A comparison of postures and gaits between two 'elands', the moose (Alces alces) and the common eland (Taurotragus oryx)

@tonyrebelo @jeremygilmore @tandala @oviscanadensis_connerties @aguilita @capracornelius @zarek @dejong @michalsloviak @christiaan_viljoen @dinofelis @maxallen @chewitt1 @saber_animal @scottdwright @calebcam @jwidness @matthewinabinett @lefebvremax @ldacosta @henrydelange @davidbygott @jandutoit @gigilaidler @sitszasadam @muir @outnabout

Also please see https://www.inaturalist.org/journal/milewski/69307-a-new-observation-on-maternal-defensive-behaviour-in-the-moose-alces-alces#

When the Dutch landed in South Africa in the late seventeenth century, they found the strange bovid, Taurotragus oryx, to be so similar to the familiar cervid, Alces alces, that they used the same name.

'Eland' is the Dutch word for Alces alces alces (https://www.inaturalist.org/observations?taxon_id=852458), which was originally indigenous to the Netherlands.

This adoption of a misnomer - which has persisted for nearly four centuries - is understandable, because both species

are the largest-bodied members of their respective categories of ruminants, namely deer vs antelopes,
habitually trot, rather than cantering/galloping, when pursued (https://www.google.com.au/search?q=Eland+bulls+fighting&sca_esv=568362559&sxsrf=AM9HkKlgG-AUfAcw_VbOYYgCRYuv81AY2w%3A1695692872768&source=hp&ei=SDgSZfTzLLXMseMPp9y1mAk&iflsig=AO6bgOgAAAAAZRJGWCnf_rbtFmhUhTBCL4oux5QgVTD9&ved=0ahUKEwj0t8-alMeBAxU1ZmwGHSduDZMQ4dUDCAs&uact=5&oq=Eland+bulls+fighting&gs_lp=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&sclient=gws-wiz#fpstate=ive&vld=cid:11586598,vid:OWcItn5k7KI,st:0),
possess noticeably high withers (https://www.dictionary.com/browse/withers), and
possess dewlaps (https://www.merriam-webster.com/dictionary/dewlap) in both sexes, which are more noticeable than the short nuchal mane present in both species.

In this Post, I compare A. alces with T. oryx with respect to postures and gaits, in light of the profound differences in their habitats and niches.

LYING

Both species rest in sternal recumbency (https://bestofthetetons.com/2014/11/18/resting-moose-a-collection-of-less-seen-lifestyle-images/ and https://www.alamy.com/stock-photo/eland-lying-down.html?sortBy=relevant).

KNEELING

A major difference is that A. alces kneels readily at all ages (https://www.azotelibrary.com/en/image/a-moose-%28alces-alces%29-kneels-to-eat-grass-in-a-meadow-in-s%C3%B6dermanland/191539), whereas T. oryx seems never to kneel (apart from transitorily when lying down in sternal recumbency, or arising from such lying, https://www.bestofthetetons.com/wp-content/uploads/2014/11/RestingMoose_GettingUpSequence1400px.jpg and https://www.shutterstock.com/image-photo/eland-antelope-rising-grass-savannah-green-2247874629).

When drinking, T. oryx sometimes spreads its forelegs to some degree, in partial emulation of Giraffa (https://www.alamy.com/giraffe-and-elands-at-a-waterhole-area-etosha-national-park-namibia-africa-image415771067.html?imageid=16AC8853-6C1A-4753-BFF4-3ABEAFFEE8E7&p=1364503&pn=1&searchId=c841ad5a0499198295bbf8dbec6d6cc6&searchtype=0 and https://africageographic.com/stories/eland/ and https://www.pinterest.com.au/pin/vertical-photo-of-huge-eland-antelopetaurotragus-oryx-standing-under-angolan-giraffe-drinking-from-waterhole-animals--664492120013532545/ and https://www.sciencephoto.com/media/389750/view/eland-antelope-drinking and https://fineartamerica.com/featured/eland-antelope-drinking-tony-camachoscience-photo-library.html?product=wood-print and https://www.alamy.com/stock-photo-eland-drinking-at-waterhole-etosha-namibia-19949416.html and https://www.gettyimages.ae/detail/photo/common-eland-drinking-at-chudop-waterhole-etosha-royalty-free-image/1032889546).

I have yet to see a similar posture in A. alces - which instead kneels in such situations.

When sparring or fighting in masculine rivalry, neither species routinely kneels (https://www.shutterstock.com/image-photo/common-eland-taurotragus-oryx-male-fight-2255105889). In the case of T. oryx, I have yet to see evidence of any kneeling at all.

There are many photos on the Web of A. alces in kneeling posture (please see examples at the end of this Post).

JUMPING

Alces alces can jump 2 m high, whereas T. oryx can jump 3 m high (https://naloolo.wordpress.com/2018/03/11/jumped-over-by-an-eland/).

The former species is a capable jumper, in line with other cervids. However, the latter is a remarkable jumper indeed. This is the converse of what might be predicted based on the length of the legs.

C R S Pitman (https://en.wikipedia.org/wiki/Charles_Pitman_(game_warden)) recorded a case where a mature male individual of T. oryx (weighing perhaps >550 kg, https://www.inaturalist.org/observations/182885490 and https://www.inaturalist.org/observations/184504263) easily vaulted an enclosure 2.4 m high.

STOTTING/LEAPING

Alces alces is not known to stot. However, it is possible that, when it trots with particularly high steps (https://www.istockphoto.com/photo/young-bull-moose-running-across-stubble-field-gm491959369-40066416?phrase=elk+running+snow), this may be a demonstration of fitness, analogous to the proud-trotting of certain other ruminants.

Also deserving further scrutiny is the juvenile 'play-trotting', while wading, seen in https://www.youtube.com/watch?v=lVPfRQjDPBE.

By contrast, T. oryx has often been photographed 'stotting', in a way different from most other bovids.

Instead of bouncing stiff-legged, individuals leap randomly, and seemingly pointlessly, while galloping in alarm in a group (https://twitter.com/Min_TourismKE/status/720195243241578496/photo/2 and https://www.zooniverse.org/projects/zooniverse/snapshot-serengeti/talk/1485/912470 and https://www.alamy.com/stock-photo-africa-kenya-masai-mara-motion-blur-of-eland-jumping-through-grass-32822773.html?imageid=B09244FD-293E-4CD1-B166-556A9E4CB550&p=95453&pn=1&searchId=6101ba2c62ae717283a6f3f1f002323c&searchtype=0 and https://www.alamy.com/stock-photo-eland-taurotragus-oryx-herd-running-laikpia-kenya-125584865.html?imageid=2DF7F368-4CD7-498D-97F8-AF1C2B837801&p=361664&pn=1&searchId=010f32aaa73c04f1f11960177dc59d66&searchtype=0). This sometimes means leaping over the back of an adjacent individual. This behaviour may function similarly to stotting, in demonstrating individual fitness to potential predators.

BIPEDALITY

Alces alces sometimes stands bipedally, both to forage (https://www.reddit.com/media?url=https%3A%2F%2Fi.redd.it%2Feot6wzys7i981.jpg) and to quarrel (https://www.gettyimages.ca/detail/video/cow-moose-standing-up-on-their-hind-legs-and-fighting-in-stock-footage/460383460 and scroll in https://www.canadiannaturephotographer.com/Joe_Desjardins.html).

By contrast, T. oryx - as far as I know - never uses any bipedal posture except transitorily (for a split-second) by males in copulation (https://www.dreamstime.com/stock-photo-common-elands-mating-pair-taurotragus-oryx-amakhala-game-reserve-eastern-cape-south-africa-image51205440). Even in copulation, males support their weight on the females (https://www.agefotostock.com/age/en/details-photo/a-pair-of-common-elands-taurotragus-oryx-mating-in-the-amakhala-game-reserve-eastern-cape-south-africa/ESY-026334176).

Taurotragus oryx compensates for its inability to stand bipedally, as follows (https://www.youtube.com/watch?app=desktop&v=tqe_qvUX43k). This species has often been observed to forage as high as 2.13 m, "frequently using their horns to break branches in order to get at the leaves" (J Posselt, African Wild Life magazine, June 1961).

Alces alces also breaks woody stems while foraging, but does so by

reaching up with its mouth and then pulling downward, or
walking against an upright stem to force it down by means of the chest (https://winterberrywildlife.ouroneacrefarm.com/2019/08/09/moose-feeding-sign-with-video/).

FORAGING NEAR GROUND LEVEL WITHOUT KNEELING

Both A. alces and T. oryx are capable of foraging, without kneeling, on terrestrial plants lower than 20 cm above ground level. This is because the former combines a short neck with a long muzzle, whereas the latter combines a relatively long neck with a relatively short muzzle.

However, foraging in this way is easier for T. oryx than for A. alces, the former grazing as a staple part of its foraging (short green grass during the rainy season), whereas the latter seldom grazes. The neck of A. alces is so proportionately short that the length of the muzzle does not quite compensate.

Thus the shoulders and elbows are more flexed in an awkward posture in the case of A. alces, with the humerus approaching the horizontal (https://www.usgs.gov/media/images/moose-alces-alces-19 and https://unsplash.com/photos/tLUKrYFh4-w and https://pixels.com/featured/bull-moose-in-velvet-grazing-edie-ann-mendenhall.html and https://www.123rf.com/photo_6921276_bull-moose-grazing.html and https://www.agefotostock.com/age/en/details-photo/alaska-moose-grazing-in-tundra-male-moose-in-green-tundra-grazing-alaska/MEV-12905729).

By contrast, T. oryx appears more at ease when grazing (https://www.inaturalist.org/observations/148679101 and https://www.inaturalist.org/observations/183667651 and https://www.inaturalist.org/observations/186183888 and https://www.inaturalist.org/observations/185385079).

To reach ground level itself with the mouth, the two species differ categorically.

Alces alces is incapable of doing so without kneeling (please see compendium of photos at the end of this Post).

By contrast, T. oryx does so by flexing the shoulders and elbows slightly, and leaving one foreleg at an angle (https://www.inaturalist.org/observations/11166225 and https://www.inaturalist.org/observations/141140285 and https://www.inaturalist.org/observations/181256408 and https://www.alamy.com/cape-eland-grazing-at-de-hoop-national-reserve-cape-overberg-south-image9879753.html?imageid=43081444-5267-444D-BC18-39F285FC4BCE&p=6945&pn=1&searchId=bebe53939638886546a1f3e44729a7e9&searchtype=0 and https://www.alamy.com/eland-taurotragus-oryx-de-hoop-nature-reserve-western-cape-south-africa-image399153412.html?imageid=B11E5FBA-723B-4A35-8CD5-F15BD837EF7F&p=12455&pn=2&searchId=5cd2bf43ed0e0cf1d243f5a9906dcb64&searchtype=0 and https://www.inaturalist.org/observations/34994877 and https://www.inaturalist.org/observations/27276823).

WALKING

Please also see https://www.inaturalist.org/journal/milewski/85161-variation-in-walking-gaits-in-ungulates-why-some-hoofed-mammals-cross-walk-whereas-others-amble#.

Please see https://www.youtube.com/watch?v=t4zdTn02PWQ

Alces alces and T. oryx seem to walk with identical gaits. This is a semi cross-walk, not an amble.

Semi cross-walking is typical of ungulates dependent on cover, whereas ambling is typical of gregarious ungulates adapted to open environments. Alces alces is more cover-dependent than T. oryx.

Alces alces is the only long-legged ungulate that semi cross-walks. For its part, T. oryx is possibly the only 'plains game' ungulate that semi cross-walks.

Alces alces fully cross-walks when walking backwards (https://www.youtube.com/watch?v=Z5eY7kVsLQA), as does the horse (Equus caballus, https://www.deviantart.com/nexu4/art/white-horse-walking-backwards-392656340 and https://kenziedysli.com/walking-backwards/).

The following show the normal semi cross-walk of A. alces. Please note that, by the time that the fore is about to be placed, the hind has already been lifted, for long enough that the lower leg has reached the vertical.

https://a57.foxnews.com/global.fncstatic.com/static/managed/img/fn2/video/876/493/edge_moose_052014.jpg?ve=1&tl=1
https://www.istockphoto.com/photo/bull-moose-gm1055977806-282201912
https://fineartamerica.com/featured/walking-bull-moose-randy-straka.html
https://www.alamy.com/bull-moose-denali-national-park-preserve-alaska-united-states-of-america-a-unique-optimised-version-of-an-image-by-nps-ranger-jw-frank-credit-npsjacob-w-frank-image454466690.html?imageid=DC229BEB-798B-4807-AFC7-245D832C911F&p=1837101&pn=1&searchId=acbc6be48e9684687a1dabcb59018a60&searchtype=0
https://pixels.com/featured/bull-moose-crossing-river-jack-bell.html?product=wood-print

The following, correspondingly, show the semi cross-walk of T. oryx.

https://www.kimballstock.com/popuppreview.asp?db=a&image=AFW+36+MH0004+01&itemw=4&itemf=0001&itemstep=1&itemx=1
https://www.istockphoto.com/photo/young-eland-walking-on-the-skyline-gm670061476-122489693
https://www.alamy.com/common-or-southern-eland-taurotragus-oryx-de-hoop-nature-reserve-western-cape-south-africa-image555774369.html?imageid=B8DBE331-4F0A-4C2E-9AA5-68B247944D26&p=824482&pn=1&searchId=bebe53939638886546a1f3e44729a7e9&searchtype=0

The following (https://www.wanyamasafaris.com/kruger-giants/) nicely shows that, when T. oryx walks briskly, the left fore hoof touches the ground at the same instant that the right hind hoof loses touch with the ground. This is the normal configuration in walking in Syncerus caffer, as well as equids.

The following (https://www.inaturalist.org/observations/70876892) is intriguing in showing a full cross-walk in A. alces, in forward motion. I surmise that this extreme gait was resorted to during a brief crossing of unstable stony ground.

Walking gaits of other ungulates, for comparison:

The following, of Alcelaphus caama, shows the difference between the semi cross-walk of tragelaphins (including T. oryx) and the semi amble of alcelaphins (including hartebeests). The hind hoof is lifted only once the fore hoof has touched the ground.

https://www.istockphoto.com/photo/wild-red-hartebeest-during-the-summer-in-beautiful-pilanesberg-national-park-south-gm1358864733-432370852?phrase=hartebeest

The following shows that Camelus dromedarius, although similar to A. alces in the proportional length of the legs, is similar in walking gait to alcelaphins, and different from A. alces.

https://www.gettyimages.com.au/detail/photo/side-view-of-camel-walking-royalty-free-image/1146381538

https://www.inaturalist.org/observations/182080028 and https://www.inaturalist.org/observations/179933097 and https://www.inaturalist.org/observations/145035064

TROTTING

Both species trot, as their preferred gait in running (https://www.youtube.com/watch?v=yxTsf51vqmE and https://www.inaturalist.org/observations/10849500 and https://www.youtube.com/watch?v=e-PA5qsOjBg and https://www.inaturalist.org/observations/25872372 and https://commons.wikimedia.org/wiki/File:Muybridge,Eadweard-Trottender_Eland%28Zeno_Fotografie%29.jpg and https://www.1stdibs.com/art/photography/black-white-photography/eadweard-muybridge-animal-locomotion-plate-696-eland-walking-1887-eadweard-muybridge/id-a_5936652/ and https://www.bridgemanimages.com/en/muybridge/eadweard-muybridge-the-trot-b-w-photo/black-and-white-photograph/asset/6015695 and https://www.flickr.com/photos/wikimediacommons/15998245714/).

Alces alces can trot with a high-stepping style, allowing it

to clear obstacles such as fallen logs, without jumping, and
to run through deep snow (https://www.youtube.com/watch?v=6GEhM2Byk7w).

Because its legs are so long, its speed while trotting corresponds to the cantering gait of other ruminants of similar body mass.

For its part, T. oryx is remarkable in that it trots even when in immediate danger from predators (https://www.youtube.com/watch?v=OWcItn5k7KI and https://www.youtube.com/watch?v=wSMoF7fDRYQ andhttps://www.youtube.com/watch?v=wSMoF7fDRYQ and https://thegreatestmaasaimara.com/?competition_data=cheetah-hunting-eland).

CANTERING

Please see https://www.youtube.com/watch?v=AoLg6aDqwUI

I have yet to see evidence that either A. alces and T. oryx ever canter.

GALLOPING

Please see https://www.youtube.com/watch?v=RV9P0w8vZi8 and

Both species are capable of galloping for limited distances.

Alces alces:
!https://www.youtube.com/watch?app=desktop&v=XlcypwnGcjQ
https://www.facebook.com/fox13seattle/videos/snowboarders-film-moose-running-alongside-them-down-mountain/10156033272534199/?locale=ms_MY
https://www.alamy.com/stock-photo-galloping-moose-on-the-run-39492457.html?imageid=5350CBC7-C4B1-4B9A-9B64-6D290EE4B904&p=34828&pn=1&searchId=9980e653e8ba1e2c918d82ab21cc4bd3&searchtype=0
https://www.alamy.com/stock-photo-galloping-moose-on-the-run-39492473.html?imageid=33DB1C68-DD8A-4C00-AD3B-DC32CDCC8420&p=34828&pn=1&searchId=9980e653e8ba1e2c918d82ab21cc4bd3&searchtype=0
https://www.alamy.com/eurasian-elk-alces-alces-alces-galloping-jaemtland-or-jamtland-sweden-image61916611.html?imageid=3BC9868E-A8B6-4863-99A7-6C3CC44A4B4A&p=75026&pn=1&searchId=9980e653e8ba1e2c918d82ab21cc4bd3&searchtype=0

Taurotragus oryx:
https://www.alamy.com/stock-photo-cape-eland-running-through-the-golden-grasses-of-the-open-serengeti-23833726.html?imageid=ED4E0569-531B-4524-BA79-6BD1B4DFE2A1&p=17446&pn=1&searchId=010f32aaa73c04f1f11960177dc59d66&searchtype=0
https://www.alamy.com/stock-photo-cape-eland-running-through-the-golden-grasses-of-the-open-serengeti-23832589.html?imageid=17B542F1-E9D2-4E86-826E-EF7E8C5BB3F9&p=17446&pn=1&searchId=010f32aaa73c04f1f11960177dc59d66&searchtype=0
https://www.alamy.com/eland-running-taurotragus-oryx-masai-mara-kenya-image181597557.html?imageid=B8F85DBD-7D48-4E42-9140-A35A91064D77&p=738418&pn=1&searchId=010f32aaa73c04f1f11960177dc59d66&searchtype=0
https://www.alamy.com/eland-taurotragus-oryx-masai-mara-kenya-image181635085.html?imageid=BA59C0DA-E616-4513-896B-D62F229A744A&p=738418&pn=1&searchId=010f32aaa73c04f1f11960177dc59d66&searchtype=0
https://www.alamy.com/stock-photo-eland-taurotragus-oryx-pair-running-through-spring-flowers-west-coast-125584310.html?imageid=86EB619E-ADDE-42BF-A3F5-11AD3FF7BAD5&p=361664&pn=1&searchId=010f32aaa73c04f1f11960177dc59d66&searchtype=0
https://www.alamy.com/stock-photo-eland-taurotragus-oryx-pair-running-through-spring-flowers-west-coast-125584310.html?imageid=86EB619E-ADDE-42BF-A3F5-11AD3FF7BAD5&p=361664&pn=1&searchId=010f32aaa73c04f1f11960177dc59d66&searchtype=
https://www.inaturalist.org/observations/126996895

SWIMMING

Alces alces is one of the most proficient swimmers among ruminants (https://northernontario.ctvnews.ca/northern-ont-logging-crew-rescues-moose-that-fell-through-the-ice-1.5774814 and https://www.youtube.com/watch?v=AuZ0znvhyeE and https://www.youtube.com/watch?v=60m6keIWNW0 and https://www.youtube.com/watch?v=tIqwUX6n1o0 and https://www.youtube.com/watch?v=59_wBH2hSBQ and https://www.youtube.com/watch?v=ZGDrNNKA30M).

It can dive so deep while foraging that it disappears completely below the surface (https://www.montanaoutdoor.com/2020/11/big-bull-moose-disappears-underwater-video/ and https://www.youtube.com/watch?v=BvVjFqgdEiI).

This is consistent with its foraging niche, which combines browsing on the foliage of trees and shrubs with holding its breath and submerging its head for aquatic algae and herbaceous plants.

By contrast, T. oryx has hardly been recorded swimming, or even foraging in wetlands. The following (https://www.youtube.com/watch?v=XTA1FmyVkDY) suggests that it is capable of swimming, which makes sense given that its tragelaphin relatives are known to be capable swimmers.

However, T. oryx

is generally associated with dry climates,
tends to avoid the dense vegetation near rivers, and
is not known to take refuge in water when pursued by predators.

DISCUSSION

Estes (1991, page 190) states:
"Elands only gallop when badly frightened (or playing) and if pursued quickly tire...a trot...is the eland's fastest gait under usual conditions; it can trot at a rate of 35 kph for several kilometres, or much further at a slower rate...Is it the eland's bulk that makes it slow, or is it simply a tragelaphine trait the eland has been unable to change in adapting to open habitats? Cows are not particularly bulky and certainly calves not at all, yet both are slower than other plains antelopes. The fact that elands, like their closest relation, the kudu, are incredible high jumpers is also against the bulk argument. Fleeing elands often display their prowess...by jumping effortlessly right over a neighbor, and youngsters can sail over a 3 m fence from a standing jump".

ILLUSTRATIONS OF ALCES ALCES IN KNEELING POSTURES

https://gf.nd.gov/magazine/2020/feb/kneeling-moose

https://www.flickr.com/photos/annkelliott/18675993910

https://stock.adobe.com/images/kneeling-moose/210074588

https://www.alamy.com/a-cow-moose-feeding-on-some-green-grass-image3090710.html?imageid=E08948EE-59AF-47DF-A106-73F150DEC73A&p=166893&pn=1&searchId=f41a66b0ac6853a16f070809256f1de5&searchtype=0

https://www.inaturalist.org/observations/46888551

http://www.silencelikethunder.com/kneeling-moose.html

https://www.reddit.com/r/alaska/comments/hhwh1k/moose_kneeling_to_eat_in_my_side_yard/

https://www.istockphoto.com/photo/moose-kneeling-to-eat-green-grass-gm859156154-142020203

https://www.alamy.com/young-moose-kneeling-on-hydroseed-by-the-dalton-highway-alaska-usa-image64756672.html?imageid=8F50A8CE-928B-4673-B30E-7C7E4BBC53F3&p=410512&pn=1&searchId=f41a66b0ac6853a16f070809256f1de5&searchtype=0

https://www.alamy.com/stock-photo-a-moose-calf-alces-alces-kneels-down-at-the-edge-of-an-alpine-lake-74641850.html?imageid=29A2FE7B-09EA-4EAB-A517-109459C96FF7&p=95964&pn=1&searchId=f41a66b0ac6853a16f070809256f1de5&searchtype=0

https://www.alamy.com/moose-calf-alves-alces-7-months-old-eating-salt-a-winter-road-canadian-image69996792.html?imageid=FE4806B9-BF29-401F-A10B-E4115B5B7C1D&p=59158&pn=1&searchId=f41a66b0ac6853a16f070809256f1de5&searchtype=0