Irish elk

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Irish elk
Temporal range: Middle
Ma
↓
Mounted skeleton
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Artiodactyla
Family: Cervidae
Subfamily: Cervinae
Genus: †Megaloceros
Species:
†M. giganteus
Binomial name
†Megaloceros giganteus
(Blumenbach, 1799)
Time averaged range of M. giganteus during the Late Pleistocene
Synonyms
  • Alce gigantea Blumenbach, 1799
  • Cervus hibernus Desmarest, 1820
  • Cervus megaceros Hart, 1825
  • Megaloceros antiquorum Brookes, 1828
  • Cervus euryceros (Aldrovandi, 1621), Hibbert, 1830
  • Cervus megaceros irlandicus Fischer, 1838
  • Cervus (Megaceros) hibernicus Owen, 1844
  • Cervus giganteus Reynolds, 1929
  • Megaceros giganteus latifrons Raven,1935

The Irish elk (Megaloceros giganteus),

radiocarbon dated to about 7,700 years ago in western Russia.[3][4] Its antlers, which can span 3.5 metres (11 ft) across are the largest known of any deer.[5] 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 (Dama).[5][6][7][8]

Taxonomy

Research history

Skeletal reconstruction from 1856

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.[9] It was first formally named as Alce gigantea by Johann Friedrich Blumenbach in his Handbuch der Naturgeschichte in 1799,[10] with Alce being a variant of Alces, the Latin 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,[10] the distance between summits of giant deer antlers may attain 14 feet (approximately 4.4 m). This particular feature mentioned by Blumenbach permitted to Roman Croitor to identify the type specimen of giant deer [11] that was figured and described for the first time in Louthiana of Thomas Wright.[12] The holotype of Megaloceros giganteus (Blumenbach, 1799) is a well-preserved male skull with exceptionally large antlers found in Dunleer environs (County Louth, Ireland).[11] The type specimen of giant deer is currently exposed in Barmeath Castle where Thomas Wright first saw and described it.[11]

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" (the most famous of all fossil ruminants).[13] 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:[14][15]

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

priority of Megaloceros over Megaceros, and Megaloceros to be the correct spelling.[15][18]

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.[6] 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.[19][20] The currently favoured view is that sexual selection was the driving force behind the large antlers rather than orthogenesis or natural selection.[20]

Evolution

Skull of M. g. antecedens

M. giganteus belongs to the genus

Praedama by other authors.[23] While the M. savini/Praedama lineage is often suggested to be closely related to M. giganteius, most authors agree that this group of deer is unlikely to be directly ancestral to M. giganteus.[11]

Outdated 1906 restoration by Charles R. Knight

The origin of M. giganteus remains unclear, and appears to lie outside Western Europe.[22] 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 (P4).[22] 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.[23] The earliest possible records of M. giganteus comes from Homersfield, England thought to be about 450,000 years ago—though the dating is uncertain.[26] 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),[27][22] 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.[26][22] 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 Ballybetagh Bog near Dublin.[28]

It has been historically thought that, because both have palmated antlers, the Irish elk and

nasal bones, and proportionally short front portion of the skull.[23] 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.[29] 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.[5] The close relationship with Dama was supported by another cytochrome b study in 2006,[6] a 2015 study involving the full mitochondrial genome,[7] and by a 2017 morphological analysis of the bony labyrinth.[8] The 2006 and 2017 studies also directly suggest that the results of the 2005 cytochrome b paper were the result of DNA contamination.[6][8]

Cladogram of Cervidae based on mitochondrial DNA:[30]

Cervidae

Hydropotes
(water deer)

Capreolus (roe deer)

Alces
(moose)

Rangifer (reindeer/caribou)

Odocoileini (brocket deer, mule deer, white tailed deer, etc)

Elaphodus
(tufted deer)

Muntiacus
(muntjacs)

Cervini

Rucervus (Schomburgk's deer and barasingha)

Axis (chital, hog deer)

Dama (Fallow deer)

†Megaloceros giganteus (Irish elk)

Elaphurus (Père David's deer)

Panolia
(Eld's deer)

Rusa alfredi
(Visayan spotted deer)

Rusa marianna
(Philippine deer)

Rusa timorensis
(Javan rusa)

Rusa unicolor
(Sambar deer)

Cervus (red deer, elk, sika deer)

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.[31]

Dama

Sinomegaceros+Megaloceros

Sinomegaceros pachyosteus (China)

Megaloceros giganteus (Russia, Belgium)

Sinomegaceros ordosianus (China, Russia)

Sinomegaceros pachyosteus (China)

Megaloceros giganteus (Europe, Russia)

Description

Life restoration

The Irish elk stood about 2 m (6 ft 7 in) tall at the shoulders,

cervine ("Old World deer");[5] 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][33] 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.[36] 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.[37] 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.[23]

Based on

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.[37]

Habitat

The Irish elk had a far-reaching range, extending from the Atlantic Ocean in the West to

Chenopodiaceae.[4]

Palaeobiology

Physiology

Drawing of cave art from Grotte de Cougnac, France showing coloured shoulder hump and lines. c. 25,000 to 19,000 years old[38]

In 1998, Canadian biologist Valerius Geist hypothesised that the Irish elk was

gnus, and reindeer have a top speed of over 80 km/h (50 mph), and can maintain high speeds for up to 15 minutes.[37]

Reproduction

Mounted skeletons of a buck (left) and a doe

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 many modern deer species. Bucks generally suffer higher mortality rates because they eat little during the autumn rut.[39] 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.[37]

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;[13][33] 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,[40] 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.[37] Irish elk antlers vary widely in form depending upon the habitat, such as a compact, upright shape in closed forest environments.[40] 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.[33]

The large antlers have generally been explained as being used for male-male battle during mating season.[41] They may have also been used for display,[13] to attract females and assert dominance against rival males.[39] 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.[42]

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.[37]

Diet and life history

Skull in front view

The mesodont (meaning neither high (

cow parsley (Anthriscus sylvestris), cow parsnip/hogweed (Heracleum), water pennywort (Hydrocotyle), Asteraeceae, Filipendula, Symphytum and grass embedded with its teeth.[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 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.[37]

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,[13] or being too taxing nutritionally when the vegetation makeup shifted.[33] In these scenarios, sexual selection by does for stags with large antlers would have contributed to decline.[49]

However, antler size decreased through the Late Pleistocene and into the Holocene, and so may not have been the primary cause of extinction.[40] 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.[40] Human hunting may have forced Irish elk into suboptimal feeding grounds.[3]

The distribution of M. giganteus is thought to have been strongly controlled by climactic conditions. The range of the Irish elk appears to have collapsed during the Last Glacial Maximum (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, where it appears to have re-colonized most of its former range, with abundant remains in the UK, Ireland, and Germany.[4] 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.[51]

While the range of the taxon was dramatically reduced after the Pleistocene-Holocene transition, it managed to survive into the middle Holocene in the central part of its range within European Russia and Western Siberia. 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][52] 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."[4]

Relationship with early humans

14C sample)
Replica of a cave painting from Lascaux
, c. 15,000 BC

A handful of Irish elk depictions are known from the

Chatelperronian levels of the Labeko Koba site in Spain are noted for bearing puncture marks, which have been interpreted as anthropogenic.[55] A terminal Pleistocene (13710-13215 cal BP) skull from Lüdersdorf, Germany is noted to have had the antler and facial part of the skull removed in a way unlikely to be due to natural causes.[56] 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".[57][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.[58][4] A ring-like mark on a shed antler beam from the similarly aged Paderborn site in Germany has been suggested to be anthropogenic.[59]

Modern significance

Sculptures in Crystal Palace

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.[60] Indeed Leeds Philosophical and Literary Society bought a full skeleton in 1847, from Glennon's in Dublin, for £38.[61] This specimen, discovered at Lough Gur near Limerick, is still on display at Leeds City Museum.[62]

See also

References

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Further reading

  • 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

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