Irish Elk
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The Irish elk (''Megaloceros giganteus''), also called the giant deer or Irish deer, is an extinct species of
deer A deer (: deer) or true deer is a hoofed ruminant ungulate of the family Cervidae (informally the deer family). Cervidae is divided into subfamilies Cervinae (which includes, among others, muntjac, elk (wapiti), red deer, and fallow deer) ...
in the genus '' Megaloceros'' and is one of the largest deer that ever lived. Its range extended across northern
Eurasia Eurasia ( , ) is a continental area on Earth, comprising all of Europe and Asia. According to some geographers, Physical geography, physiographically, Eurasia is a single supercontinent. The concept of Europe and Asia as distinct continents d ...
during the
Pleistocene The Pleistocene ( ; referred to colloquially as the ''ice age, Ice Age'') is the geological epoch (geology), epoch that lasted from to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fin ...
, from
Ireland Ireland (, ; ; Ulster Scots dialect, Ulster-Scots: ) is an island in the North Atlantic Ocean, in Northwestern Europe. Geopolitically, the island is divided between the Republic of Ireland (officially Names of the Irish state, named Irelan ...
(where it is known from abundant remains found in bogs) to
Lake Baikal Lake Baikal is a rift lake and the deepest lake in the world. It is situated in southern Siberia, Russia between the Federal subjects of Russia, federal subjects of Irkutsk Oblast, Irkutsk Oblasts of Russia, Oblast to the northwest and the Repu ...
in
Siberia Siberia ( ; , ) is an extensive geographical region comprising all of North Asia, from the Ural Mountains in the west to the Pacific Ocean in the east. It has formed a part of the sovereign territory of Russia and its predecessor states ...
. The most recent remains of the species have been radiocarbon dated to about 7,700 years ago in western Russia.Supplementary information
Erratum in
Its antlers, which can span across are the largest known of any deer. It is not closely related to either living species called the elk, with it being widely agreed that its closest living relatives are
fallow deer Fallow deer is the common name for species of deer in the genus ''Dama'' of subfamily Cervinae. There are two living species, the European fallow deer (''Dama dama''), native to Europe and Anatolia, and the Persian fallow deer (''Dama mesopotamic ...
(''Dama'').


Taxonomy


Research history

The first scientific descriptions of the animal's remains were made by Irish physician Thomas Molyneux in 1695, who identified large antlers from Dardistown—which were apparently commonly unearthed in Ireland—as belonging to the elk (known as the moose in North America), concluding that it was once abundant on the island. It was first formally named as ''Alce gigantea'' by
Johann Friedrich Blumenbach Johann Friedrich Blumenbach (11 May 1752 – 22 January 1840) was a German physician, naturalist, physiologist and anthropologist. He is considered to be a main founder of zoology and anthropology as comparative, scientific disciplines. He has be ...
in his ''Handbuch der Naturgeschichte'' in 1799,Blumenbach J. 1799.
Handbuch der Naturgeschichte
' (6th Ed.) 16: 697
with ''Alce'' being a variant of ''Alces'', the
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
name for the elk. The original Blumenbach's description of ''Alce gigantea'' provides rather scant information about the species, specifying only that this particular kind of "fossil elk" comes from Ireland and is characterized by immense body size. According to Blumenbach, the distance between summits of giant deer antlers may attain . This particular feature mentioned by Blumenbach permitted to Roman Croitor to identify the type specimen of giant deer that was figured and described for the first time in Louthiana of Thomas Wright. 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 ...
of ''Megaloceros giganteus'' (Blumenbach, 1799) is a well-preserved male skull with exceptionally large antlers found in Dunleer environs (
County Louth County Louth ( ; ) is a coastal Counties of Ireland, county in the Eastern and Midland Region of Republic of Ireland, Ireland, within the Provinces of Ireland, province of Leinster. Louth is bordered by the counties of County Meath, Meath to the ...
,
Ireland Ireland (, ; ; Ulster Scots dialect, Ulster-Scots: ) is an island in the North Atlantic Ocean, in Northwestern Europe. Geopolitically, the island is divided between the Republic of Ireland (officially Names of the Irish state, named Irelan ...
). The type specimen of giant deer is currently exposed in Barmeath Castle where Thomas Wright first saw and described it. French scientist
Georges Cuvier Jean Léopold Nicolas Frédéric, baron Cuvier (23 August 1769 – 13 May 1832), known as Georges Cuvier (; ), was a French natural history, naturalist and zoology, zoologist, sometimes referred to as the "founding father of paleontology". Cuv ...
documented in 1812 that the Irish elk did not belong to any species of mammal currently living, declaring it "''le plus célèbre de tous les ruminans fossiles''" (the most famous of all fossil ruminants). In 1827 Joshua Brookes, in a listing of his zoological collection, named the new genus ''Megaloceros'' (spelled ''Megalocerus'' in the earlier editions) in the following passage: The etymology being from
Greek Greek may refer to: Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group *Greek language, a branch of the Indo-European language family **Proto-Greek language, the assumed last common ancestor of all kno ...
: "great" + "horn, antler". The type and only species named in the description being ''Megaloceros antiquorum'', based on Irish remains now considered to belong to ''M. giganteus'', making the former a
junior synonym In taxonomy, the scientific classification of living organisms, a synonym is an alternative scientific name for the accepted scientific name of a taxon. The botanical and zoological codes of nomenclature treat the concept of synonymy differently. ...
. The original description was considered by Adrian Lister in 1987 to be inadequate for a taxonomic definition. In 1828 Brookes published an expanded list in the form of a catalogue for an upcoming auction, which included the Latin phrase "''Cornibus deciduis palmatis"'' (meaning "deciduous palmate antlers") as a description of the remains. The 1828 publication was approved by
International Commission on Zoological Nomenclature The International Commission on Zoological Nomenclature (ICZN) is an organization dedicated to "achieving stability and sense in the scientific naming of animals". Founded in 1895, it currently comprises 26 commissioners from 20 countries. Orga ...
(ICZN) in 1977 as an available publication for the basis of zoological nomenclature. Adrian Lister in 1987 judged that "the phase "''Cornibus deciduis palmatis"'' constitutes a definition sufficient under the /nowiki>International Code of Zoological Nomenclature">International Code of Zoological Nomenclature">/nowiki>International Code of Zoological Nomenclature/nowiki> (article 12) to validate ''Megalocerus''." The original spelling of ''Megalocerus'' was never used after its original publication.In 1844, Richard Owen named another synonym of the Irish elk, including it within the newly named subgenus ''Megaceros'', ''Cervus'' (''Megaceros'') ''hibernicus''. This has been suggested to be derived from another junior synonym of the Irish elk described by J. Hart in 1825, ''Cervus megaceros''. Despite being a junior synonym, ''Megaloceros'' remained in obscurity and ''Megaceros'' became the common genus name for the taxon. The combination "''Megaceros giganteus"'' was in use by 1871.
George Gaylord Simpson George Gaylord Simpson (June 16, 1902 – October 6, 1984) was an American paleontologist. Simpson was perhaps the most influential paleontologist of the twentieth century, and a major participant in the modern synthesis, contributing '' Tempo ...
in 1945 revived the original ''Megaloceros'' name, which became progressively more widely used, until a taxonomic decision in 1989 by the ICZN confirmed the priority of ''Megaloceros'' over ''Megaceros'', and ''Megaloceros'' to be the correct spelling.Lister, A M, 198
''Megaloceros'' Brookes 1828 Mammalia Artiodactyla Proposed Emendation Of The Original Spelling
''The Bulletin of zoological nomenclature''. 44 255–256
Before the 20th century, the Irish elk, having evolved from smaller ancestors with smaller antlers, was taken as a prime example of
orthogenesis Orthogenesis, also known as orthogenetic evolution, progressive evolution, evolutionary progress, or progressionism, is an Superseded theories in science, obsolete biological hypothesis that organisms have an innate tendency to evolution, evolve ...
(directed evolution), an evolutionary mechanism opposed to Darwinian evolution in which the successive species within the lineage become increasingly modified in a single undeviating direction, evolution proceeding in a straight line void of
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the Heredity, heritable traits characteristic of a population over generation ...
. Orthogenesis was claimed to have caused an evolutionary trajectory towards antlers that became larger and larger, eventually causing the species' extinction because the antlers grew to sizes which inhibited proper feeding habits and caused the animal to become trapped in tree branches. In the 1930s, orthogenesis was disputed by Darwinians led by
Julian Huxley Sir Julian Sorell Huxley (22 June 1887 – 14 February 1975) was an English evolutionary biologist, eugenicist and Internationalism (politics), internationalist. He was a proponent of natural selection, and a leading figure in the mid-twentiet ...
, who noted that antler size was not grossly large, and was proportional to body size.Zimmer, Carl
"The Allure of Big Antlers"
The Loom. Discover, ''National Geographic''. 3 September 2008. Web. 23 October 2014.
The currently favoured view is that
sexual selection Sexual selection is a mechanism of evolution in which members of one sex mate choice, choose mates of the other sex to mating, mate with (intersexual selection), and compete with members of the same sex for access to members of the opposite sex ...
was the driving force behind the large antlers rather than orthogenesis or natural selection.


Evolution

''M. giganteus'' belongs to the genus '' Megaloceros.'' ''Megaloceros'' has often been placed into the tribe Megacerini, alongside other genera often collectively referred to as "giant deer", like ''
Sinomegaceros ''Sinomegaceros'' is an extinct genus of deer known from the Late Pliocene/Early Pleistocene to Late Pleistocene of Central and East Asia. It is considered to be part of the group of "giant deer" (often referred to collectively as members of the ...
'' and '' Praemegaceros''.'''' The taxonomy of giant deer lacks consensus, with genus names used for species varying substantially between authors. The earliest possible record of the genus is a partial antler from the Early Pleistocene MN 17 (2.5–1.8 Ma) of Stavropol Krai in the
North Caucasus The North Caucasus, or Ciscaucasia, is a subregion in Eastern Europe governed by Russia. It constitutes the northern part of the wider Caucasus region, which separates Europe and Asia. The North Caucasus is bordered by the Sea of Azov and the B ...
of
Russia Russia, or the Russian Federation, is a country spanning Eastern Europe and North Asia. It is the list of countries and dependencies by area, largest country in the world, and extends across Time in Russia, eleven time zones, sharing Borders ...
, which were given the name of ''M. stavropolensis'' in 2016, however this species has been subsequently suggested to belong to '' Arvernoceros''. or ''Sinomegaceros''. The oldest generally accepted records of the genus are from the late
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 ...
. Other species often considered to belong to ''Megaloceros'' include the
reindeer The reindeer or caribou (''Rangifer tarandus'') is a species of deer with circumpolar distribution, native to Arctic, subarctic, tundra, taiga, boreal, and mountainous regions of Northern Europe, Siberia, and North America. It is the only re ...
sized ''M. savini'', which is known from early Middle Pleistocene (~700,000–450,000 years ago) localities in England, France, Spain and Germany, and the more recently described species ''M. novocarthaginiensis'', which is known from late Early Pleistocene (0.9–0.8 Ma) localities in Spain, and the small ''M. matritensis'' endemic to the Iberian peninsula during the late Middle Pleistocene (~400,000 to 250,000 years ago), which overlaps chronologically with the earliest ''M. giganteus'' records. Jan van der Made proposed ''M. novocarthaginiensis , M. savini'' and ''M. matritensis'' to be sequential
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 ...
, due to shared morphological characteristics not found in ''M. giganteus'' and gradual transition of morphological characters through time. ''M. savini'' and related species have also been suggested to comprise the separate genus '' Praedama'' by other authors. While the ''M. savini/Praedama'' lineage is often suggested to be closely related ''to M. giganteus,'' most authors agree that this group of deer is unlikely to be directly ancestral to ''M. giganteus''. The origin of ''M. giganteus'' remains unclear, and appears to lie outside Western Europe. Jan van der Made has suggested that remains of an indeterminate ''Megaloceros'' species from the late Early Pleistocene (~1.2 Ma) of Libakos in Greece are closer to ''M. giganteus'' than the ''M. novocarthaginiensis-savini-matritensis'' lineage due to the shared molarisation of the lower fourth
premolar The premolars, also called premolar Tooth (human), teeth, or bicuspids, are transitional teeth located between the Canine tooth, canine and Molar (tooth), molar teeth. In humans, there are two premolars per dental terminology#Quadrant, quadrant in ...
(P4). Croitor has suggested that ''M. giganteus'' is closely related to what was originally described as ''Dama clactoniana mugharensis'' (which he proposes be named ''Megaloceros mugharensis'') from the Middle Pleistocene of Tabun Cave in Israel, due to similarities in the antlers, molars and premolars. The earliest possible records of ''M. giganteus'' comes from Homersfield, England thought to be about 450,000 years ago—though the dating is uncertain. The oldest securely dated Middle Pleistocene records are those from Hoxne, England, which have been dated to
Marine Isotope Stage 11 Marine Isotope Stage 11 or MIS 11 is a Marine Isotope Stage in the geologic temperature record, covering the interglacial period between 424,000 and 374,000 years ago. It corresponds to the Hoxnian Stage in Britain. Interglacial periods which o ...
(424,000 to 374,000 years ago), other Middle Pleistocene early records include Steinheim an der Murr, Germany, (classified as ''M. g. antecedens'') about 400,000–300,000 years ago and
Swanscombe Swanscombe /ˈswɔnzkəm/ is a town in the Borough of Dartford in Kent, England, and the civil parish of Swanscombe and Greenhithe. It is 4.4 miles west of Gravesend and 4.8 miles east of Dartford. History Prehistory Bone fragments and to ...
, England. Most remains of the Irish elk are known from the
Late Pleistocene The Late Pleistocene is an unofficial Age (geology), age in the international geologic timescale in chronostratigraphy, also known as the Upper Pleistocene from a Stratigraphy, stratigraphic perspective. It is intended to be the fourth division ...
. A large proportion of the known remains of ''M. giganteus'' are from Ireland, which mostly date to the
Allerød oscillation Allerød may refer to: * Allerød Municipality, a municipality in Denmark ** Lillerød, also called ''Allerød'', seat of the municipality ** Allerød station, a railway station in the Danish town * Allerød oscillation, a climatic period at the en ...
near the end of the Late Pleistocene around 13,000 years ago. Over 100 individuals have been found in Ballybetagh Bog near Dublin. Some authors have proposed that Late Pleistocene ''M. giganteus'' should be divided into several subspecies including ''M. giganteus ruffii'' and ''M. giganteus giganteus,'' based primarily on differences in antler morphology''.'' It has been historically thought that, because both have palmated antlers, the Irish elk and
fallow deer Fallow deer is the common name for species of deer in the genus ''Dama'' of subfamily Cervinae. There are two living species, the European fallow deer (''Dama dama''), native to Europe and Anatolia, and the Persian fallow deer (''Dama mesopotamic ...
(''Dama'' spp.) are closely related, this is supported by several other morphological similarities, including the lack of upper canines, proportionally long braincase and nasal bones, and proportionally short front portion of the skull. In 2005, two fragments of
mitochondrial DNA Mitochondrial DNA (mtDNA and mDNA) is the DNA located in the mitochondrion, mitochondria organelles in a eukaryotic cell that converts chemical energy from food into adenosine triphosphate (ATP). Mitochondrial DNA is a small portion of the D ...
(mtDNA) from the
cytochrome b Cytochrome b is a protein found in the membranes of aerobic cells. In eukaryotic mitochondria (inner membrane) and in aerobic prokaryotes, cytochrome b is a component of respiratory chain complex III () — also known as the bc1 complex or ubiq ...
gene were extracted and sequenced from 4 antlers and a bone, the mtDNA found that the Irish elk was nested within ''
Cervus ''Cervus'' is a genus of deer that primarily are native to Eurasia, although one species occurs in northern Africa and another in North America. In addition to the species presently placed in this genus, it has included a whole range of other s ...
,'' and were inside the
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
containing living
red deer The red deer (''Cervus elaphus'') is one of the largest deer species. A male red deer is called a stag or Hart (deer), hart, and a female is called a doe or hind. The red deer inhabits most of Europe, the Caucasus Mountains region, Anatolia, Ir ...
(''Cervus elaphus''). Based on this, the authors suggested that the Irish elk and red deer interbred. However, another study from the same year in the journal ''
Nature Nature is an inherent character or constitution, particularly of the Ecosphere (planetary), ecosphere or the universe as a whole. In this general sense nature refers to the Scientific law, laws, elements and phenomenon, phenomena of the physic ...
'' utilising both fragmentary mitochondrial DNA and morphological data found that the Irish elk was indeed most closely related to ''Dama''. The close relationship with ''Dama'' was supported by another cytochrome b study in 2006, a 2015 study involving the full mitochondrial genome, and by a 2017 morphological analysis of the
bony labyrinth The bony labyrinth (also osseous labyrinth or otic capsule) is the rigid, bony outer wall of the inner ear in the temporal bone. It consists of three parts: the vestibule, semicircular canals, and cochlea. These are cavities hollowed out of the ...
. The 2006 and 2017 studies also directly suggest that the results of the 2005 cytochrome b paper were the result of DNA contamination. Cladogram of Cervidae based on mitochondrial DNA: A study of mitochondrial genomes from ''Sinomegaceros'' from the Late Pleistocene of East Asia found that the mitochondrial genomes of ''Megaloceros giganteus'' were nested within those of ''Sinomegaceros,'' suggesting that the two lineages interbred after their initial split. Cladogram of ''Megaloceros'' and ''Sinomegaceros'' mitochondrial genomes following Xiao et al. 2023.


Description

The Irish elk stood about tall at the shoulders, and had large palmate (flat and broad)
antler Antlers are extensions of an animal's skull found in members of the Cervidae (deer) Family (biology), family. Antlers are a single structure composed of bone, cartilage, fibrous tissue, skin, nerves, and blood vessels. They are generally fo ...
s, the largest of any known deer, with the largest specimens reaching over from tip to tip (though it is rare for specimens to exceed across) and in weight. The antlers are considerably larger than those of living moose, being on average over twice the volume of moose antlers. For body size, at about and up to or more, the Irish elk was the heaviest known cervine ("Old World deer"); and tied with the extant Alaska moose (''Alces alces gigas'') as the third largest known deer, after the extinct '' Cervalces latifrons'' and '' Cervalces scotti''.R. D. E. Mc Phee, ''Extinctions in Near Time: Causes, Contexts, and Consequences'' p. 262 The shape and span of the antlers varied significantly over time and space, likely reflecting some populations adaptation to forested environments. Compared to ''Alces'', Irish elk appear to have had a more robust skeleton, with older and more mature ''Alces'' skeletons bearing some resemblance to those of prime Irish elk, and younger Irish elk resembling prime ''Alces''. Likely due to different social structures, the Irish elk exhibits more marked
sexual dimorphism Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
than ''Alces'', with Irish elk bucks being notably larger than does. In total, Irish elk bucks may have ranged from , with an average of , and does may have been relatively large, about 80% of buck size, or on average. The distinguishing characters of ''M. giganteus'' include concave frontals, proportionally long braincase, proportionally short front section of the skull (orbitofrontal region), alongside the absence of upper canines and the molarisation of the lower fourth premolar (P4). The skull and mandible of the Irish elk exhibit substantial thickening (
pachyostosis Pachyostosis is a non-pathological condition in vertebrate animals in which the bones experience a thickening, generally caused by extra layers of lamellar bone. It often occurs together with bone densification ( osteosclerosis), reducing inner c ...
), with the early and complete obliteration of
cranial sutures In anatomy, fibrous joints are joints connected by Fibrous connective tissue, fibrous tissue, consisting mainly of collagen. These are fixed joints where bones are united by a layer of white fibrous tissue of varying thickness. In the skull, the ...
. Based on Upper Palaeolithic cave paintings, the Irish elk seems to have had overall light colouration, with a dark stripe running along the back, a stripe on either side from shoulder to haunch, a dark collar on the throat and a chinstrap, and a dark hump on the
withers Withers are the ridge between the shoulder blades of an animal, typically a quadruped. In many species, this ridge is the tallest point of the body. In horses and dogs, it is the standard place to measure the animal's height. In contrast, catt ...
(between the
shoulder blade The scapula (: scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on either side ...
s). In 1989, American palaeontologist Dale Guthrie suggested that, like
bison A bison (: bison) is a large bovine in the genus ''Bison'' (from Greek, meaning 'wild ox') within the tribe Bovini. Two extant taxon, extant and numerous extinction, extinct species are recognised. Of the two surviving species, the American ...
, the hump allowed a higher hinging action of the front legs to increase stride length while running. Valerius Geist suggested that the hump may have also been used to store fat. Localising fat rather than evenly distributing it may have prevented overheating while running or in rut during the summer.


Habitat

The Irish elk had a far-reaching range, extending from the Atlantic Ocean in the West to
Lake Baikal Lake Baikal is a rift lake and the deepest lake in the world. It is situated in southern Siberia, Russia between the Federal subjects of Russia, federal subjects of Irkutsk Oblast, Irkutsk Oblasts of Russia, Oblast to the northwest and the Repu ...
in the East. Irish elk do not appear have extended northward onto the open
mammoth steppe The mammoth steppe, also known as steppe-tundra, was once the Earth's most extensive biome. During glacial periods in the later Pleistocene, it stretched east-to-west, from the Iberian Peninsula in the west of Europe, then across Eurasia and thr ...
in Siberia, rather keeping to the boreal steppe-woodland environments, which consisted of scattered
spruce A spruce is a tree of the genus ''Picea'' ( ), a genus of about 40 species of coniferous evergreen trees in the family Pinaceae, found in the northern temperate and boreal ecosystem, boreal (taiga) regions of the Northern hemisphere. ''Picea'' ...
and
pine A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. ''World Flora Online'' accepts 134 species-rank taxa (119 species and 15 nothospecies) of pines as cu ...
, as well as low-lying herbs and shrubs including grasses,
sedges The Cyperaceae () are a family of graminoid (grass-like), monocotyledonous flowering plants known as sedges. The family is large; botanists have described some 5,500 known species in about 90 generathe largest being the "true sedges" (genu ...
, '' Ephedra, Artemisia'' and
Chenopodiaceae Amaranthaceae ( ) is a family of flowering plants commonly known as the amaranth family, in reference to its type (biology), type genus ''Amaranthus''. It includes the former goosefoot family Chenopodiaceae and contains about 165 genera and 2,040 ...
. The species appears to have had a degree of ecological plasticity, as during
interglacial An interglacial period (or alternatively interglacial, interglaciation) is a geological interval of warmer global average temperature lasting thousands of years that separates consecutive glacial periods within an ice age. The current Holocene i ...
periods prior to the Holocene, the species was present in temperate forested environments in Europe. During these times, the species generally had less broad antlers than during glacial periods, likely as an adaptation to moving through forested environments.


Palaeobiology


Physiology

In 1998, Canadian biologist Valerius Geist hypothesised that the Irish elk was
cursorial A cursorial organism is one that is adapted specifically to run. An animal can be considered cursorial if it has the ability to run fast (e.g. cheetah) or if it can keep a constant speed for a long distance (high endurance). "Cursorial" is often ...
(adapted for running and stamina). He noted that the Irish elk physically resembled
reindeer The reindeer or caribou (''Rangifer tarandus'') is a species of deer with circumpolar distribution, native to Arctic, subarctic, tundra, taiga, boreal, and mountainous regions of Northern Europe, Siberia, and North America. It is the only re ...
. The body proportions of the Irish elk are similar to those of the cursorial
addax The addax (''Addax nasomaculatus''), also known as the white antelope and the screwhorn antelope, is an antelope native to the Sahara Desert. The only member of the genus ''Addax'', it was first described scientifically by Henri de Blainvil ...
,
oryx ''Oryx'' ( ) is a genus consisting of four large antelope species called oryxes. Their pelage is pale with contrasting dark markings in the face and on the legs, and their long horns are almost straight and annulated. The exception is the sci ...
, and saiga antelope. These include the relatively short legs, the long front legs nearly as long as the hind legs, and a robust cylindrical body. Cursorial saiga,
gnu GNU ( ) is an extensive collection of free software (394 packages ), which can be used as an operating system or can be used in parts with other operating systems. The use of the completed GNU tools led to the family of operating systems popu ...
s, and reindeer have a top speed of over , and can maintain high speeds for up to 15 minutes.


Reproduction

At Ballybetagh Bog, over 100 Irish elk individuals were found, all small antlered bucks. This indicates that bucks and does segregated during at least winter and spring. Many modern deer species do this partly because males and females have different nutritional requirements and need to consume different types of plants. Segregation would also imply a
polygynous Polygyny () is a form of polygamy entailing the marriage of a man to several women. The term polygyny is from Neoclassical Greek πολυγυνία (); . Incidence Polygyny is more widespread in Africa than in any other continent. Some scholar ...
society, with stags fighting for control over harems during rut. Because most of the individuals found were juvenile or geriatric and were likely suffering from malnutrition, they probably died from winterkill. Most Irish elk specimens known may have died from winterkill, and winterkill is the highest source of mortality among many modern deer species. Bucks generally suffer higher mortality rates because they eat little during the autumn rut. For rut, a lean stag normally may have fattened up to , and would burn through the extra fat over the next month. Assuming a similar response to starvation as red deer, a large, healthy Irish elk stag with antlers would have had antlers under poor conditions; and an average sized Irish elk stag with antlers would have had antlers under poorer conditions, similar sizes to the moose. A similar change in a typical Irish elk population with prime stags having antlers would result in antler weights of or less in worsening climatic conditions. This is within the range of present-day wapiti/red deer (''Cervus'' spp.) antler weights. Irish elk antlers vary widely in form depending upon the habitat, such as a compact, upright shape in closed forest environments. Irish elk likely shed their antlers and re-grew a new pair during mating season. Antlers generally require high amounts of
calcium Calcium is a chemical element; it has symbol Ca and atomic number 20. As an alkaline earth metal, calcium is a reactive metal that forms a dark oxide-nitride layer when exposed to air. Its physical and chemical properties are most similar to it ...
and
phosphate Phosphates are the naturally occurring form of the element phosphorus. In chemistry, a phosphate is an anion, salt, functional group or ester derived from a phosphoric acid. It most commonly means orthophosphate, a derivative of orthop ...
, especially those for stags which have larger structures, and the massive antlers of Irish elk may have required much greater quantities. Stags typically meet these requirements in part from their bones, suffering from a condition similar to
osteoporosis Osteoporosis is a systemic skeletal disorder characterized by low bone mass, micro-architectural deterioration of bone tissue leading to more porous bone, and consequent increase in Bone fracture, fracture risk. It is the most common reason f ...
while the antlers are growing, and replenishing them from food plants after the antlers have grown in or reclaiming nutrients from shed antlers. The large antlers have generally been explained as being used for male-male battle during mating season. They may have also been used for display, to attract females and assert dominance against rival males. A
finite element analysis Finite element method (FEM) is a popular method for numerically solving differential equations arising in engineering and mathematical models, mathematical modeling. Typical problem areas of interest include the traditional fields of structural ...
of the antlers suggested that during fighting, the antlers were likely to interlock around the middle tine, the high stress when interlocking on the distal tine suggests that the fighting was likely more constrained and predictable than among extant deer, likely involving twisting motions, as is known in extant deer with palmated antlers. In deer, gestation time generally increases with body size. A doe may have had a gestation period of about 274 days. Based on this and patterns seen in modern deer, last year's antlers in Irish elk bucks were potentially shed in early March, peak antler growth in early June, completion by mid-July, shedding
velvet Velvet is a type of woven fabric with a dense, even pile (textile), pile that gives it a distinctive soft feel. Historically, velvet was typically made from silk. Modern velvet can be made from silk, linen, cotton, wool, synthetic fibers, silk ...
(a layer of blood vessels on the antlers in-use while growing them) by late July, and the height of rut falling on the second week of August. Geist, believing the Irish elk to have been a cursorial animal, concluded that a doe would have to have produced nutrient-rich milk so that her calf would have enough energy and stamina to keep up with the herd.


Diet and life history

The mesodont (meaning neither high ( hypsodont) or low ( brachydont) crowned) condition of the teeth suggests that the species was a mixed feeder, being able to both browse and graze. Pollen remains from teeth found in the North Sea around 43,000 years old were found to be dominated by ''Artemisia'' and other
Asteraceae Asteraceae () is a large family (biology), family of flowering plants that consists of over 32,000 known species in over 1,900 genera within the Order (biology), order Asterales. The number of species in Asteraceae is rivaled only by the Orchi ...
, with minor ''
Plantago ''Plantago'' is a genus of about 200 species of flowering plants in the family Plantaginaceae, commonly called plantains or fleaworts. The common name plantain is shared with the unrelated cooking plantain. Most are herbaceous plants, though a ...
'', ''
Helianthemum ''Helianthemum'' (), known as rock rose, sunrose, rushrose, or frostweed,
'',
Plumbaginaceae Plumbaginaceae is a family (biology), family of flowering plants, with a cosmopolitan distribution. The family is sometimes referred to as the leadwort family or the plumbago family. Most species in this family are perennial plant, perennial h ...
and
willow Willows, also called sallows and osiers, of the genus ''Salix'', comprise around 350 species (plus numerous hybrids) of typically deciduous trees and shrubs, found primarily on moist soils in cold and temperate regions. Most species are known ...
(''Salix''). Another earlier specimen from the Netherlands (dating to the
Eemian The Last Interglacial, also known as the Eemian, was the interglacial period which began about 130,000 years ago at the end of the Penultimate Glacial Period and ended about 115,000 years ago at the beginning of the Last Glacial Period. It cor ...
interglacial or early in the Last Glacial Period) was found to have pollen of
Apiaceae Apiaceae () or Umbelliferae is a family of mostly aromatic flowering plants named after the type genus ''Apium,'' and commonly known as the celery, carrot, or parsley family, or simply as umbellifers. It is the 16th-largest family of flowering p ...
, including cow parsley (''Anthriscus sylvestris''), cow parsnip/hogweed ('' Heracleum''), water pennywort (''Hydrocotyle''), Asteraeceae, ''
Filipendula ''Filipendula'' is a genus of 12 species of perennial plant, perennial herbaceous flowering plants in the family Rosaceae, native to the temperate regions of the Northern Hemisphere. Well-known species include meadowsweet (''Filipendula ulmaria' ...
'', '' Symphytum'' and grass embedded with its teeth. A stable isotope analysis of the terminal Pleistocene Irish population suggests a grass and
forb A forb or phorb is a herbaceous flowering plant that is not a graminoid (grass, sedge, or rush). The term is used in botany and in vegetation ecology especially in relation to grasslands and understory. Typically, these are eudicots without woo ...
based diet, supplemented by browsing during stressed periods. Dental wear patterns of specimens from the late Middle and Late Pleistocene of Britain suggest a diet tending towards mixed feeding and grazing, but with a wide range including leaf browsing. Comparisons of ''δ''15N between Irish elk and red deer at the Middle Pleistocene site of Schöningen in Germany suggest that grasses were a more important component of the former's diet relative to the latter. Examination of histological sections of their long bones suggests that the species has relatively rapid growth rates, reaching skeletal maturity by around 6 years of age. Analysis of the
cementum Cementum is a specialized calcified substance covering the root of a tooth. The cementum is the part of the periodontium that attaches the teeth to the alveolar bone by anchoring the periodontal ligament. Structure The cells of cementum are ...
layers of their teeth suggests that Irish elk reached a maximum lifespan of at least 19 years, comparable to moose. Based on the dietary requirements of red deer, a lean Irish elk stag would have needed to consume of fresh forage daily. Assuming antler growth occurred over a span of 120 days, a stag would have required 1,372 g (3 lb) of protein daily, as well as access to nutrient- and mineral-dense forage starting about a month before antlers began sprouting and continuing until they had fully grown. Such forage is not very common, and stags perhaps sought after aquatic plants in lakes. After antler growing, stags could probably satisfy their nutritional requirements in productive sedge lands bordered by willow and birch forests. Gnaw marks on found on Irish elk bones indicates that they were preyed on or scavenged by cave hyenas.


Relationship with early humans

At a number of
Middle Paleolithic The Middle Paleolithic (or Middle Palaeolithic) is the second subdivision of the Paleolithic or Old Stone Age as it is understood in Europe, Africa and Asia. The term Middle Stone Age is used as an equivalent or a synonym for the Middle P ...
sites, remains of ''M. giganteus'' have been found with cut marks indicating butchery by
Neanderthals Neanderthals ( ; ''Homo neanderthalensis'' or sometimes ''H. sapiens neanderthalensis'') are an extinction, extinct group of archaic humans who inhabited Europe and Western and Central Asia during the Middle Pleistocene, Middle to Late Plei ...
. These include Bolomor Cave in Spain, dating to around 180,000 years ago, and De Nadale Cave and Riparo del Broion in northern Italy, dating to 71-69,000 and 50-44,000 years ago, respectively. Other sites probably resulting from exploitation of Irish elk by Neanderthals include Abri du Maras in southeast France, dating to 55-40,000 years ago. A mandible from Ofatinţi, Moldova dating to either the Eemian or the early Late Pleistocene, has been noted for having "tool-made notches on its lateral side". A handful of Irish elk depictions are known from the
art of the Upper Paleolithic The art of the Upper Paleolithic represents the oldest form of prehistoric art. Figurative art is present in prehistoric Europe, Europe and Prehistoric Indonesia, Southeast Asia, beginning around 50,000 years ago. Non-figurative cave paintings, c ...
in Europe. However, these are much less abundant than the common red deer and reindeer depictions. Only a handful of examples of modern human interaction are known. Several ''M. giganteus'' bones from the Chatelperronian levels of the Labeko Koba site in Spain are noted for bearing puncture marks, which have been interpreted as anthropogenic. A terminal Pleistocene (13,710-13,215 cal BP) skull from
Lüdersdorf Lüdersdorf is a municipality in the Nordwestmecklenburg district, in Mecklenburg-Vorpommern, Germany. It is close to the cities of Lübeck, Wismar and Schwerin Schwerin (; Mecklenburgisch-Vorpommersch dialect, Mecklenburgisch-Vorpommersch Low ...
, Germany is noted to have had the antler and facial part of the skull deliberately removed. A
calcaneum In humans and many other primates, the calcaneus (; from the Latin ''calcaneus'' or ''calcaneum'', meaning heel; : calcanei or calcanea) or heel bone is a bone of the tarsus of the foot which constitutes the heel. In some other animals, it is t ...
from an associated lower hind limb from the early Holocene site of Sosnovy Tushamsky in Siberia is noted to have "two short and deep traces of cutting blows", which are interpreted as "clear evidence of butchery". The use of shed antler bases is also known, at the terminal Pleistocene ( Allerød) Endingen VI site in Germany, a shed antler base appears to have been used in a way analogous to a
lithic core In archaeology, a lithic core is a distinctive Artifact (archaeology), artifact that results from the practice of lithic reduction. In this sense, a core is the scarred nucleus resulting from the detachment of one or more lithic flake, flakes fr ...
to produce " blanks" for the manufacture of barbed projectile tips. A ring-like mark on a shed antler beam from the similarly aged
Paderborn Paderborn (; Westphalian language, Westphalian: ''Patterbuorn'', also ''Paterboärn'') is a city in eastern North Rhine-Westphalia, Germany, capital of the Paderborn (district), Paderborn district. The name of the city derives from the river Pade ...
site in Germany has been suggested to be anthropogenic.


Extinction

Outside of the Irish Late Pleistocene, remains of Irish elk are uncommon, suggesting that they were usually rare in the areas where they did occur. Historically, its extinction has been attributed to the encumbering size of the antlers, a "
maladaptation In evolution, a maladaptation ( /ˌmælædæpˈteɪʃən/) is a trait that is (or has become) more harmful than helpful, in contrast with an adaptation, which is more helpful than harmful. All organisms, from bacteria to humans, display maladapt ...
" making fleeing through forests especially difficult for males while being chased by human hunters, or being too taxing nutritionally when the vegetation makeup shifted. In these scenarios,
sexual selection Sexual selection is a mechanism of evolution in which members of one sex mate choice, choose mates of the other sex to mating, mate with (intersexual selection), and compete with members of the same sex for access to members of the opposite sex ...
by does for stags with large antlers would have contributed to decline. However, antler size decreased through the Late Pleistocene and into the Holocene, and so may not have been the primary cause of extinction. A reduction in forest density in the Late Pleistocene and a lack of sufficient high-quality forage is associated with a decrease in body and antler size. Such resource constriction may have cut female fertility rates in half. Human hunting may have forced Irish elk into suboptimal feeding grounds. The distribution of ''M. giganteus'' is thought to have been strongly controlled by climatic conditions. The range of the Irish elk appears to have collapsed during the
Last Glacial Maximum The Last Glacial Maximum (LGM), also referred to as the Last Glacial Coldest Period, was the most recent time during the Last Glacial Period where ice sheets were at their greatest extent between 26,000 and 20,000 years ago. Ice sheets covered m ...
(LGM), with few remains known between 27,500 and 14,600 years ago, and none between 23,300 and 17,500 years ago. Known remains substantially increase during the latest Pleistocene Bølling–Allerød Interstadial, where it appears to have re-colonized northern Europe, with abundant remains in the UK, Ireland, and Denmark, though its range contracted again during the following
Younger Dryas The Younger Dryas (YD, Greenland Stadial GS-1) was a period in Earth's geologic history that occurred circa 12,900 to 11,700 years Before Present (BP). It is primarily known for the sudden or "abrupt" cooling in the Northern Hemisphere, when the ...
, disappearing from northern Europe by the end of the period. A 2021 study found that ''M. giganteus'' saw a progressive decline in mitochondrial genome diversity beginning around 50,000 years ago, which accelerated during the LGM. By the early Holocene, the range of the species had been dramatically reduced, with the youngest records in the eastern part of its range near Lake Baikal dating to around 10,700–10,400 years
Before Present Before Present (BP) or "years before present (YBP)" is a time scale used mainly in archaeology, geology, and other scientific disciplines to specify when events occurred relative to the origin of practical radiocarbon dating in the 1950s. Because ...
(BP), surviving latest in central part of its range within
European Russia European Russia is the western and most populated part of the Russia, Russian Federation. It is geographically situated in Europe, as opposed to the country's sparsely populated and vastly larger eastern part, Siberia, which is situated in Asia ...
and
Western Siberia Western Siberia or West Siberia ( rus, Западная Сибирь, p=ˈzapədnəjə sʲɪˈbʲirʲ; , ) is a region in North Asia. It is part of the wider region of Siberia that is mostly located in the Russia, Russian Federation, with a Sout ...
. It is suggested that extinction was contributed to by further climatic changes transforming preferred open habitat into uninhabitable dense forest. The final demise may have been caused by several factors both on a continental and regional scale, including climate change and hunting. The youngest dates in this region from Kamyshlov in Western Siberia and Maloarkhangelsk, Oryol Oblast In European Russia date to around 7,700-7,600 years ago, and it is suggested that it likely became extinct shortly after this time. Lister and Stewart concluded in a study of the extinction of the Irish elk that "it seems clear that environmental factors, cumulatively over thousands of years, reduced giant deer populations to a highly vulnerable state. In this situation, even relatively low-level hunting by small human populations could have contributed to its extinction."


Modern significance

Due to the abundance of Irish elk remains in Ireland, a thriving trade in their bones existed there during the 19th century to supply museums and collectors. Skeletons and skulls with attached antlers were also prized ornaments in aristocratic homes. The remains of Irish elk were of high value: "In 1865, full skeletons might fetch £30, while particularly good heads with antlers could cost £15." with £15 being more than 30 weeks' wages for a low skilled worker at the time. Indeed Leeds Philosophical and Literary Society bought a full skeleton in 1847, from Glennon's in Dublin, for £38. This specimen, discovered at Lough Gur near
Limerick Limerick ( ; ) is a city in western Ireland, in County Limerick. It is in the Provinces of Ireland, province of Munster and is in the Mid-West Region, Ireland, Mid-West which comprises part of the Southern Region, Ireland, Southern Region. W ...
, is still on display at
Leeds City Museum Leeds City Museum, established in 1819, is a museum in Leeds, West Yorkshire, England. Since 2008 it has been housed in the former Mechanics' institute, Mechanics' Institute built by Cuthbert Brodrick, in Cookridge Street (now Millennium Squar ...
. In the
Coat of arms of Northern Ireland The coat of arms of the Government of Northern Ireland was granted to the Executive Committee of the Privy Council of Northern Ireland in 1924. The Government of Northern Ireland was replaced by the Northern Ireland Executive in 1974 and con ...
issued in 1924 (with
supporter In heraldry, supporters, sometimes referred to as ''attendants'', are figures or objects usually placed on either side of the shield and depicted holding it up. Historically, supporters were left to an individual's free choice and were assu ...
s added the next year) an Irish elk is the sinister supporter.Genealogical Office, Dublin, ''Register of Arms 111C (Grants M), 1920 -1929'', folio 66


See also

*


References


Further reading

* (1995): '' Dance of the Tiger''. University of California Press. . :Kurten is a paleo-anthropologist, and in this novel, he presents a theory of
Neanderthal Neanderthals ( ; ''Homo neanderthalensis'' or sometimes ''H. sapiens neanderthalensis'') are an extinction, extinct group of archaic humans who inhabited Europe and Western and Central Asia during the Middle Pleistocene, Middle to Late Plei ...
extinction. Irish elk feature prominently, under the name ''shelk'' which Kurten coins (based on the aforementioned old German ''schelch'') to avoid the problematic aspects of "Irish" and "elk" as discussed above. The book was first published in 1980 when "Giant Deer" was not yet being used widely. * ''Zoological Science'' 22: 1031–1044 (2005). * Larson, Edward J. (2004). ''Evolution: The Remarkable History of a Scientific Theory''.


External links

* * * * {{Taxonbar, from=Q312054 Prehistoric deer Pleistocene Artiodactyla Pleistocene mammals of Europe Pleistocene mammals of Asia Pleistocene first appearances Extinct animals of Ireland Extinct animals of Russia Extinct animals of China Holocene extinctions Taxa named by Johann Friedrich Blumenbach Fossil taxa described in 1799 Species that are or were threatened by habitat loss Species that are or were threatened by climate change