milewski's Journal

@capracornelius @paradoxornithidae @botswanabugs @koenbetjes @grinnin @matthewinabinett

Mediterranean-type climates, with dry summers and rainy winters, occur both in South Africa (shown in blue in https://www.researchgate.net/figure/Climatic-regions-of-South-Africa-48-note-the-climatic-variations-along-the-coastlines_fig1_334771843) and along the Mediterranean coasts of North Africa and the Levant.

And gazelles have been evolving and shifting their ranges across Africa and Asia for millions of years.

However, there is a puzzling faunistic difference:
No species of gazelle is indigenous to the southern area, whereas four species of gazelles are indigenous to the northern area, viz.

• Gazella cuvieri (https://www.inaturalist.org/taxa/42261-Gazella-cuvieri),
• Gazella dorcas massaesyla (https://www.researchgate.net/publication/351515966_The_Last-Surviving_Moroccan_Dorcas_Gazelle_Gazella_dorcas_massaesyla_Viability_Analysis_M'Sabih_Talaa_Reserve_Morocco),
• Gazella gazella gazella (https://animaldiversity.org/accounts/Gazella_gazella/ and https://www.cambridge.org/core/journals/oryx/article/plight-of-the-endangered-mountain-gazelle-gazella-gazella/EE6B2AC340EC8B04F6BC1F0AA9EB65A7), and
• Eudorcas rufina (https://en.wikipedia.org/wiki/Red_gazelle).

In South Africa there is one species of gazelle, namely the springbok (Antidorcas marsupialis, https://www.inaturalist.org/observations/35692467).

However, this species did not historically occur under the mediterranean-type climate of Western Cape Province (https://en.wikipedia.org/wiki/Western_Cape). The many observations in iNaturalist are of introduced populations.

Furthermore, even within its natural distribution in South Africa, the springbok avoids stony slopes, in contrast to

• Cuvier's gazelle in the Atlas ranges of Morocco-Algeria (https://www.inaturalist.org/observations/39161994), and
• the mountain gazelle, which is named after its association with stony slopes in e.g. Israel (https://www.inaturalist.org/observations/5305686).

Why/how have these disparities in habitat arisen?

One partial explanation involves competing members of the ruminant fauna.

A crucial difference is that the following species of South Africa have no counterparts in North Africa or the Levant:

• the grey rhebok (Pelea capreolus, see series of photos in https://animalia.bio/grey-rhebok),
• the mountain reedbuck (Redunca fulvorufula),
• the common eland (Taurotragus oryx, see http://shutterstock.puzzlepix.hu/kep/1614041191),
• the steenbok (Raphicerus campestris, see https://www.zoochat.com/community/media/steenbok-raphicerus-campestris.488693/), and
• the Cape grysbok (Raphicerus melanotis).

These species were, until recently, common in relevant environments at or near (https://www.researchgate.net/figure/Distribution-records-for-Southern-Mountain-Reedbuck-Redunca-fulvorufula-fulvorufula_fig1_325473215) the southwestern tip of Africa.

I hypothesise that, among them (i.e. collectively), these species arguably usurped the niche of gazelles.

The grey rhebok and mountain reedbuck prefer stony slopes, and the distribution of the former included most of the South African area of mediterranean-type climate. Both are similar in body size and partly similar in diet to the springbok.

The common eland is extremely large (adult female about 500 kg); the two species of Raphicerus are smaller than gazelles. However, all have diets that overlap those of gazelles enough for them to be potential competitors.

To 'rewild' the climatically similar stony slopes near Punta Almina (https://en.wikipedia.org/wiki/Peninsula_of_Almina) in the north and Cape Agulhas (https://en.wikipedia.org/wiki/Cape_Agulhas) in the south of the same continent, we would reintroduce

• to the northern area only Cuvier's gazelle, but
• to the southern area at least three species: grey rhebok, common eland, and steenbok/Cape grysbok.

It would be naive to expect that the same ecological function could be restored by simply 'reintroducing' the springbok in Agulhas National Park (see https://www.inaturalist.org/observations/3975537), even though it is by far the most similar antelope to Cuvier's gazelle that is available in the southern African fauna.

@mr_fab @geichhorn

Naturalists interested in the subspecies of the springbok (Antidorcas marsupialis) should read https://www.zobodat.at/pdf/Zeitschrift-Saeugetierkunde_46_0189-0197.pdf.

Although now forty years old, Groves (1981) is still the best reference on a topic which will probably never be resolved satisfactorily.

The basic problem is that zoologists failed to document the nominate subspecies.

The Karoo springbok (Antidorcas marsupialis marsupialis) occurred in Namaqualand, the Karoo, Eastern Cape, Free State, Gauteng, and the southern part of Northwest Province, with possible extensions into Mpumalanga, Limpopo, and even Kwazulu-Natal Provinces.

Despite the extreme abundance of this form, the museums of the world have virtually no study-skins and only a few skulls.

The Karoo springbok was, like the extinct quagga (Equus quagga quagga, https://en.wikipedia.org/wiki/Quagga), taken for granted until it was too late. Furthermore, far fewer specimens were collected scientifically than for the quagga.

We cannot know exactly what subspecies marsupialis looked like in its original, fully wild condition.

It is technically extinct because

• artificial reintroductions failed to discriminate against subspecies hofmeyri of the Northern Cape, and then
• there has been nearly a century of advertent and inadvertent selective breeding by farmers, partly to promote mutant genotypes.

Furthermore,

• body size is too adaptable to mean much, and
• the facial colouration varies individually in all the subspecies.

South Africa has thus lost its national mammal as a subspecific genetic entity. Even more unfortunately, we can never define what that entity was in the first place.

Our best guess, based on a south-north cline from the northeastern Karoo to Angola (Groves 1981), is that the springbok of the Klein Karoo and Eastern Cape

• had extremely short, smooth horns in females (https://www.alamy.com/stock-photo-springbok-ewe-with-lamb-in-the-karoo-south-africa-near-calitzdorp-21932842.html?imageid=21A477CA-80CB-4C9D-93A7-07A9E05E89AA&p=10608&pn=1&searchId=4113351e3b1f6f44ed6a2ce04ac7d525&searchtype=0), and
• was not as pale as conspecifics in the Kalahari.

Groves found only 33 study-skins of the whole species of the springbok to examine in the museums of the world. And, ironically, 39% of these came from Angola and Kaokoland (https://en.wikipedia.org/wiki/Kaokoland), which remain so remote that iNaturalist still features few photos from these areas.

More particularly, we have an odd situation:

Subspecies angolensis is the best-represented of all the subspecies, in terms of study-skins (13 of the 33 specimens examined by Groves). Subspecies hofmeyri, south of the Kunene River, in Kaokoland, is also surprisingly well-represented, by 7 study-skins.

However, ssp. angolensis is so poorly represented photographically that, to this day, no photo shows clearly the darkness of colouration that Groves found, relative to ssp. hofmeyri.

The false notion that angolensis extends to northern Namibia, including Etosha, seems to have arisen in Cain et al. (2004) Mammalian Species no. 753 (https://www.science.smith.edu/departments/Biology/VHAYSSEN/msi/pdf/753_Antidorcas_marsupialis.pdf).

Although ostensibly an authoritative summary of the springbok, Cain et al. (2004) misportrayed the subspecies by means of an erroneous distribution map.

In summary, the matter of subspecies has been botched in several different ways.

Therefore, my suggestion for iNaturalist:

• for observations from Angola, assume angolensis;
• for observations from northern Namibia, the Namib south of the Kunene River, Botswana, and the Kgalagadi Transfrontier Park, assume hofmeyri; and
• for all other locations, just identify to species-level, because the subspecies-status has been irretrievably compromised.

Also see:

https://www.inaturalist.org/posts/53665-is-the-golden-springbok-a-semi-domestic-sport-or-a-throwback-to-an-extinct-species#

https://www.inaturalist.org/journal/milewski/69433-do-wildlife-photographers-have-a-blind-spot-for-adult-females-a-test-using-the-springbok#

Rogerio Ferreira has just kindly supplied a good series of photos of the Angolan springbok (Antidorcas marsupialis angolensis), at https://biodiversidadeangola.com/mammals/scientific/9527.

As it turns out, the darkness of the pygal band and other markings, described by Groves (1981) and used as a partial justification to raise this taxon to full species-status by Groves & Grubb (2011), is not striking.

The appearance is, after all, similar to that of subspecies hofmeyri, bearing in mind that even within hofmeyri the specimens from southeastern Botswana were paler than those from Namibia (Groves 1981).

In my view, the possibility that the Angolan springbok is a separate species (Groves & Grubb 2011) has not stood up to scrutiny. And even as a subspecies it is not nearly as distinctive as the black-faced impala (Aepyceros melampus petersi) is from the common impala.

If so, how would this change our overall view of subspecific variation in the springbok?

We can accept that the female horns vary in length along a south-north cline culminating in angolensis. However, clinal variation is not the same as subspeciation.

If we assume that the geographical variation in body size and the relative length of the legs is phenotypic as much as genotypic in the springbok, what emerges is a species more noteworthy for its uniformity than for its subspeciation.

It seems possible, after all, that the original wild springbok of the Eastern Cape, near Addo, was similar enough (except for the shortness of the female horns) that most naturalists would not have noticed any differences in body size or colouration from the Angolan springbok, 2500 km and 20 degrees of latitude distant.

And what this would mean in turn is that, although Science let the nominate subspecies disappear as a 'pure' entity before it was properly described, Nature may have, as it were, let us off lightly for that lapse.

The two commonest antelopes in southern Africa are the impala (Aepyceros melampus) and the springbok (Antidorcas marsupialis).

Here are some similarities and differences, perhaps new to readers.

In flight from predators, both bound high into the air even where there are no obstacles, while being perversely reluctant to jump over fences (instead preferring to crawl under them).

However, the impala differs from the springbok in that it neither stands on its hind legs to reach food, nor habitually uses a trotting gait.

Both have complex tails with two tracts of erectile fur, unlike any other ungulate.

However, in the impala the piloerections are to the side and inwards (ventral), whereas in the springbok the piloerections are outwards (dorsal).

Both have one infant at a time and form creches.

However, the impala has a flesh-coloured udder with four teats, whereas the springbok has a black-pigmented udder with two.

Furthermore, the hiding period of infants is shorter in the impala than in the springbok.

This is the opposite of expectations, given that the impala is the more cover-dependent of the two species.

Both use urine and faeces for social communication, with similar postures.

However, of the two species, the territorial male piles its feces only in the impala.

Both

• are exceptionally noisy for antelopes in masculine display, despite the normal appearance of the larynx; and
• can utter their loudest calls while running after rival males (in contrast to deer which roar, bellow or grunt only while standing or walking).

However, only the impala intersperses its rutting vocalisations with alarm-snorts.

Both have patterns of colouration similar to gazelles.

However, that of the impala has a mainly camouflaging effect (see https://www.inaturalist.org/observations/78256383 and https://www.inaturalist.org/observations/77839882 and https://www.inaturalist.org/observations/75215507 and https://www.inaturalist.org/observations/67019115).

This is odd for a species in which even infants are gregarious.

By contrast, the colouration of the springbok is more conspicuous (see https://www.inaturalist.org/observations/47788993 and https://www.inaturalist.org/observations/43308762 and https://www.inaturalist.org/observations/40749849 and https://www.inaturalist.org/observations/37893316 and https://www.inaturalist.org/observations/37480920) than any species of Gazella.

Both stot in earnest seemingly only for the painted hunting dog (Lycaon pictus). Both the kick-stotting of the impala and the head-down bouncing of the springbok are styles unique to these species.

However, only in the springbok is the same stotting display also performed by females in courtship.

All of Biology depends on precise use of words. And much of what makes the thousands of photos of ungulates and other mammals on iNaturalist interesting is the patterns of colouration.

However, words to describe adaptive colouration have been so imprecise and inconsistent that it has been hard to write coherently on this topic. Both the anatomical parts and the features of colouration have been misnamed, or named by misleading analogy.

Describing the carpal joint - which is actually homologous with the human wrist - as the 'knee' is a gross example of this. Confusion between the withers and the shoulders is a subtler but equally important case. However, in anatomy at least the correct word already exists in most instances.

When it comes to important aspects of colouration, we may not progress in our thinking until we fill in the gaps in our lexicon. Here, I introduce three new terms: the bleeze, the flag and the semet.

Various ungulates feature a conspicuous pale patch, informally called a 'blaze'. A typical example is the blesbok (Damaliscus pygargus phillipsi), named after the Afrikaans translation of blaze.

The metaphor applies because flame can be eye-catchingly pale, lighting things up. However, should we not avoid ambiguity with combustion?

I suggest the new term 'bleeze', applicable only to the biological topic of colouration.

We would describe the face of the blesbok as featuring a bleeze, not a blaze. An additional advantage is that bleeze can be defined to include conspicuously dark features, such as that on the face of the eastern white-bearded wildebeest (Connochaetes albojubatus) - or the juvenile of the blesbok itself.

The bleeze is, by its nature, large and prominent enough to be conspicuous to scanning predators even when the animal stands still.

However, many large mammals feature pale/dark patterns which are easily overlooked until moved or raised. For these dynamic forms of advertisement, I suggest the term 'flag' instead.

The most obvious examples are the caudal flags of the many species of ungulates which raise dark/pale tails in alarm or flight.

However, in principle the term can also apply to the ears (e.g. impala, Aepyceros melampus), the feet (e.g. bush duiker, Sylvicapra grimmia), or the buttocks (e.g. steenbok, Raphicerus campestris).

The third word, 'semet', is newly derived from the Greek word for sign (think of semaphore or aposematic).

This refers to dark/pale features too small-scale to matter for detection by potential predators, but moved in certain ways for social communication within the species.

For example, many species of ungulates have subtle markings around the mouth, or at the base of the ears, which help companions to monitor each other's cud-chewing or listening - facilitating the spread of any alarm within seconds.

And then there are many social signals involving colouration, used in rivalry and courtship, which are candidates for also being called semets.

Many ungulates deal with predation partly by being gregarious. Even 'solitary' species may rest in sight of a mate or parent when ruminating, i.e. chewing the cud. A 'strength in numbers' is that the group - whether it numbers two or twenty - functions as a unit in cooperative vigilance by virtue of its collective eyes, ears and noses. The moment one individual becomes suspicious and stops chewing to listen up, the apprehension needs to be communicated efficiently across the 'white noise' of the tooth-grinding of group-members.

The point of mutual monitoring of the routine chewing of companions would be to detect any interruption immediately, and to direct attention accordingly.

Ruminants have a problem which humans do not: they spend much of their lives, both night and day, noisily grinding the cud - while at the same time maintaining constant vigilance by hearing (e.g. see https://www.youtube.com/watch?v=E_Uc0kfNvgg).

The visual system of ungulates is extremely sensitive to slight movements. However, it makes adaptive sense that various species would have evolved patterns of colouration around the mouth to accentuate, or 'amplify', the chewing motion, particularly when the light is dim. And that each species would produce its own 'signature' pattern (e.g. see https://www.shutterstock.com/nb/image-photo/cud-chewing-deer-584459 and https://www.shutterstock.com/nb/image-photo/elk-chewing-his-cud-grasslands-82571272 and https://www.shutterstock.com/nb/image-photo/close-young-bull-elk-537907132 and https://www.inaturalist.org/observations/47408137).

Is it possible that, once the naturalist has a search-image along these lines, the diverse patterns of dark and pale on the lips, chin, cheeks and muzzle of various ruminants may start to look less like random scribblings of capricious Nature and more like functional adaptation?

Here are two examples to start with, illustrating the greater kudu (Tragelaphus strepsiceros, see https://www.storyblocks.com/video/stock/a-female-kudu-chews-cud-in-kruger-national-park-south-africa-e_zttsjf2eillivjey) and common eland (Taurotragus oryx, see https://depositphotos.com/58556159/stock-video-common-eland-head.html and https://www.alamy.com/elenantilope-taurotragus-oryx-nrnberg-bayern-deutschland-image69178906.html).

Note the puzzlingly intricate patterns on the faces. The dark muzzle contrasts with the pale mouth and chin, and there are dark or pale markings on the side of the mandible. Why would such patterns have arisen?

In my last Journal Post I proposed the new term 'semet' for patterns of this small scale, functioning at the close range of social communication but not mattering much to the large-scale conspicuousness or inconspicuousness of the figure to scanning predators. Because this particular category of semet concerns movements of the mouth, I propose calling it a 'buccal semet'. We can expect the buccal semets of the greater kudu and common eland to be dark/pale patterns unique to these species.

Of course the idea is still just hypothetical that ruminants are adapted to spot interruptions in the rumination of companions. And any new technical term may sound awkward at first. But could this open a new line of investigation into the subtle messages hidden in our photos of ungulates?

One of the puzzles of adaptive colouration in ungulates is the pale patches which occur at the bases of the ears of certain species, including species with otherwise plain colouration. Among the clearest examples of this are the moose (Alces alces) and the sambar deer (Rusa unicolor). As the ears swivel with changes in attention or emotion, the pale/dark contrasts flick in and out of view, potentially helping onlookers to read the mind, as it were, of the animal in question (e.g. see https://www.fruugoaustralia.com/close-up-view-of-a-cow-moose-and-calf-in-the-chugach-mountains-near-anchorage-southcentral-alaska-autumn-posterprint/p-8952313-19287570?language=en&ac=google&gclid=CjwKCAjwmK6IBhBqEiwAocMc8tCf4ZOTKDNT15CgLk-m4Fxhxrew6BnoyEUl8tunhn7Fkt0C7yAv_BoCfL0QAvD_BwE and https://www.wallpaperflare.com/moose-cow-portrait-close-up-profile-wildlife-landscape-wallpaper-aregm and https://www.canstockphoto.com/close-up-of-head-of-wild-female-moose-40347707.html and https://www.dreamstime.com/stock-photo-female-moose-close-up-grazing-bialowieza-poland-image74453735 and https://www.dreamstime.com/stock-photo-close-up-head-wild-female-moose-elk-alces-alces-largest-extant-species-deer-family-image83773415 and https://www.dreamstime.com/stock-photo-close-up-wild-female-moose-elk-head-eye-image67541853 and https://www.dreamstime.com/stock-photo-close-up-young-moose-found-wandering-around-town-north-idaho-image87320353).

Adaptive reasons for distinct patterns of colouration at the base of the ears include: a) group-members are likely to monitor the movements of each other's ears continually, so that when an individual becomes suspicious its companions can easily detect the direction in which it is listening; and b) in those species which use their ears for facial expressions, the posture of the ears can be emphasised by the 'makeup' of the colouration.

Ungulates may pay attention to each other's ears much as we humans pay attention to each other's eyes. The direction of gaze, and the pattern of movement of the eyes, imply where someone is directing his/her attention; we are extremely perceptive of the whites of the eyes because these accentuate even slight movements of the eyeball (see 'cooperative eye hypothesis' in Wikipedia). Ungulates lack a retinal fovea, and their visual system emphasises the scanning of a wide visual field for movement, rather than focussing on any particular object. This, together with the separation of the eyes on the sides of the head, means that ungulates would learn little from observing each other's eyes even if the whites showed. Instead, it makes sense to monitor the movements of the ears, which would make the pale/dark patterns on or at the base of the ears analogous to human eye-whites.

Please see https://www.youtube.com/watch?v=nEpCz_PMUss and https://www.dreamstime.com/canada-wildlife-close-up-wild-female-moose-close-encounter-beautiful-wild-female-moose-hiking-alberta-image168563330 and https://www.dreamstime.com/moose-head-close-up-profile-view-forest-winter-season-exposing-its-head-ears-eyes-mouth-brown-fur-moose-animal-image165345950 and https://www.dreamstime.com/stock-photo-moose-alces-alces-female-adult-feeding-young-spruce-tree-dense-forest-close-up-face-side-image60098607 and https://www.dreamstime.com/moose-head-close-up-profile-view-forest-winter-season-moose-animal-stock-photos-moose-animal-head-close-up-view-image165345955 for views of the pattern of pale and dark on the ears of the moose, a species which otherwise lacks clearly-defined markings. In the case of the sambar deer, the pale markings at the anterior and posterior bases of the ears are shown in https://www.istockphoto.com/photo/sambar-deer-rest-in-the-forest-gm1223158200-359185051, and https://unsplash.com/photos/84N-_6YXM1E and https://unsplash.com/photos/mJ_R9pxXdn4 are also illustrative.

In my last Post I introduced the concept and name of the buccal semet. In the case of the ears we once again have a semet, i.e. a definite pattern with potential for communication, but one too small-scale to matter for the overall conspicuousness/inconspicuousness of the animal to scanning predators. In this case, I propose the term 'auricular semet'.

@tonyrebelo @jeremygilmore @jwidness @oviscanadensis_connerties @tandala @beartracker @matthewinabinett @michalsloviak @capracornelius @colin25 @davidbygott @giorginatsvlishvili @koenbetjes

Also see https://www.inaturalist.org/journal/milewski/53340-differences-among-gazelles-in-the-structure-and-function-of-the-tail#

Two gazelles make for an obvious comparison, because they live on similar semi-arid plains in opposite hemispheres.

These are

• the goitred gazelle (Gazella subgutturosa (https://www.istockphoto.com/video/goitered-gazelle-gm473245265-26192520 and https://www.bbc.com/azeri/azerbaijan-44417549 and https://www.pedalverde.net/azerbaijan) of the Iranian region, and
• the springbok (Antidorcas marsupialis, https://www.inaturalist.org/taxa/42283-Antidorcas-marsupialis) of similar latitudes in South Africa,

Adults, while foraging, can hardly hide in these exposed environments (except for the goitred gazelle in saxaul 'forest' in winter, see comment below).

Because the two species are species are related, as antilopin bovids, and because they live in similar ecosystems, they might be expected to have similar adaptive colouration.

And indeed the two species are similar enough to be confused at first glance (https://www.inaturalist.org/observations/55091275 and https://www.inaturalist.org/observations/143913081).

Both have

• dark/pale banding on the flanks (https://gl.wikipedia.org/wiki/Antidorcas_marsupialis#/media/Ficheiro:Springbok_etosha.jpg),
• noticeable pale on the head and buttocks (https://zooinstitutes.com/animals/persian-goitred-gazelle-izmir-wildlife-park-118543.html and https://www.alamy.com/goitered-gazelle-jeyran-in-field-wildlife-nature-reserve-image419262426.html?imageid=E277963C-20F9-467F-9866-A90409FB9CB8&p=376800&pn=1&searchId=8667346e347106d14a3d1846caddb0e4&searchtype=0 and https://depositphotos.com/635532208/stock-photo-etosha-national-park-wildlife-namibia.html and https://www.dreamstime.com/royalty-free-stock-photography-herd-springbok-plain-image15137917), and
• bold displays of the hindquarters when locomoting (https://www.researchgate.net/figure/Goitered-gazelle-Gazella-subgutturosa-in-Shirvan-National-Park_fig6_259982825 and https://www.inaturalist.org/observations/10794991).

However, there are three main differences, viz.

• the colouration of the goitred gazelle (https://www.natureinstock.com/search/preview/goitered-gazella-gazella-subgutturosa-subgutturosa-touran-protected-area/0_12273093.html) is, overall, less conspicuous than that of the springbok (https://www.inaturalist.org/observations/133752316), and
• the goitred gazelle is extreme among antilopins in emphasising the dark tail (https://www.inaturalist.org/observations/117695149), whereas the springbok de-emphasises the tail and is instead unique in the size and form of a pale crest on its back and rump (https://www.inaturalist.org/observations/127282707).

In the following account, I have used photos from not only the Iranian region and South Africa, but also - owing to the scarcity of appropriate images - the rest of the distribution of both species.

STATIONARY:

A puzzling difference in adaptive colouration is that the goitred gazelle tends to blend into its surroundings, whereas the springbok tends to stand out even at a distance, and even when motionless.

What makes the springbok so conspicuous is its boldly dark flank-band, offset by white highlights on the face, rump, flanks, and upper legs (see https://www.inaturalist.org/observations/85319775 and https://www.inaturalist.org/observations/70140445 and https://www.inaturalist.org/observations/76488627 and https://www.inaturalist.org/observations/70682974 and https://www.inaturalist.org/observations/65916271).

In the goitred gazelle, the basic pattern is similar. However, it is toned down to a semblance of camouflage (https://commons.wikimedia.org/wiki/File:Ceyran.jpg and https://www.dreamstime.com/goitered-gazelle-jeyran-field-wildlife-nature-reserve-goitered-gazelle-jeyran-field-wildlife-nature-reserve-field-image216752540 and video in https://animalia.bio/goitered-gazelle and https://www.shutterstock.com/da/image-photo/gazella-subgutturosa-goitered-gazelle-1155749653).

Both the goitred gazelle and the springbok can have conspicuously pale faces, and in both species the paleness on the face develops as the animal grows.

However, the pale face is not as consistent among adult individuals in the goitred gazelle as in the springbok (please see https://www.inaturalist.org/journal/milewski/75660-the-facial-flag-of-the-springbok-antidorcas-marsupialis#).

In the goitred gazelle the pale face is associated more with the male (https://www.shutterstock.com/it/image-photo/gazella-subgutturosa-goitered-gazelle-1155749656
https://www.mediastorehouse.com.au/nature-picture-library/2020-august-highlights/goitered-black-tailed-gazelle-gazella-20172891.html and https://www.shutterstock.com/it/image-photo/gazella-subgutturosa-goitered-gazelle-1155749653) than with the female.

Even in mature males (https://www.dreamstime.com/persian-goitered-gazelle-male-female-background-latin-name-gazella-subgutturosa-persian-goitered-gazelle-male-image133088223), some individuals retain the inconspicuous pattern on the face.

MILD ALARM:

The goitred gazelle signals mild alarm by erecting the tail (https://www.inaturalist.org/observations/130903897 and https://www.inaturalist.org/observations/77627182 and https://www.inaturalist.org/observations/796456 and https://www.agefotostock.com/age/en/details-news-photo/200607-altay-june-7-2020-goitered-gazelles-are-seen-at-the-kalamayli-nature-reserve-northwest-china-s-xinjiang-uygur-autonomous-region-june-6/HTS-ZB950_351940_0069/1).

The dark tail is offset by the white buttocks (https://www.pond5.com/stock-footage/item/130905132-herd-goitered-gazelles-running-desert-001 and https://www.pond5.com/stock-footage/item/130905144-herd-goitered-gazelles-running-desert-004).

There is no analogous display in the springbok, which lacks a caudal flag.

STOTTING:

Both species greet scanning predators with stotting displays. However, these accentuate the differences in colouration.

The goitred gazelle stots by bouncing up and down with its black tail erect (https://www.istockphoto.com/video/running-goitered-gazelle-gm473245311-26196643 and video in https://discover.hubpages.com/education/Goitered-Gazelle-History-Facts-and-Efforts-to-Recover-It and https://www.diana-hunting.com/game/goitered-gazelle).

By contrast, the springbok has an extreme performance. It bounces while blazening an uniquely extended white double-crest that is normally folded invisible on the back, rump and buttocks (https://www.inaturalist.org/observations/134967922
and https://www.youtube.com/watch?v=jMIiB9DnRXg and https://www.atlasobscura.com/articles/watch-this-gazelle-spring-across-the-african-savanna-like-a-pogo-stick and https://www.gettyimages.ca/detail/photo/springbuck-jumping-in-the-air-kgalagadi-nature-royalty-free-image/976918234?adppopup=true).

However, the springbok does not always activate the crest while stotting:
https://www.gettyimages.ca/detail/photo/lone-springbok-on-plain-royalty-free-image/71587160?adppopup=true
https://www.alamy.com/stock-photo-springbok-antelope-antidorcas-marsupialis-jumping-south-africa-72687711.html?imageid=96BE87F5-378F-4276-9015-329CDA50F252&p=70019&pn=1&searchId=1ad475034542f64d7d819bbce4f5e724&searchtype=0

FLEEING:

Whereas the goitred gazelle flees and stots with its tail rigidly erect (see https://www.inaturalist.org/observations/504792 and https://www.inaturalist.org/observations/38272914), the springbok never even raises its tail to the horizontal while fleeing (see https://www.alamy.com/springbok-antidorcas-marsupialis-running-namibia-etosha-national-park-image151881858.html) or stotting (see https://www.shutterstock.com/da/image-photo/pronking-springbok-37655779 and https://elements.envato.com/springbok-pronking-in-the-grass-X6ZPTS2).

This difference makes sense partly because in the springbok the dark tassel is too small to add much to the display (https://www.inaturalist.org/observations/73197549), and the base of the tapering tail is less flexible than in the goitred gazelle (compare https://www.alamy.com/tail-end-of-goitered-gazelle-gazella-subgutturosa-image211285398.html with https://www.alamy.com/springbuck-springbok-antidorcas-marsupialis-herd-from-behind-namibia-image5449818.html).

The following show the goitred gazelle fleeing:
https://www.agefotostock.com/age/en/details-news-photo/200314-xilingol-march-14-2020-photo-taken-on-march-13-2020-in-sunite-right-banner-of-xilingol-league-north-china-s-inner-mongolia-autonomous/HTS-ZB950_348151_0039/1
https://www.inaturalist.org/observations/65033568
https://www.inaturalist.org/observations/139354386
https://www.inaturalist.org/observations/32798804
https://www.inaturalist.org/observations/17020189
https://www.inaturalist.org/observations/504792
https://www.flickr.com/photos/130113460@N03/41071430952
https://www.flickr.com/photos/wildlife_of_mongolia/14188610261
https://www.flickriver.com/photos/132934880@N02/48884899921/
https://www.shutterstock.com/it/image-photo/goitered-gazelle-jeyran-field-wildlife-nature-1972172735
https://www.naturepl.com/stock-photo-male-goitered-gazelle-gazella-subgutturosa-running-badkhyz-reserve-image01445599.html
Scroll to fourth photo in https://birdingmongolia.wordpress.com/2020/04/01/jargalant-mountains/

It is only when the fleeing is at full speed that the tail is lowered:
https://www.inaturalist.org/observations/612402
https://www.agefotostock.com/age/en/details-news-photo/200314-xilingol-march-14-2020-photo-taken-on-march-13-2020-in-sunite-right-banner-of-xilingol-league-north-china-s-inner-mongolia-autonomous/HTS-ZB950_348151_0049/1
https://www.inaturalist.org/observations/8282800
https://www.gettyimages.com.au/detail/photo/goitered-gazelle-also-known-as-gazella-subgutturosa-royalty-free-image/933847462
https://www.agefotostock.com/age/en/details-news-photo/200314-xilingol-march-14-2020-xinhua-photo-taken-on-march-13-2020-in-sunite-right-banner-of-xilingol-league/HTS-ZB950_348151_0052/1
https://www.mindenpictures.com/stock-photo-black-tailed-gazelle-gazella-subgutturosa-male-running-gobi-desert-naturephotography-image90136801.html

The following show the springbok fleeing:
https://www.gettyimages.ca/detail/photo/side-view-of-dog-running-on-field-arad-romania-royalty-free-image/1457183102?adppopup=true
https://www.inaturalist.org/observations/131774461
https://www.inaturalist.org/observations/112872508
https://www.inaturalist.org/observations/86033851
https://www.inaturalist.org/observations/73628357
https://www.inaturalist.org/observations/11258709
https://www.inaturalist.org/observations/11304134
https://www.inaturalist.org/observations/10957814
https://www.inaturalist.org/observations/1181659
https://www.gettyimages.ca/detail/news-photo/springboks-running-on-plains-kgalagadi-transfrontier-park-news-photo/144860225?adppopup=true
https://www.gettyimages.ca/detail/news-photo/herd-of-sprinbok-grazing-on-savanna-plane-at-etosha-news-photo/481582125?adppopup=true
https://www.gettyimages.ca/detail/photo/northern-region-of-the-namib-desert-namibia-royalty-free-image/80254736?adppopup=true

Unlike the goitred gazelle, the springbok sometimes bounds. This is an action different from stotting:
https://www.gettyimages.ca/detail/photo/springbok-leaping-central-kalahari-royalty-free-image/dv029121?adppopup=true
https://www.inaturalist.org/observations/25949
https://www.gettyimages.ca/detail/photo/springbok-leaping-central-kalahari-royalty-free-image/dv029121?adppopup=true
https://www.gettyimages.ca/detail/photo/springbok-running-royalty-free-image/125528069?adppopup=true

INTERPRETATION

What is the adaptive significance of these differences between the goitred gazelle and the springbok?

Consider the different regimes of natural predation in the Iranian region and South Africa. Here, the counterintuitive principle is: the more intense the predation, the more conspicuous certain ungulates evolve to be.

In both regions, the natural predators included large cursorial canids which scanned the groups of gazelles by day to choose the least-fit member before starting a chase of endurance.

These canids are

• in the habitat of the goitred gazelle, the wolf (Canis lupus, https://www.inaturalist.org/taxa/42048-Canis-lupus), and
• in the habitat of the springbok, the painted hunting dog (Lycaon pictus, https://www.inaturalist.org/taxa/42093-Lycaon-pictus).

This preliminary scrutiny means a selective pressure for self-advertisement ('showing off'), instead of keeping a low profile.

A crucial difference was that the northern species was the only wild ruminant in most of its habitat, whereas the southern species coexisted with

• the red hartebeest (Alcelaphus caama, https://www.gettyimages.ca/detail/photo/tsessebes-royalty-free-image/520403610?adppopup=true),
• the blesbok (Damaliscus pygargus phillipsi, https://www.inaturalist.org/observations?place_id=50327&taxon_id=129724), and/or
• the black wildebeest (Connochaetes gnou, https://www.alamy.com/stock-photo-mixed-herd-of-black-wildebeest-and-springbok-31832273.html and https://videohive.net/item/black-wildebeest-and-springbok-antelopes-at-a-waterhole-/23284578).

The greater reliability of prey in South Africa is likely to have maintained a denser population of predators, thus boosting the risk of any particular group of the springbok being encountered by the painted hunting dog.

Imagine an evolutionary tipping point, at which the adaptive colouration of adults of the springbok lost any concealing features, and became thoroughly revealing.

The best chance for the individual springbok is not to self-efface (either in the landscape or in the group), but to announce itself as vitally as possible, disqualifying itself from being worth chasing.

In the case of the goitred gazelle, as for most other antelopes, encounters with predators remained infrequent enough that it made sense to bet-hedge, rather than committing to conspicuousness.

Red hartebeest (see https://www.picfair.com/pics/06662183-red-hartebeest-alcelaphus-buselaphus), blesbok (see https://www.123rf.com/photo_19680505_a-blesbok-antelope-damaliscus-pygargus-south-africa.html?vti=mcypwdomf20ai7hkf7-1-5) and black wildebeest (see https://rangerdiaries.com/diary/black-wildebeest-2/) also have dark/pale features obvious to predators in certain settings.

However, their overall colouration remains ambivalent (https://www.dreamstime.com/herd-springbok-steppe-south-africa-image195499685).

This may be because all are larger and more enduring in flight than the springbok, and thus less vulnerable to the painted hunting dog.

Various species of ungulates have pale or dark patches on their bodies which are so large that they reveal rather than conceal the whole figure of the animal in its normal surroundings by day. These are usually gregarious species of open vegetation, which have abandoned any attempt to hide as adults and instead find 'safety in numbers' by various means. Perhaps the best example is the springbok (Antidorcas marsupialis), which shows conspicuous white on the rump and, separately, on the flanks (see https://www.inaturalist.org/observations/70596244).

In a recent Post I suggested that we refer to such bold features, which are likely to be noticeable to scanning predators even when the ungulates stand still, as bleezes. 'Bleeze' is a precise-minded technical variant of the informal and ambiguous words 'blaze' (English) and 'bles' (Afrikaans).

A few species of ruminants take this specialisation further: their bleezes can be expanded by unfolding of the skin and erection of the fur, which produce what I propose we call an 'erectile bleeze'. In the case of the springbok, the anatomy of the skin-folds is so complicated that nobody has yet clearly described it.

The dark-edged white bleeze on the hindquarters, without unfolding and erection, is limited to a diamond shape running from the buttocks to the rump, encompassing the white base of the tail. With full activation this fluffs out, expands fourfold, becomes far more visible in profile, and appears for the first time on the back (see eighth photo in https://ca.finance.yahoo.com/news/springboks-pronking-joy-071753589.html). The word 'crest' falls short because most of the erectile bleeze is a double (left-right) ridge of long fur, parting widely around the tail-base (see sixth photo in https://www.africahunting.com/threads/namibia-namibia-safari-with-kowas-beginning-the-end.11723/page-3) and ending separately on the two buttocks where it has a vertical orientation at right angles to the horizontal orientation seen on the back.

Photos of the springbok can be confusing, because in some instances the unfolding/erection is only partial. For example, the section on the buutocks can be flared without white appearing on the back. And when the animal erects all of its fur after rain, the bleeze may appear 'half-mast' (see https://www.flickr.com/photos/alanjcrawfordphotography/46754844415/ and https://www.pond5.com/stock-footage/item/103409474-springbok-antidorcas-marsupialis-rain-acting-wary-namibia).

Several other species of ruminants, including other genera of gazelles, possess an erectile bleeze on the hindquarters. However, only in the springbok is it the case that the bleeze and the erectile bleeze, although overlapping in anatomical position, are so different in shape and size that the two can hardly be described without resorting to different names.

Most species of ungulates have pointed 'weapons' of some kind on their heads, be these horns, antlers or long canine teeth.

These are used in masculine rivalry. In species in which they occur also in females, they are used for other social threats in certain situations.

One might expect, then, that the horns, antlers and teeth of ungulates could be applied to stabbing predators.

Surprisingly, most species of ungulates turn out to be inept in fighting off predators. The pointed structures on their heads function far more as adornments than as weapons; and their ritual rather than violent deployment is analogous with the gentlemanly sport of fencing.

The problem seems to be that the 'hardware' lacks suitable 'software', i.e. the brains of ungulates do not seem to be programmed for aiming and striking at predators, regardless of how precise their use is in masculine sparring.

Ungulates generally rely on rapid fleeing and reproduction, not self-defence, in order to survive predation. When cornered, they lack the behavioural versatility to use their 'weapons' in more than a tokenistic way. Once wounded, they go into shock and seem to lose will and coordination.

The few exceptions appear to include

• the rocky mountain goat (Oreamnos americanus, see https://www.earthtouchnews.com/natural-world/animal-behaviour/angry-mountain-goat-confronts-a-hiker-on-the-snowy-slopes/ and https://slate.com/news-and-politics/2010/10/what-should-you-do-if-you-re-attacked-by-a-mountain-goat.html),
• the black wildebeest (Connochaetes gnou, see https://www.iol.co.za/news/south-africa/eastern-cape/wildebeest-was-out-of-control-1623561 and https://www.dispatchlive.co.za/news/2013-12-18-father-and-son-killed-by-wildebeest-/ and https://www.pressreader.com/south-africa/cape-argus/20131224/281681137703955 and https://www.iol.co.za/dailynews/pair-killed-by-wildebeest-remembered-1626163 and for another member of the same genus https://www.dailymail.co.uk/news/article-5703563/Dramatic-moment-wildebeast-fights-two-lions-Kruger-National-Park.html),
• the muskox (Ovibos moschatus, see https://www.nps.gov/cakr/planyourvisit/muskox-safety.htm),
• the grey rhebok (Pelea capreolus), and
• the bushbuck (Tragelaphus sylvaticus, see https://www.africahunting.com/threads/add-bushbuck-to-the-list-of-dangerous-game.20199/).

All of the above are ecologically and/or socially extreme in ways suggesting that fleeing and reproduction alone would not allow them to survive predation. And all have horn-designs unusually suited to stabbing and hooking.

Other ungulates with defensive reputations in the semi-popular literature, such as oryxes (https://www.inaturalist.org/observations?taxon_id=42307), are more inept than the fearsome shapes of their horns might suggest.

The African savanna buffalo (Syncerus caffer, https://www.inaturalist.org/taxa/42405-Syncerus-caffer) is certainly dangerous to humans when wounded. However, its horns are too blunt to stab an antagonist.

Various video-clips show how timorously the African buffalo behaves versus the lion (Panthera leo), even when there are opportunities to strike blows (see https://www.sabisabi.com/blog/14593/lions-vs-buffalo/ and https://www.youtube.com/watch?v=zzI8ya3ARWM and https://www.youtube.com/watch?v=SObYT06hnm4 and https://www.youtube.com/watch?v=MY6conyVLlc and https://www.youtube.com/watch?v=aCwwecrasWI). The lion is intimated more by the bulk and gregariousness of buffaloes than by the horn-tips.

The following video-clip (https://www.youtube.com/watch?v=Bu2JspPdMy0 ), taken near Kruger National Park, shows:

• how obtuse the African buffalo can be, with the mother wandering out from the group with her infant, 'just asking for trouble',
• that the mother does in fact put up a spirited defence against one adult male of the lion, but loses her infant anyway because several individuals of the lion are hunting together, and
• how poorly co-ordinated the collective defence is: there are several mature males near the infant, but they do not cooperate to defend it.

And even when the predators, in the act of eating, are vulnerable to retribution, the mature male African savanna buffalo holds back, as if knowing that if several individuals of the lion grab him, he will not be able to rely on solidarity from the rest of his own group.
 
Furthermore, here is a point that I suspect will be overlooked by most naturalists: the mother, when attacked by the lion, does not flee into the nearby water.
 
If this were the river buffalo (Bubalus bubalis) or the water buffalo (Bubalus carabanensis), I expect that the mother would take refuge in the river immediately.

I realise that in plain view was a group of the hippopotamus (Hippopotamus amphibius), and that species would not necessarily show any solidarity with a buffalo. However, this footage suggests that the African savanna buffalo is quite different from its tropical Asian counterparts in its relationship to water. It obviously sees no refuge in the river.

Not only waterbucks (Kobus ellipsiprymnus and Kobus defassa), but even antelopes such as Strepsiceros strepsiceros and Hippotragus niger that have no particular aquatic affinity, would, I expect, have gone straight into the water to save themselves (http://www.dailymail.co.uk/news/article-2609382/Kudu-tell-African-antelope-escaped-pack-hyenas-Clever-prey-conceals-pack-zebra.html).

It is in the light of this general pattern, in which most ungulates are almost as vulnerable to predators as they would be were their horns absent, that we can understand the basic nature of Spanish bullfighting: choreography rather than a real contest of impalation. The matador pierces the bull, which is incapable of reciprocating - except by accident.

to be continued in https://www.inaturalist.org/journal/milewski/86153-most-ungulates-are-surprisingly-inept-at-using-their-horns-against-predators-part-2#...