The Italian wolf (Canis lupus italicus or Canis lupus lupus), also known as the Apennine wolf, is a 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. Although not universally recognised as a distinct subspecies, it nonetheless possesses a unique mtDNA haplotype and a distinct skull morphology. 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 and Switzerland. The Italian wolf features prominently in Latin and Italian cultures, such as the She-Wolf in the legendary founding of Rome. For this reason it is unofficially considered the national animal of Italy.
Taxonomy
The modern Italian wolf was first recognised as a distinct subspecies in 1921 by zoologist Giuseppe Altobello, who noted that its colour and skull differed from that of the 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 dogs and golden jackals in appearance. Altobello's classification was later rejected by several authors, including Reginald Innes Pocock, who synonymised C. l. italicus with C. l. lupus. 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 premolars, a broad frontal shield, and shallow jugal bone. The study recommended the recognition of Canis lupus italicus. As of 2005, it is classed by the third edition of 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.
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 grey wolf (Canis lupus). The earliest remains of a wolf in Europe were found in the Middle Pleistocene site of La Polledrara di Cecanibbio, 20 km (12 mi) north-west of Rome in deposits dated 406 thousand years ago. 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. 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. 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. 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 since the extinction of the Japanese wolf. 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. 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. 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 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 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. 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.
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