km2 (23,000 sq mi) of Jammu and Kashmir in northern India, and 50 more in Himachal Pradesh. Overall, India supports about 800-3,000 wolves, scattered among several remnant populations. Although protected since 1972, Indian wolves are classed as endangered, with many populations lingering in low numbers or living in areas increasingly used by humans. Although present in Nepal and Bhutan, there is no information of wolves occurring there.[185] Wolf populations throughout Northern and Central Asia are largely unknown, but are estimated in the hundreds of thousands based on annual harvests. Since the fall of the Soviet Union, continent-wide extermination of wolves has ceased, and wolf populations have increased to about 25,000–30,000 animals throughout the former Soviet Union. In China and Mongolia, wolves are only protected in reserves. Mongolian populations have been estimated at 10,000–30,000, while the status of wolves in China is more fragmentary. The north has a declining population of an estimated 400 wolves, while Xinjiang and Tibet hold about 10,000 and 2,000 respectively.[192] In 2008, an authoritative reference stated that the gray wolf could be found across mainland China.[193] In 2017, a comprehensive study found that the gray wolf was present across all of mainland China, both in the past and today. It exists in southern China, which refutes claims made by some researchers in the Western world that the wolf had never existed in southern China.[194][195] Originally, the gray wolf occupied all of North America north of about 20°N. It occurred all over the mainland, save for the southeastern United States, California west of the Sierra Nevada, and the tropical and subtropical areas of Mexico. Large continental islands occupied by wolves included Newfoundland, Vancouver Island, southeastern Alaskan islands, and throughout the Arctic Archipelago and Greenland.[4] While Lohr and Ballard postulated that the gray wolf had never been present on Prince Edward Island,[196][197]:392 analysis of references to the island's native fauna in unpublished and published historical records has found that gray wolves were resident there at the time of the first French settlement in 1720. In his November 6, 1721 letter to the French Minister of the Marine, Louis Denys de La Ronde reported that the island was home to wolves "of a prodigious size", and sent a wolf pelt back to France to substantiate his claim. As the island was cleared for settlement, the gray wolf population may have been extirpated, or relocated to the mainland across the winter ice: the few subsequent wolf reports date from the mid-nineteenth century and describe the creatures as transient visitors from across the Northumberland Strait.[197]:386 The decline of North American wolf populations coincided with increasing human populations and the expansion of agriculture. By the start of the 20th century, the species had almost disappeared from the eastern USA, excepting some areas of the Appalachians and the northwestern Great Lakes Region. In Canada, the gray wolf was extirpated in New Brunswick and Nova Scotia between 1870 and 1921, and in Newfoundland around 1911. It vanished from the southern regions of Quebec and Ontario between 1850 and 1900. The gray wolf's decline in the prairies began with the extermination of the American bison and other ungulates in the 1860s–70s. From 1900–1930, the gray wolf was virtually eliminated from the western USA and adjoining parts of Canada, because of intensive predator control programs aimed at eradicating the species. The gray wolf was exterminated by federal and state governments from all of the USA by 1960, except in Alaska and northern Minnesota. The decline in North American wolf populations was reversed from the 1930s to the early 1950s, particularly in southwestern Canada, because of expanding ungulate populations resulting from improved regulation of big game hunting. This increase triggered a resumption of wolf control in western and northern Canada. Thousands of wolves were killed from the early 1950s to the early 1960s, mostly by poisoning. This campaign was halted and wolf populations increased again by the mid-1970s.[4] The species' modern range in North America is mostly confined to Alaska and Canada, with populations also occurring in northern Minnesota, northern and central Wisconsin, Michigan's Upper Peninsula, and small portions of Washington, Idaho, northern Oregon, and Montana. A functional wolf population should exist in California by 2024 according to estimates by state wildlife officials.[198] Canadian wolves began to naturally re-colonize northern Montana around Glacier National Park in 1979, and the first wolf den in the western U.S. in over half a century was documented there in 1986.[199] The wolf population in northwest Montana initially grew as a result of natural reproduction and dispersal to about 48 wolves by the end of 1994.[200] From 1995–1996, wolves from Alberta and British Columbia were relocated to Yellowstone National Park and Idaho. In addition, the Mexican wolf (Canis lupus baileyi) was reintroduced to Arizona and New Mexico in 1998. The gray wolf is found in approximately 80% of its historical range in Canada, thus making it an important stronghold for the species.[4] Canada is home to about 52,000–60,000 wolves, whose legal status varies according to province and territory. First Nations residents may hunt wolves without restriction, and some provinces require licenses for residents to hunt wolves while others do not. In Alberta, wolves on private land may be baited and hunted by the landowner without requiring a license, and in some areas, wolf hunting bounty programs exist.[201][202] Large-scale wolf population control through poisoning, trapping and aerial hunting is also presently conducted by government-mandated programs in order to support populations of endangered prey species such as woodland caribou.[203] In Alaska, the gray wolf population is estimated at 6,000–7,000, and can be legally harvested during hunting and trapping seasons, with bag limits and other restrictions. As of 2002, there are 250 wolves in 28 packs in Yellowstone, and 260 wolves in 25 packs in Idaho. The gray wolf received Endangered Species Act (ESA) protection in Minnesota, Wisconsin, and Michigan in 1974, and was re-classed from endangered to threatened in 2003. Reintroduced Mexican wolves in Arizona and New Mexico are protected under the ESA and, as of late 2002, number 28 individuals in eight packs.[204] A female wolf shot in 2013 in Hart County, Kentucky by a hunter was the first gray wolf seen in Kentucky in modern times. DNA analysis by Fish and Wildlife laboratories showed genetic characteristics similar to those of wolves in the Great Lakes Region.[205] Viral diseases carried by wolves include rabies, canine distemper, canine parvovirus, infectious canine hepatitis, papillomatosis, canine coronavirus,[206] and foot and mouth disease.[207] Wolves are a major host for rabies in Russia, Iran, Afghanistan, Iraq and India.[207] In wolves, the incubation period is 8–21 days, and results in the host becoming agitated, deserting its pack, and travelling up to 80 kilometres (50 mi) a day, thus increasing the risk of infecting other wolves. Infected wolves do not show any fear of humans, with 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 doesn't cause serious illness or death, though it may alter feeding behaviors. The canine coronavirus has been recorded in Alaskan wolves, with infections being most prevalent in winter months.[206] Bacterial diseases carried by wolves include brucellosis, lyme disease, leptospirosis, tularemia, bovine tuberculosis,[208]listeriosis and anthrax.[207] 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 have any significant effect on 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 once killed two wolf pups in Canada.[208] Wolves carry ectoparasites and endoparasites, with wolves in the former Soviet Union having been recorded to carry at least 50 species.[207] Most of these parasites infect wolves without adverse effects, though the effects may become more serious in sick or malnourished specimens.[209]Parasitic infection in wolves is of particular concern to people, as 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.[207] Wolves are often infested with a variety of arthropod exoparasites, including fleas, ticks, lice, and mites. The most harmful to wolves, particularly pups, is Sarcoptes scabiei (or mange mite),[209] though they rarely develop full blown mange, unlike foxes.[9] Lice, such as Trichodectes canismay cause sickness in wolves, but rarely death. Ticks of the genus Ixodes can infect wolves with Lyme disease and Rocky Mountain spotted fever.[209] The tick Dermacentor pictus also infests wolves. Other ectoparasites include biting lice, sucking lice and the fleas Pulex irritans and Ctenocephalides canis.[9] 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: IsosporaToxoplasmaSarcocystisBabesiaand Giardia.[209] Wolves may carry Neospora caninumwhich is of particular concern to farmers, as the disease can be spread to livestock, with infected animals being 3–13 times more likely to miscarry than those not infected.[210][211] Among flukes, the most common in North American wolves is Alariawhich infects small rodents and amphibians that are eaten by wolves. Upon reaching maturity, Alaria migrates to the wolf's intestine, but harms it little. Metorchis conjunctuswhich 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. 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 canisa hookworm known to infect wolf pups in utero, 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. 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 isn't known to produce clinical signs in wolves, it can cause emaciation, salivation, and crippling muscle pains in dogs.[209]Thorny-headed worms rarely infect wolves, though three species have been identified in Russian wolves: Nicolla skrjabiniMacrocantorhynchus catulinusand Moniliformis moniliformis.[209] Human presence appears to stress wolves, as seen by increased cortisol levels in instances such as snowmobiling near their territory.[212] Old English literature contains several instances of Anglo-Saxon kings and warriors taking on wulf as a prefix or suffix in their names. Examples include Wulfhere, Cynewulf, Ceonwulf, Wulfheard, Earnwulf, Wulfmǣr, Wulfstān and Æthelwulf. Wolf-related names were also common among pre-Christian Germanic warriors: Wolfhroc (Wolf-Frock), Wolfhetan (Wolf Hide), Isangrim (Grey Mask), Scrutolf (Garb Wolf), Wolfgang (Wolf Gait) and Wolfdregil (Wolf Runner).[213] Ancient Greek literature is similar: Autolycus ("the wolf itself"), Lycurgus ("wolf-work")[citation needed]. The Latin for "female prostitute" is lupaand the most famous brothel in Pompeii was the Lupanar.[citation needed] The wolf is a common motif in the foundational mythologies and cosmologies of peoples throughout Eurasia and North America (corresponding to the historical extent of the habitat of the gray wolf). The obvious attribute of the wolf is its nature of a predator, and correspondingly it is strongly associated with danger and destruction, making it the symbol of the warrior on one hand, and that of the devil on the other. The modern trope of the Big Bad Wolf is a development of this. The wolf holds great importance in the cultures and religions of the nomadic peoples, both of the Eurasian steppe and of the North American Plains. In many cultures, the identification of the warrior with the wolf (totemism) gave rise to the notion of Lycanthropy, the mythical or ritual identification of man and wolf. 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 behavior, his portrayals added to the wolf's image as a deceitful and dangerous animal.[214] Isengrim the wolf, a character first appearing in the 12th-century Latin poem Ysengrimusis a major character in the Reynard Cycle, where he stands for the low nobility, whilst his adversary, Reynard the fox, represents the peasant hero. Although portrayed as loyal, honest and moral, Isengrim is forever the victim of Reynard's wit and cruelty, often dying at the end of each story.[215] The tale of Little Red Riding Hood, first written in 1697 by Charles Perrault, is largely considered to have had more influence than any other source of literature in forging the wolf's negative reputation in the western world. The wolf in this story is portrayed as a potential rapist, capable of imitating human speech.[216] The hunting of wolves, and their attacks on humans and livestock feature prominently in Russian literature, and are included in the works of Tolstoy, Chekhov, Nekrasov, Bunin, Sabaneyev, and others. Tolstoy's War and Peace and Chekhov's Peasants both feature scenes in which wolves are hunted with hounds and borzois.[217]Farley Mowat's largely fictional 1963 memoir Never Cry Wolf was the first positive portrayal of wolves in popular literature, and is largely 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.[218][219][220] The wolf is a frequent charge in English armory. 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.[221] The wolf is featured on the flags of the Confederated Tribes of the Colville Reservation, the Oneida Nation of Wisconsin and the Pawnee.[222] 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 Grey Wolves. During the Yugoslav Wars, several Serb paramilitary units adopted the wolf as their symbol, including the White Wolves and the Wolves of Vučjak.[223] 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.[224] Some nations help offset economic losses to wolves through compensation programmes or state insurance.[225] Domesticated animals are easy prey for wolves, as they have evolved under constant human protection, and are thus unable to defend themselves very well.[226] Wolves typically resort to attacking livestock when wild prey is depleted: 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.[224] The majority of losses occur during the summer grazing period, with untended livestock in remote pastures being the most vulnerable to wolf predation.[227] The most frequently targeted livestock species are sheep (Europe), domestic reindeer (northern Scandinavia), goats (India), horses (Mongolia), cattle and turkeys (North America).[224] 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.[228] Wolves mainly attack livestock when the animals are grazing, though they occasionally break into fenced enclosures.[105] In some cases, wolves do not need to physically attack livestock to negatively affect it: the stress livestock experiences in being vigilant for wolves may result in miscarriages, weight loss and a decrease in meat quality.[210] Being the most abundant carnivores, free-ranging dogs have the greatest potential to compete with wolves. A review of the studies in the competitive effects of dogs on sympatric carnivores did not mention any research on competition between dogs and wolves.[229][230] Competition would favor the wolf, which is known to kill dogs, however wolves tend to live in pairs or in small packs in areas where they are highly persecuted, giving them a disadvantage facing large groups of dogs.[230][231] Wolves kill dogs on occasion, with some wolf populations relying on dogs as an important food source.[224] In Croatia, wolves kill more dogs than sheep, and wolves in Russia appear to limit stray dog populations. Wolves may display unusually bold behavior when attacking dogs accompanied by people, sometimes ignoring nearby humans.[224] Wolf attacks on dogs may occur both in house yards and in forests.[232] Wolf attacks on hunting dogs are considered a major problem in Scandinavia and Wisconsin.[224][232] The most frequently killed hunting breeds in Scandinavia are harriers, with 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.[232] Although the numbers of dogs killed each year are relatively low, it induces a fear of wolves entering villages and farmyards to take dogs. In many cultures, there are strong social and emotional bonds between humans and their dogs that can be seen as family members or working team members. The loss of a dog can lead to strong emotional responses with demands for more liberal wolf hunting regulations.[230] 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.[230][233] Shepherd dogs are not particularly aggressive, but they can disrupt potential wolf predation by displaying what is to the wolf ambiguous behaviors, such as barking, social greeting, invitation to play or aggression.[230][234] The historical use of shepherd dogs across Eurasia has been effective against wolf predation,[230][234] especially when confining sheep in the presence of several livestock-guarding dogs.[230][235] However, shepherd dogs are sometimes killed by wolves.[230][236] The fear of wolves has been pervasive in many societies, though humans are not part of the wolf's natural prey.[24] 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.[237] Although wolves may react aggressively under provocation, such attacks are mostly limited to quick bites on extremities, and the attacks are not pressed. Predatory attacks (attacks by wolves treating humans as food) 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,[24] though cases of non-rabid wolf attacks in winter have been recorded in Belarus, Kirov and Irkutsk oblasts, Karelia and Ukraine.[9] Also, wolves with pups experience greater food stresses during this period.[9] 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. 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, Middle East and Central Asia. Wolves apparently develop the "furious" phase of rabies to a very high degree which, coupled with their size and strength, makes rabid wolves perhaps the most dangerous of rabid animals,[24] with bites from rabid wolves being 15 times more dangerous than those of rabid dogs.[9] 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 only occur on a single day. The victims are chosen at random, though the majority of cases involve adult men. During the 50 years to 2002, there were eight fatal attacks in Europe and Russia, and more than 200 in south Asia.[24] Between 2005–2010, two people were killed in North America.[238][239] Wolves are difficult to hunt because of their elusiveness, sharp senses, high endurance, and ability to quickly incapacitate and kill hunting dogs.[240] Historic methods include killing of spring-born litters in their dens, coursing with dogs (usually combinations of sighthounds, bloodhounds and fox terriers), poisoning with strychnine, and trapping.[241][242][243] 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 smell.[243] Some hunters are able to lure wolves by imitating their calls.[243] In Kazakhstan and Mongolia, wolves are traditionally hunted with eagles and falcons, though this practise is declining, as experienced falconers are becoming few in number.[243] Shooting wolves from aircraft is highly effective, due to increased visibility and direct lines of fire,[243] but is controversial.[244] Several types of dog, including like the Borzoi, Irish wolfhound, and Kyrgyz Tajgan, have been specifically bred for wolf hunting.[230] Wild wolves are sometimes kept as exotic pets and, in some rarer occasions, as working animals. Although closely related to domesticated dogs, wolves do not show the same tractability as dogs in living alongside humans, and generally, much more work is required in order to obtain the same amount of reliability. Wolves also need much more space than dogs, about 26–39 square kilometres (10–15 sq mi), so they can exercise.[245]
Wölfe tragen im Vergleich zu anderen Canidenarten relativ große Welpen in kleinen Würfen. [94] Der durchschnittliche Wurf besteht aus 5–6 Jungen, [95] wobei die Wurfgrößen in Gebieten, in denen die Beute reichlich vorhanden ist, zunehmen, [95] treten nur 1% der Zeit auf. [96] Welpen werden in der Regel im Frühjahr geboren, was mit einer entsprechenden Zunahme der Beutepopulationen einhergeht. [93] Welpen sind blind und taub geboren und mit einem kurzen, weichen, graubraunen Fell bedeckt. Sie wiegen bei der Geburt 300 bis 500 g und beginnen nach 9 bis 12 Tagen zu sehen. Die Milchzähne brechen nach einem Monat aus. Welpen verlassen die Höhle zuerst nach 3 Wochen. Im Alter von 1,5 Monaten sind sie beweglich genug, um vor Gefahren zu fliehen. Mutterwölfe verlassen die Höhle in den ersten Wochen nicht und verlassen sich darauf, dass die Väter ihnen und ihren Jungen Nahrung geben. Welpen beginnen im Alter von 3-4 Wochen feste Nahrung zu sich zu nehmen. Welpen haben in den ersten vier Lebensmonaten eine schnelle Wachstumsrate: In dieser Zeit kann sich das Gewicht eines Welpen fast um das 30-fache erhöhen. [9] [94] Wolfswelpen beginnen im Alter von 3 Wochen mit dem Kämpfen, obwohl ihre Bisse im Gegensatz zu jungen Füchsen und Kojoten gehemmt sind. Tatsächliche Kämpfe um die Hierarchie werden normalerweise im Alter von 5 bis 8 Wochen durchgeführt. Dies steht im Gegensatz zu jungen Füchsen und Kojoten, die möglicherweise schon vor dem Beginn des Spielverhaltens zu kämpfen beginnen. [10] Bis zum Herbst sind die Welpen reif genug, um Erwachsene bei der Jagd auf große Beute zu begleiten. [93] [19469166] ] Jagd- und Fütterungsverhalten 
Communication
Visual
used during social interactions in wolves[114]Feature Aggressive Fearful Eyes Direct stare
Open wideLooking away
Closed to slitsEars Erect and forward Flattened and turned down to side Lips Horizontal contraction
("agonistic pucker")Horizontal retraction ("submissive grin") Mouth Opened Closed Teeth Canines bared Canines covered Tongue Retracted Extended ("lick intention") Nose Shortened (skin folded) Lengthened (skin smoothed) Forehead Contracted (bulging over eyes) Stretched (smoothed) Head Held high Lowered Neck Arched Extended Hair Erect (bristled) Sleeked Body Erect, tall Crouched, low Tail Held high
QuiveringTucked under body
WaggingAuditory
Olfactory
Ecology
Habitat
Diet
Wolves supplement their diet with fruit and vegetable matter. They willingly eat the berries of mountain ash, lily of the valley, bilberries, blueberries and cowberry. Other fruits include nightshade, apples and pears. They readily visit melon fields during the summer months.[9] A well-fed wolf stores fat under the skin, around the heart, intestines, kidneys, and bone marrow, particularly during the autumn and winter.[112]Enemies and competitors
Range and conservation
Europe
Decline
Recovery
Asia
Historical range and decline
Modern range
North America
Historical range and decline
Modern range
Diseases and parasites
Viral and bacterial infections
Parasitic infections
Relationships with humans
In culture
In personal names
In folklore, religion and mythology
In fable and literature
In heraldry and symbolism
Conflicts
Wolf predation on livestock
Conflicts with dogs
Wolf predation on humans
Human predation on wolves
As pets and working animals
See also
Notes
References
sheep and goat losses doubled in the past five years to nearly 6,000 in 2012
Bibliography
Further reading
External links
Wikiquote has quotations related to: Wolf
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