milewski's Journal

September 23, 2023

A preliminary photo-guide to subspecies of the moose (Alces alces)

Also see https://www.inaturalist.org/journal/milewski/84595-adaptive-colouration-in-the-largest-living-cervid-the-moose-alces-alces#

There are currently many thousands of photos of Alces alces on the Web, particularly in iNaturalist.

However, remarkably few are labelled as to subspecies.

This is mainly because available information on the Web, on the differences in appearance among the nine recognised subspecies, is scant and confusing.

The distinctions I mention here are quantitative, the various subspecies overlapping broadly in features. No subspecies shows any categorically distinct feature, and every subspecies shows individual variation.

Instead, the variation is mainly

clinal from west to east, and, to a lesser degree,
latitudinal, following Bergmann's rule (https://en.wikipedia.org/wiki/Bergmann%27s_rule).

One subspecies, viz. caucasicus, became extinct before its colouration could be studied. Another, viz. cameloides, has seldom been photographed.

I list the eight extrant subspecies in longitudinal order, from Europe through Asia and Beringea, then through northwestern North America, to easternmost North America.

I assume that buturlini, of easternmost Asia, is similar to gigas, of Alaska, despite the sea barrier of the Holocene. This is based on the pattern seen in Cervus canadensis, which likewise invaded North America from Asia at the end of the Pleistocene (https://www.inaturalist.org/posts/53495-the-wapiti-instant-megafauna-for-america-made-in-china).

ALCES and PFIZENMEYERI

maximal development of pale on legs (= pedal flag), extending to fibular surface and, in some individuals, inner surface of buttocks
the fibular and pedal flags tend to be subsumed into a single extensive (pedofibular) pattern
ground-colour relatively pale

https://www.inaturalist.org/observations/74824385
https://www.inaturalist.org/observations/40353294
https://www.inaturalist.org/observations/99207826
https://www.inaturalist.org/observations/154930634
https://www.inaturalist.org/observations/40295954
https://www.inaturalist.org/observations/36579437

CAMELOIDES

proportionately small antlers

https://www.inaturalist.org/observations/100833248

BUTURLINI and GIGAS

torso and haunches tend to be two-toned
antlers proportionately large, and conspicuously pale (= cornual flag)

https://www.alamy.com/stock-photo-moose-alces-alces-gigas-bull-shedding-velvet-in-fall-colour-denali-43929349.html?imageid=B7C5FC8C-124E-45F2-8F36-BBECD7DF7619&p=82357&pn=1&searchId=acbc6be48e9684687a1dabcb59018a60&searchtype=0
https://www.alamy.com/stock-photo-moose-alces-alces-gigas-bull-shedding-velvet-in-fall-colour-denali-43929346.html?imageid=7FE5B5FD-8DD8-41F7-B30B-56A0F3B06FA5&p=82357&pn=1&searchId=acbc6be48e9684687a1dabcb59018a60&searchtype=0
https://create.vista.com/unlimited/stock-photos/343027438/stock-photo-cow-alaska-yukon-moose-denali-national-park-autumn/
https://www.flickr.com/photos/richardmcguire/27569540250
https://create.vista.com/unlimited/stock-photos/321158754/stock-photo-alaska-yukon-bull-moose-denali-national-park-alaska-autumn/
https://create.vista.com/unlimited/stock-photos/403606204/stock-photo-alaska-yukon-bull-moose-autumn/
https://create.vista.com/unlimited/stock-photos/347164868/stock-photo-alaska-yukon-bull-moose-autumn-denali-national-ark/
https://www.alamy.com/stock-photo-bull-moose-in-rut-near-powerline-pass-in-chugach-state-park-with-the-37569209.html?imageid=45F0C1B5-ABCD-4D02-9CE7-F9CA3C24FA8E&p=228600&pn=1&searchId=acbc6be48e9684687a1dabcb59018a60&searchtype=0
https://www.alamy.com/stock-photo-a-large-bull-moose-near-jewel-lake-road-in-anchorage-southcentral-125340208.html?imageid=9FBF37D1-13CB-4243-ADC6-9AA1AB5E2564&p=228198&pn=2&searchId=34e410ab9894add09ba6f8a15ae051be&searchtype=0

ANDERSONI and SHIRASI

plain-coloured, with minimal incidence of flags/large-scale patterns other than, perhaps, cornual flag
subspecies shirasi is essentially a relatively small form of andersoni, presumably owing to low latitude

https://www.inaturalist.org/observations/34440227
https://pngtree.com/freebackground/bull-moose-alberta-national-moose-meadow-photo_4533013.html
https://www.inaturalist.org/observations/24376876
https://www.inaturalist.org/observations/124213587
https://www.inaturalist.org/observations/87482894
https://www.inaturalist.org/observations/59045574
https://www.inaturalist.org/observations/50346121
https://www.inaturalist.org/observations/142792524
https://www.inaturalist.org/observations/97397248
https://www.agefotostock.com/age/en/details-photo/big-bull-moose-alces-alces-in-alberta-canada/ACX-acp82420

AMERICANA

maximal contrast between darkness of body and neck, and paleness of face (= facial flag), except for males in hard antler
minimal development of pale on legs and fibular surface; fibular flag/pedal flag absent in most individuals
ground-colour relatively dark

https://www.inaturalist.org/observations/199608

Posted on September 23, 2023 03:19 AM by

milewski | 4 comments | Leave a comment

September 21, 2023

The remarkable - and perhaps unnatural - success of the moose (Alces alces shirasi) in Colorado, as evidenced by observations in iNaturalist

@muir @matthewinabinett @davidbygott @aguilita

The moose (Alces alces) was virtually absent from Colorado (https://en.wikipedia.org/wiki/Colorado), at the time in the late 1800's when there was general depletion of large mammals in North America.

The appropriate subspecies, reintroduced/introduced to Colorado about 45 years ago, has become common here, at the southern extreme of the distribution of the species in the world.

Please see https://coloradooutdoorsmag.com/2021/03/03/colorado-moose-reintroduction/ and https://k99.com/colorado-has-one-of-the-fastest-growing-moose-populations-in-u-s/ and https://cpw.state.co.us/documents/wildlifespecies/mammals/moosereintroductionfactsheet.pdf and https://coloradoencyclopedia.org/article/moose#:~:text=In%20the%201970s%2C%20there%20were,to%20other%20parts%20of%20Colorado. and https://cpw.state.co.us/learn/Pages/MooseReintroductionProgram.aspx and https://www.denverpost.com/2017/06/27/colorado-moose-coexisting-people/ and http://peaktocreekfilms.com/comoosefilm and https://www.aspendailynews.com/moose-flourishing-in-colorado/article_56681247-d99a-5e80-bfc1-effeed27b741.html and https://www.steamboatpilot.com/news/many-more-moose-local-population-has-increased-dramatically-in-20-years/ and https://art19.com/shows/colorado-outdoors/episodes/899ceb83-518c-466f-9c27-76207e1ca166 and https://www.coloradovirtuallibrary.org/resource-sharing/state-pubs-blog/viewing-moose-in-colorado/ and https://cpw.state.co.us/learn/Pages/LivingwithWildlifeMoose.aspx.

The population in Colorado is reported to have reached 3,000 in 2023. Hunting of this, the largest game animal in the state, is now routinely permitted.

What is particularly remarkable about this project is the degree to which it has been apparent to naturalists. The photos of the moose in Colorado, posted in iNaturalist, already exceed 3,000 (https://www.inaturalist.org/observations?page=27&place_id=34&taxon_id=522193).

This means that, in a sense, there is one photo for every individual of the moose in Colorado. How could both the moose and iNaturalist have been more successful?

However, this spectacular success is open to interpretation.

Is this a case of restoration of the natural occurrence of the moose, or an anthropogenic extension of the distribution of the species?

Comments from Readers would be welcome.

Posted on September 21, 2023 01:59 AM by

milewski | 3 comments | Leave a comment

September 19, 2023

Adaptive colouration in the largest living cervid, the moose (Alces alces)

@muir @matthewinabinett @aguilita @tonyrebelo @jeremygilmore @beartracker @nyoni-pete @oviscanadensis_connerties @tandala @capracornelius @paradoxornithidae @wjcrins @hereinthewild @bluewhalenowhead @marshall20 @nat_zouieva @marina_gorbunova

At first sight, Alces alces seems nondescript in colouration (https://www.inaturalist.org/observations/19029155 and https://www.inaturalist.org/observations/4336204).

Valerius Geist, on pages 229-232 in Grzimek's Animal Life Encyclopedia (https://en.wikipedia.org/wiki/Grzimek%27s_Animal_Life_Encyclopedia), exaggerates somewhat when he states:
"European moose are dark brown with white legs, and American moose are black with a light saddle patch on the back, light-brown legs, and facial markings that vary between the sexes. In the female the face and nose are red-brown, while the bull has a black nose".

The following verify that the face tends to be paler than the neck and torso, in females of A. alces in North America (https://www.inaturalist.org/observations/178372818 and https://www.dreamstime.com/bull-moose-water-image259482733 and https://www.inaturalist.org/observations/8288949) and to some degree also in Europe (https://www.inaturalist.org/observations/175016790 and https://www.inaturalist.org/observations/174910182).

However, the difference in tone between the face and the rest of the figure is noticeable only in A. a. americana (see details in comment below, titled FACIAL FLAG IN ALCES ALCES AMERICANA).

Sexual differences in the colouration of the pelage of A. alces, beyond the face, are remarkably limited (https://www.inaturalist.org/observations/42138516 and https://www.youtube.com/watch?v=bujmogxgtp4 and https://www.youtube.com/watch?v=tsZUaGIt-r8 and https://www.google.com.au/search?q=Moose+mating&sca_esv=567242663&sxsrf=AM9HkKnsy5ar25NAM_QxNryBAc4ELWtrPA%3A1695294882706&source=hp&ei=oiUMZd3UJtixoATxmJzQAg&iflsig=AO6bgOgAAAAAZQwzssn9atv3I45ihgDieVv4Z3QxBVPr&ved=0ahUKEwjdrZPKybuBAxXYGIgKHXEMByoQ4dUDCAs&uact=5&oq=Moose+mating&gs_lp=Egdnd3Mtd2l6IgxNb29zZSBtYXRpbmcyBRAuGIAEMgUQABiABDIFEAAYgAQyBRAAGIAEMgUQABiABDIFEAAYgAQyBRAAGIAEMgUQABiABDIFEAAYgARImklQzg5Y-j5wAXgAkAEAmAGuA6AB_BiqAQkwLjEuOS4xLjG4AQPIAQD4AQGoAgrCAgcQIxjqAhgnwgIEECMYJ8ICCxAAGIAEGLEDGIMBwgIREC4YgAQYsQMYgwEYxwEY0QPCAgsQLhiDARixAxiABMICCxAAGIoFGLEDGIMBwgIHECMYigUYJ8ICCBAuGIoFGJECwgIIEAAYgAQYsQPCAggQABiKBRiRAsICCxAuGIoFGLEDGJECwgILEC4YgAQYxwEYrwE&sclient=gws-wiz#fpstate=ive&vld=cid:25246631,vid:YvNAJLygu9w,st:0).

Infants, although noticeably paler than adults, have colouration so plain that it, too, is nondescript (https://www.inaturalist.org/observations/4097272).

Seasonal changes in colouration are limited. There is a single annual molt (https://www.inaturalist.org/observations/108839229 and https://www.inaturalist.org/observations/125310240 and https://www.inaturalist.org/observations/22740621 and https://www.inaturalist.org/observations/20313689) in spring/early summer. The pelage, after being worn and weathered for a year, fades somewhat (https://www.inaturalist.org/observations/8449050).

However, the pelage remains dark enough in winter to be conspicuous against snowy backgrounds, even at distance (https://www.inaturalist.org/observations/148799385 and https://www.inaturalist.org/observations/135953868).

Despite the initial nondescript impression, my close scrutiny has revealed several noteworthy patterns of colouration, which deserve names, in A alces.

For example, the following shows a fibular flag, anterior auricular semet, and buccal semet in an adolescent female individual in the spring season, in Alces alces gigas: https://www.inaturalist.org/observations/163382249.

Although these features are subtle, individually variable, and perhaps seasonally variable, most of them occur in most of the nine subspecies.

CORNUAL FLAG

The antlers of A. alces, borne seasonally by males, tend to be conspicuously pale on the upper (dorsal) surface, in the subspecies with the largest antlers.

The cornual flag is derived essentially from the natural paleness of dry bone (https://www.inaturalist.org/observations/127384241).

However, in A. a. gigas and presumably A. a. buturlini,

the antlers are paler than those of other cervids,
the palmate surface of the antlers is far larger than in other cervids (https://www.cbc.ca/news/canada/north/possible-world-record-size-moose-shot-in-yukon-1.2075442), and
the paleness of the antlers contrasts with the darkness of the rest of the figure.

This makes the antlers conspicuous, even at distance (https://www.inaturalist.org/observations/4612251).

The cornual flag in A. gigas

is apparent mainly in autumn (https://www.josephfiler.com/photo/moose-alaska-2833/ and https://create.vista.com/unlimited/stock-photos/313125682/stock-photo-alaska-yukon-bull-moose-autumn-denali-national-park-alaska/), and
hypothetically functions intraspecifically, aiding sexual attraction, male advertisement, and masculine rivalry.

Cornual flag in Alces alces gigas:
https://www.inaturalist.org/observations/183421987
https://www.mediastorehouse.com/nature-picture-library/danny-green/moose-bull-alces-alces-walking-forest-clearing-15316906.html
https://www.inaturalist.org/observations/183421709
https://www.inaturalist.org/observations/458438
https://create.vista.com/unlimited/stock-photos/314205726/stock-photo-alaska-yukon-bull-moose-autumn/
scroll in https://www.akwildlife.org/news/wildlife-contest-2022

Cornual flag in Alces alces andersoni:
https://www.inaturalist.org/observations/61216754
https://www.mediastorehouse.com.au/design-pics/animals/bull-moose-peter-lougheed-provincial-park-18135905.html

Possible cornual flag in Alces alces shirasi:
https://www.inaturalist.org/observations/147438040

Possible cornual flag in Alces alces americana:
https://visitsunsetcountry.com/hunting/moose

Incipient cornual flag in Alces alces alces:
https://www.mediastorehouse.com.au/nature-picture-library/2009-highlights/moose-alces-alces-taiga-woodland-15260457.html

FIBULAR FLAG

https://www.inaturalist.org/observations/8660028
https://www.behance.net/gallery/99847037/Bull-Moose-Bearspaw-Alberta?tracking_source=search_projects_recommended|Bull+Moose
https://www.closertothewild.com/gallery/p/style-01-slgxj-738cj-bnn5s-3e26g-rewyc-36zgb
https://www.shutterstock.com/image-photo/winter-sunny-day-young-moose-cow-2125295291
https://www.dreamstime.com/stock-photography-bull-moose-image21621952
https://www.dreamstime.com/wet-cow-moose-standing-forest-image183436539
https://www.inaturalist.org/observations/61913264
https://www.inaturalist.org/observations/134851345
https://www.alamy.com/moose-in-jasper-canada-image348236851.html?imageid=D433EBFE-8D95-490C-BEAE-57C59F4AD201&p=466062&pn=5&searchId=59e69530b8d11356f2f4edcf3f5ed079&searchtype=0
https://www.alamy.com/young-elk-age-about-one-year-male-on-the-edge-of-the-spring-forest-image259781936.html?imageid=A982D41F-5A22-489F-99A6-9C17CFE32440&p=270423&pn=7&searchId=17a6b40e8c8fbef3da4fa2fc3209a279&searchtype=0
https://www.inaturalist.org/observations/35513964
https://www.inaturalist.org/observations/137101323

Please see https://www.inaturalist.org/journal/milewski/84573-a-new-feature-of-adaptive-colouration-in-ungulates-the-fibular-flag-part-2-alces-alces-cervidae#

The fibular flag is best-developed in the nominate subspecies, A. a. alces, of Europe and western Russia, in which it tends to extend to the inner surface of the buttock (https://www.inaturalist.org/observations/149814632) and/or the hock.

The role of sheen deserves investigation.

The following shows the extreme development of the pale pelage in A. alces, in which it encompasses most of the hindleg (https://www.inaturalist.org/observations/112176729).

PEDAL FLAG

A case can be made that A. alces possesses a pedal flag, in some individuals. This applies particularly to the nominate A. a. alces (https://www.inaturalist.org/observations/142224245 and https://www.inaturalist.org/observations/142020077).

This pedal flag consists of conspicuous pale on the hocks and carpals (particularly on the posterior surface of the carpals), extending to varying extent down the lower limbs towards the hooves, and connected to the fibular flag where the latter is present (https://www.alamy.com/stock-photo-bull-moose-crossing-the-road-denali-national-park-alaska-usa-29001311.html?imageid=984BF44C-2D52-4F90-BF1B-9372BC0185A4&p=29229&pn=1&searchId=acbc6be48e9684687a1dabcb59018a60&searchtype=0).

The fetlocks and pasterns themselves are not whitish (https://www.shutterstock.com/image-photo/front-split-hoofs-moose-standing-on-727556374 and https://www.inaturalist.org/observations/134620144 and https://www.inaturalist.org/observations/35655176 and https://www.inaturalist.org/observations/55865091), except in some individuals of the nominate A. a. alces (https://www.mostphotos.com/en-us/25919107/moose-or-european-elk-alces-alces-hooves).

A factor undermining the validity of a pedal flag in A. alces is that, for much of the year, the pale on the legs is inconspicuous against a background of snow (https://www.inaturalist.org/observations/42170238 and https://www.inaturalist.org/observations/143176855).

https://www.inaturalist.org/observations/51436631
https://www.inaturalist.org/observations/33064841
https://www.dreamstime.com/large-brown-black-bull-moose-standing-park-image193883277
https://www.dreamstime.com/stock-photo-moose-bull-elk-standing-road-image66248282

To the degree that the pedal flag is valid in A. alces, it is linked to, and subsidiary to, the fibular flag (https://www.inaturalist.org/observations/142257592). In A. a. alces, the joint pattern can perhaps be called a pedofibular flag.

ANTERIOR AURICULAR SEMET

Please see https://www.inaturalist.org/journal/milewski/55694-ten-best-illustrations-of-auricular-semets-in-deer#

One of the most consistent patterns of colouration in A. alces is a small-scale dark/pale contrast on the anterior base of the ear pinna (https://www.dreamstime.com/stock-photo-moose-north-america-elk-europe-alces-alces-largest-extant-species-deer-family-moose-distinguished-image50713041).

The location of this pattern is such that it accentuates the movements of the ears, in vigilance and emotional expression. Such accentuation is hypothetically adaptive in social (intraspecific) interactions, and may additionally function in anti-predator, defensive displays.

Several other genera of cervids possess auricular semets. However, that of A. alces is restricted to the anterior surface, and has its own particular configuration.

The following show the anterior auricular semet in various subspecies of A. alces:

A. a. alces:
https://www.inaturalist.org/observations/122347911
https://www.inaturalist.org/observations/119099452

A. a. americana:
https://www.inaturalist.org/observations/183781173
https://www.inaturalist.org/observations/165912205
https://www.inaturalist.org/observations/82792870

A. a. gigas:
https://www.inaturalist.org/observations/172965903
https://www.inaturalist.org/observations/15093287

A. a. andersoni:
https://www.inaturalist.org/observations/170210903
https://www.inaturalist.org/observations/166029819

A. a. shirasi:
https://www.inaturalist.org/observations/174251418

In Alces alces, unlike another large cervid partly sympatric with it, the auricular semet does not disappear in mature males (see https://colombia.inaturalist.org/journal/milewski/69281-intriguing-new-sexual-dimorphism-in-the-wapiti).

The following show that the anterior auricular semet, although absent in newborns, develops before the infantile colouration is lost, and before the muzzle develops:
https://www.inaturalist.org/observations/133091834
https://www.dreamstime.com/stock-photo-small-moose-few-days-standing-woods-next-to-pine-tree-image53403214.

BUCCAL SEMET

The peculiar muzzle of A. alces is so distracting that the colouration around the mouth may go unnoticed. Furthermore, this colouration is subtle and individually variable.

However, there tends to be a pattern in which the broadly dark lower lip is offset by pale ventral to the gape. The clearest illustration is https://www.inaturalist.org/observations/65908659.

This pattern of colouration hypothetically accentuates the chewing movements (https://www.inaturalist.org/observations/88438853). This hypothetically aiding vigilance during rumination, when two or more adult/adolescent individuals rest within sight of each other (https://www.dreamstime.com/stock-photos-moose-image20463023).

Buccal semet in Alces alces buturlini:
https://www.alamy.com/female-moose-alces-alces-in-profile-kronotsky-zapovednik-nature-reserve-kamchatka-peninsula-russian-far-east-december-image263189582.html?imageid=8337E994-E81B-42C7-9D06-32050AEFBBE0&p=216449&pn=1&searchId=693543c6c0b82d61e979d27d23f7d17f&searchtype=0

Buccal semet in Alces alces gigas:
https://www.inaturalist.org/observations/153275621
https://www.inaturalist.org/observations/2646616
https://www.inaturalist.org/observations/13172080

Variation in this feature deserves further investigation. The pattern seems clearest in spring, when the worn, weathered pelage has faded but the dark on the lower lip has not faded (https://www.inaturalist.org/observations/5155147). The following shows the minimal expression of the buccal semet (https://www.inaturalist.org/observations/114831099), but it is unclear whether this is because of the summer season or individual variation.

Of all the features described here, the buccal semet is the most precocial, in the sense that the lower lip is dark even in infants (https://www.inaturalist.org/observations/29864809 and https://www.123rf.com/photo_188787653_portrait-of-a-walking-cow-moose-and-her-calf-alces-alces.html and https://www.inaturalist.org/observations/7440096).

Posted on September 19, 2023 12:46 AM by

milewski | 26 comments | Leave a comment

September 18, 2023

A new feature of adaptive colouration in ungulates: the fibular flag, part 2: Alces alces (Cervidae)

...continued from https://www.inaturalist.org/journal/milewski/70368-a-new-feature-of-adaptive-colouration-in-ungulates-the-fibular-flag-part-1-raphicerus-bovidae#

The moose (Alces alces, https://www.inaturalist.org/taxa/522193-Alces-alces) possesses a fibular flag.

In this large-bodied cervid, the fibular flag covers the inner, and often also the outer, surfaces of the hindleg above the hock.

https://www.inaturalist.org/observations/107522426

The fibular flag of the moose is individually and seasonally variable, and also depends on illumination.

In the summer pelage, the fibular flag is poorly-developed (https://www.inaturalist.org/observations/179584604 and https://www.inaturalist.org/observations/170204579 and https://www.inaturalist.org/observations/79012704 and https://www.inaturalist.org/observations/31202401).

The following illustrate the fibular fag in Alces alces shirasi:
https://www.inaturalist.org/observations/22730337
https://www.inaturalist.org/observations/165808747
second photo in https://www.inaturalist.org/observations/183755724
https://www.inaturalist.org/observations/73121671
https://www.inaturalist.org/observations/4482611
https://www.inaturalist.org/observations/4308293

Fibular flag in Alces alces alces:
https://www.inaturalist.org/observations/35274716
https://www.inaturalist.org/observations/183855765
https://www.inaturalist.org/observations/183819576
https://www.inaturalist.org/observations/153934473
https://www.inaturalist.org/observations/46595843
https://www.inaturalist.org/observations/2922305

In nominate A. a. alces, the pale feature in question can be so expansive that the term 'fibular flag' no longer applies (https://www.inaturalist.org/observations/112176729).

ADDITIONAL ILLUSTRATIONS OF FIBULAR FLAG IN ALCES ALCES

https://www.inaturalist.org/observations/116777704

https://www.inaturalist.org/observations/115702446

https://www.inaturalist.org/observations/115325363

https://www.inaturalist.org/observations/115314501

https://www.inaturalist.org/observations/114790758

https://www.inaturalist.org/observations/112619055

https://www.inaturalist.org/observations/135414451

https://www.inaturalist.org/observations/115470987

https://www.inaturalist.org/observations/125843310

https://www.inaturalist.org/observations/121343112

https://www.inaturalist.org/observations/120257251

https://www.inaturalist.org/observations/129646111

https://www.inaturalist.org/observations/119626107

https://www.inaturalist.org/observations/119092478

https://www.inaturalist.org/observations/116521066

https://www.inaturalist.org/observations/134851345

https://www.inaturalist.org/observations/116789745

https://www.alamy.com/stock-photo-moose-from-behind-142718462.html

https://www.inaturalist.org/observations/131879614

https://www.inaturalist.org/observations/126552983

https://www.inaturalist.org/observations/125993345

Why A. alces is unusual, in possessing a fibular flag, is worthy of further investigation.

Posted on September 18, 2023 10:08 AM by

milewski | 13 comments | Leave a comment

September 14, 2023

The hyperfecal-hypofecal continuum in consumers of plant matter

(writing in progress)

Faeces exist, almost as surely as eating exists.

The basic principles behind the production of faeces are of costs/benefits, and diminishing returns.

Various components of food have various digestibilities. The least digestible components are - depending on the species of consumer and its particular digestive system - discarded, as the biological equivalent of rubbish.

This occurs despite

the investment of time and energy made in acquiring the food, and
the risks taken as part of this foraging.

It would be naive to expect members of the 'food-chain' to be thorough processors, simply converting all of the matter and energy they consume.

However, what are remarkable are

the degree to which various animals differ in the thoroughness of their digestive process,
the quantity of faeces produced relative to food consumed, and
the finding that it is not necessarily the most fibrous diets that produce the most faeces.

Some animals 'overproduce' faeces in the sense that they seem to extract little of what they eat, whereas others 'underproduce' faeces in the sense that they extract virtually all of what enters the digestive process. Most animals are intermediate.

It is the range in this continuum - which I dub the 'hyperfecal/hypofecal continuum' - that is greater than appreciated by many or most naturalists.

In this Post, I illustrate the hyperfecal/hypofecal continuum by examining its extremes, in the form of

the giant panda (Ailuropoda melanoleuca, https://www.inaturalist.org/taxa/41659-Ailuropoda-melanoleuca), and
fungus-culturing termites in genus Macrotermes (https://www.inaturalist.org/observations?place_id=any&taxon_id=346639&view=species).

Both the giant panda and Macrotermes forage on extremely fibrous matter, composed largely of cellulose (https://en.wikipedia.org/wiki/Cellulose) and lignin (https://en.wikipedia.org/wiki/Lignin).

However,

the former acts as a 'fecal factory', whereas the latter recycles the fibre repeatedly within the system, until virtually nothing emerges; and
the former seems to defecate for a living, whereas the latter - at least at the scale of the whole colony in this eusocial insect - hardly produces anything that can be called faeces.

Indeed, the digestive process of Macrotermes is such that it treats the faeces of various herbivores as part of the diet.

Crucial for understanding the hyperfecal/hypofecal continuum are

the role of microbes, particularly fungi (https://en.wikipedia.org/wiki/Fungus#:~:text=Fungi%20include%20symbionts%20of%20plants,and%20exchange%20in%20the%20environment.), and particularly aerobic fungi (https://en.wikipedia.org/wiki/Termitomyces), and
the repeated reingestion of food, slowing incremental breakdown of every particle of fibre.

In the giant panda, the gut is short and simple. There is minimal development of gut-chambers in which food is delayed for breakdown by bacteria and other microbes.

The result is that

virtually all of the fibre emerges as faeces, and
even the digestible proteins, such as chlorophyll, are only partly digested (https://www.pdxwildlife.com/does-panda-poop-stink/).

In other words, the hyperfecality of the giant panda is such that even the greenness of the plant matter consumed seems not to be converted to brown.

(writing in progress)

Posted on September 14, 2023 11:25 PM by

milewski | 3 comments | Leave a comment

September 08, 2023

Is there a system of eating in which the systemically toxic excess of carbohydrate can be practically mitigated?

(writing in progress)

The main aim of this Post is to suggest a practical way of 'having one's cake and eating it too', in terms of healthy diets and systems of eating, in the human species (Homo sapiens).

The particular topic is the mitigation of the effects of carbohydrate (https://en.wikipedia.org/wiki/Carbohydrate), which tends to be consumed in excessive quantity and with excessive frequency in the modern world.

CONTEXT

In industrial economies today, so much carbohydrate is available to the human species that this substance has become, effectively, an environmental toxin.

This contrasts with the ancestral condition. Hunter-gatherers lived in an environment in which carbohydrate was seasonal and relatively scarce. The physiological nature of the human body is such that the former regime remains the healthier one.

Many chronic illnesses in modern populations can be ascribed to excess carbohydrate.

The mechanism is that concentrations of glucose in the blood tend to be excessive, with excessive frequency. This causes disruption of the normal functions of the pancreas (https://en.wikipedia.org/wiki/Pancreas), mainly via the insulin (https://en.wikipedia.org/wiki/Insulin) secreted by this gland.

The results include, amongst others,

obesity (https://en.wikipedia.org/wiki/Obesity),
diabetes (https://en.wikipedia.org/wiki/Diabetes),
metabolic syndrome (https://en.wikipedia.org/wiki/Metabolic_syndrome#:~:text=Metabolic%20syndrome%20is%20a%20clustering,Dysmetabolic%20syndrome%20X), and
insulin resistance (https://en.wikipedia.org/wiki/Insulin_resistance#).

It seems safe to say that if all added sugar were eliminated from human diets, populations would be healthier.

Furthermore, it seems likely that most human individuals would be healthier on a diet lacking starchy foods - on the basis that starch is rapidly digested to glucose in the mouth and small intestine, causing metabolic stress similar to that resulting from consumption of sugar itself.

However, for most individuals in modern societies, it seems impractical - for many and diverse reasons - to eliminate sugar and starch entirely from the diet.

For example, the category of 'healthy vegetables' includes various tubers and fleshy fruits, which are starchy enough to boost glucose in the bloodstream, within minutes of consumption.

Examples of tubers include

carrot (https://en.wikipedia.org/wiki/Carrot),
beetroot (https://en.wikipedia.org/wiki/Beetroot), and
sweet potato (https://en.wikipedia.org/wiki/Sweet_potato).

Starchy examples of fleshy fruits are the various domestic species of Cucurbita (https://en.wikipedia.org/wiki/Pumpkin) and https://en.wikipedia.org/wiki/Cucurbita and https://en.wikipedia.org/wiki/Butternut_squash).

Furthermore, this undesirable boost would remain to some extent even if the foods mentioned above were eaten raw. This is because their glycemic index (https://en.wikipedia.org/wiki/Glycemic_index) exceeds x.

THE PROBLEM

It may be practicable to avoid added sugar, as well as sweet fleshy fruit-pulp and -juice, from one's diet.

However, for most individuals, it is impractical to avoid carbohydrates altogether. This would mean a ketogenic diet, which is physiologically possible for many or most individuals, but not sustainable given the social and economic realities of our lives.

So, how can one continue to consume, on a routine basis, some quantity of starchy food without ill effects?

THE CRUCIAL PRINCIPLES

I suggest that three crucial principles are that

the rate at which glucose enters the bloodstream depends largely on the dilution of carbohydrate in proteins, lipids, fibre, and water, in the stomach,
it is as important to reduce the frequency of 'insulin spikes' as it is to reduce the sizes of these 'spikes', and
mildly sweet food (e.g. wild berries) can safely be eaten provided that this is at most once per day, reducing the frequency of any 'insulin spikes' to one per day or less.

The best illustration of the first of the above principles is that peanut (Arachis hypogoea, https://en.wikipedia.org/wiki/Peanut) and chickpea (Cicer arietinum, https://en.wikipedia.org/wiki/Chickpea) contain about a third starch, but are non-insulinogenic. The starch is safely 'smuggled' into then food by virtue of being thoroughly mixed with protein and, in the case of peanut, lipids.

Posted on September 08, 2023 08:55 PM by

milewski | 0 comments | Leave a comment

September 04, 2023

A comparison of the plains viscacha (Lagostomus maximus) and the black-tailed prairie dog (Cynomys ludovicianus)

(writing in progress)

Two rodents invite comparison, viz.

the plains viscacha (Lagostomus maximus, https://www.inaturalist.org/taxa/43824-Lagostomus-maximus) of South America, and
the black-tailed prairie dog (Cynomys ludovicianus, https://www.inaturalist.org/taxa/46179-Cynomys-ludovicianus) of North America,

The former is a hystricomorph (https://en.wikipedia.org/wiki/Hystricomorpha) belonging to the Chinchillidae (https://en.wikipedia.org/wiki/Chinchillidae), whereas the latter is a sciuromorph (https://en.wikipedia.org/wiki/Sciuromorpha) belonging to the Sciuridae (https://en.wikipedia.org/wiki/Squirrel).

The two species are comparable because both are

the largest rodents in their habitats (which are mainly mesic and treeless in both cases),
herbivorous (with green grass as a staple), to the degree of being regarded as unwelcome competitors for domestic livestock,
burrowing (mainly for the excavation of permanent living quarters)
colonial (with several family groups sharing a single system of burrows),
sexually dimorphic (with males more massive than females),
precocial (with large, well-developed newborns), and
unusually vocal for rodents (with various calls used socially and in reaction to predators),

However, the plains viscacha is the more specialised species in its role as a herbivore, and the more aberrant w.r.t. rodents in general.

This is because the plains viscacha

has about five-fold the body mass of the black-tailed prairie dog, as well as being the more sexually dimorphic of the two,
has a diet more strictly herbivorous (and graminivorous) than that of the black-tailed prairie dog,
forms and maintains lawns,
is caecotrophic (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC8208658/), as opposed to having hindgut fermentation without caecotrophy in the case of the black-tailed prairie dog, and
forages throughout the year, as opposed to spending the winter partly in torpor in the case of the black-tailed prairie dog.

with caecotrophic digestion

Nocturnal vs diurnal

Unlike the black-tailed prairie dog, the plains viscacha forages gregariously.

Posted on September 04, 2023 12:32 AM by

milewski | 11 comments | Leave a comment

September 01, 2023

Surprisingly, the tiger exceeds the lion in expressing bipedality

@paradoxornithidae @beartracker @matthewinabinett @maxallen @dinofelis @zarek @botswanabugs @simontonge @ldacosta @marcelo_aranda @tonyrebelo @jeremygilmore

The lion (Panthera leo) and the tiger (Panthera tigris)

are closely related (https://en.wikipedia.org/wiki/Panthera),
have a similarly limited ability to climb the vertical boles of trees, and
have hindlegs of similar length relative to the length of the torso (https://www.123rf.com/photo_43556506_lion-walking-in-profile.html and https://stock.adobe.com/au/images/lioness-walking-at-sun-rise/144599399 and https://www.picxy.com/photo/991738 and https://www.researchgate.net/figure/One-of-the-female-tigers-walking-on-a-track-plot-TP-during-the-sampling-period-in_fig2_263702911).

Therefore, one would not expect these species to differ in their ability to adopt bipedal postures.

Yet there does seem to be a difference. The lion has not been observed to adopt bipedal postures to the extent and degree observed in the tiger.

The photographic evidence falls into three categories, namely

circus acts,
intraspecific antagonism, and
territorial marking.

CIRCUS ACTS

Please see http://www.chinadaily.com.cn/china/2007-11/12/content_6247657.htm.

I have yet to see a similar photo for the lion.

INTRASPECIFIC ANTAGONISM

Both in play and in combat, the lion and the tiger rear up on their hindlegs to some degree.

However, the lion differs considerably from the tiger, as evident from the many photos (and a few video clips) shown at the end of this Post.

In combat/sparring, the tiger adopts a ritualised transitory posture, in which the animal stands/locomotes briefly on its hindlegs. At the same time, it spreads its forelegs - mirrored by its antagonist - in a way yet to be photographed/filmed in the lion.

TERRITORIAL MARKING

The tiger often stands bipedally at the base of trees, in order to

rub its jowls on the bole, and/or
make conspicuous scratch-marks on the bark:

This is shown in the following:

https://www.reddit.com/media?url=https%3A%2F%2Fi.redd.it%2Fuoahj1bjnqo71.jpg

https://www.alamy.com/stock-photo-siberian-tiger-panthera-tigris-altaica-scent-marking-tree-20328118.html?imageid=BE2CBD26-4D9F-412D-9760-8A936B782357&p=3663&pn=1&searchId=ae1ddc7afd7c91c5b34e5ea40340ae72&searchtype=0)

https://www.alamy.com/bengal-tiger-standing-on-rear-legs-and-stretching-up-to-scratch-a-tree-trunk-a-way-of-marking-territory-ranthambore-national-park-sawai-madhopur-rajasthan-india-asia-image415610844.html?imageid=434A55DB-E1B5-454A-9AFB-0ABBE0E03AEC&p=148099&pn=1&searchId=ac5e77b5654481dc9dd3cfd10b494232&searchtype=0

https://www.alamy.com/stock-photo-a-bengal-tiger-making-scratch-marks-on-a-tree-ranthambore-tiger-reserve-19805813.html?imageid=A6A19F20-867F-4798-9C5E-288C81120E73&p=57430&pn=1&searchId=ac5e77b5654481dc9dd3cfd10b494232&searchtype=0

https://www.alamy.com/stock-image-17-months-old-bengal-tiger-cub-scratch-marking-tree-early-morning-168781557.html?imageid=B611EA12-E9FC-4024-9BB7-5842A5928E0E&p=571810&pn=1&searchId=ac5e77b5654481dc9dd3cfd10b494232&searchtype=0

https://www.alamy.com/siberian-tiger-panthera-tigris-altaica-male-tree-marking-captive-e-russia-fl011327-s-image552952737.html?imageid=AC955E3E-81F4-4076-B908-6C1AC79EC785&p=2130853&pn=1&searchId=ac5e77b5654481dc9dd3cfd10b494232&searchtype=0

https://www.alamy.com/stock-photo-siberian-tiger-panthera-tigris-altaica-immature-male-scratching-tree-47790412.html?imageid=52BBCBD9-A25D-4D79-AAFB-5A24EA412833&p=11592&pn=1&searchId=ac5e77b5654481dc9dd3cfd10b494232&searchtype=0

https://www.alamy.com/stock-photo-siberian-tiger-panthera-tigris-altaica-adult-male-scratching-tree-47790443.html?imageid=2ED629B6-5141-43A0-81BF-F4CBFC7A666D&p=11592&pn=1&searchId=ac5e77b5654481dc9dd3cfd10b494232&searchtype=0

https://www.gettyimages.com.au/detail/photo/tiger-sharpening-claws-on-tree-standing-on-hind-royalty-free-image/200329297-001

https://www.quora.com/What-sort-of-tree-does-a-tiger-choose-to-scratch-in-the-wild-and-why

https://www.gettyimages.co.uk/detail/news-photo/siberian-tiger-adult-standing-up-to-scratch-claws-on-tree-news-photo/1143011570?adppopup=true

https://www.gettyimages.co.uk/detail/news-photo/asie-inde-rajasthan-parc-national-de-rathambore-tigre-du-news-photo/953401420?adppopup=true

https://dailywildlifephoto.nathab.com/contests/1-2016-wildlife-photo-contest/photos/1966-tiger-marking-the-tree

https://www.stevebloom.com/index.php?page=single&id=503140-BS1

https://www.dreamstime.com/stock-photo-tiger-climbing-tree-image38107520

I have yet to find any photo of the lion adopting a similar posture, in this context.

The closest are https://www.123rf.com/photo_100367973_lion-standing-on-his-hind-legs-at-a-tree-in-morning-light.html and https://www.stevebloom.com/index.php?page=single&id=506649-BS1 and https://www.istockphoto.com/photo/female-white-lion-scratching-tree-bark-gm1195489631-340768364 and https://www.shutterstock.com/image-photo/african-lion-felis-leo-marking-territory-528839812.

DISCUSSION

It would be particularly surprising if the tiger exceeds the lion in bipedal ability, given that it is the more massive of the two species (https://www.diffen.com/difference/Lion_vs_Tiger and https://www.quora.com/Are-tigers-and-lions-the-same-size).

Perhaps the difference can be explained partly by the fact that the lion typically inhabits open vegetation, whereas the tiger typically inhabits wooded vegetation.

These findings should raise a search-image, among naturalists, for bipedal modes in

the various other species of Panthera (https://www.pinterest.co.uk/pin/118430665190308481/ and https://i.imgur.com/cj8aznU.jpg), as well as
Puma concolor (https://www.alamy.com/puma-felis-concolor-standing-on-its-back-paws-image6401372.html and https://www.agefotostock.com/age/en/Stock-Images/cougar-scratching-tree.html), and Acinonyx jubatus (https://www.artflakes.com/en/products/gepard-in-namibia-african-cheetah-standing and https://www.stevebloom.com/index.php?page=single&id=500915-BS1).

The leopard (Panthera pardus) has repeatedly been photographed sitting bolt-upright in vigilance (https://www.dailymail.co.uk/news/article-2143766/Simples-Leopard-surprises-safari-goers-stands-legs-like-meerkat-stalk-dinner.html and https://m.facebook.com/IntenseWalls/photos/a.527980250611783/3363912873685159/?type=3&_rdr). However, this is different from standing.

Bipedal standing during marking on trees is the least important of the three lines of evidence considered here. This is because the animal does not free-stand, but instead uses the bole to support itself with its forepaws.

I offer the hypothesis that it is the lion, rather than the tiger, that is aberrant, among 'big cats', in its (lack of) expression of bipedality.

INTRASPECIFIC ANTAGONISM: PANTHERA LEO:

https://depositphotos.com/photo/lions-fighting-4018529.html

https://www.gettyimages.com.au/detail/photo/young-male-lions-playing-with-each-other-jumping-royalty-free-image/171116536?adppopup=true

https://www.gettyimages.com.au/detail/news-photo/two-lioness-panthera-leo-sparring-masai-mara-national-news-photo/1371855370?adppopup=true

https://www.gettyimages.com.au/detail/photo/lion-and-lioness-fighting-fighting-masai-mara-royalty-free-image/1027619214?adppopup=true

https://www.flickr.com/photos/jaffles/24851778939

https://www.dreamstime.com/two-lions-fiercely-fighting-their-pride-creating-using-generative-ai-tools-pair-engage-territorial-fight-image280293785

https://www.mediastorehouse.com.au/nature-picture-library/2020-april-highlights/lion-panthera-leo-males-mock-fighting-play-19960056.html

https://www.flickr.com/photos/sheldrickfalls/32953986190

https://www.facebook.com/thewildestafrica/photos/a.2245536979094388/2405427793105305/?type=3&locale=zh_CN

https://www.youtube.com/watch?app=desktop&v=Q4Bevh-jnaQ