Irish elk

The Irish elk (Megaloceros giganteus),[1][2] also called the giant

deer or Irish deer, is an extinct species of deer in the genus Megaloceros Irish elk

and is one of the largest deer that ever lived. Its range extended across Temporal range: Middle Pleistocene
Eurasia during the Pleistocene, from Ireland to Lake Baikal in Siberia. The to Middle Holocene,
most recent remains of the species have been carbon dated to about 7,700
years ago in western Russia.[3][4]

The Irish elk is known from abundant skeletal remains which have been
found in bogs in Ireland. It is not closely related to either of the living
species currently called elk: Alces alces (the European elk, known in North
America as the moose) or Cervus canadensis (the North American elk or
wapiti). For this reason, the name "giant deer" is used in some
publications, instead of "Irish elk".[5][6][7][8][9] Although one study
suggested that the Irish elk was closely related to the red deer (Cervus
elaphus),[10] most other phylogenetic analyses support the thesis that their
closest living relatives are fallow deer (Dama).[8][11][12][13]
Mounted skeleton

Scientific classification
Contents
Kingdom: Animalia
Taxonomy
Research history Phylum: Chordata
Evolution Class: Mammalia
Description Order: Artiodactyla
Habitat Family: Cervidae
Palaeobiology Subfamily: Cervinae
Physiology
Reproduction Genus: †Megaloceros
Life history Species: †M. giganteus
Extinction Binomial name
Cultural significance †Megaloceros giganteus

See also (Blumenbach, 1799)

References
Further reading
External links

Taxonomy

Research history
Time averaged range of M. giganteus
during the Late Pleistocene
 The first scientific descriptions of the Synonyms
animal's remains were made by Irish
physician Thomas Molyneux in 1695, Alce gigantea Blumenbach,
who identified large antlers from
1799
Dardistown, Dublin—which were
apparently commonly unearthed in Cervus hibernus Desmarest,
Ireland—as belonging to the elk 1820
(known as the moose in North
America), concluding that it was once Cervus megaceros Hart, 1825
abundant on the island.[14] It was first Megaloceros antiquorum
formally named as Alce gigantea by Brookes, 1828
Johann Friedrich Blumenbach in his
Handbuch der Naturgeschichte in Cervus euryceros
Skeletal reconstruction from 1856 1799,[15] with Alce being a variant of (Aldrovandi, 1621), Hibbert,
Alces, the Latin name for the elk. The 1830
original Blumenbach's description of
Alce gigantea provides rather scanty information about the species, Cervus megaceros irlandicus
specifying only that this particular kind of "fossil elk" comes from Ireland Fischer, 1838
and is characterized by immense body size. According to Blumenbach,[15] Cervus (Megaceros)
the distance between summits of giant deer antlers may attain 14 feet
(approximately 4.4 m). This particular feature mentioned by Blumenbach hibernicus Owen, 1844
permitted to Roman Croitor to identify the type specimen of giant deer [16] Cervus giganteus Reynolds,
that was figured and described for the first time in Louthiana of Thomas 1929
Wright.[17] The holotype of Megaloceros giganteus (Blumenbach, 1799) is
a well-preserved male skull with exceptionally large antlers found in Megaceros giganteus latifrons
Dunleer environs (County Louth, Ireland).[16] The type specimen of giant Raven,1935
deer is currently exposed in Barmeath Castle where Thomas Wright first
saw and described it

French scientist Georges Cuvier 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".[18] 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:[19][20]

Amongst other Fossil Bones, there [are] ... two uncommonly fine Crania of the Megalocerus
antiquorum (Mihi). (Irish), with unusually fine horns, (in part restored)

— Joshua Brookes, Brookesian Museum. The Museum of Joshua Brookes, Esq. Anatomical and
Zoological Preparations, p 20.

The etymology being from Greek: μεγαλος megalos "great" + κερας keras "horn, antler".[21] 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. The original description was considered by Adrian
Lister in 1987 to be inadequate for a taxonomic definition.[2] 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" as a
description of the remains. The 1828 publication was approved by International Commission on Zoological
Nomenclature (ICZN) in 1977 as an available publication for the basis of zoological nomenclature.[2] Adrian
Lister in 1987 judged that "the phase "Cornibus deciduis palmatis" constitutes a definition sufficient under the
[International Code of Zoological Nomenclature] (article 12) to validate Megalocerus."[2] The original spelling of
Megalocerus was never used after its original publication.[20]
 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.[2]
Despite being a junior synonym, Megaloceros remained in obscurity and
Megaceros became the common genus name for the taxon.[20] The
combination "Megaceros giganteus" was in use by 1871.[22] George Gaylord
Simpson in 1945 revived the original Megaloceros name, which became
Outdated 1897 reconstruction of
progressively more widely used, until a taxonomic decision in 1989 by the
doe and stag Irish elk by Joseph
ICZN confirmed the priority of Megaloceros over Megaceros, and
Smit
Megaloceros to be the correct spelling.[20][23]

Before the 20th century, the Irish elk, having evolved from smaller ancestors
with smaller antlers, was taken as a prime example of orthogenesis (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. 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.[8] In the 1930s, orthogenesis
was disputed by Darwinians led by Julian Huxley, who noted that antler size was not grossly large, and was
proportional to body size.[24][25] The currently favoured view is that sexual selection was the driving force
behind the large antlers rather than orthogenesis or natural selection.[25]

Evolution

M. giganteus belongs to the genus Megaloceros. Megaloceros is a member of

the possibly polyphyletic (invalid) tribe "Megalocerini" or "Megacerini",
alongside Megaceroides, Praemegaceros, Eucladoceros and Sinomegaceros,
which are often collectively referred to as "giant deer". The taxonomy of giant
deer lacks consensus, with genus names used for species varying
substantially between authors.[26][27] 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 of Russia, which were given the name
of M. stavropolensis in 2016,[28] however these have been subsequently
suggested to belong to Arvernoceros instead.[26][27] The antler construction
suggest a close relationship between M. stavropolensis and Asian Skull of M. g. antecedens
Sinomegaceros.[16] The oldest generally accepted records of the genus are
from the late Early Pleistocene.[29] Other species often considered to belong
to Megaloceros include the reindeer 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
these species to be chronospecies, due to shared morphological characteristics not found in M. giganteus and
gradual transition of morphological characters through time.[26] M. savini has also been suggested to comprise
the separate genus Praedama by some scholars, though they are often considered closely related. Roman Croitor
has suggested closer affinities to Eucladoceros for M. savini and related species.[27]

The origin of M. giganteus remains unclear, and appears to lie outside Western Europe.[26] 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-matritensis lineage due to the
shared molarisation of the lower fourth premolar (P4).[26] Croitor has suggested that M. giganteus is closely
related to what was originally described as Dama clactoniana mugharensis (which he proposes be named

Outdated 1906 restoration by
Charles R. Knight
Ballybetagh Bog near Dublin.[32]
Megaloceros mugharensis) from the Middle Pleistocene of Tabun Cave in
Israel, due to similarities in the antlers, molars and premolars.[27] The
earliest possible records of M. giganteus comes from Homersfield, England
thought to be about 450,000 years ago—though the dating is uncertain.[30]
The oldest securely dated Middle Pleistocene records are those from Hoxne,
England, which have been dated to Marine Isotope Stage 11 (424,000 to
374,000 years ago),[31][26] 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, England.[30][26] Most
remains of the Irish elk are known from the Late Pleistocene. A large
proportion of the known remains of M. giganteus are from Ireland, which
mostly date to the Allerød oscillation near the end of the Late Pleistocene
around 13,000 years ago. Over 100 individuals have been found in

It has been historically thought that, because both have palmated antlers, the Irish elk and fallow deer (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.[27] In
2005, two fragments of mitochondrial DNA (mtDNA) from the cytochrome b gene were extracted and sequenced
from 4 antlers and a bone, the mtDNA found that the Irish elk was nested within Cervus, and were inside the
clade containing living red deer (Cervus elaphus). Based on this, the authors suggested that the Irish elk and red
deer interbred.[10] However, another study from the same year in the journal Nature utilising both fragmentary
mitochondrial DNA and morphological data found that the Irish elk was indeed most closely related to Dama.[11]
The close relationship with Dama was supported by another cytochrome b study in 2006,[8] a 2015 study
involving the full mitochondrial genome,[12] and by a 2017 morphological analysis of the bony labyrinth.[13] The
2006 and 2017 studies also directly suggest that the results of the 2005 cytochrome b paper were the result of
DNA contamination.

Description
The Irish elk stood about 2.1 m (6 ft 11 in) tall at the shoulders and carried
the largest antlers of any known deer, a maximum of 3.65 m (12.0 ft) from tip
to tip and 40 kg (88 lb) in weight. For body size, at about 450–600 kg (990–
1,320 lb) and up to 700 kg (1,540 lb) or more,[33][34][35][36] the Irish elk was
the heaviest known cervine ("Old World deer");[11] and tied with the extant
Alaska moose (Alces alces gigas) as the third largest known deer, after the
extinct Cervalces latifrons and Cervalces scotti.[34][35] Nonetheless,
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 than Alces, with Irish elk bucks being notably larger than
does.[37] In total, Irish elk bucks may have ranged from 450–700 kg (990–
1,540  lb), with an average of 575  kg (1,268  lb), and does may have been
relatively large, about 80% of buck size, or 460 kg (1,010 lb) on average.[38]
Life restoration
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), with the early and complete obliteration of cranial sutures.[27]
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 (between the shoulder blades). In 1989, American palaeontologist
Dale Guthrie suggested that, like bison, 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.[38]

Habitat
It was not exclusive to Ireland. Rather, it was so named because the most well-known and best-preserved fossil
specimens have been found in lake sediments and peat bogs in Ireland. The Irish elk had a far-reaching range,
extending from the Atlantic Ocean in the West to Lake Baikal in the East. They do not appear to have extended
northward onto the open mammoth steppe, rather keeping to the boreal steppe-woodland environments, which
consisted of scattered spruce and pine, as well as low-lying herbs and shrubs including grasses, sedges, Ephedra,
Artemisia and Chenopodiaceae.[4]

Palaeobiology

Physiology

In 1998, Canadian biologist Valerius Geist hypothesised that the Irish elk was
cursorial (adapted for running and stamina). He noted that the Irish elk
physically resembled reindeer. The body proportions of the Irish elk are
similar to those of the cursorial addax, oryx, 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, gnus, and reindeer have a Drawing of cave art from Grotte de
top speed of over 80 km/h (50 mph), and can maintain high speeds for up to Cougnac, France showing coloured
15 minutes.[38] shoulder hump and lines. c. 25,000
to 19,000 years old[39]

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 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
Mounted skeletons of a male (left) and female many modern deer species. Bucks generally suffer higher
mortality rates because they eat little during the autumn
rut.[40] For rut, a lean stag normally 575 kg (1,268 lb) may
have fattened up to 690 kg (1,520 lb), and would burn through the extra fat over the next month.[38]
Assuming a similar response to starvation as red deer, a large, healthy Irish elk stag with 40 kg (88 lb) antlers
would have had 20-to-28 kg (44-to-62 lb) antlers under poor conditions;[18][35] and an average sized Irish elk
stag with 35  kg (77  lb) antlers would have had 18 to 25  kg (40 to 55  lb) antlers under poorer conditions,[41]
similar sizes to the moose. A similar change in a typical Irish elk population with prime stags having 35 kg (77 lb)
antlers would result in antler weights of 13 kg (29 lb) or less in worsening climatic conditions. This is within the
range of present-day wapiti/red deer (Cervus spp.) antler weights.[38] Irish elk antlers vary widely in form
depending upon the habitat, such as a compact, upright shape in closed forest environments.[41] Irish elk likely
shed their antlers and re-grew a new pair during mating season. Antlers generally require high amounts of
calcium and phosphate, 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 while the antlers are growing, and replenishing them from
food plants after the antlers have grown in or reclaiming nutrients from shed antlers.[35]

The large antlers have generally been explained as being used for male-male battle during mating season.[42]
They may have also been used for display,[18] to attract females and assert dominance against rival males.[40] A
finite element analysis 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.[43]

In deer, gestation time generally increases with body size. A 460  kg (1,010  lb) 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
(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.[38]

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, with minor Plantago, Helianthemum,
Plumbaginaceae and Salix.[44] A stable isotope analysis of the terminal
Pleistocene Irish population suggests a grass and forb based diet,
supplemented by browsing during stressed periods.[45] Dental wear patterns
of specimens from the late Middle and Late Pleistocene of Britain suggest a Skull in front view
diet tending towards mixed feeding and grazing, but with a wide range
including leaf browsing.[46]

Based on the dietary requirements of red deer, a 675  kg (1,488  lb) lean Irish elk stag would have needed to
consume 39.7 kg (88 lb) 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.[38]

The Irish elk may have been preyed upon by the large carnivores of the time, including the cave lion,[47] and the
cave hyena.[48]
Extinction
Outside of the Irish Late Pleistocene, remains of Irish elk are uncommon,
suggesting that they were usually rare in the areas that they did occur.[4]

Historically, its extinction has been attributed to the encumbering size of the
antlers, a "maladaptation" making fleeing through forests especially difficult
for males while being chased by human hunters,[18] or being too taxing
nutritionally when the vegetation makeup shifted.[35] In these scenarios,
sexual selection by does for stags with large antlers would have contributed
to decline.[49]
Sculptures in Crystal Palace
However, antler size decreased through the Late Pleistocene and into the
Holocene, and so may not have been the primary cause of extinction.[41] 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.[50] Such resource constriction may have cut female fertility rates in half.[41]
Human hunting may have forced Irish elk into suboptimal feeding grounds.[3]

The range of the taxon appears to have collapsed during the Last Glacial Maximum, with few remains known
between 27,500 and 14,600 years ago, and none between 23,300 and 17,500 years ago. The remains of the taxon
substantially increase during the latest Pleistocene, where it appears to have re-colonised most of its former
range, with abundant remains in the UK, Ireland, and Germany.[4]

While the range of the taxon was dramatically reduced after the Pleistocene-Holocene transition, it managed to
survive into the early Northgrippian in the eastern part of its range within European Russia and Siberia, in a belt
extending from Maloarchangelsk in the East to Preobrazhenka in the west. It is suggested that extinction was
contributed to by further climatic changes transforming preferred open habitat into uninhabitable dense
forest.[4] The final demise may have been caused by several factors both on a continental and regional scale,
including climate change and hunting.[50][51] 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."[4]

Cultural significance
A handful of Irish elk depictions are known from the art of
the Upper Paleolithic in Europe. However, these are much
less abundant than the common red deer and reindeer
depictions. The bones of the Irish elk are uncommon in
localities where they are found, and only a handful of
examples of human interaction are known.[4] A mandible
from Ofatinţi, Moldova dating to either the Eemian or the
early Late Pleistocene, "is peculiar because it has ancient
tool-made notches on its lateral side".[52] Several M.
giganteus bones from the Chatelperronian levels of the
Labeko Koba site in Spain are noted for bearing puncture 36,000 year old Irish Replica of a cave painting from
marks, which have been interpreted as anthropogenic. [53] A elk cave painting at Lascaux, c. 15,000 BC
terminal Pleistocene (13710-13215 cal BP) skull from Chauvet Cave, France
Lüdersdorf, Germany is noted to have had the antler and (dot indicates 14C
facial part of the skull removed in a way unlikely to be due sample)
to natural causes. [54] A calcaneum 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".[55][4] 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 to produce "blanks" for the manufacture of barbed projectile tips.[56][4] A ring-like mark on a shed
antler beam from the similarly aged Paderborn site in Germany has been suggested to be anthropogenic.[57]

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.[58] Indeed Leeds Philosophical and Literary Society bought a full skeleton in 1847,
from Glennon's in Dublin, for £38.[59] This specimen, discovered at Lough Gur near Limerick, is still on display
at Leeds City Museum.[60]

A folk memory of the Irish elk was once thought to be preserved in the Middle High German word Schelch, a
large beast mentioned in the 13th-century Nibelungenlied along with the then-extant aurochs (Dar nach schluch
er schiere, einen Wisent und einen Elch, Starcher Ure vier, und einen grimmen Schelch / "After this he
straightway slew a Bison and an Elk, Of the strong Wild Oxen four, and a single fierce Schelch.") This opinion is
no longer widely held.[18] The Middle Irish word segh was also suggested as a reference to the Irish elk.[61][62]
Turf cutters of Clooney and Tulla in County Clare, Ireland referred to the Irish elk as the Fiaghmore (Fia
meaning Deer, Mór meaning "great", or "large").[63] However, these interpretations are not conclusive.[64]

See also
 Paleontology portal

References
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Further reading
Kurten, Bjorn (1995): Dance of the Tiger. University of California Press. ISBN 0-520-20277-5.

Kurten is a paleo-anthropologist, and in this novel, he presents a theory of Neanderthal 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
"Extinct Giant Deer Survived Ice Age, Study Says" (http://news.nationalgeographic.com/news/2004/10/1006
_041006_giant_deer.html). National Geographic.
"CGI picture from Walking with Beasts" (https://web.archive.org/web/20060217212625/http://dsc.discovery.c
om/convergence/beasts/photo/slide_05.html). Discovery Channel. Archived from the original (http://dsc.disco
very.com/convergence/beasts/photo/slide_05.html) on 17 February 2006. Retrieved 14 February 2006.
"Megaloceros, Irish elk, Giant deer" (http://www.bbc.co.uk/nature/wildfacts/factfiles/461.shtml). BBC.
Retrieved 25 October 2005.
"The Case of the Irish Elk" (https://web.archive.org/web/20051111051611/http://www.ucmp.berkeley.edu/ma
mmal/artio/irishelk.html). University of California at Berkeley. Archived from the original (http://www.ucmp.ber
keley.edu/mammal/artio/irishelk.html) on 11 November 2005. Retrieved 25 October 2005.

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