Walking gaits in Cervidae: all species of deer semi cross-walk, as opposed to the ambling seen in many bovids
INTRODUCTION
Ruminants walk with subtle gaits, difficult to classify and name.
As good an approach as any is initially to distinguish between an amble and a semi cross-walk.
(Readers, please note: watch all the videos linked in this Post in slow motion, by adjusting the 'playback speed' to 0.5 or 0.25.)
In an amble, the hind foot is lifted only after the opposite fore is placed (watch from 40 seconds in slow motion in https://www.youtube.com/watch?v=9EawG4Gu3Js).
By contrast, in a semi cross-walk, the hind foot is lifted before the opposite fore is placed (https://www.youtube.com/watch?app=desktop&v=hxkM96MYlb8).
The footfall-sequence, in reality, varies along a continuum, rather than dichotomously.
However the distinction between amble and semi cross-walk is significant because it potentially affects the balance/stability of the locomotion, as follows:
- in an amble, there is a moment in which two feet are off the ground on the same side (left or right) - which is relatively unstable because it is conducive to tipping to one side. whereas
- in a cross-walk (including a semi cross-walk), the two feet off the ground are on different sides, viz. left fore with right hind, or right fore with left hind - which is relatively stable because it maintains diagonal propping.
In the following video clips, adjust the playback speed to show the footfall sequence in slow motion:
- amble in Giraffa: https://www.youtube.com/watch?v=Vh2yveXTKaU
- semi cross-walk in Strepsiceros: https://www.youtube.com/watch?v=Rky362tL5kE
RESULTS FOR CERVIDAE
CERVUS
Cervus elaphus
semi cross-walking:
cross-walking (on unstable substrate):https://www.dreamstime.com/red-deer-hind-crossing-stream-water-close-up-winter-image138050863
semi cross-walking in masculine strutting:
https://www.shutterstock.com/da/video/clip-1017907225-wild-red-deer-stag-walking-bright-morning
https://www.shutterstock.com/image-photo/red-deer-rutting-season-737170840
cross-walking (on unstable substrate):
https://www.dreamstime.com/red-deer-hind-crossing-stream-water-close-up-winter-image138050863
Cervus canadensis
semi cross-walking
gait approaching an amble in Cervus canadensis:
ELAPHURUS
Elaphurus davidianus
semi cross-walking
https://www.naturepl.com/stock-photo-nature-image01617635.html
nearly ambling
https://www.staffanwidstrand.se/image/I0000NS9g.SH5Xug
https://www.shutterstock.com/da/video/clip-20558605-pere-davids-deer-herd-migrate-graze-elaphurus
DAMA
Dama dama
semi cross-walking:
https://www.sciencephoto.com/media/625669/view/male-fallow-deer-walking
https://www.youtube.com/watch?v=dBaW7gt4Z4M
https://www.shutterstock.com/da/video/clip-3399380867-lonely-little-deer-walking-forest-reserve-near
reaching the threshold of ambling, in the case of mature males, 'strutting' in rutting rivalry:
https://www.youtube.com/watch?v=5DuKpftCyYQ
RUCERVUS
Rucervus duvaucelii duvaucelii
semi cross-walk
https://www.shutterstock.com/da/video/clip-9660908-barasingha-buck-male-adult-lone-walking-dry
Rucervus duvaucelii branderi
a particularly clear illustration of semi cross-walking: https://www.youtube.com/watch?v=EBigKO906QY
RUSA
Rusa unicolor
semi cross-walking
https://www.shutterstock.com/da/video/clip-1088508487-two-sambar-deer-walking-tall-grass-chitwan
https://elements.envato.com/beautiful-male-sambar-rusa-unicolor-deer-walking-i-7B59BYH
https://www.shutterstock.com/da/video/clip-1092275273-full-shot-alert-male-sambar-deer-rusa
https://www.shutterstock.com/da/video/clip-3408587259-wide-shot-wild-male-sambar-deer-rusa
https://www.shutterstock.com/da/video/clip-1086299906-medium-shot-alert-male-sambar-deer-rusa
https://www.shutterstock.com/da/video/clip-1061061169-sambar-deer-rusa-unicolor-walking-camping-site
nearly ambling
https://www.shutterstock.com/da/video/clip-1043561428-male-sambar-rusa-unicolor-deer-walking-forest
AXIS
Axis axis
semi cross-walking
https://www.shutterstock.com/da/video/clip-1086407951-spotted-deer-walking-by-lake-grazing-on
https://www.shutterstock.com/da/video/clip-1106064753-young-fawn-walking-family-chital-deer-axis
ambling when walking slowly and intermittently
https://www.shutterstock.com/da/video/clip-1076048012-chital-axis-known-spotted-deer-walks-along
https://www.shutterstock.com/da/video/clip-1044612139-young-female-chital-spotted-deer-walking-fresh
CAPREOLUS
Capreolus capreolus
cross-walking
https://www.shutterstock.com/da/video/clip-1107586537-roe-deer-walking-grassy-field-on-cloudy
https://www.shutterstock.com/da/video/clip-1103477551-roe-deer-buck-walking-on-grass-leaving
https://www.shutterstock.com/da/video/clip-1083644467-roe-deer-buck-walking-away-on-meadow
ODOCOILEUS
Odocoileus hemionus columbianus
semi cross-walking
https://dpa730eaqha29.cloudfront.net/myedmondsnews/wp-content/uploads/2023/05/DSC_5193.jpg
Odocoileus virginianus
semi cross-walking
https://www.shutterstock.com/da/video/clip-1025463410-large-swamp-buck-white-tail-deer-walks-across
https://stock.adobe.com/images/walking-whitetail-deer-in-morning-sun/276397346
https://es.123rf.com/photo_32305362_deer-walking.html
https://www.fieldandstream.com/hunting/whitetail-deer-travels-200-miles/
https://dpa730eaqha29.cloudfront.net/myedmondsnews/wp-content/uploads/2023/05/DSC_5193.jpg
BLASTOCEROS
Blastoceros dichotomus
semi cross-walking
ALCES
Alces alces
semi-cross-walking
https://www.youtube.com/watch?v=VUVBr_jDBcQ
https://www.deviantart.com/nini1965/art/A-Young-Bull-Moose-Walking-in-a-Field-917220578
https://www.shutterstock.com/image-photo/profile-bull-moose-walking-onto-dirt-116525839
https://www.istockphoto.com/photo/bull-moose-gm1055977806-282201912
https://fineartamerica.com/featured/walking-bull-moose-randy-straka.html
https://pixels.com/featured/bull-moose-crossing-river-jack-bell.html?product=wood-print
RANGIFER
Rangifer tarandus
semi cross-walking
https://www.shutterstock.com/da/image-photo/sweden-beautiful-deer-midst-spring-tundra-1265271187
https://www.alamy.com/stock-photo-reindeer-walking-on-snow-83046876.html
https://www.alamy.com/reindeer-walking-on-the-road-in-norway-image339946013.html
https://www.sciencephoto.com/media/956135/view/reindeer-male-walking-across-beach-in-winter-iceland
DISCUSSION
I have found that - in contrast to bovids (Bovidae) - there is a surprising consistency in the walking gaits of deer (Cervidae).
In bovids, the tendency is that cover-dependent species semi cross-walk, whereas 'plains game' amble.
For example:
- cover-dependent bovids: https://www.istockphoto.com/video/kudu-walking-gm472607841-9062520 and https://www.inaturalist.org/observations/200510219
- 'plains game' bovid:
This is behaviourally and ecologically correlated, as follows:
- Cover-dependent bovids tend to be nocturnal, solitary, and camouflage-coloured, with eyes facing somewhat forward.
- By contrast, 'plains game' are adapted to living in the open, with diurnal habits, gregariousness, conspicuous colouration, and eyes so far on the sides of the head that they can scan behind and in front at the same time.
The pronghorn (Antilocapra americana) belongs to a different family, viz. Antilocapridae. It conforms to the 'plains game' category, and ambles accordingly (https://es.123rf.com/photo_30436381_pronghorn-antelope-p%C3%A9rez.html).
Impalas (Aepycerotini) are a particularly interesting tribe of bovids, combining a superficially deer-like appearance with multifaceted peculiarities (https://www.inaturalist.org/posts/89229-an-index-to-my-posts-about-the-impala-aepyceros-melampus#). They seem cover-dependent, but actually conform in various ways to the 'plains game'.
Accordingly, impalas consistently amble:
- Aepyceros melampus (https://www.africaimagelibrary.com/media/e65dbf22-af1f-4b33-aea2-6e62423423c9-impala-ram-aepyceros-melampus-gondwana-game-reserve-south-af and https://www.istockphoto.com/video/beautiful-and-graceful-impala-antelope-walking-in-savannah-gm932522106-255537892)
- Aepyceros petersi (https://www.alamy.com/black-faced-impala-aepyceros-melampus-petersi-male-walking-on-savanna-etosha-national-park-namibia-image396537387.html?imageid=130B7753-D07F-4AAF-B8A8-6326847D55ED&p=157727&pn=1&searchId=2cfc4de9b1aacee4e55653a80467551f&searchtype=0 and https://www.alamy.com/male-black-faced-impala-aepyceros-melampus-petersi-by-a-waterhole-etosha-national-park-namibia-with-a-marabou-stork-in-the-background-image214783551.html?imageid=C85EC920-7966-4CB3-97DA-59AF08E61176&p=564807&pn=1&searchId=2cfc4de9b1aacee4e55653a80467551f&searchtype=0 and https://www.alamy.com/stock-photo-black-faced-impala-103307129.html?imageid=59C58C12-F2FE-449E-B7D9-860AD06CF8CC&p=310217&pn=1&searchId=2cfc4de9b1aacee4e55653a80467551f&searchtype=0).
The bovid Kobus defassa is comparable in body size and shape to the cervid Cervus canadensis. However, the former sometimes ambles (https://front.motionarray.com/stock-video/common-waterbuck-walking-between-trees-1179445/ and https://www.alamy.com/stock-photo-walking-african-waterbuck-in-wild-78946808.html), whereas the latter barely achieves an amble under any circumstances:
In masculine display in the rutting season, Cervus elaphus and Cervus canadensis 'strut' by semi cross-walking, not ambling. By contrast, the corresponding gait in Dama dama is close to an amble, despite the fact that this species normally uses a semi cross-walk.