Ημερολόγιο της/του milewski

Stripes, whether pale or dark, can make animals hard to tell from their surroundings. However, horizontal stripes along the torso (as opposed to the classical vertical pattern seen in e.g. the tiger, Panthera tigris) are unusual in large mammals. Why is the gerenuk (Litocranius walleri) - the lankiest of all gazelles and indeed of all antelopes or deer - odd among ungulates in this way?

The gerenuk, in both sexes and at all ages from infancy, has a pale horizontal stripe running from the dorsal base of the neck across the flank to the rump (see https://www.canstockphoto.com/gerenuk-standing-upright-to-reach-leaves-59579540.html and http://christofftravel.com/Africa/Pages/Gerenuk.html). The colouration is otherwise unusually plain for a gazelle. Presumably the stripe breaks up the figure, helping the gerenuk to hide in the sparsely woody vegetation it inhabits.

The only other gazelles with a similar stripe are the female and juvenile blackbuck (Antilope cervicapra, see https://www.shutterstock.com/ja/image-photo/profile-female-blackbuck-antilope-cervicapra-known-1435032149) and some populations of the goitred gazelle (Gazella subgutturosa), typically in Azerbaijan (see http://cannundrum.blogspot.com/2017/10/goitered-or-persian-gazelle.html). In the latter, the stripe is merely a local variation of what, in most populations, is a pale band.

Horizontal pale stripes occur also in several genera of deer (Dama, see https://www.inaturalist.org/guide_taxa/889020; Cervus, see https://lightfieldstudios.net/3752748/stock-photo-sika-deer.html; Axis, see https://www.thehindu.com/thread/chasing-chitals-in-chinnar/article28249624.ece; and Rangifer, see https://www.latimes.com/travel/la-tr-tipsheet-where-to-see-reindeer-20181210-story.html) and in several species of the antelope genus Tragelaphus (see https://www.biolib.cz/en/image/id217766/ and https://www.zoochat.com/community/media/sitatunga-whipsnade-01-aug-2020.498179/). However, in these animals they tend to be mere details of complex patterns of spotting and striping.

Perhaps the unusual striping of the gerenuk arose partly because its torso is so often viewed in the vertical. Although various species of deer and antelopes can stand on the hindlegs to reach high on plants, the gerenuk is exceptionally able to remain free-standing for minutes without propping the forefeet on branches. And only the gerenuk rises bipedally so frequently that this seems to be its main posture in foraging. In the upright stance, the stripe tends to align with the main stems of the tall shrubs and short trees typical of the habitat of the gerenuk (https://www.alamy.com/stock-photo-gerenuk-litocranius-walleri-standing-on-the-hind-legs-at-a-shrub-and-76081293.html).

Two slightly less lanky species of gazelles, the dibatag (Ammodorcas clarkei) and the dama gazelle (Nanger dama), lack any stripe on the torso despite also foraging to some extent bipedally (e.g. see https://www.youtube.com/watch?v=psqN8oUycxQ). In the case of the dibatag, the typical habitat is dominated not by 'acacia' (Vachellia species such as tortilis) but instead by sundry tall shrubs in a distinctive vegetation type on sand, called 'gedguwa'. This form of 'open thicket' tends to be more cluttered with foliage at about one metre above ground than is the case in 'acacia scrub', leading to a subtle difference in visibility and thus in adaptive colouration of the gazelles in question. In the case of the dama gazelle, the habitat is seasonally far more open than that of the gerenuk, and the overall colouration is conspicuous, not camouflaging.

Most gazelles (genera Gazella, Eudorcas, Nanger, Litocranius, Antilope, Ammodorcas, Antidorcas and Procapra) are only moderately sexually dimorphic. Adult males are not strikingly larger than adult females but possess horns, or at least larger horns.

However, the blackbuck (Antilope cervicapra, see https://www.gettyimages.com.au/detail/video/female-blackbuck-antelope-walks-through-pool-on-stock-video-footage/918341978?adppopup=true) presents an intriguing comparison with the gerenuk (Litocranius walleri, see https://www.inaturalist.org/observations/62024073).

In both, adult females (compare https://www.alamy.com/stock-photo-female-indian-blackbuck-antelope-antilope-cervicapra-138074045.html with https://www.alamy.com/stock-photo-gerenuk-female-walking-48605348.html) are hornless and weigh about 30 kg, and female colouration is the least conspicuous among gazelles. The dark flank-band typical of gazelles has been lost; there is a pale horizontal stripe along the upper flank; there is pale around the eye but the pale facial stripe and dark malar stripe, both typical of gazelles, are minimal; and on the hindquarters the bold effect of most gazelles is lost because the pygal band is minimal, and the white on the buttocks is restricted (compare https://www.sciencephoto.com/media/389667/view or https://thewolfintelligencer.com/antelope-indian-blackbuck-antilope-cervicapra/#jp-carousel-10113 with https://www.inaturalist.org/observations/58316540).

Despite this uniformity of females and juveniles, the mature males could hardly differ more. In the gerenuk, males have modest horns and feminine colouration except on the crown of the head (see https://www.dreamstime.com/female-male-gerenuk-ong-necked-antelope-samburu-national-park-kenya-image137690107). In the blackbuck males grow extremely long, corkscrew horns; and their colouration is so converted into a whole-body dark-and-pale 'beacon' (see https://www.alamy.com/blackbuck-antelope-antilope-cervicapra-image279138319.html) that little remains of the pattern of gazelles. Even the face becomes so showily dark-and-pale (see https://www.gettyimages.com.au/detail/video/male-blackbuck-antelope-looks-around-on-grassland-stock-video-footage/918471144?adppopup=true) that it seems unrelated to the gerenuk and most gazelles.

This correlates with the ecological and social differences. The blackbuck is a specialised grazer which drinks frequently and concentrates in large groups, whereas the gerenuk is a specialised browser which can forgo drinking for years and often appears solitary. Both species are territorial, but in divergent ways.

In the blackbuck, territories are so small, crowded and hectically defended that the competing males show off to each other for most of the time. They alternate this with visual appeasement, because they can forage only by trespassing their way to nearby, untrampled pastures, excusing themselves gesturally along the way there and back. In the gerenuk, the territories are so large that males seldom even see each other. Not only do they not need to trespass, but they only ever patrol a limited central part of the territory - using smell rather than sight.

What this means is that - despite females being so similar - males have social modes so different that males of the blackbuck make no attempt to hide from predators, whereas those of the gerenuk remain as secretive as possible.

One would not expect the gerenuk (Litocranius walleri) to be particularly frequently photographed. It lives in remote areas, its populations are sparse, it is shy, its appearance is rather dull apart from a graceful lankiness, and it is uncommon in zoos.

In the sixties and seventies, few photos of the gerenuk were available. Pierre Dandelot and Helmut Diller, painting the species for the best field guidebooks of the seventies and eighties, erred considerably in their depictions, presumably because they had little material to examine. Yet today, photos on the Web are too many to keep track of. The gerenuk is surpassed only by the springbok (Antidorcas marsupialis) as the most frequently-photographed gazelle in the wild.

The blackbuck (Antilope cervicapra) has, understandably, been photographed extremely frequently. This gregarious and spectacular gazelle still occurs widely in semi-wild conditions in India, where a rising tech-savvy Middle Class, combining the regard for animals of Hinduism with that of the colonial English, has produced more wildlife photographers than expected in a poor country. The blackbuck is the most successful gazelle in zoos worldwide. It is also kept on hunting ranches in the USA and Argentina; there are more photos on the Web of the blackbuck in Texas alone than there are of most species of gazelles in any situation.

Such explanations hardly apply to the gerenuk. Instead, its obvious appeal is anthropomorphic: extreme bipedality for an ungulate while foraging (see http://tallday.co.uk/2015/09/17/gerenuk-almost-on-stilts/ and https://www.inaturalist.org/observations/66279951).

People love, in animals, not only upright bipedality (penguins) but also other anthropomorphic features such as flattish faces (many primates but not e.g. baboons), binocular placement of the eyes (cats and primates), and apparently smiling mouths (dolphins). Hypothetically any animal combining several of these features would be particularly photogenic, and the popularity of the suricate (Suricata suricatta) can largely be explained by its combination of bipedal standing/sitting, binocular placement of the eyes and a suggestion of a smile (see https://www.publicdomainpictures.net/en/view-image.php?image=39396&picture=suricate-or-meerkat-sitting). Penguins feature only bipedality, but they score in that they actually walk bolt-upright, and they also have the attractive dark/pale contrasts of a jacket and shirt.

The gerenuk does not walk bipedally, its eyes are on the sides of its head, and its facial profile is pinched and pointed. However, even this head, when viewed directly from the front, can appear somewhat childlike in perspective with the eyes large relative to the small face, and a hint of a smile (see https://www.dreamstime.com/royalty-free-stock-photography-antelope-close-up-detail-african-gerenuk-face-big-ears-image30627197). Thus anthropomorphism, more than the particular biological interest of a 'giraffe-gazelle', may help to explain the proliferation of photos of the gerenuk.

For some strange reason, the gerenuk (Litocranius walleri) and the impala (Aepyceros melampus) have similar colouration (see https://www.flickr.com/photos/helenehoffman/49518633298 and https://www.alamy.com/stock-photo-two-baby-impala-antelopes-in-the-african-bush-29057562.html). This is true despite the fact that the two species are unrelated phylogenetically and differ ecologically, seldom occurring together.

The gerenuk is specialised to forage with upright bipedality, can forgo drinking for years on end, and tends to be solitary. By contrast, the impala has not been observed even to prop its forelegs on a plant stem, must drink nearly daily, and is gregarious. In the narrow zone where the two species share the same landscapes in Kenya, the gerenuk prefers thorn scrub while the impala prefers grassland.

Perhaps the gerenuk has come to mimic the colouration of the impala for protection against predators. The impala tends to be common where it occurs, while the gerenuk is everywhere scarce. Based on the likelihood that the gerenuk is not as enduring a runner as the impala, a naive predator might be misled to turn down hunting opportunities after spotting the gerenuk.

However, there are obvious problems with this explanation. The gerenuk is so much lankier than the impala that the colouration could hardly disguise its identity (see https://megapixl.com/impalas-aepyeros-melampus-and-gerenuk-or-waller-s-gazelle-litocranius-walleri-samburu-park-in-kenya-stock-photo-197574302). Most of the distribution of the gerenuk, on the Horn of Africa, is hundreds or thousands of kilometres from the nearest impala. And there is scant convergence between the two species in their behaviour in alarm. For example, the gerenuk habitually trots (see https://www.istockphoto.com/photo/female-gerenuk-running-gm1143342081-307019225) while the impala is remarkable in how reluctant it is to trot; and the stotting behaviours, in display to scanning predators, are extremely different (see https://www.youtube.com/watch?v=ru-lwzg-rPk and https://www.youtube.com/watch?v=xjb6hStBahg).

Here is a frontier of understanding in Biology, where new hypotheses are needed. I can help by describing in detail the subtle similarities and differences in colouration, which will be the topic of my next post...

In my last Post I mentioned the oddly convergent patterns of colouration in the gerenuk and the impala. The following list shows that the similarities are enough that, were it not for the obviously long neck and small face of the gerenuk, it would be hard to tell the two species apart at any distance (see https://www.alamy.com/stock-photo-male-gerenuk-walking-past-impala-52531218.html and https://www.alamy.com/impalas-aepyceros-melampus-and-gerenuk-or-wallers-gazelle-litocranius-walleri-samburu-park-in-kenya-image344576107.html).

Both species have dark fawn on the dorsal surface of the torso, giving way to paler fawn on the flanks and then white on the belly, the borders between the zones being oddly crisply defined. The main difference is that in the gerenuk the back/flank border is accentuated enough to look like a pale horizontal stripe in its own right. However, another difference is that only the impala possesses a dark spot of bare skin at the stifle-fold.

The pattern of double-white on the chest is remarkably similar between the gerenuk and the impala, while different from other ruminants (see https://i.redd.it/7zntk0098v221.jpg and https://www.inaturalist.org/observations/6484988 and https://depositphotos.com/44113279/stock-photo-impala.html and https://www.amazon.com/African-Impala-Antelope-River-Journal/dp/1540354911).

The face, like the chest, has patterns too similar in detail to have arisen merely by chance. However, the back of the head differs in that only the impala possesses dark posterior ear-tips and a sheeny-haired crown which switches from fawn to silvery in some lights (https://www.youtube.com/watch?v=9CGoEJtE9X8).

The legs have similar colouration except, for example, that the dark gland-tuft is on the foreleg of the gerenuk, vs the hindleg of the impala. Only the impala has white pasterns, visible when the animal stands on bare ground or close-cropped lawn (https://www.facebook.com/keith.ladzinski/posts/2708823022476328/ and https://depositphotos.com/150252040/stock-photo-young-impala-baby-stands-and.html).

As for the hindquarters, the patterns are different in detail (e.g. only the impala has dark pygal stripes) but give a similar overall impression of inconspicuous vertical bars of whitish near the tail (see https://stock.adobe.com/cy_en/search/images?k=gerenuk&asset_id=30423210 and https://commons.wikimedia.org/wiki/File:Impala_ewe_behind.jpg).

The tails differ in various inconspicuous ways (see https://animalworld.tumblr.com/post/189703937600/gerenuk-male-preening-tail-litocranius-walleri and https://www.alamy.com/stock-photo-impala-aepyceros-melampus-tail-south-africa-krueger-national-park-76137298.html). Whereas the terminal, insect-swishing tassel is white and relatively large in the impala vs blackish and small in the gerenuk, only the gerenuk can flare the white fur on the posterior haunch.

Given that the specialised proportions of the gerenuk should make it easy for predators to recognise, what could the adaptive advantages of the similarities in colouration possibly be?

There have been many historical attempts to domesticate both sheep (Ovis) and gazelles (Gazella). What many naturalists may not realise is that several types of ostensibly wild gazelles seem to have originated, at least partly, by selective breeding in captivity.

The species/subspecies bilkis, dareshurii, erlangeri, farasani, hamishi and muscatensis seem never to have been found in wild populations, although three of these now occur free-range on small islands in the Red Sea or Persian Gulf (e.g. see https://www.researchgate.net/figure/Photographs-of-gazelles-from-the-Farasan-Islands-A-male-and-C-female-and-mainland_fig2_262386010). I suspect that all are anthropogenically modified variants of the mountain gazelle (Gazella gazella), transported by humans to their locations and at least partly bred in captivity in the past. Even in its original range (in the Levant), the mountain gazelle has long lived somewhat commensally with tolerant farmers because no truly wild situations have remained in the Biblical Lands for hundreds of years.

The main effects of quasi-domestication in the above gazelles seem to be darkened colouration (e.g. see https://awwp.alwabra.com/?p=1177 and https://awwp.alwabra.com/wp-content/uploads/2014/04/Adult_Male_Erlangeri_Gazelles_01.jpg and https://www.biolib.cz/en/taxon/id726062/) and a reduction in the size of the brain.

Four North African species (cuvieri, dorcas, pelzelni and leptoceros) seem to have been kept in captivity, for example in oases in Tunisia and elsewhere in the Maghreb, for thousands of years. Probably in most cases the animals were caught as wild infants and hand-reared, to be kept as pets but not selectively bred. However, an odd aspect of Cuvier's gazelle, in addition to a dark colouration in some populations, is that in some individuals there are irregular whitish markings on the face (see https://kaymeclark.photoshelter.com/image/I0000lvhyKY5PUks and https://www.biolib.cz/en/image/id315675/), reminiscent of the asymmetrical colouration so often produced inadvertently by domestication.

Certain traditions in India have long cared for the chinkara (Gazella bennettii), which lives somewhat commensally as well as often being raised as a pet. However, no aspects of colouration suggest that this species has been modified by this relationship.

Why did domestication of gazelles prove so unsuccessful that most naturalists assume them to be purely wild animals? Possibly because all gazelles, unlike all wild sheep, have a territorial social system, which limits their amenability to herding. None of the twelve species of domestic hoofstock originating in Eurasia have territorial wild ancestors.

Which leaves us with an odd thought. Had things turned out differently and Gazella domestica arisen in place of sheep, would the Bible have spoken of gazherds rather than shepherds?

Antilopin bovids range widely in dry climates in Africa and Asia, but their greatest concentration of genera and species occurs in arid to semi-arid northeastern Africa (Somalia, Somaliland, Djibouti, Eritrea and parts of Sudan, Ethiopia and Kenya). In an area the size of France plus Spain located just north of the equator, seven species of gazelles, the beira (Dorcatragus megalotis, see https://www.researchgate.net/figure/Adult-beiras-photographed-in-the-study-area-male-on-the-left-hand-female-on-the-right_fig2_229180699), and eight species/subspecies of dikdiks (Madoqua) occur. This is more than all the species of bovids, antilocaprids and deer in the whole of the United States of America.

One clue to the reasons for such diversity is the poverty of antilopins in the similarly extensive arid to semi-arid climate in southern Africa, just south of the tropic of Capricorn, where only two species of antilopins occur in the relevant parts of Namibia, Botswana and South Africa. These are the springbok (Antidorcas marsupialis, which is a type of gazelle, see https://www.123rf.com/photo_46595145_springbok-in-desert-land-in-etosha-national-park-namibia.html) and the steenbok (Raphicerus campestris, which is intermediate in size between dikdiks and the beira, see https://www.alamy.com/male-steenbok-antelope-raphicerus-campestris-kalahari-desert-south-africa-image210577561.html).

That is a dozen species versus a couple, in ostensibly similar environments on the same continent.

The antilopins of arid to semi-arid northeastern Africa include some of the most peculiar and specialised of ruminants. The gerenuk (Litocranius, see https://parody.fandom.com/wiki/Gerenuk?file=Gerenuk_%2528Buck%2529.jpg) has an exceptionally small face, an unrivalled ability to free-stand upright on its hind hooves, and such extreme economy of water that it refuses to drink even when raised in zoos. A small species of dikdik (see https://naturerules1.fandom.com/wiki/Salt%27s_Dik-dik?file=93360711.lkzJWrfQ.jpg) is the smallest of all ungulates, worldwide, that live in semi-desert. And Speke's gazelle (Gazella spekei) has an oddly inflatable nose even compared to its closest relatives within the same genus (see https://naturerules1.fandom.com/wiki/Speke%27s_Gazelle?file=3e12ca6933ee751da51ada5379c67733.jpg).

Both parts of Africa have similar mean annual rainfall, but a crucial difference is this. Whereas the dry parts of southern Africa have a single rainy season each year (see https://www.safaribookings.com/karoo/climate), those of northeastern Africa tend to have two rainy seasons each year (see https://cdn.hikb.at/charts/meteo-average-weather/garissa-meteo-average-weather.png) owing to the East African Monsoon. This means that plant growth tends to be more reliable in the northeast than in the southwest of the continent, allowing greater specialisation of body sizes and shapes, diets, and foraging heights, and thus ecological niches. Whereas in general arid climates mean that 'beggars cannot be choosers' and survival depends on versatility, the antilopins of northeastern Africa include some of the choosiest species of ungulates known with respect to type of terrain, vegetation and diet.

Everyone knows that the impala (https://en.wikipedia.org/wiki/Impala) bounds in a striking way (https://upload.wikimedia.org/wikipedia/commons/8/8f/Impala_AdeFrias.jpg and https://www.storytrender.com/114774/antelope-jumps-so-high-it-reaches-the-height-of-an-elephant/).

However, how many realise that this species - looking like a normal antelope but with an ancient and distinctive origin - is more aberrant in other aspects of its locomotion and postures?

The impala is puzzlingly reluctant to trot.

This standard gait is taken for granted in most unguligrade and digitigrade mammals (e.g. https://www.gettyimages.com.au/detail/video/with-grants-gazelle-trotting-along-shoreline-with-stock-video-footage/691461859 and https://www.youtube.com/watch?v=V1mlwzMH4kM), and in some of them exaggerated into a form of stotting called 'style-trotting' (e.g. https://www.youtube.com/watch?v=-8R2Lp_S9Es).

One of the few times when the impala trots - and then for only a few steps - is when a courting male approaches a female over a short distance (see https://www.youtube.com/watch?v=deTFxRWrnKM).

The reluctance of the impala to trot is more odd than its bounding. This is because an ecological counterpart in India, the blackbuck (Antilope cervicapra, https://en.wikipedia.org/wiki/Blackbuck), frequently trots (https://www.dreamstime.com/black-buck-adult-male-portrait-close-up-green-bucks-resident-species-gujarat-india-found-many-places-big-image184881075) in addition to bounding high and far (see https://www.gettyimages.com.au/detail/video/pronking-blackbuck-females-run-and-leap-on-indian-stock-video-footage/1B02605_0001 and https://www.reddit.com/r/NatureIsFuckingLit/comments/ax7t1s/jumping_skills_of_this_black_buck_is_on_point/).

What truly is distinctive of the impala is a gait that I call kick-stotting (https://www.youtube.com/watch?v=JOAGylDP18g and https://www.alamy.com/stock-photo-impalas-aepyceros-melampus-leaping-16555737.html).

Many types of antelopes and deer stot (e.g. https://www.dreamstime.com/black-buck-baby-jumping-mid-air-greenery-bucks-resident-species-gujarat-india-found-many-places-big-groups-image184881477 and https://www.birdsoutsidemywindow.org/2014/06/17/stotting/ and https://www.shutterstock.com/nb/video/clip-5775500-hartebeest-pronking-side-view) in response to the approach of predators. These include the kob (Kobus kob, https://www.youtube.com/watch?v=80d1K5FSilg) - which ecologically replaces the impala north of the equator - and gazelles (e.g. https://www.reddit.com/r/Awwducational/comments/2mzdfk/stotting_is_a_behavior_amongst_gazelles_in_this/).

However, the kick-stotting of the impala differs in form and has yet to be explained in function.

As it runs, the impala flings its hind legs high in unison - in some cases so high that it seems to risk somersaulting - while waving its tail high as well (see https://www.youtube.com/watch?v=xjb6hStBahg and https://www.youtube.com/watch?v=0PFq4l_v1iI).

Many naturalists have watched kick-stotting in social play, but few have seen it in serious situations. Since social play is rehearsal, there is presumably a real, life-or-death purpose to stotting in the impala as in other species.

I have noticed that another of the few times when the impala trots is in slowing down to a halt after a bout of playful kick-stotting (see https://www.youtube.com/watch?v=X6Gtjcl6sm4).

When charged by most types of predator, the impala does not stot. The limited evidence hints that kick-stotting in earnest may be reserved for the African hunting dog (Lycaon pictus, https://en.wikipedia.org/wiki/African_wild_dog).

M Burton, in an article titled 'Impala behaviour' (Black Lechwe 4(4), pp. 46-48) states: "Impala sometimes use a similar action (to kick-stotting), as when one is chased by a dog. This it soon outdistances, and then it will proceed for a short distance bouncing on stiff legs before resuming the normal method of progression...the conspicuous black and white markings on the rump...are more prominently displayed in moments of excitement".

All bovids and deer can swim, but the impala is among the most inept in the water (see https://www.youtube.com/watch?v=onAE9aJi9qU and https://www.youtube.com/watch?v=DXQc_v5qjS4). This was first noticed in the mission to rescue animals stranded on islands during the filling of Kariba Dam on the Zambezi River (https://www.facebook.com/watch/?v=204822168221559 and https://m.facebook.com/watch/?v=2910628739265279&_rdr).

The impala often lives along river banks where it must risk being chased into the water by predators. So it seems odd that gazelles that spend their lives far from rivers can - if needs be - swim more confidently than the impala (e.g. https://tenor.com/view/gazelle-swimming-escape-gazelles-croc-gif-9565007).

The maximum competence of the impala when immersed can be seen in https://www.youtube.com/watch?v=Yp4P3mxhomc and https://www.youtube.com/watch?v=WjEmeqrka88.

The impala seems unwilling to rise on its hind legs to forage, even in drought when the only remaining food is high on branches. The blackbuck specialises more on herbaceous plants and is thus less likely than the impala to seek the foliage of shrubs for food. Yet females of the blackbuck sometimes rear up on their hind legs to flail at each other with their hooves, which has not been observed in the impala.

Once the suckling juvenile reaches a certain size, it needs either to kneel or to splay its fore legs to reach the teats. In the impala the posture adopted is splaying (http://www.africaimagelibrary.com/media/29045c02-d8e0-480f-af5d-8c67d32dc7c4-impala-aepyceros-melampus-lake-mburo-national-park-uganda) - which is unremarkable because various bovids and cervids do the same. However, this posture undermines the idea that the impala is related to alcelaphins (https://www.canstockphoto.com/red-hartebeest-and-suckling-calf-56774538.html), which kneel while suckling in common with hippotragins and e.g. the nilgai (https://www.youtube.com/watch?v=IteLEYGUKAU).

Finally: even in the case of lying down to chew the cud (https://www.inaturalist.org/observations/97303706), the impala seems odd.

Most other antelopes and deer are easy enough to spot lying down by day. However, the adult impala tends to remain standing during its midday rest, reserving its recumbency for the secrecy of night - which it tends to spend in certain open places away from vegetation. Perhaps this explains why there are few photos in iNaturalist of the impala in a lying position?

to be continued in https://www.inaturalist.org/journal/milewski/67632-locomotory-and-postural-peculiarities-of-the-impala-part-2...

It has long been realised that the impala (Aepyceros melampus) is something of a 'living fossil', unrelated to antelopes of superficially similar appearance. However, what seems to have been overlooked is the versatility of the tail relative to other bovids.

The impala normally hides its tail, tucking the tassel between the legs more than occurs in other ungulates including gazelles and the blackbuck (Antilope cervicapra). This is consistent with the peculiar striped pattern on the posterior of the haunches (see https://www.naturepl.com/stock-photo-impala-tail-pattern-aepyceros-melampus-kruger-np-south-africa-nature-image01068237.html), which helps to make the whole animal inconspicuous in the sense of disruptive colouration (roughly equivalent to camouflage). Many ungulates have boldly-marked hindquarters, conspicuous from a distance, and many others have plain hindquarters which blend into the surroundings, but the impala is unusual in blending into the surroundings by means of hindquarters marked similarly to the stripes of the tiger (Panthera tigris). The habitual hiding of the white tassel makes sense in this context.

The impala does frequently display its tail in certain behaviours, but in doing so reveals the caudal anatomy to be unlike that in any other ruminant. Firstly, the long white hairs are piloerected either laterally (as in masculine displays in which the tail looks like a white fan, see https://www.alamy.com/stock-photo-impala-buck-with-tail-raised-and-making-sounds-territorial-behavior-48605342.html) or vertically (as when the tail is flicked up synchronously with the kicking of the hind legs in kick-stotting, see https://www.gettyimages.com.au/detail/photo/impala-leaping-royalty-free-image/10190009 and https://www.inaturalist.org/observations/3739299 and https://www.flickr.com/photos/wild_images/50904109398/). Secondly, it is the ventral surface of the tail on which the vertical piloerection occurs - unlike the tails of various antelopes, including gazelles, on which any vertically-arranged tomahawk-like hairs (usually black) are on the dorsal side.

The impala also shows the white tassel when shooing insects and when passing urine or faeces, but in these cases there is no piloerection in either of the orientations described above (e.g. see https://www.wikiwand.com/en/Impala).

Surprisingly, the length of the tassel differs between the two main subspecies of the impala. Many species of ungulates show subspecific variation in various features, but it is rare for the tails to vary much within a given species. In the black-faced impala (Aepyceros melampus petersi) the tail is so large (eg. see https://m.facebook.com/ongavalodge/photos/a.914545575258908/2515148435198606/?type=3 and https://www.canstockphoto.com/black-faced-impala-ram-showing-its-tail-87852614.html) that it seems to possess an additional curve in its shape as seen in kick-stotting. When tucked between the legs, the tail-tip reaches as far as the prepuce, instead of merely the scrotum.

The impala (Aepyceros melampus) is a strange antelope appearing as an ordinary antelope. This is partly because its peculiarities are small-scale anatomically, such as the nature and arrangement of its fur and the grooming apparatus of its teeth and gums.

Little-known is how odd it is that the impala can roar without obvious modification of the larynx (see https://www.youtube.com/watch?v=IqzBw9EWhjU and https://www.youtube.com/watch?v=X5ifYPaDGXo and https://www.youtube.com/watch?v=B0Kxm5kgLMA and https://www.youtube.com/watch?v=6fBYZDOUJas and https://www.youtube.com/watch?v=Ywnn2TPMyDQ and http://ourlifeinkruger.co.za/2019/05/21/the-rutting-season-when-impalas-get-busy/).

Several ruminants are, like the impala, capable of roaring or loud grunting during masculine display, i.e. during the rut (https://onlinelibrary.wiley.com/doi/10.1111/j.1469-7580.2011.01361.x). These are the goitred (Gazella subgutturosa) and mongolian (Procapra gutturosa, see https://link.springer.com/chapter/10.1007/978-4-431-76933-0_1 and http://www.bioacoustica.org/expedition/dauria2017_eng.html) gazelles and the red (Cervus elaphus, see https://www.youtube.com/watch?v=RduhVcBn-0M and https://www.youtube.com/watch?v=K4aG93vZImg) and fallow (Dama dama, see https://www.youtube.com/watch?v=oxHvwrwyuso and https://www.youtube.com/watch?v=I9enG3Tz96E) deer.

However, all have obvious modifications of the larynx of the male in season. In the goitred gazelle and red deer the larynx descends so far that it can abut the sternum. In the goitred and mongolian gazelles the larynx is so enlarged that the very names of the species refer to the swelling. And in all these species, the larynx recoils far down the neck during roaring.

The rutting male impala shows none of these specialisations (see https://onlinelibrary.wiley.com/doi/full/10.1111/joa.13114 and https://pxhere.com/en/photo/1600574), yet manages to excel in several ways. It roars more loudly than the goitred gazelle, it roars as loudly while running as deer roar while standing, and it somehow intersperses its roars with loud snorts made non-vocally.

The result is that the impala is one of the loudest of ruminants while retaining a larynx which looks no different in the rutting male from that in the female. And the female impala has not, as far as I know, been recorded vocalising loudly, although she - like the male - certainly snorts loudly when alarmed by predators (see https://www.youtube.com/watch?v=U_GbVH8AOEY).