View source for Italian wolf

{{Short description|Subspecies of carnivore}}
{{Subspeciesbox
| image = C. l. italicus in MNP.jpg
| image_caption = 
| status = VU
| status_system = IUCN3.1
| status_ref = {{sfn|Boitani|Phillips|Jhala|2023}}
| genus = Canis
| species = lupus
| species_link = Wolf
| subspecies = italicus
| authority = Altobello, 1921{{sfn|Altobello|1921}}
| range_map = Canis_lupus_italicus_areale_2019.gif
| range_map_caption = Italian wolf range: red: stable presence; yellow: irregular presence
}}

The '''Italian wolf''' ('''''Canis lupus italicus'''''<ref>{{harvp|Altobello|1921}}; {{harvp|Nowak|Federoff|2002}}; {{harvp|Castelló|2018}}; {{harvp|Boitani|Phillips|Jhala|2023}}</ref><ref>{{cite web|author=IUCN/SSC Canid Specialist Group ( Claudio Sillero)|title=Grey wolf ''Canis lupus''|website=International Union for the Conservation of Nature - Canid Specialist Group|url=https://www.canids.org/species/view/PREKLD895731}}</ref> or '''''Canis lupus lupus'''''<ref name=msw3/>), also known as the '''Apennine wolf''',{{sfn|Salari|Achino|Gatta|Petronio|2017}}<ref name=sardella2014>{{cite journal |last1=Sardella |first1=Raffaele |doi=10.1016/j.quaint.2013.11.016 |title=The wolf from Grotta Romanelli (Apulia, Italy) and its implications in the evolutionary history of Canis lupus in the Late Pleistocene of Southern Italy |journal=[[Quaternary International]] |volume=328–329 |pages=179–195 |year=2014 |last2=Bertè |first2=Davide |last3=Iurino |first3=Dawid Adam |last4=Cherin |first4=Marco |last5=Tagliacozzo |first5=Antonio |bibcode=2014QuInt.328..179S}}</ref> is a [[Subspecies of Canis lupus|subspecies of the grey wolf]] native to the [[Italian Peninsula]]. It inhabits the [[Apennine Mountains]] and the [[Western Alps]], though it is undergoing expansion towards the north and east. As of 2022 the wolf population within Italy is estimated to be 3,307 individuals.<ref>{{cite web |url=https://www.isprambiente.gov.it/it/attivita/biodiversita/monitoraggio-nazionale-del-lupo/risultati |title=Risultati: I risultati del Monitoraggio nazionale del lupo |language=it |trans-title=Results: The results of the National Wolf Monitoring |website=Istituto Superiore per la Protezione e la Ricerca Ambientale |date=12 May 2022}}</ref> Although not universally recognised as a distinct subspecies,<ref name=msw3/> it nonetheless possesses a unique [[mtDNA]] [[haplotype]]{{sfn|Wayne|Lehman|Allard|Honeycutt|1992}}{{sfn|Randi|Lucchini|Christensen|Nadia|2000}}<ref>{{cite journal|doi=10.1016/j.biocon.2016.01.003 |title=Why do wolves eat livestock? |journal=[[Biological Conservation]] |volume=195 |pages=156–168 |year=2016 |last1=Imbert |first1=Camille |last2=Caniglia |first2=Romolo |last3=Fabbri |first3=Elena |last4=Milanesi |first4=Pietro |last5=Randi |first5=Ettore |last6=Serafini |first6=Matteo |last7=Torretta |first7=Elisa |last8=Meriggi |first8=Alberto|s2cid=87399626 }}</ref> and a distinct skull [[Morphology (biology)|morphology]].{{sfn|Nowak|Federoff|2002}}

It has been strictly protected in Italy since the 1970s, when the population reached a low of 70–100 individuals. The population is increasing in number, though illegal hunting and persecution still constitute a threat. Since the 1990s, the Italian wolf's range has expanded into southeastern France{{sfn|Monnier|Figuet|2013}} and Switzerland.{{sfn|Glenz|Massolo|Kuonen|Schlaepfer|2001}}

The Italian wolf features prominently in Latin and Italian cultures, such as the [[She-wolf (Roman mythology)|She-Wolf]] in the legendary [[founding of Rome]].<ref name="Baker">{{cite book |publisher=Printed for A. Strahan |last=Livy |others=George Baker (trans.) |title=The history of Rome |year=1797}}</ref> For this reason it is unofficially considered the [[national animal]] of Italy.<ref>{{cite book |last1=Minahan |first1=James B. |title=The Complete Guide to National Symbols and Emblems |date=2009 |publisher=[[ABC-CLIO]] |page=436 |isbn=9780313344978 |url=https://books.google.com/books?id=jfrWCQAAQBAJ&pg=PA436}}</ref><ref>{{cite book |last1=Blashfield |first1=Jean F. |title=Italy |date=2009 |publisher=Scholastic |page=33 |isbn=9780531120996 |url=https://archive.org/details/italy0000blas/page/33/mode/2up}}</ref>

== Taxonomy ==
[[File:Vertebrati del Molise e dell'Abruzzo (1925) C. l. lupus & C. l. italicus.jpg|left|thumb|Giuseppe Altobello's comparative illustration of the skulls and dentition of ''[[Canis lupus lupus|C. l. lupus]]'' ('''a''') and ''C. l. italicus'' ('''b''')]]
[[File:La bellezza dei lupi Parco Nazionale d'Abruzzo l'Aquila Italy 11.png|thumb|Italian wolf pack in the [[Abruzzo, Lazio and Molise National Park]]]]

The modern Italian wolf was first recognised as a distinct subspecies in 1921 by zoologist [[:it:Giuseppe Altobello|Giuseppe Altobello]],{{sfn|Altobello|1921}} who noted that its colour and skull differed from that of the [[Eurasian wolf|common European wolf]]. He described the Italian population's skull as being rounder in form than that of the typical European wolf, with smaller teeth closely approaching those of [[dog]]s and [[golden jackal]]s in appearance.<ref name="altobello1925">{{in lang|it}} Altobello, G. (1925), ''[http://www.storiadellafauna.it/scaffale/testi/alto/forme.htm Vertebrati del Molise e dell’Abruzzo. Forme locali]'', Annuario dell’Istituto Tecnico Provinciale "Leopoldo Pilla", Campobasso, pp. 231-255</ref> Altobello's classification was later rejected by several authors, including [[Reginald Innes Pocock]], who [[Synonym (taxonomy)|synonymised]] ''C. l. italicus'' with ''C. l. lupus''.{{sfn|Nowak|Federoff|2002}} In 2002, the noted paleontologist R.M. Nowak reaffirmed the morphological distinctiveness of the Italian wolf in a study on grey wolf skulls from Italy, other Eurasian localities, and dog skulls. The results of this assessment showed no overlap in the skull morphology of Italian wolves and other grey wolves and dogs. Among the discovered characteristics distinguishing the Italian wolf were its relatively narrow [[palate]] between the first [[premolar]]s, a broad [[Frontal bone|frontal shield]], and shallow [[jugal bone]]. The study recommended the recognition of ''Canis lupus italicus''.{{sfn|Nowak|Federoff|2002}}

{{As of|2005}},<ref name=msw3>{{MSW3 Wozencraft |pages= |id=14000739}}</ref> it is classed by the third edition of [[MSW3|Mammal Species of the World]] as synonymous with ''C. l. lupus''. Nevertheless, the [[National Center for Biotechnology Information]] does list and publish research papers recognising its distinctiveness.<ref>{{cite web|title=NCBI search Canis lupus italicus |url=https://pubmed.ncbi.nlm.nih.gov/?term=canis+lupus+italicus}}</ref><ref>{{cite journal|pmid=24450075 |year=2013 |last1=Zeira |first1=O. |title=Clinical and diagnostic imaging findings in an Italian wolf (Canis lupus italicus) with discospondylitis |journal=Journal of Zoo and Wildlife Medicine |volume=44 |issue=4 |pages=1086–9 |last2=Briola |first2=C. |last3=Konar |first3=M. |last4=Plonek |first4=M. |last5=Papa |first5=V. |doi=10.1638/2013-0007R1.1}}</ref><ref>{{cite journal|pmid=26042204 |pmc=4443502 |year=2015 |last1=Gori |first1=F. |title=The occurrence of taeniids of wolves in Liguria (northern Italy) |journal=International Journal for Parasitology: Parasites and Wildlife |volume=4 |issue=2 |pages=252–5 |last2=Armua-Fernandez |first2=M. T. |last3=Milanesi |first3=P. |last4=Serafini |first4=M. |last5=Magi |first5=M. |last6=Deplazes |first6=P. |last7=MacChioni |first7=F. |doi=10.1016/j.ijppaw.2015.04.005|bibcode=2015IJPPW...4..252G }}</ref>

=== Lineage ===
The last specimen of the Mosbach wolf ''[[Canis mosbachensis]]'' in Europe dates to 456–416 thousand years ago, when it gave rise to the wolf ''Canis lupus''. The earliest remains of a wolf in Europe were found in the [[Middle Pleistocene]] site of La Polledrara di Cecanibbio, {{cvt|20|km}} north-west of Rome in deposits dated 406 thousand years ago.<ref>{{Cite journal|last1=Iurino |first1=D.A. |last2=Mecozzi |first2=B. |last3=Iannucci |first3=A. |author3-link=Armando Iannucci |last4=Moscarella |first4=Alfio |last5=Strani |first5=Flavia |last6=Bona |first6=Fabio |last7=Gaeta |first7=Mario |last8=Sardella |first8=Raffaele |title=A Middle Pleistocene wolf from central Italy provides insights on the first occurrence of Canis lupus in Europe |journal=[[Scientific Reports]] |volume=12 |number=2882 |date=2022 |page=2882 |doi=10.1038/s41598-022-06812-5|pmid=35217686 |pmc=8881584 |bibcode=2022NatSR..12.2882I }}</ref>{{sfn|Anzidei|Bulgarelli|Catalano|Cerilli|2012}} The genetic analysis of Apennine wolves indicates that they went through a population decline of 100–1,000 fold between the past 4,700–23,800 years, which indicates genetic isolation south of the alps from other wolf populations for many thousands of years.{{sfn|Randi|2007}}

In 1992, an examination of the [[mitochondrial DNA]] (mDNA) of 26 grey wolf populations worldwide revealed that the Italian wolf has a unique mitochondrial [[haplotype]] (a mutation) not shared by any other grey wolf population.{{sfn|Wayne|Lehman|Allard|Honeycutt|1992}} Further tests on grey wolf mDNA revealed that, unlike several European grey wolf populations, Italian wolves do not share haplotypes with either other grey wolves or domestic dogs.{{sfn|Randi|Lucchini|Christensen|Nadia|2000}}{{sfn|Randi|2007}} In 2010, a study compared the mDNA [[haplotypes]] of 24 ancient wolf specimens from Western Europe dated between 44,000 and 1,200 YBP with those of modern gray wolves. The [[phylogenetic tree]] indicated that the haplotypes represented two [[haplogroups]] that were separated by five mutational steps. The ancient wolf samples from Western Europe all belonged to haplogroup 2, indicating haplogroup 2 predominance in this region for over 40,000 years before and after the [[last glacial maximum]]. A comparison of current and past frequencies indicated that in Europe, haplogroup 2 became outnumbered by haplogroup 1 over the past several thousand years, but in North America, haplogroup 2 became extinct and was replaced by haplogroup 1 after the last glacial maximum. The Italian wolf is the only remaining grey wolf subspecies included in this ancient haplogroup{{sfn|Pilot|Branicki|Jędrzejewski|Goszczyński|2010}} since the extinction of the [[Honshu wolf]].<ref name=matsumura2014>{{cite journal |doi=10.1016/j.ympev.2014.08.004 |pmid=25132126 |title=Reconstructing the colonization history of lost wolf lineages by the analysis of the mitochondrial genome |journal=Molecular Phylogenetics and Evolution |volume=80 |pages=105–12 |year=2014 |last1=Matsumura |first1=Shuichi |last2=Inoshima |first2=Yasuo |last3=Ishiguro |first3=Naotaka|bibcode=2014MolPE..80..105M }}</ref>

In 2016, a study of mDNA sequences of both modern and ancient wolves indicated that in Europe, the two most genetically distinct haplotypes form the [[Iberian wolf]] and separately the Italian wolf. The phylogenetic tree generated from the sequences showed the Italian wolf positioned close to the ancient wolves of the Late Pleistocene.<ref>{{cite journal |doi=10.3389/fevo.2016.00134 |title=From the Past to the Present: Wolf Phylogeography and Demographic History Based on the Mitochondrial Control Region |journal=[[Frontiers in Ecology and Evolution]] |volume=4 |year=2016 |last1=Ersmark |first1=Erik |last2=Klütsch |first2=Cornelya F. C. |last3=Chan |first3=Yvonne L. |last4=Sinding |first4=Mikkel-Holger S. |last5=Fain |first5=Steven R. |last6=Illarionova |first6=Natalia A. |last7=Oskarsson |first7=Mattias |last8=Uhlén |first8=Mathias |last9=Zhang |first9=Ya-Ping |last10=Dalén |first10=Love |last11=Savolainen |first11=Peter |doi-access=free}}</ref> In 2017, a study found a second mDNA haplotype that belonged to the Italian wolf, and called for the morphologically and genetically distinct Italian wolf to be considered as a subspecies.<ref>{{cite journal|doi=10.1016/j.mambio.2017.01.005 |title=A new mitochondrial haplotype confirms the distinctiveness of the Italian wolf (Canis lupus) population |journal=Mammalian Biology - Zeitschrift für Säugetierkunde |volume=84 |pages=30–34 |year=2017 |last1=Montana |first1=Luca |last2=Caniglia |first2=Romolo |last3=Galaverni |first3=Marco |last4=Fabbri |first4=Elena |last5=Randi |first5=Ettore|bibcode=2017MamBi..84...30M }}</ref>

In 2019, an mDNA study of 19 Late Pleistocene-Holocene wolf samples from northern Italy found that these fell within mitochondrial haplogroup 2 except for one sample. Four out of the six detected haplotypes matched ancient [[Beringian wolf|Beringian wolves]], ancient wolves from northern Europe, some modern European and Chinese wolf populations, and are closely related to the two haplotypes currently found in the Italian wolves. The Italian wolf haplotypes were only one or two mutations away from those of the Pleistocene wolves, indicating mutation in their Italian glacial refuge. The Italian wolf population represents genetic uniqueness highlighted in several mitochondrial and nuclear DNA studies. It is the only remaining wolf population in Europe which belongs exclusively to an mDNA haplogroup that was once widespread in central and western Europe for over 40,000 years, and in North America until the [[Last Glacial Maximum]]. Additionally, one canid specimen from the [[:it:Cava a Filo|Cava Filo]] archaeological site of [[San Lazzaro di Savena]], [[Bologna]] fell within the domestic dog clade A [[haplotype]] — it was radio-carbon dated to be 24,700 years old.{{sfn|Ciucani|Palumbo|Galaverni|Serventi|2019}}

In 2020, a genomic study of Eurasian wolves found that the populations of the [[Dinaric Alps]]-[[Balkan Mountains]] region, the Iberian peninsula, and Italy diverged from each other 10,500 years ago followed by negligible gene flow between them. Their long-term isolation may explain the morphological and genetic differences between them.

[[File:Wolf_mandible_diagram.jpg|thumb|right|Wolf mandible diagram showing the names and positions of the teeth]]
{| class="wikitable"
|+ Italian wolf mean carnassial tooth measurements (millimeters) across years (thousands) before present<ref>{{cite journal |doi=10.1016/j.quascirev.2014.12.009 |title=Evolutionary trends and stasis in carnassial teeth of European Pleistocene wolf Canis lupus (Mammalia, Canidae) |journal=[[Quaternary Science Reviews]] |volume=110 |pages=36–48 |year=2015 |last1=Sansalone |first1=Gabriele |last2=Bertè |first2=Davide Federico |last3=Maiorino |first3=Leonardo |last4=Pandolfi |first4=Luca}}</ref>
! Tooth variable
! 90
! 55
! 50
! 45
! 40
! 30
! 20
! Today
|-
| lower m1 length
| 24.5
| 24.0
| 28.0
| 28.5
| 27.5
| 28.0
| 29.0
| 27.0
|-
| lower m1 width
| 10.0
| 9.5
| 11.25
| 11.75
| 11.25
| 11.5
| 12.0
| 11.0
|-
| upper P4 length
| 22.0
| 25.0
| 24.75
| 26.0
| 24.5
| 24.0
| 25.5
| 24.0
|-
| Upper P4 width
| 8.75
| 8.9 
| 9.5
| 10.0
| 10.5
| 9.5
| 10.0
| 9.75
|-
|}

{{clade |style=font-size:75%; line-height:75%;width:500px;
 |label1=Canis
 |1={{clade
  |1=[[Golden jackal]] 1.9 million YBP{{sfn|Koepfli|Pollinger|Godinho|Robinson|2015}}[[File:Dogs, jackals, wolves, and foxes (Plate X).jpg|50 px]]
  |2={{clade
  |1=[[Coyote]] 1.1 million YBP{{sfn|Koepfli|Pollinger|Godinho|Robinson|2015}}[[File:Dogs, jackals, wolves, and foxes (Plate IX).jpg|50 px]]
  |label2=Canis lupus
  |2={{clade
  |1=[[Himalayan wolf]] 630,000 YBP{{sfn|Aggarwal|Kivisild|Ramadevi|Singh|2007}}[[File:Dogs, jackals, wolves, and foxes (Plate III).jpg|50 px]]
  |2={{clade
  |1=[[Indian wolf|Indian grey wolf]] 270,000 YBP{{sfn|Aggarwal|Kivisild|Ramadevi|Singh|2007}}[[File:Dogs, jackals, wolves, and foxes (Plate IV).jpg|50 px]]
  |2={{clade
  |1={{clade
  |label1=Haplogroup-2
  |1={{clade
  |1='''Italian wolf''' (Apennine Peninsula only){{sfn|Pilot|Branicki|Jędrzejewski|Goszczyński|2010}}[[File:Dogs, jackals, wolves, and foxes (Plate III) C. l. italicus mod.jpg|50 px]]
  |2=clade B (pockets of Eurasia only){{sfn|Pilot|Branicki|Jędrzejewski|Goszczyński|2010}}[[File:Beringian wolf reconstruction.jpg|50 px]]
  |3=clade C (pockets of Eurasia only){{sfn|Pilot|Branicki|Jędrzejewski|Goszczyński|2010}}
      }}
     }}
  |2={{clade
  |1=[[Dog]] 40,000 YBP{{sfn|Skoglund|2015}}[[File:Dogs, jackals, wolves, and foxes (Plate XXXVII).jpg|50 px]]
  |label2=Haplogroup-1
  |2=Holarctic [[Gray wolf]] 40,000 YBP{{sfn|Skoglund|2015}}[[File:Dogs, jackals, wolves, and foxes (Plate I).jpg|50 px]]
     }}
    }}
   }}
  }}
 }}
}}
 }}

=== ''Canis lupus maximus'' ===
''Canis lupus maximus'' (Boudadi-Maligne, 2012) was a species larger than all other known fossil and extant wolves from Western Europe. The fossilized remains of this Late Pleistocene subspecies were found across a wide area of south-western France at Jaurens cave, [[Nespouls]], [[Corrèze]] dated 31,000 YBP; Maldidier cave, [[La Roque-Gageac]], [[Dordogne]] dated 22,500 YBP; and Gral pit-fall, [[Sauliac-sur-Célé]], [[Lot (department)|Lot]] dated 16,000 YBP. The wolf's long bones are 10% longer than those of extant European wolves and 20% longer than its probable ancestor, ''C. l. lunellensis''. The teeth are robust, the posterior denticules on the lower premolars p2, p3, p4 and upper P2 and P3 are highly developed, and the diameter of the lower carnassial (m1) were larger than any known European wolf. Wolf body size in Europe has followed a steady increase from their first appearance up to the peak of the last glacial maximum. The size of these wolves is thought to be an adaptation to a cold environment ([[Bergmann's rule]]) and plentiful game, as their remains have been found in association with reindeer fossils.<ref>{{cite journal|doi=10.1016/j.crpv.2012.04.003 |title=Une nouvelle sous-espèce de loup (Canis lupus maximus nov. Subsp.) dans le Pléistocène supérieur d'Europe occidentale [A new subspecies of wolf (Canis lupus maximus nov. subsp.) from the upper Pleistocene of Western Europe] |journal=Comptes Rendus Palevol |volume=11 |issue=7 |pages=475–484 |year=2012 |last1=Boudadi-Maligne |first1=Myriam}}</ref>

In a 2017 study, the dimensions of the upper and lower carnassial teeth of the Italian wolf are close to those of ''C. l. maximus''. Fluctuations in the size of ''C. lupus'' carnassial teeth correlate with the spread of megafauna. The Italian wolf underwent a reduction in body size with the loss of the red deer during the [[Italian Renaissance]] only centuries ago.{{sfn|Salari|Achino|Gatta|Petronio|2017}}

== Description ==
[[File:La bellezza dei lupi Parco Nazionale d'Abruzzo l'Aquila Italy 01.png|thumb|Italian wolf]]
The Italian wolf typically weighs {{convert|25|-|35|kg|abbr=on}}, though some big males have been weighed at {{convert|40|-|45|kg|abbr=on}}. It measures {{cvt|110–148|cm}} in body length and {{cvt|50–70|cm}} in shoulder height.<ref name="lupo">{{cite book|first1=Alessia |last1=Viviani |first2=Andrea |last2=Gazzola |first3=Massimo |last3=Scandura |chapter=Il Lupo: Un predatore sociale ed adattabile |trans-chapter=The Wolf: A social and adaptable predator |editor-first=Marco |editor-last=Apollonio |editor2-first=Luca |editor2-last=Mattioli |title=Il Lupo in provincia di Arezzo |trans-title=The Wolf in the Province of Arezzo |language=it |year=2006 |location=Montepulciano (SI) |publisher=Editrice Le Balze |pages=29–43}}.</ref> The pelt is generally of a grey-fulvous colour, which reddens in summer. The belly and cheeks are more lightly coloured, and dark bands are present on the back and tail tip, and occasionally along the fore limbs. [[Black wolf|Black wolve]]s have been reported in the north-central Apennines, though their origin is unknown, as some [[melanistic]] individuals show no sign of [[wolf-dog hybrid]]isation. It typically lives in [[Pack (canine)|pack]]s of two to seven individuals.<ref name="ciuccu2003">{{cite book|last1=Ciucci |first1=P. |last2=Boitani |first2=L. |author2-link=Luigi Boitani |date=2003 |chapter=Il Lupo ''Canis lupus'' Linnaeus, 1758 |trans-chapter=The Wolf 'Canis lupus' Linnaeus, 1758 |editor1-first=L. |editor1-last=Boitani |editor1-link=Luigi Boitani |editor2-first=S. |editor2-last=Lovari |editor3-first=A. Vigna |editor3-last=Taglianti |title=Fauna d'Italia: Mammalia III, Carnivora-Artiodactyla |trans-title=Fauna of Italy: Mammalia III, Carnivora-Artiodactyla |publisher=Calderini |location=Bologna |pages=20–47 |language=it}}</ref>

== Range and legal status ==
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 [[Nordic countries|Northern Europe]]. Their population decline continued until the 1960s, with isolated populations surviving in Italy, Spain, Portugal, Greece, and Finland. Wolf populations have commenced recovering naturally since then.{{sfn|Randi|2007}}

=== Italy ===
[[File:Areale dei canidi italiani.png|thumb|Current range of the Italian wolf (red) alongside the range of the [[red fox]] and [[European jackal]]]]
[[File:Antonio Mina & Lupo (1929).jpg|thumb|Grey wolf killed in Malga Campo Bon ([[Comelico]]) on 24 May 1929]]
The Italian wolf was widespread in the Italian Peninsula, including [[Sicily]], until the mid-1800s. The extermination of the grey wolf in Italy was not as complete as in Northern Europe, due to greater cultural tolerance of the species.<ref name="bocedi2004">{{cite journal|language=it |last1=Bocedi |first1=R. |last2=Bracchi |first2=P.G. |date=2004 |url=http://www.unipr.it/arpa/facvet/annali/2004/403.pdf |title=Evoluzione demografica del lupo (Canis Lupus) in Italia: cause storiche del declino e della ripresa, nuove problematiche indotte e possibili soluzioni |trans-title=Demographic evolution of the wolf (Canis Lupus) in Italy: historical causes of decline and recovery, new problems and possible solutions |journal=Ann. Fac. Medic. Vet. Di Parma |volume=XXIV |pages=403–415}}</ref> It was largely extirpated in the [[Alps]] during the 1920s, and disappeared from Sicily in the 1940s. Its range along the south-central Apennines was still relatively continuous by the 1950s, though this population was reduced in the decades after [[World War II]] because of widespread poisoning campaigns. At least 400 wolves were killed between 1960 and 1970, with the population reaching an all-time low in the early 1970s. The last documented wolf in the northern Apennines was killed in [[Santo Stefano d'Aveto]], [[Genoa]], in 1946, though this was an isolated individual, as the local wolf population had long been extinct.<ref name="PB2010">{{cite book|language=it |author=Corpo Di Polizia Provinciale |date=2010 |url=http://www.ecosistema.it/sangherardo/PDF/lupo_SFFP_quaderno%202.pdf |title=Il Lupo (Canis Lupus L. 1758) nell'Appenino Bolognese |trans-title=The Wolf (Canis Lupus L. 1758) in the Bolognese Apennines |publisher=[[Province of Bologna]] |series=Quaderno |number=2}}</ref>

The Italian wolf was first given legal protection on 23 July 1971, with a nationwide population census being taken in 1973. This census was funded by the Italian branch of the [[World Wide Fund for Nature]] as part of a conservation plan dubbed "Operazione San Francesco” (Operation Saint Francis, linked to the traditional legend of peace set by the saint between wolf and men of Gubbio in Umbria region). The census revealed that the Italian wolf population consisted of 100–110 individuals distributed throughout a fragmented range in the main mountainous areas of south-central Italy, from the [[Sibillini Mountains|Sibillini]] to [[La Sila]]. In 1983, the population had reached 200–220 individuals inhabiting two unconnected areas in central and southern Italy. By the late 1990s, the Italian wolf population had increased to an estimated 400–500 individuals with a continuous distribution along the entire Apennines, from [[Aspromonte]] to the [[Maritime Alps]], with some isolated populations in [[Tuscany]] and [[Lazio]]. In 2008, a wolf carcass was discovered in the [[Fiemme Valley]] in [[Trentino]], and by 2010, 45–55 wolves were estimated to have recolonised [[Piedmont]].

[[International Union for Conservation of Nature|IUCN]] assessment information about peninsular population is as follows: “The Italian wolf population is estimated to be 500-800 individuals distributed along the Apennines. The shape of the range is narrow and elongated, restricted to the Apennines. The population has limited exchanges with the population of the Western Alps and recent genetic evidence indicates a flux of genes only in the direction toward the Alps. In spite of the recent increase in numbers and range, the Italian wolf population is still highly vulnerable to local extermination from human pressures (poison, shooting, car accidents) and the stochastic nature of these events suggest to maintain a cautionary assessment. The population does not qualify for the category Endangered, but it may easily reverse its current favorable status.”

=== France ===
In the early 1990s Italian wolves began to cross from Italy into France, where they have become established in approximately one third of its contintental territories, particularly in the French Alps and [[Provence]], but also throughout the [[Massif Central]].{{sfn|Bobbé|2000}}{{sfn|Meuret|Moulin|Bonnet|Garde|2020}} Even before the presence of wolves was publicly reported in France, some farmers around the [[Mercantour National Park]] had reported unusual stock depredation which was at the time attributed by authorities to uncontrolled domestic dogs.{{sfn|Meuret|Moulin|Bonnet|Garde|2020}} Wolves are protected in France;{{refn|group=note|In France, illegally killing a wolf is punishable by two years' imprisonment and a €150,000 fine.{{sfn|Meuret|Moulin|Bonnet|Garde|2020}} }} in order to protect the livelihoods of farmers from wolf predation, from the late 1990s the French government has subsidised various methods of protecting flocks from depredation, including electrified pasture fencing, secured electrified night pens, hiring of additional farm hands, and the purchase, training and upkeep of livestock guardian dogs.{{sfn|Meuret|Moulin|Bonnet|Garde|2020}}

The Italian wolf was first sighted in southwestern France in 1992. In the two decades following its initial recolonisation, the wolf has expanded its range at the west of the [[Rhône]], in the Massif Central, the eastern [[Pyrenees]], and the [[Jura Mountains|Jura]] and [[Vosges Mountains]].{{sfn|Monnier|Figuet|2013}}

=== Spain ===
At least 13 transient Italian wolves (12 males and a female) were counted in [[Catalonia]] between 2000 and 2011, a century after the local [[Iberian wolf]] (''C. l. signatus'') was extirpated from the area.<ref>{{cite journal|last=Garcia-Lozano |first=C. |others=et al. |date=2015 |title=Análisis de la disponibilidad de hábitat adecuado para el lobo (Canis lupus) en Cataluña y en los Pirineos Orientales. |language=es |trans-title=Analysis of the availability of suitable habitat for the wolf (Canis lupus) in Catalonia and in the Eastern Pyrenees. |journal=Pirineos. Revista de Ecología de Montaña |volume=170 |pages=1–13}}</ref> One of these was reported in the media to have been killed on the road and identified in [[Baix Empordà]], Catalonia in 2018. Technicians and veterinarians of the Torreferrussa Wildlife Center identified the body as being of the subspecies ''Canis lupus italicus''.<ref>{{cite news|url=http://www.lavanguardia.com/natural/20180302/441196233217/muere-atropellado-un-lobo-en-una-carretera-en-el-baix-emporda.html |title=Muere atropellado un lobo en una carretera en el Baix Empordà |language=es |trans-title=A wolf dies on a road in the Baix Empordà |work=[[La Vanguardia]] |date=3 February 2018}}</ref>

=== Switzerland ===
The first evidence of grey wolf expansion into Switzerland occurred in 1995–1996 in the southern [[Canton of Valais]], where around 100 sheep were killed. In 1998–1999, 40 sheep were killed and two wolves were found dead from poaching and car collisions.{{sfn|Glenz|Massolo|Kuonen|Schlaepfer|2001}} The first wolf pack formed 17 years after the return of the first wolves, in 2012 in the canton of [[Grisons]]. The reproducing female F07 of this first pack was confirmed to be still alive in 2023, making it 13–14 years old.<ref>{{Cite web |last=Kora |title=F07 – die älteste Wölfin der Schweiz |url=https://www.kora.ch/de/aktuell/f07-–-die-aelteste-woelfin-der-schweiz--532 |access-date=2023-04-25 |website=www.kora.ch |language=de |trans-title=F07 – the oldest she-wolf in Switzerland}}</ref> Since the formation of the first pack, the Swiss population is growing rapidly.<ref name="Kora">{{Cite web |last=Kora |title=BESTAND |url=https://www.kora.ch/de/arten/wolf/bestand |access-date=2023-04-25 |website=www.kora.ch |language=de}}</ref> In 2023, there were 240 wolves, 18 wolf packs entirely in Switzerland and five on the border to France or Italy.<ref name="Kora"/> Most of these wolf packs roam in the cantons of Grisons, Valais and [[Ticino]].<ref>{{Cite web |title=Rudel - Gruppe Wolf Schweiz |url=https://www.gruppe-wolf.ch/Rudel.htm |access-date=2023-04-25 |website=www.gruppe-wolf.ch}}</ref>

Italian, French and Swiss wolves share the same mtDNA haplotype,{{sfn|Valière|Fumagalli|Gielly|Miquel|2003}} a haplotype that has never been found in any other wolf population worldwide,<ref>{{cite journal|doi=10.1101/gr.116301.110 |pmid=21566151 |title=A genome-wide perspective on the evolutionary history of enigmatic wolf-like canids |journal=[[Genome Research]] |volume=21 |issue=8 |pages=1294–305 |year=2011 |last1=Vonholdt |first1=Bridgett M. |last2=Pollinger |first2=John P. |last3=Earl |first3=Dent A. |last4=Knowles |first4=James C. |last5=Boyko |first5=Adam R. |last6=Parker |first6=Heidi |last7=Geffen |first7=Eli |last8=Pilot |first8=Malgorzata |last9=Jedrzejewski |first9=Wlodzimierz |last10=Jedrzejewska |first10=Bogumila |last11=Sidorovich |first11=Vadim |last12=Greco |first12=Claudia |last13=Randi |first13=Ettore |last14=Musiani |first14=Marco |last15=Kays |first15=Roland |last16=Bustamante |first16=Carlos D. |last17=Ostrander |first17=Elaine A. |last18=Novembre |first18=John |last19=Wayne |first19=Robert K. |pmc=3149496}} refer Figure 3</ref> which corroborate the scenario of a natural expansion of wolves from the Italian source population.{{sfn|Valière|Fumagalli|Gielly|Miquel|2003}} Although the Italian and Dinaric wolf populations have remained distinct for a long time, their ranges are beginning to overlap as they expand. In the canton of Glarus, Switzerland, the first hybrid offspring has been documented, born to parent wolves originating from both of these populations.<ref>{{Cite web |title=Rudel-Schilt-Karpf - Gruppe Wolf Schweiz |url=https://www.gruppe-wolf.ch/Rudel/Rudel-Schilt-Karpf.htm |access-date=2023-04-25 |website=www.gruppe-wolf.ch}}</ref>

== Cultural significance ==
[[File:Lupa Capitolina, Rome.jpg|thumb|right|According to legend, [[founding of Rome|Rome was founded]] in 753&nbsp;BC by [[Romulus and Remus]], who were raised by a [[She-wolf (Roman mythology)|she-wolf]].]]
The animal features prominently in pre-Roman, [[Roman culture|Roman]], and later [[Italian culture]]s. In [[Roman mythology]], the wolf played a role in the [[founding of Rome]] by suckling the twins [[Romulus and Remus]]. According to [[Terry Jones]], "The Romans did not see [the tale of Romulus, Remus and the she-wolf] as a charming story; they meant to show that they had imbibed wolfish appetites and ferocity with their mother's milk".<ref>Jones, T. (2007), ''Terry Jones' Barbarians: An Alternative Roman History'', BBC Books, {{ISBN|056353916X}}</ref> The wolf was also considered sacred to [[Mars (mythology)|Mars]], and to see a wolf before going into battle was considered a good omen.<ref name="impelluso2004">Impelluso, L. (2004), ''Nature and its Symbols'', Getty Publications, p. 212, {{ISBN|0892367725}}</ref> The origin of the myth can be traced back to a wolf cult among the neighbouring [[Sabines]]. The Sabines had two words for wolf: ''hirpus'' (used in religious contexts) and ''lupus'', the latter of which was incorporated into [[Latin language|Latin]].{{sfn|Boitani|1995}}

Although the Romans did not worship wolves, killing them was likely considered [[taboo]]; unlike the [[Etruscans]], the Romans very rarely sacrificed wolves in rituals, and no records have been found of wolves being used in the [[Roman amphitheatre|amphitheatre]]s, despite being more numerous and easily accessible compared to other, more exotic animals used. The use of wolves in Roman [[folk medicine]], while attested by [[Pliny the Elder]], was minimal compared to other animals such as snakes or bears and, contrary to popular imagery, [[Imaginifer|Roman standard bearer]]s did not wear wolf skins, with the only units attested to have worn them being the [[velites]], who were the poorest and youngest warriors using the wolf skins to distinguish themselves. Wolves entering cities or temples were usually only killed when the animal had no means of escape, unlike the case with wasps, oxen, and owls, which were quickly eliminated if they entered sacred areas.{{sfn|Rissanen|2014}} Negative attitudes towards wolves in Italy largely began with the [[Kingdom of the Lombards|invasion of the Lombards]], who [[Zoomorphism|zoomorphically]] described their raids and invasions as wolf raids, bringing wolves into disrepute.{{sfn|Boitani|1995}} The belief in [[Werewolf|werewolves]] was still widespread in Italy during the early 1920s, and covering their faces when resting outside at night was once traditional among rural people, as sleeping whilst facing the [[full moon]] was thought to transform the sleeper into a wolf. The wolf also featured prominently in Italian folk medicine. Tying a sack filled with a piece of wolf gut around the child's neck was thought to treat [[baby colic]] and tying a wolf's intestine around the mother's abdomen was thought to prevent [[miscarriage]]. Wolf fat was considered a treatment for [[rheumatism]] and [[tonsillitis]], while a tooth or tuft of fur was sometimes worn as a [[talisman]] against the [[evil eye]].{{sfn|Altobello|1921}}

The Romans apparently did not consider wolves overly dangerous to people, with the only references to them attacking people being proverbial or mythological.{{sfn|Rissanen|2014}} Although Italy has no records of [[wolf attacks on humans]] after [[World War II]] and the eradication of [[rabies]] in the 1960s,<ref>{{cite book|url=http://www.wnmuoutdoors.org/PDF/Linnell2002_Wolves.pdf |title=The Fear of Wolves: A Review of Wolf Attacks on Humans |publisher=NINA |last1=Linnell |first1=J.D.C. |last2=Andersen |first2=R. |author2-link=:no:Reidar Andersen (biolog) |last3=Andersone |first3=Z. |last4=Balciauskas |first4=L. |author4-link=:lt:Linas Balčiauskas |last5=Blanco |first5=J.C. |last6=Boitani |first6=L. |author6-link=Luigi Boitani |last7=Brainerd |first7=S. |last8=Breitenmoser |first8=U. |last9=Kojola |first9=I. |last10=Liberg |first10=O. |last11=Løe |first11=J. |last12=Okarma |first12=H. |author12-link=Henryk Okarma |last13=Pedersen |first13=H. C. |last14=Promberger |first14=C. |last15=Sand |first15=H. |last16=Solberg |first16=E. J. |last17=Valdmann |first17=H. |last18=Wabakken |first18=P. |year=2002 |isbn=978-82-426-1292-2 |url-status=dead |archive-url=https://web.archive.org/web/20131109205016/http://www.wnmuoutdoors.org/PDF/Linnell2002_Wolves.pdf |archive-date=2013-11-09}}</ref> historians examining church and administrative records from northern Italy's central [[Po Valley]] region (which includes a part of modern-day Switzerland) found 440 cases of wolves attacking people between the 15th and 19th centuries. The 19th-century records show that from 1801 to 1825, 112 attacks occurred, 77 of which resulted in death. Of these cases, only five were attributed to rabid animals.<ref name="cagnolaro1992">{{cite conference |last1=Cagnolaro |first1=L. |last2=Comincini |first2=M. |last3=Martinoli |first3=A. |last4=Oriani |first4=A. |url=http://www.storiadellafauna.it/scaffale/testi/cagnolaro/antropofagia.htm |title=Dati Storici sulla Presenza e su Casi di Antropofagia del Lupo nella Padania Centrale |language=it |trans-title=Historical data on the presence and cases of anthropophagy of the wolf in central Padania |conference=atti del convegno nazionale "Dalla parte del lupo" |location=Parma |date=9–10 October 1992 |publisher=Atti & Studi del WWF Italia |number=10 |pages=1–160 |postscript=,F. Cecere (a cura di), 1996, Cogecstre Edizioni |access-date=2 May 2014 |archive-date=9 November 2013 |archive-url=https://web.archive.org/web/20131109210643/http://www.storiadellafauna.it/scaffale/testi/cagnolaro/antropofagia.htm |url-status=dead }}</ref>

== Notes ==
{{reflist|group=note}}

== References ==
{{reflist}}

=== Bibliography ===
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{{refend}}

== Further reading ==
* {{cite book |last1=Apollonio |first1=Marco |last2=Mattioli |first2=Luca |title=Il Lupo in Provincia di Arezzo |trans-title=The Wolf in the Province of Arezzo |publisher=Editrice Le Balze |year=2006 |isbn=978-88-7539-123-2 |language=it}}
* {{cite book |last1=Boitani |first1=Luigi |author-link=Luigi Boitani |title=Dalla parte del lupo: la riscoperta scientifica e culturale del mitico predatore |trans-title=On the side of the wolf: the scientific and cultural rediscovery of the mythical predator |publisher=Mondadori |year=1987 |language=it}}
* {{cite book |last1=Comincini |first1=Mario |title=L'uomo e la "bestia antropofaga": storia del lupo nell'Italia settentrionale dal XV al XIX secolo |trans-title=Man and the "anthropophagous beast": history of the wolf in northern Italy from the 15th to the 19th century |publisher=Unicopoli |year=2002 |isbn=978-8840007748 |language=it}}
* {{cite book |last1=Diviacco |first1=Giovanni |last2=Campora |first2=Massimo |last3=Cottalasso |first3=Renato |title=Sulle tracce del lupo in Liguria |trans-title=On the trail of the wolf in Liguria |publisher=Centro Stampa Offset |year=2009 |language=it}}
* {{cite book |last1=Ferrari |first1=Marco Albino |author-link=:it:Marco Albino Ferrari |title=La via del lupo |trans-title=The way of the wolf |publisher=Editori Laterza |year=2012 |isbn=9788842056003 |language=it}}
* {{cite book |editor-last=Genovesi |editor-first=P. |date=2002 |url=http://www1.nina.no/lcie_new/pdf/634989762488267659_Italian%20wolf%20action%20plan%202002.pdf |title=Piano d'azione nazionale per la conservazione del Lupo (''Canis lupus'') |trans-title=National action plan for the conservation of the wolf (''Canis lupus'') |publisher=Quaderni di Conservazione della Natura, 13, Min. Ambiente - Istituto Nazionale per la Fauna Selvatica |language=it}}
* {{cite book |last1=Marucco |first1=Francesca |title=Il lupo: Biologia e gestione nelle Alpi ed in Europa |trans-title=The wolf: Biology and management in the Alps and in Europe |publisher=Il Piviere |year=2014 |isbn=978-8896348239 |language=it}}
* {{cite book |last1=Marucco |first1=Francesca |title=I lupi delle Alpi Marittime: Storie e curiosità sui branchi che per primi hanno ricolonizzato le Alpi |trans-title=Wolves of the Maritime Alps: Stories and curiosities about the packs that first re-colonized the Alps |publisher=Blu Edizioni |year=2015 |isbn=978-8879042000 |language=it}}		
* {{cite journal |last1=Meriggi |first1=Alberto |last2=Brangi |first2=Anna |last3=Matteucci |first3=Carlo |last4=Sacchi |first4=Oreste |year=1996 |title=The feeding habits of wolves in relation to large prey availability in northern Italy |journal=[[Ecography]] |volume=19 |issue=3 |pages=287–295 |doi=10.1111/j.1600-0587.1996.tb00238.x}}
* {{cite journal |last1=Rissanen |first1=M. |url=https://www.academia.edu/2177407 |title=The Hirpi Sorani and the Wolf Cults of Central Italy |journal=Acta Philologica Fennica |volume=XLVI |date=2012}}

== External links ==
{{Wikispecies|Canis lupus italicus}}
{{Commons category|Canis lupus italicus}}

{{grey wolf subspecies}}
{{National symbols of Italy}}
{{Italy topics}}
{{Taxonbar|from=Q1188023}}

[[Category:Subspecies of Canis lupus]]
[[Category:Fauna of Italy|Italian Wolf]]
[[Category:Mammals described in 1921]]
[[Category:Wolves]]
[[Category:National symbols of Italy]]