The unique sexual dimorphism of the northern oribi (Ourebia montana), an ungulate combining polygamy with male diminution

In iNaturalist, Ourebia (https://www.inaturalist.org/taxa/42380-Ourebia-ourebi) seems to be photographed in pairs (https://www.alamy.com/stock-photo-oribi-ourebia-ourebi-kafue-national-park-zambia-11846018.html?imageid=857A4ABE-7DBB-44E5-A844-B03268BAA80C&p=74587&pn=3&searchId=b18d9220987fab88ffb57be4cfa504ab&searchtype=0 and https://www.inaturalist.org/observations/110886912) more frequently than is any other ruminant.

Several small-bodied bovids are monogamous. In these spp., males tend to be smaller-bodied than females (please see comment below for details in the southern African fauna).

So, at first sight, it would seem that Ourebia is yet another example of this syndrome.

However, on closer scrutiny, Ourebia is revealed as odd, in the following ways.

Firstly, it is not monogamous. Territorial males consort with up to four individual females (https://www.inaturalist.org/observations/135350368). Therefore, it is anomalous that males are smaller-bodied than females, in Ourebia montana.

(Please see https://www.inaturalist.org/journal/milewski/70050-the-three-main-types-of-oribi-ourebia-at-a-glance# for the differences between Ourebia montana, the common oribi, and Ourebia ourebi, the southern oribi.)

Indeed, it seems possible that O. montana is the only polygamous ungulate, on Earth, in which males are smaller-bodied than females.

Secondly, adult males of O. montana not only weigh less than adult females, they also have a disproportionately shorter body and neck.

What seems to be unique, in Ourebia, is the way in which adult females differ in body shapes from adult males.

Exaggerating somewhat, in order to give readers a search-image:

Females in O. montana have body shapes similar to reduncins (https://en.wikipedia.org/wiki/Reduncinae).

By contrast, males in Ourebia montana have body shapes similar to those of 'neotragin dwarf antelopes' (https://en.wikipedia.org/wiki/Neotragini), i.e. the smallest-bodied antilopins.

So, when one views a sexual pair of O. montana, it can look like a 'marriage' of two different tribes of bovids, somehow combined within a single species (https://www.inaturalist.org/observations/7215985).

Females of O. montana resemble a diminutive version of reedbucks (Reduncini: Redunca), while males resemble the largest-bodied of the neotragin 'dwarf antelopes'.

To be more precise about the sexual difference in body shapes:

Adult females have a long torso and neck (https://www.ourbreathingplanet.com/oribi/ and https://www.shutterstock.com/image-photo/oribi-ourebia-ourebi-1243168594 and https://www.shutterstock.com/image-photo/portrait-oribi-ourebia-ourebi-murchison-falls-2188507957).

This makes females of O. montana look somewhat like a scaled-down version of adult females of the common reedbuck (Redunca arundinum, https://en.wikipedia.org/wiki/Southern_reedbuck), which actually weigh threefold more than females of O. montana.

Meanwhile, adult males have the unremarkable proportions of a generalised 'dwarf antelope' (https://www.alamy.com/oribi-ourebia-ourebi-masai-mara-national-park-kenia-image60113750.html?imageid=AD7C8970-246D-4C7D-B8E4-36915DA3494F&p=191343&pn=3&searchId=b18d9220987fab88ffb57be4cfa504ab&searchtype=0).

The body mass of O. montana is about 15 kg, with males (about 14-15 kg) at least 2 kg lighter than females (about 16-17 kg). (This is different from Ourebia ourebi ourebi, in which both sexes average 14 kg, and females are, if anything, slightly smaller-bodied than males, https://www.inaturalist.org/observations/8778172.)

However, when one views the sexes apart from each other,

I have chosen the following to show the sexes separately, but in similar postures and perspectives.

Female https://www.inaturalist.org/observations/66999204
Male https://www.inaturalist.org/observations/66999205

Female https://www.inaturalist.org/observations/91161425
Male https://www.inaturalist.org/observations/8490581

Female https://www.inaturalist.org/observations/8490569
Male https://www.inaturalist.org/observations/8490590

Female https://www.inaturalist.org/observations/6577553
Male https://www.inaturalist.org/observations/6441964

Female https://www.inaturalist.org/observations/49107418
Male https://www.inaturalist.org/observations/8490545

Female https://www.inaturalist.org/observations/7103895
Male https://www.inaturalist.org/observations/77578819

Female https://www.inaturalist.org/observations/67005615
Male https://www.inaturalist.org/observations/6183578

Female https://www.inaturalist.org/observations/8490553
Male https://www.inaturalist.org/observations/67005621

Female https://www.inaturalist.org/observations/8490547
Malehttps://www.inaturalist.org/observations/8490577

Female https://www.inaturalist.org/observations/8490546
Male https://www.inaturalist.org/observations/8490548

Female https://www.inaturalist.org/observations/34293509
Male https://www.inaturalist.org/observations/6560837

Female https://www.inaturalist.org/observations/44892361
Male https://www.inaturalist.org/observations/8490568

Female https://www.inaturalist.org/observations/16097725
Male https://www.inaturalist.org/observations/38669966

It is unlikely that this dimorphism in O. montana is directly related to differences between the sexes in the height of foraging.

This is because Ourebia

  • depends on low, herbaceous plants throughout the year, being the smallest specialised grazer among all the ungulates of the world, and
  • forages only quadrupedally, and does not rise to bipedal postures to seek food above its normal reach.

However, there are potential advantages of the absolutely and relatively longer neck of females of O. montana, in vigilance.

This would be understandable, given that

A possible explanation for the absolutely longer torso in females than in males is as follows.

Grass-eating ruminants tend to have capacious stomachs, compared to those eating mainly the foliage of shrubs, which tends to be more nutritious than grass.

Ourebia montana is at the lower limit of body mass at which specialisation on grazing can be sustained. Therefore, there is likely to be particular pressure on the gestating and lactating sex, to accommodate the necessary 'bulk and roughage' in the gut.

The additional bulk could be accommodated by either an increase in the girth of the torso, or an increase in its length. The latter may possibly be more compatible with the extreme locomotion in this gracile, lightly-built antelope.

The following illustrations reinforce the impression that, in O. montana, the neck is not only absolutely, but also relatively, longer, in females than in males.

https://www.inaturalist.org/observations/7994502

scroll in https://www.zoochat.com/community/threads/ungulate-taxonomy-revisited-the-evidence-for-the-splits-of-g-g.467230/page-2#post-1046851

https://www.shutterstock.com/image-photo/sexual-dimorphism-oribi-antelope-only-bucks-2040799799
https://www.shutterstock.com/image-photo/sexual-dimorphism-oribi-antelope-only-bucks-2040800168

https://www.alamy.com/stock-photo-oribi-ourebia-ourebi-maze-national-park-ethiopia-85989749.html?imageid=6645CFFF-4910-460B-86C0-CC35B462628D&p=22059&pn=3&searchId=b18d9220987fab88ffb57be4cfa504ab&searchtype=0

https://www.alamy.com/stock-photo-oribi-ourebia-ourebi-murchison-falls-national-park-uganda-92320381.html?imageid=0589FEED-640D-432F-BA8E-3EF832819C00&p=22059&pn=3&searchId=b18d9220987fab88ffb57be4cfa504ab&searchtype=0

https://www.inaturalist.org/observations/131699606

In the past, it was assumed that Ourebia belonged to Antilopini. This was partly because it seemed to provide a link, in body size and habits, between 'neotragin' antilopins and gazelle-like antilopins.

However, there has been a growing realisation that Ourebia deserves its own tribe (Ourebini), among Bovidae.

My findings, in this Post, seem to support the distinctiveness of Ourebia.

This is partly because one of its spp., namely O. montana, seems to emerge as unique in two, related ways, viz.

  • it shows masculine diminution, despite retaining the polygamy typical of specialised grazers, and
  • this diminution is in not only body mass, but also the lengths of torso and neck relative to body mass.

Furthermore, I have noticed another odd combination, in Ourebia, corroborating its phylogenetic distinctiveness.

This is the combination of stotting behaviour by adults and extreme hiding by juveniles.

Ourebia is small-bodied among ungulates that stot. However, Madoqua kirkii damarensis is half as massive (body mass 5 kg) and stots, as follows: "In reaction to sudden fright they may...bound away stiff-legged, stotting, their legs tucked up under their bodies, at each contact with the ground giving a short explosive whistle" (Smithers and Chimimba 2005). What is noteworthy is that, in M. kirkii, infants hide for only three weeks.

Most bovids that show stotting behaviour tend to have infants that hide for only a few weeks. Furthermore, in these spp., it is infants and juveniles that are most likely to stot. (Eudorcas and Antidorcas are typical examples.)

In Ourebia, the hiding period is extended beyond infancy into the juvenile phase, until up to three months old. I assume that juveniles forage at night. However, their hiding by day remains noteworthy, given that adults forage mainly diurnally.

So, does this make Ourebia the only grazing ungulate, on Earth, that hides like an infant until it is nearly half-grown?

Furthermore, it seems possible that, in Ourebia, juveniles are less likely to stot - even in play - than adults are. Does this make Ourebia the only ungulate, on Earth, in which adults stot more frequently than juveniles do?

In summary, here are the anomalous aspects of Ourebia, particularly O. montana, starting with those most supported by the evidence, so far:

  • smallest-bodied ungulate on Earth that is specialised for grazing,
  • only ungulate with male diminution despite polygamy,
  • only ungulate with infantile hiding extended to the juvenile phase,
  • only ungulate with male diminution in lengths of torso and neck, relative to body mass, and
  • only ungulate in which adults stot more frequently than juveniles do.
Posted on September 15, 2022 12:37 AM by milewski milewski

Comments

I did not know the adult female was larger than the male in Ourebia, very distinct then amongst artiodactyls.

Posted by paradoxornithidae over 1 year ago

@paradoxornithidae

Sexual dimorphism in body mass, with females > males, occurs also in some of the other bovid bambis.

In the fauna of southern Africa, mean adult body masses (rounded to nearest 0.5 kg) are as follows, with females first in each case):

Raphicerus melanotis 10.5, 10
Raphicerus sharpei 7.5, 7
Raphicerus campestris 11.5, 11
Madoqua damarensis 5, 5
Neotragus moschatus 5.5, 5
Oreotragus 'oreotragus' 13, 11
Sylvicapra grimmia caffra 16.5, 16
Cephalophus natalensis 12, 12
Philantomba monticola 4.5, 4

Ourebia ourebi ourebi 14, 14

Ref.: Skinner and Chimimba (2005)

Several bovid bambis, belonging to several subfamilies and tribes, have this direction of sexual dimorphism.

Possibly for this reason, nobody has previously scrutinised Ourebia. It is too easy to assume that, as in other bovid bambis, the difference in body size involves no significant difference in body proportions.

However, something new emerges here.

Firstly, there seems to be a difference between Ourebia ourebi (14 kg in both sexes) and Ourebia montana, supporting my suggestion that we call these different species. According to my definition of bambis as having adult female body mass 15 kg or less, O. ourebi qualifes as a bambi, but O. montana does not.

Secondly, the northern oribi is a particularly sexually dimorphic species (female > male) among bambis or near-bambis, in both body mass and body shape.

Thirdly, Ourebia montana may possibly be the only ruminant on Earth in which females are more massive than males at an overall mean body mass exceeding 15 kg (the larger-bodied spp. of Cephalophus need checking).

Posted by milewski over 1 year ago

@tonyrebelo @oviscanadensis_connerties @jeremygilmore @beartracker @capracornelius @dejong @tandala @michalsloviak @matthewinabinett @paradoxornithidae @simontonge @jakob

I have rewritten much of this Post, after realising that my first version undersold the manifold oddness of Ourebia. Please do have another look.

Posted by milewski over 1 year ago

If females of Ourebia have superior vigilance by virtue of the height of their eyes, this may perhaps help to explain why so many photos give the false impression of monogamy. The adult male may perhaps seek out the company of one of the females in his territory, while resting around midday, in order to benefit from its advantage in vigilance.

Posted by milewski over 1 year ago

It has long been known that, within ruminants, there is variation in behavioural precociality.

Certain species have precocial infants, which never hide, but instead follow the mother from birth. This is epitomised by Connochaetes mearnsi.

Other species have 'behaviourally altricial' infants, which hide for various periods after birth, being visited by mothers for suckling. The periods vary from a few days to a few months, but in general the hiding phase is confined to infancy.

Ourebia seems as specialised as wildebeests on this spectrum, but at the opposite extreme.

Posted by milewski over 1 year ago

Another possibly odd aspect of Ourebia:

Most spp. of 'neotragin' dwarf antelope nave no distinctive colouration in infants; and the same seems true for Redunca.

However, in Ourebia the ground-colour of infants is significantly different from that of adults, being darker and duller-hued (https://www.alamy.com/stock-photo-oribi-ourebia-ourebi-senkele-game-sanctuary-ethiopia-85939030.html?imageid=E948BFFA-B280-4274-BA93-83A6C67DD514&p=22059&pn=1&searchId=f7d66f1874b5dfa1e536bf9cbab6379c&searchtype=0 and https://www.alamy.com/stock-photo-female-oribi-ourebia-ourebi-with-nursing-calf-54880619.html?imageid=E19322FB-4C31-42BC-AD44-FD23D62DF262&p=164715&pn=1&searchId=b51320e4aa8f23d8475c9a13e60a54fb&searchtype=0 and https://www.pinterest.com.au/pin/mom-and-baby-oribi--57350595226601377/ and https://www.alamy.com/stock-photo-oribi-ourebia-ourebi-fawn-47353319.html?imageid=88E5E1E4-7F19-47C8-9B97-3CDF3E4CA2B8&p=53483&pn=1&searchId=b51320e4aa8f23d8475c9a13e60a54fb&searchtype=0).

This colouration is said to remain for five weeks.

This seems consistent with an extended reliance on hiding in the offspring of Ourebia.

Note the relatively dark back on this juvenile male individual: https://www.gettyimages.ie/detail/photo/beautiful-oribi-royalty-free-image/566355631?adppopup=true and https://www.gettyimages.ie/detail/photo/oribi-royalty-free-image/573793019?adppopup=true.

Is this the juvenile colouration, changing into the adult colouration?

Posted by milewski over 1 year ago

BODY MASS OF OUREBIA IN ZAMBIA

Ref.: Robinette W L (1963) Weights of some of the larger mammals of Northern Rhodesia, The Puku, pp. 207-215.

Males:

Bangweulu 10 kg
Kalichero 13.6 kg
Kalichero 17.7 kg
Kalichero 14.1 kg

Females (immature):

Mwinilunga 7.7 kg
Balovale 10.9 kg

Females (pregnant):

Kalichero 16.8 kg

Commentary:

These data, although too few (total sample size = 7, including immature specimens), suggest that, in Ourebia ourebi rutila, body masses are similar (about 15 kg) in the two sexes.

Posted by milewski over 1 year ago

interesting

Posted by paradoxornithidae over 1 year ago

@paradoxornithidae

There are two possible ways in which females can come to exceed males in body mass, torso length relative to body mass, and neck length relative to body mass.

One is for females to grow faster than males, so that the different body sizes are achieved at the same age (say, about two years).

Another is for females and males to grow at the same rate to the point of sexual maturity, followed by females continuing to grow certain parts of the body, particularly the torso and neck.

The implication of the latter is that the full size and shape of the body of females would be reached only later in life, say about four years old.

In most of the sexually dimorphic ruminants, males are more massive than females, with brawnier necks (https://www.dreamstime.com/stock-photo-common-elands-mating-pair-taurotragus-oryx-amakhala-game-reserve-eastern-cape-south-africa-image51205440).

Males achieve this difference by a combination of growing more rapidly than females as adolescents, and continuing to grow for several years after adulthood.

If Ourebia montana really does continue to grow after adulthood is reached, then this would - as far as I know - be unprecedented for females among ungulates.

Such prolonged growth would not merely be a case of 'androgenisation', such as that seen in Crocuta crocuta, in which females are larger-bodied than males (https://www.superstock.com/asset/spotted-hyena-crocuta-crocuta-pair-mating-serengeti/4201-16389 and https://www.shentonsafaris.com/blog/daytime-hyaenas-mating/ and scroll in https://africafreak.com/hyena-birth).

Instead, it would represent a previously unrecognised pattern of sexual dimorphism, in which the neck and torso continue to grow from adulthood to full 'social maturity', but in a way making the figure less, not more, brawny.

If both sexes of O. montana reach mature body size and shape at the same age, then 'male diminution' seems an apt description of the odd form of sexual dimorphism in this species. However, if females show extended growth, then perhaps 'female enlargement' might be more apt.

Posted by milewski over 1 year ago

@tonyrebelo @jeremygilmore @ludwig_muller @kevinatbrakputs @henrydelange @rion_c @lwnrngr @justinhawthorne @iambebop @callumevans @johnnybirder @douglasriverside @christiaan_viljoen @klauswehrlin @koosretief

Have you ever seen a grysbok stotting (https://en.wikipedia.org/wiki/Stotting)?

Austin Roberts (1951, pp. 336-337 in The Mammals of South Africa) states of Ourebia ourebi ourebi: "...when disturbed bound along in hops, displaying their tails conspicuously...This hopping habit is also peculiar to Grysboks..."

I have never observed stotting in Raphicerus melanotis, which, in my experience, simply rushes off under cover.

I suspect that what Roberts may be referring to is the stamping sound of the hooves. He states of O. o. ourebi: "Probably the stamping sound made by the hopping gives the alarm to others nearby..."

However, if any of you have ever seen a grysbok jumping up when fleeing, then I will stand corrected.

Posted by milewski over 1 year ago

I have conducted several net capture and translocation operations of Cape Grysbok (for population management purposes) and have therefore been able to observe their behaviour when flushed from cover. To date I have not observed any type of stotting or hopping when disturbed. They tend to duck into cover, or when approached further, they may dart from cover to cover trying to evade capture. Ultimately, if they have no way to evade capture, they go into a flat-out run. But never have I seen any stotting or hopping display in Cape Grysbok.

*Note, capturing and translocation of wildllife for population management purposes (and indeed any form of disturbing, for that matter) may only be done in accordance with permission by the relevant authority in terms of the laws of the jurisdiction.

Posted by koosretief over 1 year ago

@koosretief
Many thanks for your helpful reply.

Posted by milewski over 1 year ago

Sexual dimorphism in Raphicerus campestris:

scroll in https://www.sabisabi.com/blog/20563/a-week-in-the-bush-vol-267/

https://stock.adobe.com/search?k=steenbok&asset_id=96029539

https://stock.adobe.com/search?filters%5Bcontent_type%3Aphoto%5D=1&filters%5Bcontent_type%3Aillustration%5D=1&filters%5Bcontent_type%3Azip_vector%5D=1&filters%5Bcontent_type%3Avideo%5D=1&filters%5Bcontent_type%3Atemplate%5D=1&filters%5Bcontent_type%3A3d%5D=1&filters%5Bcontent_type%3Aaudio%5D=0&filters%5Binclude_stock_enterprise%5D=0&filters%5Bis_editorial%5D=0&filters%5Bfree_collection%5D=0&filters%5Bcontent_type%3Aimage%5D=1&k=steenbok&order=relevance&safe_search=1&limit=100&search_page=9&search_type=pagination&get_facets=0&asset_id=438251069

https://stock.adobe.com/search?filters%5Bcontent_type%3Aphoto%5D=1&filters%5Bcontent_type%3Aillustration%5D=1&filters%5Bcontent_type%3Azip_vector%5D=1&filters%5Bcontent_type%3Avideo%5D=1&filters%5Bcontent_type%3Atemplate%5D=1&filters%5Bcontent_type%3A3d%5D=1&filters%5Bcontent_type%3Aaudio%5D=0&filters%5Binclude_stock_enterprise%5D=0&filters%5Bis_editorial%5D=0&filters%5Bfree_collection%5D=0&filters%5Bcontent_type%3Aimage%5D=1&k=steenbok&order=relevance&safe_search=1&limit=100&search_page=9&search_type=pagination&get_facets=0&asset_id=437447742

Sexual dimorphism in Oreotragus saltatrixoides transvaalensis:

https://stock.adobe.com/search?filters%5Bcontent_type%3Aphoto%5D=1&filters%5Bcontent_type%3Aillustration%5D=1&filters%5Bcontent_type%3Azip_vector%5D=1&filters%5Bcontent_type%3Avideo%5D=1&filters%5Bcontent_type%3Atemplate%5D=1&filters%5Bcontent_type%3A3d%5D=1&filters%5Bcontent_type%3Aaudio%5D=0&filters%5Binclude_stock_enterprise%5D=0&filters%5Bis_editorial%5D=0&filters%5Bfree_collection%5D=0&filters%5Bcontent_type%3Aimage%5D=1&k=steenbok&order=relevance&safe_search=1&limit=100&search_page=3&search_type=pagination&get_facets=0&asset_id=141757142

Posted by milewski over 1 year ago

@koosretief

Kingdon (2015, page 549 in The Kingdon Field Guide to African Mammals, second edition) states: "At any disturbance, Sharpe's grysbok departs rapidly, with short, stamping hops (but otherwise silently).".

Posted by milewski over 1 year ago

@koosretief @beartracker @tonyrebelo

Roberts (1951), on page 339, states:

Raphicerus sharpei:

"When alarmed during daylight I have repeatedly noticed how it starts off in short stamping hops, evidently to give the alarm to others, before darting off through the tangles."

Raphicerus melanotis:

"It has not been recorded that it hops away at the start, as I have frequently seen done by its congener of the tropics, but I have no doubt it does so as well."

My commentary:

Roberts seems to have been proven wrong about R. melanotis. This species differs from R. sharpei, in that it does not 'hop-stamp' when fleeing.

What this means is that we can now summarise the anti-predator 'flight-announcements' in the three spp. of Raphicerus as follows:

melanotis: none

sharpei: 'hop-stamping', which is a non-vocal audial signal

campestris: activation of a buttock flag, which is a visual signal (see https://www.inaturalist.org/journal/milewski/70293-the-bambis-part-9-bleezes-flags-and-semets-in-the-bovid-genus-raphicerus#)

This adaptive differentiation, within the genus, makes sense in view of the differences in habitat, viz. dense, evergreen cover in melanotis, open vegetation in campestris, and sharpei intermediate (dry-season deciduous).

Posted by milewski over 1 year ago

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