A test of evolutionary convergence: bush duiker vs gray brocket, part 1

The principle of biological adaptation is that organisms are shaped by selective pressures. Given similar environments and ecological niches, unrelated animals on different continents should, theoretically, come to look similar.

This idea can be tested by comparing two species of small ruminants, the bush duiker (Sylvicapra grimmia) in Africa and the gray brocket (Mazama gouazoubira) in South America. These are ecological counterparts: both have body mass about 15 kg, tend to hide from view, are widespread in various forms of savanna, and eat mainly greens and fruits.

Comparing the appearance of these two species is particularly interesting because they belong to different families: Bovidae vs Cervidae. And one of the immediately noticeable convergences is that in the gray brocket the antlers are so modified to short spikes that they resemble horns.

However, in other respects the differences remain more noteworthy than the similarities. The African species looks more specialised than the South American species in various ways.

The following photo-pair shows that in the bush duiker the eyes are larger and placed more laterally on the head than is the case for the gray brocket: bush duiker https://www.robertharding.com/preview/764-6047/common-duiker-grey-duiker-bushduiker-sylvicapragrimmia-kruger-national/ vs gray brocket https://www.inaturalist.org/observations/91193009. This suggests that the predatory pressures are greater, and/or that the habitat is more open, in African than in South American savannas.

Compared to the African species, the South American species looks primitive and undifferentiated, similar to how I imagine an ancestral odocoileine to have looked: bush duiker https://animalia.us-east-1.linodeobjects.com/animals/photos/full/original/common-duiker-sylvicapra-grimmia-just-before-the-light-was-gone.jpg vs gray brocket
https://www.inaturalist.org/observations/61562922 and https://www.inaturalist.org/observations/67207093 and https://www.inaturalist.org/observations/71846673.

The muzzles and ears of the two species have different shapes: bush duiker https://www.euronews.com/green/2021/04/07/first-ever-images-of-mysterious-walter-s-duiker-captured-in-the-wild vs gray brocket https://www.inaturalist.org/observations/90656839 and https://www.inaturalist.org/observations/73497350.

Both species have a bare, moist rhinarium, but the form of the nose and mouth seems more specialised in the African species. There is a long preorbital gland in the African species, whereas there is no obvious facial gland in the South American species: bush duiker https://en.wikipedia.org/wiki/Common_duiker#/media/File:Common_Duiker_(Sylvicapra_grimmia)_(6011670419).jpg vs gray brocket https://www.inaturalist.org/observations/83213332.

The African species possesses a pedal flag by virtue of the dark/pale contrast on the feet. There is no such feature in the South American species: bush duiker http://janesjaunts.com/wp-content/uploads/2016/10/Grey-duiker.jpg vs gray brocket https://www.inaturalist.org/observations/9072964.

Both species have small tails with only modest potential for flagging. However, the tails are hardly convergent. The African species flags the dark/pale contrast on the narrow tail by flicking it intermittently while walking (https://www.inaturalist.org/observations/91165836), but does not flag the tail while fleeing. The South American species shows the white undersurface of the tail to some extent, but this lacks dark/pale contrast and any flagging of the tail while fleeing is limited by the shortness of the tail: bush duiker https://mldlect2rr41.i.optimole.com/vNyw-1Y-XKT679hT/w:auto/h:auto/q:auto/https://wildmoz.com/wp-content/uploads/2013/11/Duiker-Ewe.jpg and https://www.artranked.com/topic/Duiker#&gid=1&pid=22 vs gray brocket https://www.inaturalist.org/observations/72941220 and https://www.inaturalist.org/observations/48054069 and https://www.inaturalist.org/observations/7036758.

The following provide additional illustrations of the points made above: bush duiker https://www.destinationuganda.com/travel-guide/mammals/gray-duiker/ vs gray brocket https://www.inaturalist.org/observations/59383130 and https://www.inaturalist.org/observations/65598410.

Are these failures to converge owing to 'phylogenetic inertia', i.e. the inherent genetic limitations of being a bovid vs a cervid, or to the more intense predatory regime in Africa than in South America? In a thought-experiment in which the large carnivores in South America were as numerous and diverse as in Africa, would the gray brocket have evolved to resemble the bush duiker more closely?

To be continued...

הועלה ב-אוגוסט 16, 2021 10:53 לפנה"צ על ידי milewski milewski

תגובות

פורסם על-ידי milewski לפני כמעט 3 שנים
פורסם על-ידי milewski לפני כמעט 3 שנים

That's an interesting paper linked above -- am I reading that figure right that suggests mule and white-tailed deer are offshoots within the red brocket species group?! Wild.

I don't know if it's relevant to your comparison of gray brockets and bush duikers, but seems worth noting that bush duikers have a much larger range and, according to Red List assessors, have 14 recognized subspecies, while gouazoubira has no recognized subspecies (again according to its RL assessment team). Still, they have this to say about individual and regional variation: "In Brazil, regional, ecological and individual variations in coloration of M. gouazoubira have frequently been reported (Duarte 1996, Pinder and Leeuwenberg 1997, Rossi 2000). Light and brownish colors prevail in individuals living in grasslands, while forest populations tend to be darker and grayer, significant individual variation exists within a single population. M. gouazoubira can be readily distinguished from the most similar species, the Amazonian Brown Brocket Mazama nemorivaga, by its orange rump and dorsal side of the tail, larger rounded ears, smaller eyes and orbital cavities, and wider auditory bulla rather than chocolate brown rump and dorsal side of the tail, small pointed ears, large eyes, large orbital cavities, and narrow auditory bulla of the latter species (Duarte 1996, Rossi 2000)."

פורסם על-ידי muir לפני כמעט 3 שנים

@muir I've done my best to lay out subspeciation in the bush duiker in a Post of June 28, 2021, titled 'A succinct photo-guide to subspecies of the bush duiker' (https://www.inaturalist.org/journal/milewski?page=6). This species is variable enough, over its extensive range, that it could - with analogy to the splitting of Mazama gouazoubira and M. nemorivaga - also be split into at least two species, and indeed that is what Groves & Grubb (2011) did. So I don't see one form as much more widespread than the other. More soon...

פורסם על-ידי milewski לפני כמעט 3 שנים

@muir It is puzzling that Mazama gouazoubira differs from the bush duiker in defying any classification into subspecies. It varies, but along different lines. I also don't know what to make of the genetic analysis, other than to suspect that further genetic analyses will show different patterns.

פורסם על-ידי milewski לפני כמעט 3 שנים

@muir More evidence of the capacity of Mazama to surprise: https://www.youtube.com/watch?v=R7E8YGgep4E.

פורסם על-ידי milewski לפני כמעט 3 שנים

@muir At face value the following suggests that Mazama can resemble Odocoileus in flagging the tail in flight. However, I suspect that the label is incorrect and that this is actually a Colombian subspecies of O. virginianus. My reasons: hue too bright for Mazama nemorivaga, and the dark/pale contrast at the mouth is visible even in blurry footage at a distance. Please see https://www.youtube.com/watch?v=yGhaU8O_oV8.

פורסם על-ידי milewski לפני כמעט 3 שנים

The following shows that the tail is inert as Mazama gouazoubira walks around: https://www.youtube.com/watch?v=20ZMMtCdBxg. This contrasts with the bush duiker, which flags its tail intermittently while walking: https://www.youtube.com/watch?v=m3g4vFoNkLA.

פורסם על-ידי milewski לפני כמעט 3 שנים

The following illustrate the distress-call of the bush duiker: https://www.youtube.com/watch?v=eRc4aV76g_U and https://www.youtube.com/watch?v=VQ1B4Df3y_0 and https://www.youtube.com/watch?v=qKgTxf9wz5c. This is important because if the victim can call up other, larger carnivores before it dies it has a chance to escape in the confusion. Note how long, in the first video, it takes the cheetah (Acinonyx jubatus) to strangle the bush duiker, possibly because it is choosing to clamp the larynx rather than the trachea.

Odocoleine deer, possibly including Mazama, do utter a similar distress-call: https://www.youtube.com/watch?v=sNo-Qttru2Y and https://www.youtube.com/watch?v=LCXFX8d-4QY and http://www.whitetailsunlimited.com/i/p/bk_deertalk.pdf. However, is the larynx in this clade as resistant to compression as in cephalophin bovids?

פורסם על-ידי milewski לפני כמעט 3 שנים

The following shows how long it can take even the leopard (Panthera pardus) to strangle the bush duiker: https://www.youtube.com/watch?v=EaqW7xDzlJg. Again, I suspect that the strangling is inefficient because the leopard has chosen to clamp down on the larynx (which is sturdy) rather than the trachea below the larynx (which is more easily compressed but which might allow the victim to continue to utter its loud distress-call, risking the attraction of the spotted hyena, Crocuta crocuta, to the scene).

פורסם על-ידי milewski לפני כמעט 3 שנים

הוספת תגובה

כניסה או הרשמה להוספת הערות