milewski's Journal

Naturalists are used to thinking that animals tend to have adaptive colouration that blends into the surroundings.

However, in open environments where it is hard to hide, certain animals opt instead to have conspicuous colouration.

One well-recognised pattern of conspicuous colouration in medium-size to large animals is the pied pattern (https://www.tripadvisor.com/LocationPhotoDirectLink-g469397-d469470-i381557535-Bontebok_National_Park-Swellendam_Overberg_District_Western_Cape.html and https://www.flickr.com/photos/myplanetexperience/50867088327).

This consists of a dark/pale patchwork, too bold to function disruptively. I.e. pied colouration is too conspicuous for the figure to be camouflaged.

However, there is another conspicuous pattern, less familiar but obvious in certain mammals. I provisionally call this 'caleonic', because it deserves a name, but has not previously been described.

In the caleonic pattern, it is as if the figure is 'highlit' by a flame below.

Caleonic colouration seems to have arisen repeatedly in lineages as diverse as

• bovids (https://www.alamy.com/male-critically-endangered-dama-gazelle-nanger-dama-northern-africa-image364952487.html and https://mammiferesafricains.org/2022/02/oryx-algazelle/),
• antilocaprids (https://www.inaturalist.org/observations/122793376 and https://www.inaturalist.org/observations/122107163 and https://www.inaturalist.org/observations/120998764 and https://www.inaturalist.org/observations/119233203),
• camelids (https://theconversation.com/when-rewilding-isnt-mad-guanacos-can-transform-the-espinal-of-chile-24248),
• equids (https://www.sciencephoto.com/media/1249170/view/two-male-indian-wild-asses and https://www.sandysharkey.com/blog/2018/10/21/saving-przewalskis-horse), and
• canids (https://www.agefotostock.com/age/en/details-photo/arctic-fox-alopex-lagopus-adult-female-summer-coat-playing-with-cub-nunavut-canada-july/FHR-10406-00211-849 and https://www.alamy.com/stock-photo-arctic-fox-alopex-lagopus-adult-female-summer-coat-standing-on-tundra-52965457.html).

Animals with caleonic colouration have, in a sense, ‘stretched’ the principle of countershading (https://en.wikipedia.org/wiki/Countershading) to breaking-point. This extension of pale pelage, upwards, has achieved whole-body conspicuousness instead of crypsis (https://en.wikipedia.org/wiki/Crypsis).

Caleonic colouration thus differs from pied colouration in how countershading has been modified. In the latter, pale ventral surfaces have been converted into parts of the large-scale, dark/pale contrast in the pigmentation of the pelage.

It is normal for the ventral parts of the body to be pale, as part of the inconspicuous pattern called cryptic colouration. However, the extension of this pale switches the effect to conspicuousness. This is because the pale, encroaching upwards towards the dorsal side, tends to catch the sunlight at

• all seasons, and
• most times of day.

In caleonic colouration, this ‘lateralisation’ of the pale parts of the pelage occurs variously on the

• cheeks,
• neck,
• shoulders,
• flanks, and/or
• hindquarters.

Caleonic colouration thus achieves whole-body conspicuousness.

In extreme cases, the 'lateralisation' of pale pelage reaches the dorsal surfaces of the rump, neck, and even back.

In this series of Posts, I hypothesise that a widespread species, the caribou (Rangifer tarandus, https://www.inaturalist.org/taxa/42199-Rangifer-tarandus) has

• certain subspecies with pied colouration (https://www.alamy.com/carribu-reindeer-rangifer-tarandus-caribou-image2513439.html?imageid=FAF39066-8FA2-4FFD-A129-1DC298D8D84F&p=17185&pn=12&searchId=1665bd569435214806da60bc576d4bf8&searchtype=0 and https://www.alamy.com/portrait-of-a-reindeer-walking-in-the-snow-alaska-usa-image364557768.html?imageid=DEE9244F-8F74-4688-BC41-A7D75C47B398&p=1311224&pn=1&searchId=8ebe633421a99c35728cae9a8c7eab8c&searchtype=0), and
• other subspecies with caleonic colouration.

This arguably makes R. tarandus the only species of mammal in which both patterns (pied and caleonic) occur, depending on the region/subspecies.

The only other species possessing caleonic colouration in only certain populations seems to be the domestic horse (Equus caballus). However, this may have arisen by hybridisation among several wild congeners, during the process of selective breeding (see https://www.inaturalist.org/posts/66546-a-hyperconspicuous-horse-hiding-in-plain-sight#).

The following show pied colouration in R. tarandus:

https://www.alamy.com/caribou-barren-ground-bull-autumn-denali-park-alaska-image219057000.html?imageid=CA708987-0B78-4D58-A301-7570ACD15211&p=365985&pn=3&searchId=8ef83f2f1de0431123cf39602bbd6277&searchtype=0 and https://www.alamy.com/stock-photo-caribou-rangifer-tarandus-bull-with-female-calf-on-migration-south-41499424.html?imageid=24FDBEC7-2515-4910-A5C6-57A0352BEF7C&p=54193&pn=3&searchId=8ef83f2f1de0431123cf39602bbd6277&searchtype=0 and https://www.alamy.com/stock-photo-caribou-rangifer-tarandus-bull-female-in-snow-on-migration-south-through-41499304.html?imageid=013FADBA-BE26-48E8-96CC-B83A1BEDFC74&p=54193&pn=4&searchId=53fe7e947085511a49c95d7dac229e54&searchtype=0 and https://www.alamy.com/caribou-barren-ground-bull-autumn-denali-park-alaska-image219057056.html?imageid=8BFBDFF1-5FE8-4504-BC54-0C1ED673978D&p=365985&pn=4&searchId=53fe7e947085511a49c95d7dac229e54&searchtype=0 and https://www.adfg.alaska.gov/static/home/library/pdfs/wildlife/caribou_trails/caribou_trails_2014.pdf.

The following seem to show caleonic colouration in R. tarandus:

https://www.mindenpictures.com/stock-photo-woodland-caribou-rangifer-tarandus-caribou-male-newfoundland-canada-naturephotography-image00596574.html and https://m.facebook.com/NewfoundlandLabradorTourism/photos/a.111088443781/111092763781/?type=3 and https://www.inaturalist.org/observations/9039103 and https://www.inaturalist.org/observations/91586926 and https://www.inaturalist.org/observations/38196369.

Pied colouration occurs in the autumn coat of most of the subspecies of R. tarandus, including R. t. groenlandicus and R. t. granti. The darkest features are arranged to provide dark/pale contrast, not crypsis or disruption of the outline of the animal.

These dark parts in this pied colouration are

• muzzle,
• forelegs,
• brisket/chest/lower shoulders, and
• lower flanks.

In the same figures, the palest features are

• nose (rhinarium),
• neck,
• beard/dewlap,
• tail, and
• the narrow area of pale on the rump.

In R. tarandus, caleonic colouration seems to occur in the subspecies found on two systems of islands, far apart geographically (https://en.wikipedia.org/wiki/Boreal_woodland_caribou#/media/File:Rangifer_tarandus_Map_NA.svg).

Firstly, on the island of Newfoundland (https://en.wikipedia.org/wiki/Newfoundland_(island)) occurs the subspecies R. t. terranovae (https://www.naturepl.com/stock-photo-woodland-caribou-nature-image00596572.html and https://www.shutterstock.com/de/image-photo/caribou-adult-female-rangifer-tarandus-avalon-1911012163).

Secondly (see part 2 of this series), on the Arctic islands of Canada occurs the subspecies R. t. pearyi (https://www.natureconservancy.ca/en/what-we-do/resource-centre/featured-species/mammals/peary-caribou.html and https://www.inaturalist.org/guide_taxa/1119027 and https://www.enr.gov.nt.ca/en/services/caribou-nwt/peary-caribou and https://www.alamy.com/stock-photo-bull-caribou-in-velvet-antlers-stands-in-the-colorful-autumn-tundra-75290355.html?imageid=89F3DD5C-43FB-48AC-B540-F92A8006B30D&p=228679&pn=25&searchId=9892403e79eeaa448727cb13a1e36c01&searchtype=0).

I agree with Valerius Geist that R. tarandus on the island of Newfoundland is quite distinct from other forms of ‘woodland caribou’, and that taxonomy took a wrong turn when R. t. terranovae was lumped with R. t. caribou.

The ‘woodland caribou’, in the western part of the boreal zone of North America, is unusually dark for the species (e.g. according to Valerius Geist). By contrast, R. t. terranovae is unusually pale - particularly considering that it lives at a far lower latitude than R. t. pearyi.

The following further illustrate R. t. terranovae: https://www.wildandexposed.com/journal/2018/10/17/marks-adventures-in-newfoundland
and https://mobile.twitter.com/NFLDdesigns/status/1339255210749865986/photo/1 and http://www.nlnature.com/Newfoundland-Canada-Nature/1520.aspx and http://birdingnewfoundland.blogspot.com/2009/12/southern-avalon-peninsula-birds-and.html and second photo in https://www.cbc.ca/news/canada/newfoundland-labrador/reindeer-deer-lake-video-1.4918560.

In the pied pattern, the lower flanks, chest, lower shoulders, and brisket are the darkest parts of the figure. This differs from the caleonic pattern in R. t. terranovae, in which these are the palest parts of the figure. Furthermore, a distinctive feature of R. t. terranovae is an individually variable diagonal border between relative dark and relative pale on the haunch (https://www.natureinstock.com/search/preview/woodland-caribou-rangifer-tarandus-caribou-bull-in-town-newfoundland-canada/0_10121446.html and http://conservationbiologynewfoundland.blogspot.com/2012/04/newfoundlands-wilderness-reserves.html).

Detracting from the caleonic pattern is the fact that some individuals of R. t. terranovae have a faint version of the flank-banding seen in other subspecies of R. tarandus.

However, this may perhaps be owing to some degree of anthropogenic mixing with another subspecies.

I hypothesise that the original appearance of the Newfoundland form was truly caleonic, to the exclusion of the flank-banding.

Wilkerson (2010, https://www.mun.ca/biology/scarr/Wilkerson%202010,%20excerpt.pdf) states that the domestic reindeer (R. t. tarandus) was introduced to the island of Newfoundland early in the twentieth century, and that there was indeed contact between this subspecies (which possesses flank-banding) and the indigenous populations, viz R. t. terranovae. I infer the possibility of hybridisation.

It is hard to see how the two patterns of colouration in R. tarandus, namely pied and caleonic, can be represented as extremes on a continuum. Instead, what seems to have happened is that, in an ancestral form,

• the whitish at the belly has spread so far up that it has replaced the entire flank-band complex of the pied pattern, while at the same time
• the legs have gone from basically dark to basically pale, and
• the neck has acquired a darkish dorsal (nuchal) zone.

It is this combination of pale flanks, pale legs, and a dark nuchal area that makes R. t. terranovae a candidate for caleonic colouration. The combination of extension of pale pelage on to flank and haunch, plus an overall pallour, set this subspecies apart from all other subspecies, besides R. t. pearyi.

However, the following three aspects detract from any simple characterisation of the colouration in R. t. terranovae as caleonic:

• some individuals do retain a faint version of the flank-banding typical of most subspecies of R. tarandus;
• the pale of the neck tends to be disjunct from the pale of the shoulders and flanks, separated by a more-or-less vertical tract of pale greyish-fawn pelage; and
• the pale of the lower haunch tends to give way, ventrally, to a darker tone on the upper hindleg, in some individuals.

What this means is that the pattern in R. t. terranovae - at least in its remaining, hypothetically hybridised form - is not categorically different from that in other subspecies.
   
In R. tarandus, any description of adaptive colouration must account for the seasonal cycle, in which the pelage moults. The moulting leads from this appearance (https://www.inaturalist.org/observations/137363786) to this (https://www.inaturalist.org/observations/73867876).

In winter, the pigmentation fades, owing to wear and weathering. The pelage then falls out in early summer, and re-grows from the darkly-pigmented skin. The colouration initially looks fairly uniform, before the full length of the hairs is achieved and the pigmentation differentiates.

Thus, the fully-differentiated colouration tends to be expressed in autumn, corresponding to the rutting season.

The following two male individuals, probably in September, show some features linking the ‘typical’ pattern above to the pattern typical of other subspecies of R. tarandus: https://www.alamy.com/stock-photo-two-caribou-rangifer-tarandus-forage-during-autumn-in-gros-morne-national-43870484.html. These include the relative darkness of the legs and the faint banding on the flank. These detractions notwithstanding, the colouration remains different from those of any other forms of ‘woodland caribou’ at the same season.

I take the following to be an adolescent male individual in autumn: http://www.hww.ca/en/wildlife/mammals/caribou.html. The side of the body shows a faint version of the banding seen in most subspecies of R. tarandus.

WINTER:
January: https://www.inaturalist.org/observations/85723506
March: https://www.inaturalist.org/observations/97411894
April: https://www.inaturalist.org/observations/43119657 and https://www.inaturalist.org/observations/73404224

SPRING:
Early May: https://www.inaturalist.org/observations/4829891 and https://www.inaturalist.org/observations/12796122 and https://www.inaturalist.org/observations/127414114 and https://www.inaturalist.org/observations/147676351
Late May: https://www.inaturalist.org/observations/1535227 and https://www.inaturalist.org/observations/120545240
Early June: https://www.inaturalist.org/observations/59202799 and https://www.inaturalist.org/observations/127339998 and https://www.inaturalist.org/observations/120600128 and https://www.inaturalist.org/observations/30141537 and https://www.inaturalist.org/observations/26703702 and https://www.inaturalist.org/observations/136693452

SUMMER:
Mid-June: https://www.inaturalist.org/observations/72014363 and https://www.inaturalist.org/observations/135072586
Late June: https://www.inaturalist.org/observations/87895806 and https://www.inaturalist.org/observations/69215843
Early July: https://www.inaturalist.org/observations/28248163 and https://www.inaturalist.org/observations/17045792 
Mid-July: https://www.inaturalist.org/observations/14903618 and https://www.inaturalist.org/observations/128037050 and https://www.inaturalist.org/observations/130969801 and https://www.inaturalist.org/observations/2870518 and https://www.inaturalist.org/observations/146311257 and https://www.inaturalist.org/observations/28836472
Late July: https://www.inaturalist.org/observations/8063943
Early August: https://www.inaturalist.org/observations/91586926 and https://www.inaturalist.org/observations/73867876
Mid-August: https://www.inaturalist.org/observations/38196369 and https://www.inaturalist.org/observations/38195578
Late August: https://www.inaturalist.org/observations/33067919

AUTUMN:
Early September: https://www.inaturalist.org/observations/134114763 and https://www.inaturalist.org/observations/65690022
Mid-September: https://www.inaturalist.org/observations/65690664 and https://www.inaturalist.org/observations/75381992 and https://www.inaturalist.org/observations/99568979

The following (https://www.inaturalist.org/observations/28836472) seems to conform to caleonic colouration. However, once one understands the cycle of moult and regrowth of the pelage, it is obvious that the effect results mainly from the fact that the faded coat of the previous year has fallen out from the dorsal part of the figure first (as seen more clearly in https://www.inaturalist.org/observations/126510207).

to be continued in https://www.inaturalist.org/journal/milewski/67534-caleonic-colouration-in-the-caribou-part-2-rangifer-tarandus-pearyi-in-context#...

https://www.inaturalist.org/observations/38739255

https://www.inaturalist.org/observations/38702920

https://www.inaturalist.org/observations/66795801

https://www.inaturalist.org/observations/45111163

https://escholarship.org/content/qt7xs9j2zs/qt7xs9j2zs.pdf

https://www.inaturalist.org/observations/15178773

https://www.inaturalist.org/observations/18851909

https://www.inaturalist.org/observations/36832325

https://www.inaturalist.org/observations/47665170

https://www.inaturalist.org/observations/39408908

https://www.inaturalist.org/observations/61325069

https://www.inaturalist.org/observations/69950772

https://www.inaturalist.org/observations/68894248

https://www.inaturalist.org/observations/68809821

https://www.inaturalist.org/observations/67319990

https://www.inaturalist.org/observations/107312217

https://www.inaturalist.org/observations/106889318

https://www.inaturalist.org/observations/121600503

https://www.inaturalist.org/observations/121887136

https://www.inaturalist.org/observations/117843874

https://www.inaturalist.org/observations/141842865

https://www.inaturalist.org/observations/141842864

https://www.inaturalist.org/observations/141842863

I have previously shown that what I call the western wildebeest

• should be distinguished from the blue wildebeest (Connochaetes taurinus taurinus), and
• already has a taxonomically legitimate name, viz. Connochaetes taurinus mattosi, which for mainly historical reasons has been overlooked.

Please see https://www.inaturalist.org/journal/milewski/67992-photo-pair-summarising-the-distinction-between-blue-wildebeest-and-western-wildebeest# and https://www.inaturalist.org/journal/milewski/54306-how-one-of-the-most-familiar-of-african-large-mammals-came-to-be-unrecognised#.

It is also known that the two South African spp. of wildebeests, namely C. taurinus and the black wildebeest (Connochates gnou), are capable of producing fertile hybrids (https://link.springer.com/article/10.1007/s10592-018-1071-x and https://link.springer.com/article/10.1007/s10344-011-0567-1).

The context in which hybridisation has been discovered is the holding of both spp. together, under artificial conditions, on game ranches. However, the natural distributions did overlap even under natural conditions, in and near what is now Free State province of South Africa, at the time of European arrival (https://www.researchgate.net/figure/Overlap-of-species-distribution-of-blue-and-black-wildebeest-Map-was-redrawn-from-IUCN_fig2_265128890 and https://www.researchgate.net/figure/Historical-distribution-ranges-of-blue-light-blue-and-black-wildebeest-greyThe_fig3_322539832).

In this Post, I hypothesise that the natural origin of C. t. mattosi has been by virtue of past (Pleistocene) hybridisation between C. taurinus and C. gnou.

I base this on the observation that most of the ways in which C. t. mattosi differs from C. t. taurinus are in line with the nature of C. gnou.

These are as follows.

Brindling is well-developed in C. t. taurinus (http://www.safari-club.co.uk/photo-galleries/kenya-familiarisation-trip-2010/dsc_0062/), minimally developed in C. gnou, and intermediate in C. t. mattosi.

The mane is lax in C. t. taurinus, vs stiff in C. gnou. In C. t. mattosi, the mane is stiff, making this the only form in the taurinus-complex that possesses a stiff mane.

The beard is lax and short in C. t. taurinus (https://www.gettyimages.com.au/detail/photo/close-up-of-blue-wildebeest-greater-kruger-national-royalty-free-image/81897934?phrase=blue%20wildebeest%20south%20africa&adppopup=true), vs stiff and relatively long in C. gnou (https://www.istockphoto.com/photo/black-wildebeest-gm887262280-246261777?phrase=wildebeest%20fur%20close%20up). In C. t. mattosi, the beard is far better-developed than in C. t. taurinus, and is particularly stiff (https://www.gettyimages.com.au/detail/photo/stripe-gnu-in-the-etosha-national-park-in-namibia-royalty-free-image/1297158239?phrase=blue%20wildebeest%20south%20africa&adppopup=true).

Sheen on the rump is noticeable in C. t. taurinus (albeit not as well-developed as in mearnsi or albojubatus), vs absent in C. gnou. In C. t. mattosi, sheen on the rump is hardly noticeable.

Paleness (depigmentation) on the proximal part of the mane (i.e. at the base of the mane) is well-developed in C. gnou, vs poorly-developed in C. t. taurinus. In C. t. mattosi, this feature seems to have an intermediate occurrence.

Infants tend to be paler (apart from the feet) in C. t. taurinus (https://www.gettyimages.com.au/detail/photo/side-view-of-buffalo-grazing-on-field-bushbuckridge-royalty-free-image/1347710373?phrase=blue%20wildebeest%20south%20africa&adppopup=true and https://www.alamy.com/stock-photo-baby-blue-wildebeest-connochaetes-taurinus-with-mother-21318646.html?imageid=B9A71E5D-12A8-4A62-992C-AAE26EA50A2A&p=50730&pn=1&searchId=4093b7b4151429bdd59b4c6faf51ee5c&searchtype=0 and https://www.reddit.com/r/wildlifephotography/comments/107gvvp/blue_wildebeest_with_suckling_calf_black_rhino/ and https://www.flickr.com/photos/antonettevdb/24818598341) than in C. gnou (https://www.youtube.com/watch?v=YQ-JJyS9HSo and https://www.irishnews.com/magazine/daily/2019/08/27/news/rare-baby-wildebeest-born-at-newquay-zoo-1696156/ and https://www.zooborns.com/zooborns/2009/07/newquay-welcomes-a-leggy-wildbeest.html).

In C. t. mattosi, the tone of infants seems intermediate (https://cloudfront.safaribookings.com/library/zambia/xxl/Liuwa_Plain_National_Park_041.jpg).

I can think of two alternative hypotheses for the origin of C. t. mattosi.

Firstly, C. t. mattosi may be a paedomorphic version of C. t. taurinus (https://www.inaturalist.org/journal/milewski/54384-the-western-wildebeest-as-an-example-of-paedomorphic-evolution#). This hypothesis is not mutually exclusive with the approach taken in the present Post.

Secondly, C. t. mattosi may have arisen by natural selection, free of any hybridisation, under ecological conditions different from those to which C. t. taurinus is adapted. South of the Kunene and Okavango Rivers, this seems to make sense, because C. t. mattosi is associated with Kalahari sand and semi-arid climates. However, it is undermined by the fact that C. t. mattosi extends deep into mesic south-central Africa, in the countries of Angola and Zambia.

Here I test the idea that, whereas bovids tend to have bold whole-body markings, cervids tend to have conspicuous erectile tails.

I have searched for the three most boldly-marked species/subspecies of first bovids, then cervids. Then I have searched for the three species/subspecies, in each family, that have tails that tend to be displayed in conspicuous form when raised.

The three most boldly-marked bovids, at a whole-body scale:

https://www.krugerpark.co.za/africa_bontebok.html

https://www.mindenpictures.com/stock-photo-sable-antelope-hippotragus-niger-in-hwange-national-park-zimbabwe-naturephotography-image90037248.html

"How rapidly do kangaroos breed" is an important question, if we are to understand the nature of Australia.

This continent has lacked any terrestrial wild predator larger than a dog (https://en.wikipedia.org/wiki/Dingo), for thousands of years.

Carnivores depend on the productivity of their prey. Why did no carnivore specialise on kangaroos, in the way the cheetah (Acinonyx jubatus, https://www.inaturalist.org/taxa/41955-Acinonyx-jubatus) specialises on gazelles in Africa and Asia?

The answer lies partly in an understanding of the reproductive system of kangaroos.

It is true that an individual female can have three dependent offspring, in various stages of growth at the same moment. However, this is evidence of slow, rather than fast, growth. An overlap of this kind may amount to an insurance policy, allowing at least one offspring to survive under conditions that are generally unfavourable for growth.

How to compare reproductive rates?

Kangaroos can be compared with the most similar herbivores under similar climates on other continents. Body size is a prime factor. Reproduction is likely to be more rapid in small-bodied than in large-bodied species, and the corollary is that lifespan is shorter in the former than in the latter.

Adult female kangaroos have body masses of 25-35 kg. Terrestrial herbivores of similar body size, under similar climates, consist of various ruminants and one large rodent. Since males of kangaroos eventually grow to double the body mass of females in middle age (older than 10 years), other herbivores of up to 65 kg can also be considered comparable.

Reproductive rates can be measured as either the rate of increase of populations free of predators, or the rate at which an established population can be culled with no risk of extermination. The prime factors are how fast females grow to breeding age, and how many offspring they pack into a single litter. Periods need to be calculated from conception , not from birth, because pregnancy in kangaroos is extremely brief.

Populations of the red kangaroo (Macropus rufus, https://www.inaturalist.org/taxa/42885-Macropus-rufus and https://australian.museum/learn/animals/mammals/red-kangaroo/) can be culled at an average of up to 15% per year. An annual offtake rate of 3 out of every 20 in the population seems to be sustainable in the long term.

The corresponding figures for grey kangaroos (Macropus fuliginosus https://www.inaturalist.org/taxa/42881-Macropus-fuliginosus and Macropus giganteus https://www.inaturalist.org/taxa/42888-Macropus-giganteus) and the common wallaroo (Macropus robustus, https://www.inaturalist.org/taxa/42872-Macropus-robustus) seem to be 13% and 11%, respectively. These rates of reproduction are modest for mammals of this body size.

No match for ruminants:

Kangaroos are often compared to ruminants. However, the many spp. of ruminants maintain rapid reproduction and growth over a wide range of body sizes.

Species the size of kangaroos generally outbreed kangaroos two-to-one. Only the largest-bodied of ruminants, or those living on mountaintops or in the Arctic, reproduce as slowly as kangaroos do.

The domestic sheep (Ovis aries, https://www.inaturalist.org/taxa/121578-Ovis-aries) is very variable, according to breed and plane of nutrition. Sustainable offtake in sheep varies from 15% in the case of the merino breed that coexists with kangaroos, to 300% in the case of the most prolific breeds on other continents.

In the domestic sheep, growth from conception to sexual maturity takes at least 5 months, and usually 15 months, which is less than in the red kangaroo. Females of the domestic sheep - despite being the more massive - have a shorter lifespan than that of females of kangaroos.

Wool breeds allocate protein to fleece instead of milk, and females wean only one offspring each per year. Nevertheless, the merino breed (https://en.wikipedia.org/wiki/Merino) in Australia outbreeds the red kangaroo, relative to body mass.

Average reproductive rates are similar, but females of the ungulate (65 kg) grow to double the body mass of females of kangaroos. Therefore, even wool-producing breeds of sheep can produce food for meat-eaters more rapidly than can any of the four spp. of kangaroos mentioned so far.

Please note the following:

• The D'man breed of sheep, of the northern edge of the Sahara, is extremely prolific (https://www.researchgate.net/figure/Reproductive-performance-of-Moroccan-sheep-breeds-1_tbl1_233381669). This is because it bears 2-5 offspring per litter, bears two litters per year, and produces no wool.
• The small-tailed han breed of China (similar in body size to merino, https://en.wikipedia.org/wiki/Small-tail_Han) can bear up to 7 newborns in one litter.
• The finnsheep breed (https://en.wikipedia.org/wiki/Finnsheep) of northern Europe can grow to 50 kg by 7 months, and can give birth by one year, after its own birth. This means that finnsheep grow from a single cell to 50kg in one year, whereas the red kangaroo takes three-fold longer to reach this mass.

Kangaroos are renowned for nurturing up to 3 offspring simultaneously. However, the finnsheep does the same for up to 10 offspring (5 in the womb, 5 at the udder).

Thus, the most prolific breeds of sheep reproduce up to 20-fold more rapidly than kangaroos do, if comparisons are made on a long-term basis with correction for differences in body mass of females.

The domestic goat (Capra hircus, https://www.inaturalist.org/taxa/123070-Capra-hircus) reproduces several-fold more rapidly than the coexisting common wallaroo does. The feral goat normally bears twins in semi-arid Australia, whereas the common wallaroo has the slowest reproduction among kangaroos.

Relatives of camels breed as slowly as kangaroos:

The largest ruminants, giraffes (Giraffa spp., https://www.inaturalist.org/taxa/42157-Giraffa-camelopardalis), sustain an average offtake of 15% of the population per year, on cattle ranches in Africa where they are conserved and culled. Like kangaroos, giraffes bear a single newborn. This means that kangaroos breed no more rapidly than a ruminant 25-fold their body mass (females: 30 kg vs 800 kg; males: 65 kg vs 1300 kg).

Other large ruminants err to one side or the other of the value of 15%. For example, Bison bison (https://www.inaturalist.org/taxa/42408-Bison-bison) has one offspring per birth, breeds in only 2 of every 3 years, and sustains an average offtake of 10%. The moose (Alces alces, https://www.inaturalist.org/taxa/522193-Alces-alces) often has twins, and sustains an average offtake of more than 20%.

Camels, unlike giraffes, deer, and other true ruminants, reproduce relatively slowly. The vicugna (Vicugna vicugna, https://www.inaturalist.org/taxa/42236-Vicugna-vicugna) is the smallest living member of the camel family, and approaches kangaroos in body mass as well as its diet of grass. Whereas the domestic sheep is pregnant for 5 months, the like-size vicugna is pregnant for 11 months. The sustainable rate of offtake for the vicugna is likely to be less than 10% per year.

The vicugna is restricted to the Andes in South America, at double the altitude of the Australian Alps (https://en.wikipedia.org/wiki/Australian_Alps). It is less prolific than kangaroos, probably because its food supply is always limited.

Prolific ruminants of arid climates:

The springbok (Antidorcas marsupialis, https://www.inaturalist.org/taxa/42283-Antidorcas-marsupialis) is a ruminant of similar body mass and diet to the red kangaroo, restricted to dry climates in southern Africa, and likewise culled on sheep pastures.

Please note the following:

• Although the springbok bears only 1 offspring per birth, this species of gazelle can sustain double the offtake rate of the kangaroo (about 30% per year).
• Females of the springbok usually mature sexually in less than half the time taken by females of the red kangaroo: 13 months vs 31 months after conception.
• Under good conditions, females of the springbok can reach sexual maturity less than 1 year after being conceived, a record unmatched by any kangaroo.
• The springbok can bear 2 offspring per year after good rainfall, because it is pregnant for less than 6 months, and the mother is prepared to mate shortly after giving birth.
• The only gazelle commonly bearing twins is a relatively small-bodied species, Gazella subgutturosa (https://www.inaturalist.org/taxa/568727-Gazella-subgutturosa), which is able to survive the cold winters of Asia.

It is true that the red kangaroo can reproduce rapidly after rainfall, with each mother having up to 3 offspring in various stages of development: 1 freely suckling, 1 attached inside the pouch, and 1 in the womb. However, this does not compensate for the relatively slow growth of these offspring. Even in the best of seasons, the red kangaroo mother can wean no more than 3 offspring in 2 consecutive years, compared to 4 in the case of the springbok.

Herbivores in the extremely arid Sahara Desert are also prolific. Two spp. of gazelles just manage to survive in the central Sahara, after decades of persecution with sophisticated firearms and all-terrain vehicles.

Gazella leptoceros (https://en.wikipedia.org/wiki/Rhim_gazelle), similar in body mass to kangaroos, has always been restricted to sandy desert in North Africa. Remnant populations of the cheetah have been recorded in the Sahara (https://en.wikipedia.org/wiki/Northeast_African_cheetah#:~:text=Once%20existing%20in%20Egypt%2C%20the,blindfolded%2C%20and%20kept%20on%20leashes.).

The cheetah is likely to be particularly dependent on the fecundity of its prey in desert, where all herbivores are naturally rare. This felid, twice as massive as the extinct thylacine (Thylacinus cynocephalus, https://en.wikipedia.org/wiki/Thylacine) of Australia, may have survived to this day in a habitat more barren than the heart of Australia.

Giant rodents and grazing apes:

The capybara (Hydrochoerus hydrochaeris, https://www.inaturalist.org/taxa/74442-Hydrochoerus-hydrochaeris) is the largest rodent on Earth, similar in body mass to kangaroos. It is culled on cattle ranches in South America, with a sustainable offtake of about 35%. The capybara reproduces far more rapidly than kangaroos do, because it bears 1-8 (usually 2-6) offspring per litter.

The gelada (Theropithecus gelada, https://www.inaturalist.org/taxa/43530-Theropithecus-gelada) of the Ethiopian Highlands is the only living primate that relies on green grass for food, and it inhabits treeless grassland. This is the most strictly herbivorous and terrestrial of all monkeys, and is therefore comparable with kangaroos.

Primates in general breed slowly, and the gelada is no exception. It reproduces below par for kangaroos, because females take twice as long to reach sexual maturity, and give birth only once every two years. Like the vicugna, the gelada is adapted to altitudes beyond the ranges of most comparable ruminants.

Why are kangaroos not prolific?

Marsupials should be capable of rapid reproduction, because the small size of newborns potentially allows many offspring to be packed into each litter. For example, 56 maggot-size individuals emerged from one birth of the North American opossum (Didelphis virginiana, https://www.inaturalist.org/taxa/42652-Didelphis-virginiana).

This marsupial has a short natural lifespan for a mammal of its size (3 kg), dying of old age at 3 years even if it has survived predators and winter snow. Females wean 9-12 offspring in the second hear of life, and largely rely in this one summer's maternal effort for the propagation of the species. Therefore, the consistent restriction of kangaroos to 1 offspring per birth, and their extended lifespan (more than 20 years), are not merely the result of their genetic heritage as marsupials.

Drought does not seem to be the limiting factor. The original numbers of kangaroos were modest, even after a series of years with more than average rainfall.

The tropical north of Australia has copious rainfall and perennial rivers. However:

• Far from supporting large populations of kangaroos, its savannas are remarkably empty of indigenous herbivorous mammals.
• Red and grey kangaroos are scarce/marginal.
• The only species of kangaroos present are wallaroos (https://en.wikipedia.org/wiki/Wallaroo), which are either restricted to scattered boulder outcrops, or are small for kangaroos (antilopine wallaroo, Macropus antilopinus, https://www.inaturalist.org/taxa/42880-Macropus-antilopinus, less than 20 kg).

In southern Australia, the red kangaroo of the semi-arid interior outreproduces the grey kangaroos of coastal woodlands, if long-term averages are compared. However, the fecundity of the red kangaroo remains modest, even where bore water is provided.

Originally, kangaroos were remarkably scarce in the treeless mitchell grassland (dominated by Astrebla, a grass restricted to Australia) that covers an area the size of Britain (450000 square kilometres) in the northern half of Australia (https://en.wikipedia.org/wiki/Mitchell_Grass_Downs and https://www.agric.wa.gov.au/rangelands/mitchell-grass-alluvial-plain-pastures-pilbara-western-australia#:~:text=Curly%20Mitchell%20grass%20(Astrebla%20lappacea,)%20of%2010%20to%2025%25.).

Mitchell grassland is as extensive as the Highveld (https://en.wikipedia.org/wiki/Highveld) in South Africa, and is the closest equivalent in Australia to prairies and steppes on other continents. However, it lacked the expected herds of herbivores.

Research by Alan Newsome (https://www.eoas.info/biogs/P005736b.htm), together with traditional aboriginal knowledge, revealed that kangaroos were not migratory, and failed to graze mitchell grassland even after rain, before the advent of domestic livestock.

Kangaroos are adapted to nutrient-poverty:

Australia is the nutrient-poorest continent. Virtually all of its soils are poor in macronutrients (e.g. phosphorus) or micronutrients (e.g. cobalt), or unbalanced in their combinations of nutrients.

On other continents, herbivores with slow reproduction are marginalised to the most challenging environments. In Australia, they are prevalent in the form of kangaroos, owing to continent-wide nutrient-poverty, compounded by intense wildfire.

Nutrient-poverty, and a lack of succulent plants other than halophytes, explains why kangaroos have a limited ability to exploit arid environments.

All spp. of kangaroos are absent from parts of the Simpson and Great Sandy Deserts (maps by Graeme Caughley, https://www.goodreads.com/book/show/6819726-kangaroos and https://en.wikipedia.org/wiki/Graeme_Caughley and https://www.science.org.au/fellowship/fellows/biographical-memoirs/graeme-james-caughley-1937-1994). Even the spp. of the arid interior need drinking water or shade.

The red kangaroo depends mainly on alluvial woodlands, which retain a few pools evening droughts. The common wallaroo depends on boulder outcrops far from water, which provide shade even at noon.

By contrast, the springbok is extremely adapted for drought, and resides year-round in the Namib Desert, which is more arid (average less than 100 mm of rainfall per year) than the Simpson and Great Sandy Deserts. Although unlikely to reproduce in severe drought, adults of the springbok need neither drinking water nor shade to survive until the next episode of rain. The springbok seldom drinks even when water is available, although it visits pans (salinas, https://journals.co.za/doi/abs/10.10520/AJA03794369_3442) to eat succulent plants and to eat nutrient-rich earth.

Adaptation to nutrient-poverty would explain the paradox of limited reproductive rates of kangaroos, and their absence from large areas. Even the clay soils of mitchell grassland seem too poor to support the succulent plants capable of sustaining herbivores in drought. Unlike ruminants, kangaroos are unknown to eat earth as a nutritional supplement. The possible reason for this that such supplements are unavailable on the ancient, deeply weathered continent of Australia.

Whatever the reason, the contrast between semi-arid Australia and southern Africa was extreme. Although red kangaroo and springbok both prefer palatable, small grasses such as Enneapogon (a genus indigenous to both continents, https://www.inaturalist.org/observations?place_id=any&taxon_id=72116&view=species), the springbok was prolific to a degree unmatched in Australia.

Colonists of South Africa observed irruptions of tens of millions of individuals walking shoulder-to-shoulder, sweeping sheep before them, and denuding the vegetation over areas exceeding 100 km by 10 km.

Sceptical readers will realise that I have understated these 'springbok treks', on referring to eye-witness accounts (see J D Skinner and G N Louw, 1996, Transvaal Museum Monographs, no. 10, https://www.biblio.com/book/springbok-antidorcas-marsupialis-zimmermann-1780-transvaal/d/938429382). The springbok multiplied its populations rapidly, despite being prey to at least 4 spp. of carnivores larger than the dingo, and many other predators.

Irruptions were repeatedly recorded, most recently in 1896 (https://www.tandfonline.com/doi/abs/10.1080/00359199309520276?journalCode=ttrs20 and https://www.jstor.org/stable/29734323).

@tonyrebelo @jeremygilmore @matthewinabinett @paradoxornithidae @davidbygott @dejong @jakob @jwidness @bobby23 @marcelo_aranda @maxallen @aguilita @tfrench @beartracker @douglasriverside @saber_animal @biohexx1 @chewitt1 @simontonge @adamwelz @jimsinclair @russellcumming

Acknowledgement: I thank Elizabeth A. Carrie-Wilson for help in brainstorming a formal term in improvement of 'hitting the funny bone'.

Everyone knows what it is to 'hit your funny bone'.

It seems natural and inevitable that one will bump the point of the elbow, which is the proximal end of the ulna (the larger of the two bones of the forearm).

However, bumping a nerve in the region of the elbow is puzzling from a viewpoint of biological form and function. Furthermore, it may possibly be an experience unique to humans (Homo).

This phenomenon deserves a proper term, because

• 'hitting your funny bone' is informal, inept, and ambiguous (https://www.google.com.au/search?q=Define+funnybone&sxsrf=ALiCzsZSkx0H0sCQS5ZNFChfAxTGMebX7A%3A1672361426883&source=hp&ei=0jWuY7TjM4Lv4-EPu-uV6AU&iflsig=AJiK0e8AAAAAY65D4ml5WDtqpz8_6XFf3d_Wku6OoACz&ved=0ahUKEwi0lKLmj6D8AhWC9zgGHbt1BV0Q4dUDCAo&uact=5&oq=Define+funnybone&gs_lcp=Cgdnd3Mtd2l6EAMyBwgAEIAEEA0yCAgAEB4QDxANMgoIABAFEB4QDxANMg8IABAIEB4QDxDxBBANEAoyCggAEAgQHhAPEA0yCggAEAgQHhAPEA0yBQgAEIYDMgUIABCGAzIFCAAQhgM6BwgjEOoCECc6BAgjECc6BQgAEJECOhQILhCABBCxAxCDARDHARDRAxDUAjoRCC4QgAQQsQMQgwEQxwEQ0QM6CwgAEIAEELEDEIMBOgQIABBDOgUIABCABDoKCAAQkQIQRhD5AToKCAAQgAQQRhD5AToMCAAQgAQQDRBGEPkBOggIABAWEB4QClDACVj_M2DRO2gBcAB4AIABzAKIAewdkgEIMC4zLjEwLjOYAQCgAQGwAQo&sclient=gws-wiz),
• the experience is painful, not enjoyable as implied by the adjective 'funny',
• the relevant impact is not on a bone, but on a major nerve (the ulnar nerve, https://en.wikipedia.org/wiki/Ulnar_nerve and https://www.physio-pedia.com/images/a/a7/Nerves_of_the_left_upper_extremity.gif), and
• the verb 'hit' implies active striking, whereas the accidental action involved is invariably passive.

'Hitting the funny bone' is relevant to natural history, because

• anatomy is generally subject to the biological principles of adaptation,
• Homo sapiens is by far the most dexterous species on Earth, mainly owing to the neurological sophistication of the forearm, and
• it is puzzling that the evolutionary process has allowed the persistence of a particular point of external vulnerability to a major nerve.

First, let us correct the lack of a technical term for the phenomenon. Then, let us begin to think about the adaptive significance of this ostensible flaw in the design of the human body.

The Latin term for elbow is 'cubitus'. The vulnerable section of the ulnar nerve runs through the cubital tunnel (https://en.wikipedia.org/wiki/Cubital_tunnel).

The cubital tunnel is located between two bony projections, one on a bone of the forearm and the other on a bone of the upper arm.

These are, respectively

• the olecranon (https://en.wikipedia.org/wiki/Olecranon and https://en.wikipedia.org/wiki/Olecranon and https://www.shutterstock.com/image-vector/medical-illustration-anatomy-elbow-joint-on-1989821303), which is part of the ulna and is the most prominent bone of the elbow, and
• the medial epicondyle of the humerus (https://en.wikipedia.org/wiki/Medial_epicondyle_of_the_humerus and https://en.wikipedia.org/wiki/File:Human_arm_bones_diagram.svg).

(The cubital tunnel is not to be confused with the cubital fossa (https://teachmeanatomy.info/upper-limb/areas/cubital-fossa/), which is the 'pit of the elbow'.)

The word 'bump' conveys the correct meaning in this context, but is unsatisfactory, partly because of ambiguity between the noun and the verb, and partly because 'bumpage' is a particularly awkward word.

Therefore, the term 'allision' (https://en.wiktionary.org/wiki/allision#:~:text=allision%20(plural%20allisions),dashing%20against%20or%20striking%20upon.), although seldom used, is preferable.

On this basis, I might suggest that we replace 'hitting the funny bone' with 'cubital neural allision'.

However, this remains somewhat awkward.

The ancient Greek word for elbow is 'olene' (from which 'ulnar' is derived in the first place). Hence a more satisfactory new term might be OLENEURAL ALLISION.

In explanation of the peculiar vulnerability of the ulnar nerve, at the cubital tunnel, to accidental and passive impact, I hypothesise as follows.

Firstly, there is a principle that any evolutionary modification has both advantages and disadvantages. Thus, an adaptative syndrome as valuable as human dexterity can be expected to have some sort of 'downside'.

Ideally, the evolutionary process ensures that the whole anatomical configuration minimises this downside. However, some 'cost', in the sense of persistent risk and inconvenience, may be unavoidable.

Secondly, human dexterity depends partly on the particularly wide axial rotation of the forearm. This range of movement (in the action used to wield a screwdriver) is - in conjunction with opposability of the thumb - exceptional even among anthropoid primates.

This may necessitate a certain minimum distance between the bones to which the muscles attach at the elbow, particularly the medial epicondyle of the humerus and the head of the radius (https://www.ortho.wustl.edu/content/Patient-Care/3151/Services/Shoulder-Elbow/Overview/Elbow-Arthroscopy-Information/The-Anatomy-of-the-Elbow.aspx#:~:text=The%20elbow%20is%20a%20hinged,that%20form%20the%20joint%20capsule.).

For illustrations of the greater width of the head of the humerus in Homo than in chimpanzees (Pan) and baboons (Papio), please see Figures 1-2 and 1-3 in https://musculoskeletalkey.com/phylogeny/.

Thirdly, the exceptional dexterity of the human forearm demands three nerves of exceptionally large diameter. The ulnar nerve is one of these.

Please see:
https://upload.wikimedia.org/wikipedia/commons/5/51/Front_of_Sensory_Homunculus.gif
https://en.wikipedia.org/wiki/Cortical_homunculus#/media/File:Side-black.gif
https://en.wikipedia.org/wiki/Cortical_homunculus#/media/File:Rear_of_Sensory_Homunculus.jpg
https://upload.wikimedia.org/wikipedia/commons/0/0a/Sensory_and_motor_homunculi.jpg
https://en.wikipedia.org/wiki/Cortical_homunculus#/media/File:Sensory_Homunculus-en.svghttps://en.wikipedia.org/wiki/Cortical_homunculus#/media/File:Motor_homunculus.svg
https://upload.wikimedia.org/wikipedia/commons/d/df/Sensory_Homunculus-en.svg
https://en.wikipedia.org/wiki/Cortical_homunculus.

Putting these three lines of thinking together:

We seem to have a combination of a wide setting of the bony processes of the elbow (which produces a gap wide enough to warrant the term 'cubital tunnel') and a large ulnar nerve.

This 'spread of the bones' plus 'enlargement of the nerve' may make it unavoidable that some risk persists of accidental impact, when the arm occasionally bumps - at a certain angle - against objects in the environment.

Can readers improve on either the term 'oleneural allision', or my explanatory hypothesis?

@tonyrebelo @jeremygilmore @ellurasanctuary @benjamin_walton @adriaan_grobler @wynand_uys @dnicolle @slowplants @alan_dandie @reiner @porcoespinho15 @iancastle @eremophila @hillsflora @terra_australis @arthur_chapman @guillaume_papuga

Western Australia is unique on Earth in having extensive woodlands under a semi-arid, temperate climate (https://www.youtube.com/watch?v=-racshg-_u8 and https://www.researchgate.net/publication/236335929_The_Extraordinary_Nature_of_the_Great_Western_Woodlands and https://cdn.wilderness.org.au/archive/files/the-great-western-woodlands-report.pdf).

Please see

• https://www.dpaw.wa.gov.au/management/off-reserve-conservation/the-great-western-woodlands/77-visiting-the-great-western-woodlands
• https://en.wikipedia.org/wiki/Great_Western_Woodlands
• https://www.dpaw.wa.gov.au/management/off-reserve-conservation/the-great-western-Woodlands
• https://gondwanalink.org/where-we-work/great-western-woodlands/
• https://www.wilderness.org.au/iconic-places/great-western-woodlands
• https://en.wikipedia.org/wiki/Western_Australian_mulga_shrublands#:~:text=The%20Western%20Australian%20Mulga%20shrublands,areas%20of%20grassland%20and%20scrub.

I have hypothesised (https://www.jstor.org/stable/2844553 and https://www.jstor.org/stable/2844553) that part of the reason for this anomaly is a climatic pattern in which tropical cyclones degenerate southeastwards from the northwestern coast of Australia, becoming rain-bearing systems that episodically drench otherwise arid southeastern parts of Western Australia (please see second map in http://www.bom.gov.au/cyclone/climatology/wa.shtml).

This explanation might have seemed far-fetched, because of the remoteness of the hypothesised 'cause'. The distance from Exmouth (https://en.wikipedia.org/wiki/Exmouth,_Western_Australia) to Kalgoorlie (https://en.wikipedia.org/wiki/Kalgoorlie) is about 1200 km (https://upload.wikimedia.org/wikipedia/commons/0/06/WAHighways.png).

What I learned from the case of tropical cyclone Glenda (http://www.bom.gov.au/cyclone/history/pdf/glenda.pdf and https://en.wikipedia.org/wiki/Cyclone_Glenda) is:
It is not just a degenerating cyclone, happening to travel southeastwards, that brings rain to the Great Western Woodlands and the mulga region of Western Australia.

Instead, this rain can set in before the cyclone enters the continent, and seems to take the pattern of extending southeastwards, regardless of the happenstance trajectory of the cyclone itself.

This directionality seems intrinsic/inherent in the pattern. The rain can occur additionally to the cyclone, as part of a far more broadly-defined, manifold weather-event - extending over a week and 1500 km - for which we have no name.

Thus, we had a situation, 29-30 March 2006, in which the Bureau of Meteorology (http://www.bom.gov.au/wa/) was still forecasting risk to towns in the Pilbara (https://en.wikipedia.org/wiki/Pilbara), from a cyclone that was well offshore in the ocean north of the Pilbara and of uncertain trajectory, while already reporting the start of rain over a wide area of the mulga region (https://en.wikipedia.org/wiki/Western_Australian_mulga_shrublands#/media/File:Ecoregion_AA1310.svg).

What was particularly interesting is that

• one day before this cyclone crossed the coast, there was already an arc-shaped cloud-band, extending southeastwards to the northeastern part of the Eastern Goldfields (https://en.wikipedia.org/wiki/Eastern_Goldfields), and
• rain was already expected, or had already fallen, in the Gascoyne (https://en.wikipedia.org/wiki/Gascoyne_bioregion) and Murchison (https://en.wikipedia.org/wiki/Murchison_bioregion) - and even in the Eastern Goldfields, up to 1500 km away.

The following is a more detailed account of the events.

Tropical cyclone Glenda affected Western Australia at the end of March 2006.

It intensified in the sea off the Pilbara (https://earth.esa.int/web/earth-watching/natural-disasters/cyclones/cyclone-events/-/asset_publisher/4Lfz/content/cyclone-glenda-australia-march-2006/ and http://www.bom.gov.au/cyclone/history/glenda.shtml), then turned south to approach the Pilbara coast, as expected.

Alarm was sounded for coastal towns (https://en.wikinews.org/wiki/Cyclone_Glenda_closes_in_on_Western_Australia) - which, as it transpired, were fortunately spared.

Tropical cyclone Glenda struck the coast fast and hard, but then collapsed.

The intense system degenerated immediately after crossing the coast, into a system of low atmospheric pressure that posed no further risk beyond flooding in e.g. the Gascoyne region (https://en.wikipedia.org/wiki/Gascoyne).

Despite the fact that the cyclone degenerated in the northern half of Western Australia - and indeed before it deeply penetrated even the Gascoyne region - it produced widespread, soaking rain over a wide area. This included the Eastern Goldfields, where the Great Western Woodlands occur.

These events show that the rain-effect may not be merely a consequence of the spent cyclone travelling southeastwards - which is a typical pattern (http://www.bom.gov.au/cyclone/history/steve.shtml and https://upload.wikimedia.org/wikipedia/commons/a/a2/Steve_2000_track.png and https://en.wikipedia.org/wiki/Cyclone_Steve and https://www.ausstormscience.com/tropical-cyclones/historic-tropical-cyclones/ and http://www.bom.gov.au/cyclone/climatology/wa.shtml).

Instead, the rain may be an inherent part of a larger-scale system. This is perhaps partly owing to the effect of the Coriolis force (https://en.wikipedia.org/wiki/Coriolis_force) on the tropical warm, moist air, and the interaction of the latitudinal and longitudinal 'heat toughs' in northern and western Australia.

Rain already reached the Eastern Goldfields well before the cyclone actually entered the Australian landmass, and continued well after the cyclone degenerated not far from where it entered. For this reason, the normally semiarid Eastern Goldfields were bestowed with several days of rain.

This shows that copious rain in the Eastern Goldfields can occur despite the remoteness and short duration of a tropical cyclone itself.

The extension of the rain to the Eastern Goldfields in this way was despite the steep gradient up which the system had to climb, as it were, from minimal atmospheric pressure off the Pilbara to maximal atmospheric pressure in the Great Australian Bight (https://en.wikipedia.org/wiki/Great_Australian_Bight).

Because of the northwest-southeast band of cloud resulting from the Coriolis force, rain affected the eastern part of southern Western Australia two days before it affected Perth (https://en.wikipedia.org/wiki/Perth). It affected the northern Eastern Goldfields one day before it crossed the coast, but affected Perth one day after it crossed the coast.

This was the third such episode of rain to occur in southern Western Australia during the warm season of 2005-2006. The first one thoroughly wetted the Lake Grace area (https://en.wikipedia.org/wiki/Lake_Grace,_Western_Australia).

This was then added to by the next two systems. Thus, trees would have had an excellent opportunity to regenerate germinatively, just north of Fitzgerald River National Park (https://en.wikipedia.org/wiki/Fitzgerald_River_National_Park).

Much of the water sustaining the Great Western Woodlands may originate in the Timor Sea (https://en.wikipedia.org/wiki/Timor_Sea), more than 2000 km to the north - if not in the Pacific Ocean (https://en.wikipedia.org/wiki/Pacific_Ocean), even farther away.

This evaporative origin follows the 'heat trough' of the northern Australian coast, westwards, recharges itself over the sea off the Pilbara, and turns southwards and then southeastwards, because of a 'corner' that it meets.

This corner lies at the intersection of the band of low atmospheric pressure, running west-east in tropical northern Australia, with the coastal heat trough of the west coast of Australia (http://www.bom.gov.au/climate/about/?bookmark=westtrough).

The season of tropical cyclones in Western Australia is in autumn, rather than summer. This is exemplified by the fact that, in 2006, the last major cyclone threatening Darwin occurred at the same time as the first proper cold-frontal winter rain in Perth, i.e. on Anzac Day (https://en.wikipedia.org/wiki/Anzac_Day) in late April 2006.

My overall comment:
I stand by the basic conceptual framework of my climatic hypothesis, more than 40 years after publishing my explanation for the growth of trees in semi-arid Western Australia.

However, since than I have been able to track several tropical cyclones over the decades. Of these cyclonic events, Glenda was perhaps the most chronologically revealing, in showing the larger nature of the meteorological systems involved.

At first glance, there seems to be nothing special about the location of the Great Western Woodlands, which merely occupy part of the vast low-lying plain of Australia (https://upload.wikimedia.org/wikipedia/commons/5/52/Great_Western_Woodlands_location_within_Australia.jpg).

However, this location happens to be at the receiving end of climatic systems determined by the coastal configuration of the whole western section of the continent, from the tropics to the temperate zone.

The climatic factors promoting trees cannot be understood in a regional context alone. They must be framed in systems that are unusually extensive, owing to the lack of topographical barriers in Western Australia.

Various animals, when attacked by a predator, scream.

I have surmised that a main adaptive function of this behaviour is to attract other predators, boosting the chance that the victim may be able to escape in the ensuing confusion and conflict between the original attacker and a subsequent, more powerful one.

Interspecies antagonism and competition, and freeloading, seems common among predators. Thus, I interpret distress-calling not according to the intuitive human explanation, viz. calling conspecifics to defence, but in an alternative light, i.e. directed not at conspecifics but at even more formidable predators.

With this idea in mind, I note the following comments on Oryctolagus cuniculus (https://www.inaturalist.org/taxa/43151-Oryctolagus-cuniculus) and Lepus europaeus (https://www.inaturalist.org/taxa/43128-Lepus-europaeus), made by Brian Vesey-Fitzgerald (https://en.wikipedia.org/wiki/Brian_Vesey-Fitzgerald) in his book 'British Game' (https://www.harringtonbooks.co.uk/pages/books/58907/brian-vesey-fitzgerald/british-game-the-new-naturalist-library-2).

This provides food for deep thought on the adaptive value of distress-calling in the wild rabbit.

"Only the doe tends the young, and timid as she is, she will fight fiercely - even attacking a stoat in defence of the litter...Normally excessively timid, there have been records of bucks (males of the rabbit) turning upon and routing stoats. The kick of a big buck rabbit is powerful, and I have known a large tom-cat put to flight by a doe in defence of her young. But this sort of thing is exceptional. It is much more usual for a rabbit hunted byna stoat to run only a few yards and then sit down and wait for death, screaming piteously the while. A 'stoated' rabbit, indeed, seems to be hypnotised, and it is noteworthy that other rabbits seem to know that they are not in danger. I well remember hearing a rabbit scream very close to me in Dorset once, and on looking over a bank I saw plenty of rabbits playing about unconcernedly and one moving very slowly screaming loudly. In due course a stoat appeared, the playful rabbits made way (giving it a wide berth and showing no sign of fear), and killed the screaming rabbit. It seemed quite evident that those rabbits knew they were in no danger...hares if not killed outright scream in an almost human manner, like small children that have been hurt. And so I hate shooting hares".

(writing in progress)

"We see the world not as it is, but as we are"

Many mammals have colouration adapted for inconspicuousness.

Seasonal changes in the colouration of the pelage are common in mammals. Why not green in the green season?

Furthermore, certain species of mammals are colour-polymorphic. A green morph would seem to be a particularly good idea for many mammals.

Mammals in which it is particularly puzzling that the pelage is never green, at least in part, include

• Giraffa,
• spotted forms and growth-stages of deer (particularly infants, and small species in summer pelage, e.g. Capreolus capreolus),
• striped and spotted bovids such as Tragelaphus, and
• diurnal rodents in marshes, e.g. Otomys,
• voles (e.g. Microtus) in summer pelage, and
• sloths.

Equids are generally surrounded by green, and have eyesnthat are prooirtiinately the largest of any land animal. Yet they are unable to see green and are not coloured to blend into green backgrounds. Instead, equids see the wavelengths known to humans as blue and yellow.

The answer is partly that

• there is no green pigment in animals
• most mammals do not see green (viz electromagnetic wavelength 500-550 nanometres)
• many mammals are nocturnal, active when all colours are difficult to see
• most mammals have a strong sense of smell, so hardly need to see all the colours as we do
• most mammals are so active that crypsis is of limited value.

The key to understanding the lack of green in mammals is that the failure to see green is more than made up by other wavelengths of light that mammals can see, but humans cannot see.

For example, ultraviolet is probably visible to many mammals other than humans. Mammals are preoccupied with colours other than free, and they may even see some of these as green.

We could investigate this, beginning with 'colour-blind' persons. Some human individuals are born with dichromatic vision. It is easy to assume that dichromats see less than trichromats, but perhaps they just see differently.

So, most mammals may as well be green, because to mammalian predators they blend into the green background anyway, and to avian and reptilian predators they will be detected anyway, by other means.

(writing in progress)

(writing in progress)

Also see https://www.inaturalist.org/posts/67881-carpobrotus-a-plant-paradoxically-combining-succulence-with-fire-proneness#

The fleshy fruit of Carpobrotus edulis (Aizoaceae, https://en.wikipedia.org/wiki/Carpobrotus_edulis and https://pza.sanbi.org/carpobrotus-edulis#:~:text=Carpobrotus%20edulis%20is%20an%20easy,the%20herb%20or%20kitchen%20garden.) is noteworthy for several reasons.

These are as follows:

• This presents an example of 'plasticfruits', because it seems unlikely that the genus would be separated from Mesembryanthemum were it not for the shift of its fruits to endozoochory.
• The species, C. edulis, is unusual in that its flowers and fleshy fruits advertise themselves by means of the same hue (yellow).
• Carpobrotus has a disjunct distribution on southern continents, with the various spp. retaining the ability to hybridise, despite having been isolated long enough to speciate.

However, describing the fruit of Carpobrotus is complicated, because of

• unusual involvement of the calyx, in the form of persistent, leaf-like sepals,
• difficulty of determining which stage of the fruit is most aptly described as 'ripe', and
• recent hybridisation; two spp. have widely invaded the Northern Hemisphere for anthropogenic reasons, and there has been blurring of species-specific features by interbreeding among various congeners.

The following shows the fruit of Carpobrotus edulis, broken open at a stage when the pericarp and sepals are still succulent: https://www.inaturalist.org/observations/42120429.

The following show the progression of development and ripening of the fruit of C. edulis.

Before the fruits are full-size, they (including persistent, succulent sepals) start to turn yellowish.

https://www.inaturalist.org/observations/115872051
https://www.inaturalist.org/observations/89895230
https://www.inaturalist.org/observations/88236776
https://www.inaturalist.org/observations/87772815
https://www.inaturalist.org/observations/106596528
https://www.inaturalist.org/observations/123335213
https://www.inaturalist.org/observations/32491481
https://www.inaturalist.org/observations/124909927
https://www.inaturalist.org/observations/132469845
https://www.inaturalist.org/observations/131339813
https://www.inaturalist.org/observations/87294043
https://www.inaturalist.org/observations/130757305
https://www.inaturalist.org/observations/106394290
https://www.inaturalist.org/observations/93731117
https://www.inaturalist.org/observations/93894400
https://www.inaturalist.org/observations/99695747

The persistent sepals may be reddish. However, this does not necessarily make the fruits more conspicuous, because the leaves also tend to feature reddish hues:

https://www.inaturalist.org/observations/104813471
https://www.inaturalist.org/observations/108365096
https://www.inaturalist.org/observations/124730681
https://www.inaturalist.org/observations/104689416

When the fruits are full-size, the yellow is conspicuous to the human eye. At this stage, the fruits (including the bright-hued, persistent sepals) are as still as succulent as the leaves:

https://www.inaturalist.org/observations/103899006
https://www.inaturalist.org/observations/94488192
https://www.inaturalist.org/observations/90143530
https://www.inaturalist.org/observations/18837165
https://www.inaturalist.org/observations/61602265
https://www.inaturalist.org/observations/107171583
https://www.inaturalist.org/observations/132570222
https://www.inaturalist.org/observations/87784339
https://www.inaturalist.org/observations/129976088
https://www.inaturalist.org/observations/129498972

Then the sepals begin to wither:

https://www.inaturalist.org/observations/127908784

Then the rest of the fruit also starts to wither and turn brown:

https://www.inaturalist.org/observations/101685251
https://www.inaturalist.org/observations/125799262

Finally, the whole fruit and pedicel become brown, with no part of the complex structure remaining succulent, although the moist, pasty, translucent mass in which the seeds are embedded remains fluid:

https://www.inaturalist.org/observations/58832483
https://www.inaturalist.org/observations/42874622

(writing in progress)