Nesiotites

Nesiotites Temporal range: Early Pliocene–Holocene PreꞒ Ꞓ O S D C P T J K Pg N
Size comparison of the Balearic shrew Nesiotites hidalgo (top) with a water shrew (Neomys, below)
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Eulipotyphla
Family:	Soricidae
Tribe:	Nectogalini
Genus:	†Nesiotites Bate, 1945
Species
†N. hidalgo Bate, 1945 (type) †N. meloussae Pons and Moyà, 1980 †N. ponsi Reumer, 1979 †N. rafelinensis? Rofes et al, 2012

Nesiotites is an extinct genus of large red-toothed shrews belonging to the tribe Nectogalini that inhabited the Balearic Islands from the latest Miocene/Early Pliocene (from around 5 million years ago) up until the arrival of humans on the islands during the late Holocene (around 2500-2300 BC). It was present on Mallorca and Menorca. It represented one of only 3 native land mammals to the islands at the time of human arrival, alongside the goat-antelope Myotragus and the giant dormouse Hypnomys. The genus is closely related to the also recently extinct Corsican-Sardinian shrews belonging to the genus Asoriculus, with their closest living relatives being the Himalayan shrews of the genus Soriculus.

Nesiotites was originally described by Dorothea Bate in 1945, with the type species being Nesiotites hidalgo.^[1] Originally, two species from the islands of Corsica and Sardinia (N. corsicanus and N. similis) were included in the genus, but these are now rejected from the genus, and Nesiotites is now generally used exclusively for the Balearic species, as otherwise the genus would likely be polyphyletic.^[2]

The genus includes the chronospecies N. rafelinensis (earliest Pliocene)^[3] (the validity of this species disputed, as some authors contend that it is not morphologically distinct from N. ponsi^[4][5]) N. ponsi (Late Pliocene), N. meloussae/N. aff. ponsi (Early Pleistocene) and N. hidalgo (Middle Pleistocene-Holocene). These are largely distinguished by differences in body size and characters of the teeth.^[3][6]

Based on a mitochondrial genome from Nesiotites hidalgo, their closest living relative is the terrestrial Soriculus shrews to the Himalayas and surrounding areas, and related to other terrestrial nectogaline shrews known from Asia (Episoriculus and Chodsigoa), rather than to the nectogaline water shrews (Chimarrogale, Nectogale and Neomys). A molecular clock analysis suggests that Himalayan shrews and Balearic shrews genetically diverged approximately 6.44 million years ago. Based on morphological data, it is thought that Nesiotites is closely related and likely descended from the extinct genus Asoriculus, known from the Late Miocene-Holocene of Europe and North Africa, which now includes the Corsican and Sardinian species formerly included in Nesiotites.^[2]

Position of Nesiotites within Nectogalini based on DNA and morphological characters after Bover et al (2018):^[2]

Members of Nesiotites exhibited a large body size compared to their likely mainland ancestor, Asoriculus gibberodon (estimated to weigh 8.85 grams (0.312 oz)), as well to most other members of Nectogalini, an example of island gigantism. The species of the genus showed an increase in body size over time, with the estimated weight of N. ponsi being 14.58 grams (0.514 oz) while the last species, N. hildalgo being estimated at 26.63–29.31 grams (0.939–1.034 oz), being exceeded in size amongst Nectogalini only by Asiatic water shrews (Chimarrogale).^[7]

Nesiotites originally colonised Mallorca during the Messinian stage of the Late Miocene, when during the Messinian salinity crisis (5.96–5.33 million years ago) the Mediterranean sea evaporated allowing animals from the Iberian Peninsula to disperse to the Balearics. Later becoming isolated on the island when the Mediterranean refilled as result of the Zanclean Flood around 5.3 million years ago, at the beginning of the Pliocene. Nesiotites later spread to Menorca during the Pliocene-Pleistocene transition, when episodes of low sea level connected the two islands.^[8][6] During most of its existence, it represented only one of three terrestrial mammal lineages native to the Balearic islands, alongside the giant dormouse Hypnomys and the dwarf goat-antelope Myotragus. The last Nesiotites chronospecies, N. hidalgo, became extinct shortly after human settlement of the Balearics, which occurred sometime prior to 2282 BC, with the youngest radiocarbon date for the species dating to approximately 3027 BC.^[9]