''Canis etruscus'' (the Etruscan wolf), is an extinct species of
canine that was
endemic
Endemism is the state of a species being found only in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also foun ...
to
Mediterranean Europe
Southern Europe is also known as Mediterranean Europe, as its geography is marked by the Mediterranean Sea. Definitions of southern Europe include some or all of these countries and regions: Albania, Andorra, Bosnia and Herzegovina, Bulgaria, C ...
and
Crimean peninsula
Crimea ( ) is a peninsula in Eastern Europe, on the northern coast of the Black Sea, almost entirely surrounded by the Black Sea and the smaller Sea of Azov. The Isthmus of Perekop connects the peninsula to Kherson Oblast in mainland Ukrai ...
during the
Early Pleistocene
The Early Pleistocene is an unofficial epoch (geology), sub-epoch in the international geologic timescale in chronostratigraphy, representing the earliest division of the Pleistocene Epoch within the ongoing Quaternary Period. It is currently esti ...
.
[ The Etruscan wolf is described as a small wolf-like dog.] It is widely agreed to be the ancestor of '' Canis mosbachensis'', and thus ultimately the modern grey wolf
The wolf (''Canis lupus''; : wolves), also known as the grey wolf or gray wolf, is a canine native to Eurasia and North America. More than thirty subspecies of ''Canis lupus'' have been recognized, including the dog and dingo, though gr ...
(''Canis lupus'').
Taxonomy
The fossil record
A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
for ancient vertebrates
Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain.
The vertebrates make up the subphylum Vertebra ...
is composed of rarely occurring fragments, from which it is often impossible to obtain genetic material. Researchers are limited to morphologic analysis but it is difficult to estimate the intra-species and inter-species variations and relationships that existed between specimens across time and place. Some observations are debated by researchers who do not always agree, and hypotheses that are supported by some authors are challenged by others.[ Several species of ]Caninae
Caninae (whose members are known as canines () is the only living subfamily within Canidae, alongside the extinct Borophaginae and Hesperocyoninae. They first appeared in North America, during the Oligocene around 35 million years ago, subsequent ...
from the Pleistocene of Europe have been described. Most of their systematic
Systematic may refer to:
Science
* Short for systematic error
* Systematic fault
* Systematic bias, errors that are introduced by an inaccuracy inherent to the system
Economy
* Systematic trading, a way of defining trade goals, risk control ...
and phylogenetic
In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
relationships have not been resolved because of their similar morphology.
Upper Valdarno
The Valdarno is the valley of the river Arno, from Florence to the sea. The name applies to the entire river basin, though usage of the term generally excludes Casentino and the valleys formed by major tributaries.
Some towns in the area:
* R ...
is the name given to that part of the Arno
The Arno is a river in the Tuscany region of Italy. It is the most important river of central Italy after the Tiber.
Source and route
The river originates on Monte Falterona in the Casentino area of the Apennines, and initially takes a sou ...
valley situated in the provinces of Florence
Florence ( ; ) is the capital city of the Italy, Italian region of Tuscany. It is also the most populated city in Tuscany, with 362,353 inhabitants, and 989,460 in Metropolitan City of Florence, its metropolitan province as of 2025.
Florence ...
and Arezzo
Arezzo ( , ; ) is a city and ''comune'' in Italy and the capital of the Province of Arezzo, province of the same name located in Tuscany. Arezzo is about southeast of Florence at an elevation of Above mean sea level, above sea level. As of 2 ...
, Italy. The region is bounded by the Pratomagno
The Pratomagno is a mountain range, which has the Arno River on both sides: to the west is the upper Valdarno and to the east is the Casentino. It lies north-west of the city of Arezzo, in Tuscany, Italy
Italy, officially the Italian Repu ...
mountain range to the north and east and by the Chianti
Chianti is an Italian red wine produced in the Chianti (region), Chianti region of central Tuscan wine, Tuscany, principally from the Sangiovese grape. It was historically associated with a squat bottle enclosed in a straw basket, called a ''fia ...
mountains to the south and west. The Upper Valdarno Basin has provided the remains of three fossil canid species dated to the Late Villafranchian era of Europe, 1.9–1.8 million years ago, that arrived with a faunal turnover around that time. The Swiss paleontologist Charles Immanuel Forsyth Major discovered two species in this region, these being the Falconer's wolf ('' Canis falconeri'' Forsyth Major 1877) that was later reclassified as ''Lycaon falconeri'', and the smaller Etruscan wolf (''C. etruscus'' Forsyth Major 1877).[ Forsyth Major did not publish a complete description of the Etruscan wolf,] and later Domenico Del Campana worked on expanding Forsyth Major's descriptions when he recognized among the specimens a smaller, jackal
Jackals are Canidae, canids native to Africa and Eurasia. While the word has historically been used for many canines of the subtribe Canina (subtribe), canina, in modern use it most commonly refers to three species: the closely related black-b ...
-sized species. This he named the Arno River dog[ (''C. arnensis'' Del Campana 1913) in honour of the nearby ]Arno
The Arno is a river in the Tuscany region of Italy. It is the most important river of central Italy after the Tiber.
Source and route
The river originates on Monte Falterona in the Casentino area of the Apennines, and initially takes a sou ...
river.
Type Specimen
The lectotype
In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally associated. In other words, a type is an example that serves to anchor or centralizes ...
is a cranium from an unrecorded locality of Upper Valdarno. It is housed in the Montevarchi
Montevarchi is a town and ''comune'' in the province of Arezzo, Tuscany, Italy.
History
The town of Montevarchi sprang up around 1100, near to a fortified Benedictine monastery, founded by bishop Elempert (986–1010) of Arezzo. At first the cas ...
Paleontological Museum. The specimen has been designed as lectotype by the Italian paleontologist Danilo Torre in 1967 from the sample described by Forsyth Major.[
]
''Canis apolloniensis''
One specimen of ''C. apolloniensis'' (Koufos and Kostopoulos, 1997) was found in site of Apollonia-1 near the village of Nea Apollonia, Macedonia
Macedonia (, , , ), most commonly refers to:
* North Macedonia, a country in southeastern Europe, known until 2019 as the Republic of Macedonia
* Macedonia (ancient kingdom), a kingdom in Greek antiquity
* Macedonia (Greece), a former administr ...
, in northern Greece
Greece, officially the Hellenic Republic, is a country in Southeast Europe. Located on the southern tip of the Balkan peninsula, it shares land borders with Albania to the northwest, North Macedonia and Bulgaria to the north, and Turkey to th ...
. The holotype
A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
consists of the rostral portion of a skull and a mandible
In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla).
The jawbone i ...
. In 2011, a study compared all of the fifty-five Early Pleistocene
The Early Pleistocene is an unofficial epoch (geology), sub-epoch in the international geologic timescale in chronostratigraphy, representing the earliest division of the Pleistocene Epoch within the ongoing Quaternary Period. It is currently esti ...
wolf-like specimens found across Europe and suggested that their morphometric variation was no different to that of modern wolf populations, with their difference in size representing male and female specimens. However, the study proposed two lineages. One lineage is '' C. arnensis'' which includes ''C. accitanus'' and ''C. senezensis'', and the other lineage being ''C. etruscus'' that includes ''C. apolloniensis''.[ Other studies] contrast with this biometric interpretation. Considering morphological features retained by ''C. apolloniensis'', this species should be considered as a synonym of '' C. mosbachensis'' or a very affine taxon.
Lineage
The large wolf-sized ''Canis'' first appeared in the Middle Pliocene
The Pliocene ( ; also Pleiocene) is the epoch (geology), epoch in the geologic time scale that extends from 5.33 to 2.58[Shanxi
Shanxi; Chinese postal romanization, formerly romanised as Shansi is a Provinces of China, province in North China. Its capital and largest city of the province is Taiyuan, while its next most populated prefecture-level cities are Changzhi a ...](_blank)
Province, China. 2.5 million years ago its range included the Nihewan Basin in Yangyuan County, Hebei
Hebei is a Provinces of China, province in North China. It is China's List of Chinese administrative divisions by population, sixth-most populous province, with a population of over 75 million people. Shijiazhuang is the capital city. It bor ...
, China, and Kuruksay, Tajikistan.[ In Europe, ''C. etruscus'' first appeared 1.9-1.8 million years ago.][ The lineage from ''C. etruscus'' to the Mosbach wolf (''C. mosbachensis'' Soergel, 1925) to the grey wolf (''C. lupus'') is widely accepted in the European scientific literature.][ The French archaeologist Jean-Philip Brugal proposes ''C. mosbachensis'' as a subspecies of ''C. etruscus'',][ and another French archaeologist, Henry de Lumley, considers ''C. mosbachensis'' to be a subspecies of the grey wolf ''C. lupus mosbachensis''.][
In 2020, researchers named a new species ''C. borjgali'' that was found in ]Dmanisi
Dmanisi ( ka, დმანისი, tr, , ) is a town and archaeological site in the Kvemo Kartli region of Georgia (country), Georgia approximately 93 km southwest of the nation’s capital Tbilisi in the river valley of Mashavera.
Abandoned i ...
, Georgia in a site dated 1.8—1.75 million years ago. This specimen did not show the peculiarities of ''C. etruscus'' (as shown by Cherin et al., 2014) but appears to be closer to a primitive form of ''C. mosbachensis'' and is proposed as the ancestor of ''C. mosbachensis'', with ''Canis etruscus'' being the ancestor of the Eurasian-African jackal
Jackals are Canidae, canids native to Africa and Eurasia. While the word has historically been used for many canines of the subtribe Canina (subtribe), canina, in modern use it most commonly refers to three species: the closely related black-b ...
.[ The wolf ''C. borjgali'' is probably the ancestor of the wolf-like crown-clade species ('' C. lupus'', '' C. latrans'', '' C. lupaster'').]
Paleoecology
The dispersal of carnivoran species occurred approximately 1.8 million years ago and this coincided with a decrease in precipitation and an increase in annual seasonality which followed the 41,000-year amplitude shift of Milankovitch cycles
Milankovitch cycles describe the collective effects of changes in the Earth's movements on its climate over thousands of years. The term was coined and named after the Serbian geophysicist and astronomer Milutin Milanković. In the 1920s, he pr ...
. First to arrive was ''C. etruscus'', which was immediately followed by ''C. arnensis'' and ''Lycaon falconeri'', and then the giant hyena ''Pachycrocuta
''Pachycrocuta'' is an extinct genus of hyena. The largest and most well-researched species is ''Pachycrocuta brevirostris'', colloquially known as the giant short-faced hyena as it stood about at the shoulder and it is estimated to have average ...
brevirostris''. These were all better adapted to open, dry landscapes than the two more primitive canini '' Eucyon'' and '' Nyctereutes'' that they replaced in Europe.
Description
A description of the Etruscan wolf appears below:
Medium-sized dog (average size of a small ''C. lupus''); elongated snout; marked constriction of the snout beyond the infraorbital foramina; elongated nasal bones extending beyond the maxillofrontal suture; well-developed sagittal
The sagittal plane (; also known as the longitudinal plane) is an anatomical plane that divides the body into right and left sections. It is perpendicular to the transverse plane, transverse and coronal plane, coronal planes. The plane may be in ...
and nuchal crests; laterally enlarged occipital region; P1-P2-P3 laterally compressed; P1 with lingual cingulum; P3 normally with both posterior and accessory cusp (=modified posterior cingulum); large relative length of the upper molar row (in comparison with ''C. arnensis''); M1 with paracone more elevated than metacone; labial basin of M1 as deep as but larger than the lingual one; M1 and M2 with a continuous cingulum; reduced contact area between M1 and
M2. Lower dentition characterized by larger dimensions in respect to ''C. arnensis'' and by a wolf-like M1/M2 ratio. The lower carnassial
Carnassials are paired upper and lower teeth modified in such a way as to allow enlarged and often self-sharpening edges to pass by each other in a shearing manner. This adaptation is found in carnivorans, where the carnassials are the modified f ...
(M1) is distinguished by main trigonid
The molars or molar teeth are large, flat teeth at the back of the mouth. They are more developed in mammals. They are used primarily to grind food during chewing. The name ''molar'' derives from Latin, ''molaris dens'', meaning "millstone tooth ...
cusps (protoconid
Many different terms have been proposed for features of the tooth crown in mammals.
The structures within the molars receive different names according to their position and morphology. This nomenclature was developed by Henry Fairfield Osborn i ...
and paraconid) relatively small (in comparison with ''C. lupus'') and by talonid cusps ( hypoconid and entoconid) linked by a sinuous cristid.
Describing ''C. etruscus'' as wolf-like and ''Canis arnensis'' as jackal-like is therefore an over-simplification, because ''C. arnensis'' is more similar to ''C. lupus'' than is ''C. etruscus'' in some cranial characters. ''C. etruscus'' shows a set of peculiar features.[
]
Range
Definitive specimens are only known from Mediterranean Europe
Southern Europe is also known as Mediterranean Europe, as its geography is marked by the Mediterranean Sea. Definitions of southern Europe include some or all of these countries and regions: Albania, Andorra, Bosnia and Herzegovina, Bulgaria, C ...
and Crimean peninsula
Crimea ( ) is a peninsula in Eastern Europe, on the northern coast of the Black Sea, almost entirely surrounded by the Black Sea and the smaller Sea of Azov. The Isthmus of Perekop connects the peninsula to Kherson Oblast in mainland Ukrai ...
during the Early Pleistocene
The Early Pleistocene is an unofficial epoch (geology), sub-epoch in the international geologic timescale in chronostratigraphy, representing the earliest division of the Pleistocene Epoch within the ongoing Quaternary Period. It is currently esti ...
, though there is one potential skull material identified as ''C.'' cf.
The abbreviation cf. (short for either Latin or , both meaning 'compare') is generally used in writing to refer the reader to other material to make a comparison with the topic being discussed. However some sources offer differing or even contr ...
''etruscus'' from the Yushe Basin of China.
Extinction
The Etruscan wolf and ''Canis arnensis'' both disappeared from the fossil record in Italy after the end of the Tasso Faunal Unit and were replaced by the mid-Pleistocene era Mosbach wolf (''C. mosbachensis'' Soergel, 1925) by 1.5 million years ago.[
]
References
{{Taxonbar, from=Q30682478
Extinct wolves
Pleistocene carnivorans
Pleistocene mammals of Europe
Prehistoric canines
Fossil taxa described in 1877