יומן של milewski

The Arabian peninsula is bigger than western Europe, and ecologically similar to North Africa. However, an unexplained fact is that most of its species of large terrestrial animals are diminutive.

All of the wild ungulates and the major Carnivora of the Arabian peninsula are smaller than conspecifics, or their closest ecological counterparts, in Africa or mainland Asia. The same was true for the extinct Arabian form of the ostrich (Struthio camelus). This generalisation applies to Gazella saudiya (extinct), Gazella arabica, Gazella marica, Capra nubiana, Oryx leucoryx (which narrowly escaped extinction), Arabitragus jayakari, Panthera pardus, Caracal caracal, Acinonyx jubatus (locally extinct), Canis lupus, Vulpes vulpes and Hyaena hyaena.

One of the reasons why this puzzle is poorly appreciated is that most of the species listed above do not have the body proportions usually associated with miniature varieties of mammals, such as proportionately large heads and eyes and proportionately short legs and horns. The most familiar example is the Arabian oryx, which looks as well-proportioned as its Saharan relative, Oryx dammah, despite being only about half the body mass.

Reduction in body size is a familiar pattern on islands, and is usually explained by the limitation on resources in confined areas. This rationale can theoretically apply to peninsulas too, to the degree that peninsulas produce isolation from mainlands. And a further parallel with islands is the tendency of the fauna to be driven rapidly to extinction by human hunters.

However, isolation is an unsatisfactory explanation for the remarkably consistent diminution in the Arabian fauna, because this peninsula is larger than any island on Earth. Even if it is borne in mind that aridity limits the productivity and reliability of resources, how can the effects have been so different from the Sahara?

The mystery of the Arabian miniatures remains an intriguing puzzle for naturalists to ponder.

True gazelles, of the genus Gazella, are bewilderingly speciose in Arabia and North Africa. Each species is individually variable. Many of the species have local subspecies. Photographers tend to focus on males. As a result, identification from photos - particularly in the case of females - can be difficult even for naturalists with plenty of experience with ungulates.

In the case of two species of sand gazelles inhabiting the desert dunes, the confusion is aggravated by the indiscriminate use of similar common names derived from the Arabic. 'Rhim' refers to the Saharan Gazella leptoceros while 'rheem' refers to the Arabian Gazella marica, and the two seem to be mislabelled interchangeably on the internet even when the specimens are in zoos.

The name 'slender-horned gazelle' for G. leptoceros hardly helps because both species have long, slender, somewhat asymmetrical horns in most females.

In reality, the two can easily be told apart if you know what to look for, as follows.

All species of true gazelles have are more or less fawn with whitish ventral parts separated from the fawn by a relatively dark flank-band. All also have intricate markings on the face.

First, look at the fawn of the body, neck and legs. In G. leptoceros, this is the most uniform of any species of true gazelle, whereas in G. marica it is clearly carved into a pale upper flank-band, a pale lower-haunch, and pale legs. Another difference which I have never seen mentioned in field guides is that the feet of G. leptoceros are marked with subtle dark/pale contrasts near the hooves, whereas those of G. marica are the same plain pale as the legs.

Now look at the face. In G. leptoceros there is just the standard, inconspicuous pattern typical of gazelles, except for a small whitish patch above the nose in about half of all individuals. In G. marica, the whole face tends to be conspicuously whitened.

The rocky mountain goat (Oreamnos americanus) is the only ungulate species on Earth which possesses all-white pelage, in both sexes and at all ages. What is the adaptive reason for this extreme specialisation?

The obvious answer is 'nivicolous crypsis', i.e. hiding in the snow and ice in the way epitomised by the polar bear (Ursus maritimus). However, the sheer number of photos of the rocky mountain goat in iNaturalist suggests that this interpretation is incorrect. How could it possibly be that a prey-animal coloured to blend into its typical environment is so easy to spot, and so extremely photogenic?

Instead, it should be clear from the thousands of photos of the rocky mountain goat on the internet that its white pelage advertises it in its normal surrounds, that it is particularly gregarious in winter, that it does not attempt to crouch or 'freeze' when alarmed, and that even the newborns do not hide in the way of most ungulates. The terrain chosen by the rocky mountain goat - even in winter - tends to be rocky and dark rather than snowy and pale, making the animals conspicuous.

The rocky mountain goat is one of the most extreme examples, among ungulates, of the female emulating the male in muscularity, weaponry (deadly horns), adornment (e.g. beard, mane, pantaloons), aggressiveness, and self-advertisement. A similar syndrome occurs in adaptively conspicuous bovids such as wildebeests (Connochaetes) and oryxes (Oryx). However, in the case of the rocky mountain goat the male is so intimidated by the female that his courtship requires juvenile-like postures of submission, and any naturalist familiar with the species knows how hard it can be to distinguish the males in any group.

The rocky mountain goat is so specialised for taking deliberate, sure-footed refuge from predators on cliffs that it has abandoned any attempt to hide, even at night or in infancy, and it has nearly abandoned running when alarmed. It seems to be compatible with this strategy that it has also feminised the bravado usually associated with masculinity in mammals, extending the silhouette by means of furry adornments to make the female figure big and noticeable.

I suggest that it is view of this combination of specialisations - anti-predator reliance on cliffs and maternal defence, going together with extreme 'feminist' self-assertiveness - that the conspicuous monotonal whiteness of the rocky mountain goat can best be understood. This species is all-white not to hide but instead to show off.

At first sight, it might seem that Gazella marica is the form of the widespread Gazella subgutturosa which occurs on the Arabian peninsula. So there was some surprise when a genetic study a decade ago found the two forms to be different.

However, even without genetic information marica can be seen to be more than just an extremely pale, desertic subspecies of gutturosa.

Firstly, marica is considerably smaller than subgutturosa, the adult females weighing respectively about 19 kg and 25 kg. Gazella marica is not just desertic but also rather diminutive.

Secondly, the larynx is different in the two forms. In marica the larynx seems normal for gazelles and ruminants generally, whereas subgutturosa possesses what is perhaps the oddest laryngeal anatomy of any ungulate.

Thirdly, the horns are different in the female: Extremely well-developed in marica but extremely poorly developed in subgutturosa.

And fourthly, the arrangement of the flank-banding is surprisingly different, as follows. In marica, the flank-banding conforms with that in gazelles generally, in that the darkest, most ventral flank-band runs obliquely towards the shoulder, leaving enough space for the paler flank-band above it to be triangular. By contrast, in subgutturosa the flank-bands run horizontally, the darkest, most ventral band running towards the scapula and the paler band being reduced to a line instead of a long triangle.

As far as I know, the difference in pattern of colouration has never been pointed out before.

Furthermore, it is hard to see the extreme pallor of marica as essentially arid-adapted because subspecies yarkandensis of species subgutturosa occurs in the extreme aridity of the Gobi desert without similar pallor.

The colouration of the Dorcas gazelle is ambivalent. On one hand, it could be that the adaptive function is to hide the animal in plain sight in its environment, which is open because this is a species of the desert. On the other hand, it could be that gazelles, far from trying to hide, advertise themselves for the sake of gregariousness and the display of fitness, which are arguably their best strategy versus predators given their inevitable exposure.

In the past, almost all authors have assumed that the Dorcas gazelle's colouration blends into the desert, following the rationale of camouflage. However, I see the animal differently: Gleaming out like a beacon in most illuminations. In my view, it is not that this gazelle is pale to match the pale sand; instead, it is so pale that it becomes a living highlight. The most important element in its pattern, ensuring conspicuousness, is the extension of the white underparts to a level so high on the flank and rump that, even at noon, the white is likely to draw attention to the figure.

So which of these opposing interpretations is true?

What is not readily apparent from photos is that the Dorcas gazelle continually wags its tail, which is particularly conspicuous in being the only black part of the body. Because the eyes of ungulates and predators alike are extremely sensitive to motion, this continual movement is a real giveaway as to the antipredator strategy of this gazelle. Instead of 'freezing' in alarm and crouching low as camouflaged antelopes would do in dense vegetation, the Dorcas gazelle advertises itself and its fitness, relying on its ability to outrun the predator.

For its part, the predator relies on spotting any individual which is sick, old or injured, as betrayed by any slight failure to stot vigorously, to alarm-sneeze loudly or to keep up the flagging of the tail.

In reality, the colouration of most ungulates is probably some complex combination representing an adaptive compromise between hiding and self-advertisement. For example, the colouration of the large ears of the Dorcas gazelle suggests camouflage of that part of the figure at least. And it is possible that the same overall pattern which gleams by day actually hides the whole body when the light is dim. One way to investigate this would be to see whether gazelles behave differently when alarmed by night, freezing, keeping quiet and facing the threat rather than presenting the bold hindquarters as they typically do by day.

The grey rhebok (Pelea capreolus) has long been recognised as unrelated to other living antelopes despite a superficial resemblance to reedbucks. However, up to now an obvious aspect of this peculiarity has been 'hiding in plain sight'.

The colouration of the grey rhebok is a prime example of crypsis, i.e. plainness designed to blend into the surroundings. Few African antelopes have such uncomplicated markings, so that even a group of six can be overlooked as they stand among the low shrubs of the fynbos.

However, when alarmed the grey rhebok does something not done to the same degree by any other antelope or, indeed, any other bovid. It transforms itself in the simplest but most effective possible way by curling its tail over its rump, producing an unnaturally luminescent, handkerchief-sized patch of white which can be spotted by the naturalist at ranges up to several hundred metres.

Although many field guidebooks state or imply that reedbucks similarly display themselves when fleeing, such is not the case. And any displays of the white underside of the tail by bushbucks, kudus and their relatives are less noteworthy than that of the grey rhebok because the tails are not consistently raised and do not flash much white, and these antelopes have complicated rather than plain colouration in the first place.

The springbok (Antidorcas marsupialis) has an equally spectacular but far more complicated display of white on its hindquarters, produced by unfolding a fur-lined fold of skin on the rump. But this is categorically different from the case of the grey rhebok, because the overall colouration of the springbok (as in gazelles generally) is designed to be conspicuous, not cryptic or camouflaged. So, the springbok when fleeing transforms a flag on its already flag-like body, but does not emulate the grey rhebok in transforming itself from effective invisibility into a white flag.

In this ability to transform its appearance, no bovid and perhaps not any deer rivals the grey rhebok. And this extreme adaptation has been configured in a deceptively simple way, by deploying a tail, in itself similar in its white underside to that of reedbucks and bushbucks, on the plainnest of bodies. All that is needed is for the tail to be raised and held raised in flight.

it is stated repeatedly in the literature that reedbucks (genus Redunca) raise the tail in alarm, this displaying the white underside. I have never observed this in the field for any of the three species and I cannot recall seeing any photo showing this. So I wonder how this apparently misleading information arose.

The white-tailed deer (Odocoileus virginianus) is the wild ruminant most familiar to North Americans, and in summer coat it happens to resemble reedbucks. The tails have similar size and bushiness, and similarly are white underneath. So anyone knowing the white-tailed deer might understandably tend to assume that reedbucks would emulate the deer in displaying the tail.

In fact, the difference could hardly be greater. Firstly, the white-tailed deer not only raises it's tail, it fully erects it and pilo-erects the long white hairs on it's underside, and then waves the upright, fanned tail from side to side as it runs. Secondly and simultaneously, it flares out the white fur on the posteriormost haunch, more than doubling the 'blaze' of extremely conspicuous white on the hindquarters.

Far from having converged evolutionarily with this display, reedbucks do not seem even to possess the anatomical structures needed for it. They do advertise themselves when alarmed, but in a peculiar and poorly-understood way involving the production of popping sounds from hindquarters.

(By the way, the white-tailed deer produces a nasal sound comparable to the whistling of reedbucks, while standing alarmed and watching the intruder. But here again the difference is that the deer flicks its tail up synchronously with each 'sneeze', in this case without any erection of white fur so that the display is one of dramatic motion rather than whiteness.)

To dispel any misconception that reedbucks display the tail in flight, here is a challenge to naturalists. Can anyone show us a video or photo of any reedbuck displaying its tail in any way in alarm? And if not should we begin to write, in field-guides on African wildlife, that a remarkable aspect of reedbucks is the inertness of tails that seem bushy and white enough to be suitable for flagging alarm?

I have just spent six weeks perusing the thousands of photos of Sylvicapra grimmia available on the internet. I have also reviewed the account of this species by Groves and Grubb (2011).

The common duiker is one of the most widespread of ungulates in Africa, but its distribution and habitat are puzzling in certain ways. Here are the five most important puzzles.

Firstly, why has S. grimmia not occurred in Mediterranean North Africa? And which species has replaced it ecologically, there?

Secondly, why does subspecies (treated as a full species by Groves & Grubb 2011) pallidior not occur west of Chad despite being adapted to the Sahel?

Thirdly, why is S. grimmia absent from most of the Horn of Africa, given that it is widespread in similarly dry climates in southern Africa and the eastern Sahel?

Fourthly, why does subspecies (treated as a full species by Groves & Grubb 2011) coronata occur only in extreme western West Africa?

Fifthly, why is it that, to this day, not one photo of the common duiker has been published from the whole Serengeti Ecosystem (including Ngorongoro Conservation Area and the Maasai Mara National Reserve)?

[Footnote: a main reason why Groves & Grubb (2011) gave pallidior species-status is that this form is extremely small and seems distinct despite overlapping in range with subspecies campbelliae. A main reason why these authors gave coronata species-status is that this form has an odd skull-shape. Groves & Grubb (2011) describe this skull as peculiarly narrow but the few photos show me that it also has oddly wide orbits.]

written August 2020