prehistoric
Prehistory, also known as pre-literary history, is the period of human history between the use of the first stone tools by hominins 3.3 million years ago and the beginning of recorded history with the invention of writing systems. The use o ...
barn-owl
Barn-owls (family Tytonidae) are one of the two families of owls, the other being the true owls or typical owls, Strigidae. They are medium to large owls with large heads and characteristic heart-shaped faces. They have long, strong legs with po ...
. It lived at what is now
Monte Gargano
Gargano (, Gargano Apulian Italo-Romance arˈgæːnə is a historical and geographical sub-region in the province of Foggia, Apulia, southeast Italy, consisting of a wide isolated mountain massif made of highland and several peaks and forming ...
in Italy, and was an island throughout much of the
Neogene
The Neogene ( ), informally Upper Tertiary or Late Tertiary, is a geologic period and system that spans 20.45 million years from the end of the Paleogene Period million years ago (Mya) to the beginning of the present Quaternary Period Mya. ...
when
sea level
Mean sea level (MSL, often shortened to sea level) is an average surface level of one or more among Earth's coastal bodies of water from which heights such as elevation may be measured. The global MSL is a type of vertical datuma standardise ...
s were higher. The owl's remains date back to the
Miocene
The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recent" ...
-
Pliocene
The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58barn owl
The barn owl (''Tyto alba'') is the most widely distributed species of owl in the world and one of the most widespread of all species of birds, being found almost everywhere except for the polar and desert regions, Asia north of the Himala ...
, giving a total length of about 50–65 cm for ''T. robusta''. This owl provides an interesting case study of
evolution
Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
and
insular gigantism
Island gigantism, or insular gigantism, is a biological phenomenon in which the size of an animal species isolated on an island increases dramatically in comparison to its mainland relatives. Island gigantism is one aspect of the more general Fos ...
.
Evolution, taxonomy and systematics
It was found
sympatric
In biology, two related species or populations are considered sympatric when they exist in the same geographic area and thus frequently encounter one another. An initially interbreeding population that splits into two or more distinct species s ...
ally with the gigantic relative, ''
Tyto gigantea
''Tyto gigantea'' is an extinct barn owl from what is now Gargano, Italy, dating back to the late Miocene. From its remains, ''T. gigantea'' is suggested to have been as large as or larger than the Eurasian eagle-owl (''Bubo bubo'').
This spec ...
'', which at some 220% of the size of the
barn owl
The barn owl (''Tyto alba'') is the most widely distributed species of owl in the world and one of the most widespread of all species of birds, being found almost everywhere except for the polar and desert regions, Asia north of the Himala ...
grew to some 70–85 cm in length—larger than a
Eurasian eagle-owl
The Eurasian eagle-owl (''Bubo bubo'') is a species of eagle-owl that resides in much of Palearctic, Eurasia. It is also called the Uhu and it is occasionally abbreviated to just the eagle-owl in Europe. It is one of the largest species of owl, ...
and maybe up to twice as heavy. Like other
apex predator
An apex predator, also known as a top predator, is a predator at the top of a food chain, without natural predators of its own.
Apex predators are usually defined in terms of trophic dynamics, meaning that they occupy the highest trophic le ...
s they can be presumed to have occurred at low population densities. Being apparently
endemic
Endemism is the state of a species being found 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 found els ...
to Gargano Island and perhaps some other islands in today's
Apennines
The Apennines or Apennine Mountains (; grc-gre, links=no, Ἀπέννινα ὄρη or Ἀπέννινον ὄρος; la, Appenninus or – a singular with plural meaning;''Apenninus'' (Greek or ) has the form of an adjective, which wou ...
, where one or two huge
eagle
Eagle is the common name for many large birds of prey of the family Accipitridae. Eagles belong to several groups of genera, some of which are closely related. Most of the 68 species of eagle are from Eurasia and Africa. Outside this area, j ...
s and at least at times a normal-sized barn-owl also occurred (though no large mammalian carnivores), these owls raise the question how
minimum viable population
Minimum viable population (MVP) is a lower bound on the population of a species, such that it can survive in the wild. This term is commonly used in the fields of biology, ecology, and conservation biology
Conservation biology is the study of ...
s could be maintained long enough for them to live and prosper.
It is extremely commonplace for larger
predator
Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill ...
y birds such as
Falconiformes
The order Falconiformes () is represented by the extant family Falconidae (falcons and caracaras) and a handful of enigmatic Paleogene species. Traditionally, the other bird of prey families Cathartidae (New World vultures and condors), Sagit ...
and Strigiformes to evolve differently-sized sexes. Namely, the females being considerably larger with little or no overlap in size between sexes. This allows them to hunt for different-sized prey, allowing a larger, more resilient population to live off a land. But this was apparently not the case here, as there is a reasonably large sample of bones of each of the 3 owl
taxa
In biology, a taxon ( back-formation from '' taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular n ...
to display the natural and sexual variation within each.
In the oldest fossil samples, no giant owls were found to accompany them, whereas the latest seemed to a range of owls attributed to all 3 species. The small owls supposedly were replaced by or evolved into ''T. robusta'' from some point onwards, and ''T. gigantea'' appeared later, supposedly alongside it as well as a recolonization by Continental European stock. That this was by no means a very parsimonious assumption was already noted in the original descriptions of the remains assigned to ''T. robusta'':
"If one assumes that the large barn-owls on Gargano would have arisen from multiple colonizations .. ''T. robusta'' would have to be separated into two distinct species. The medium-sized form of the first colonist cannot be the same as the middle-sized form of the second colonist. As there are no morphological findings to support this assumption, it was refrained from separating
he ''T. robusta'' material into
He or HE may refer to:
Language
* He (pronoun), an English pronoun
* He (kana), the romanization of the Japanese kana へ
* He (letter), the fifth letter of many Semitic alphabets
* He (Cyrillic), a letter of the Cyrillic script called ''He'' ...
two species. In how far one can actually expect morphological differences in forms so closely related is debatable."
More recently, it was noted that the smaller barn-owl of Gargano was not ''Tyto sanctialbani'', which inhabited Europe north of the
Alps
The Alps () ; german: Alpen ; it, Alpi ; rm, Alps ; sl, Alpe . are the highest and most extensive mountain range system that lies entirely in Europe, stretching approximately across seven Alpine countries (from west to east): France, ...
in the latter Miocene, but its
Mediterranean
The Mediterranean Sea is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean Basin and almost completely enclosed by land: on the north by Western and Southern Europe and Anatolia, on the south by North Africa, and on th ...
relative ''Tyto balearica'', which was a
sister species
In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree.
Definition
The expression is most easily illustrated by a cladogram:
Taxon A and ...
or
chronospecies
A chronospecies is a species derived from a sequential development pattern that involves continual and uniform changes from an extinct ancestral form on an evolutionary scale. The sequence of alterations eventually produces a population that is p ...
of or a southward divergence from ''T. sanctialbani'' and decidedly more robust. In addition, it appeared as if the ''Tyto'' of the youngest samples fell into 2, not 3 size groups.
Presumably, the island was initially colonized by ''T. balearica'' whose core range eventually extended from the Apennines westwards to
Iberia
The Iberian Peninsula (),
**
* Aragonese and Occitan: ''Peninsula Iberica''
**
**
* french: Péninsule Ibérique
* mwl, Península Eibérica
* eu, Iberiar penintsula also known as Iberia, is a peninsula
A peninsula (; ) is a la ...
; they might have become cut off from the main population when the Mediterranean reflooded at the end of the
Messinian salinity crisis
The Messinian salinity crisis (MSC), also referred to as the Messinian event, and in its latest stage as the Lago Mare event, was a geological event during which the Mediterranean Sea went into a cycle of partial or nearly complete desiccation (dr ...
6 million years ago. The entire population evolved into larger ''T. robusta'', which were still close enough in size to