Diario de milewski

Many of the large animals of the Mesozoic (https://en.wikipedia.org/wiki/Mesozoic) walked with cantilevered bipedality. Why is it that the only animal locomoting this way today is one species of pangolin?

Our human bias makes it easy to assume that bipedality is, per se, a meaningful category of posture and locomotion. Actually, bipedality is so heterogeneous that using it as an overall description can be misleading (see https://en.wikipedia.org/wiki/Bipedalism and https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1571302/ and https://owlcation.com/stem/Animals-that-are-Bipedal-two-legs).

When the torso is held horizontal, balanced by head and neck on one side and tail on the other, this forms a level beam pivoting on the hind limbs ((https://en.wikipedia.org/wiki/Cantilever). Balance is maintained - despite the instability of just two supporting feet - because the mass of the beam anterior to the hips equals that posterior to the hips.

Of course, this needs a sufficiently long and massive tail. However, the potential adaptive advantage is economical walking, because moving one pair of limbs costs less energy than moving two.

Keeping cantilevered balance while running should be relatively easy for the same reason that riding a bicycle hands-off is easier at some speed than slowly. It is the animals able to keep anterior/posterior balance while walking bipedally that are of particular interest.

Cantilevered bipedality is so different from upright (mainly human) bipedality that the two modes have little in common beyond the weight being placed solely on the hind feet.

In Mesozoic times, cantilevered bipedality was pervasive. It evolved independently in at least two major clades of 'dinosaurs' (https://en.wikipedia.org/wiki/Dinosaur), the saurischians and the ornithischians, that configured the hips in different ways.

It also occurred over a wide range of families and body sizes, and including flying, feathered forms (https://en.wikipedia.org/wiki/Dromaeosauridae and https://www.mcgill.ca/newsroom/channels/news/some-dinosaurs-could-fly-they-were-birds-323548). Even some sauropods (https://en.wikipedia.org/wiki/Sauropoda), although walking on all four limbs, probably held the tail horizontal to balance the long neck (https://itotd.com/articles/4723/the-argentinosaurus/).

Today, various lizards can run bipedally (https://www.youtube.com/watch?v=XAo09yYOpCU), but none walks with cantilevered bipedality.

Birds differ categorically from theropods (https://en.wikipedia.org/wiki/Theropoda) in the bony tail being so short (https://www.fossilhunters.xyz/geography-of-life/into-thin-air-the-origin-of-birds.html), with any long caudal feathers so light (https://es.123rf.com/photo_95366001_peacock-walking-around-on-the-grass.html), that a cantilever is out of the question.

In this sense birds - despite being the descendants of theropods - are posturally in a category of their own. No bird walks with cantilevered bipedality, the emu (Dromaius novaehollandiae) perhaps coming closest (https://www.youtube.com/watch?v=cJPNiXOV2XE) because it is odd in depositing fat on its tail.

Various mammals have independently evolved towards emphasis of the hind limbs, but few of them walk bipedally.

Some rodents (e.g. https://en.wikipedia.org/wiki/Pedetes), lagomorphs and macropodid marsupials hop slowly instead of walking, using gaits unknown in the Mesozoic, or 'walk' by moving first the fore and then the hind feet synchronously (https://www.youtube.com/watch?v=8hEKqUG-WVc).

Few bipedally-inclined mammals other than kangaroos have tails massive enough to act as cantilevers. And kangaroos - far from walking bipedally - cumbersomely use all fours plus the muscular tail as a fifth limb (https://www.youtube.com/watch?v=Mi53VlMA31I).

Ground sloths (https://en.wikipedia.org/wiki/Ground_sloth) had a bipedal tendency, but it is unlikely that they walked with cantilevered bipedality. Their tails were muscular enough to act as a prop but not long or massive enough to act as a cantilever (e.g. see https://www.gettyimages.com.au/detail/news-photo/mounted-prontoterium-skeleton-of-an-extinct-ground-sloth-news-photo/543654805).

The same applies to the extant giant armadillo (Priodontes maximus, https://en.wikipedia.org/wiki/Giant_armadillo and https://www.thedodo.com/in-the-wild/rare-giant-armadillo-rescued-brazil).

The only living mammal known to use cantilevered bipedality as its most frequent mode of walking is the second largest-bodied of the eight species of pangolins: the Cape pangolin (Smutsia temminckii, see https://fascinatingafrica.com/species/ground-pangolin/ and https://sites.psu.edu/shanetheman/files/2020/03/pangolin-2019.jpg and https://www.inaturalist.org/observations/16236845 and https://pangolindiamondscorp.files.wordpress.com/2014/04/cape-pangolin-from-youtube-video.jpg and second photo in https://www.dogcatplace.com/wildanimals/list-of-animals-that-can-walk-move-freely-on-two-legs).

In all species of pangolins the tails are unusually heavy, partly because they are exceptionally armoured and partly because they are, in arboreal species, exceptionally long (and prehensile). Furthermore, the fore claws of pangolins are so prominent that they tend to obstruct walking on the fore feet.

Since all pangolins have particularly economical metabolism and walk slowly, all are candidates for cantilevered bipedality.

However, the small species of pangolins walk on all fours, in some cases placing weight on the knuckles instead of the sole or claws. The smallest species of all (see https://www.inaturalist.org/journal/milewski/59232-tamanduas-have-converged-with-african-pangolins-except-in-anti-predator-defences#) is so arboreally specialised that it rarely walks on the ground in the first place.

The giant pangolin (Smutsia gigantea) is probably capable of bipedal walking but seems usually to use all fours, possibly because its tail has a different proportional size to that of the Cape pangolin, and its fore limbs are particularly muscular.

This leaves - seemingly almost by accident - the Cape pangolin (https://en.wikipedia.org/wiki/Ground_pangolin) as an afterthought of a mode of locomotion which once ruled the terrestrial world.

For the last 30 years, Sonchus oleraceus (https://en.wikipedia.org/wiki/Sonchus_oleraceus) has been an important - and perhaps the healthiest - part of my diet.

This herbaceous member of the daisy family is a common weed (https://www.researchgate.net/publication/334503437_Biology_impact_and_management_of_common_sowthistle_Sonchus_oleraceus_L) near my home, which has a mediterranean-type climate. The only way I have eaten it is pre-hominid, simply plucking the green leaves (https://en.wikipedia.org/wiki/Sonchus_oleraceus#/media/File:Leaf_of_Sonchus_oleraceus.png) and chewing them raw on the spot. I do not accept the flowers, even in the form of the buds.

I usually eat about 50 leaves at a time, on average several times per week during the cool and rainy half of the year. That totals about 5,000 leaves per year, with a break in the summer and autumn when S. oleraceus is largely unavailable.

I value salad in my diet, but I have found S. oleraceus so reliable and satisfactory that I have not bought any leafy greens from any sort of shop for the last 15 years. I simply browse when I am out and about, finding my favourite weed easily near sidewalks and in neglected gardens and vacant lots in this metropolitan area.

Herbicides are frequently used to kill weeds here, but it has been easy to find unsprayed plants, and in all these years I have never had the slightest negative reaction from my herbivory.

The leaves are less fibrous than those of dandelion (Taraxacum officinale, https://en.wikipedia.org/wiki/Taraxacum_officinale, which is scarcer here because of its preference for slightly neglected but frequently watered lawns). However, they are similarly bitterish and moderately astringent. I eat S. oleraceus not for its taste but for its salubrious effects.

The genera Sonchus and Lactuca both have a tendency for the leaves to be somewhat prickly. Sonchus oleraceus can be thought of as a 'relaxed thistle', in which the potentiality for prickliness is seldom realised and the leaves are therefore palatable.

Sonchus has hardly been subjected to selective breeding (https://jasonpadvorac.com/projects/sonchus-breeding/ and https://www.researchgate.net/publication/225167218_Crop_domestication_in_the_Compositae_A_family-wide_trait_assessment), but it presents, in its weedy natural form, something approaching the perfect leafy green from the viewpoint of human health.

At times over the past decades I have grown chicory leaves (https://en.wikipedia.org/wiki/Chicory) in my garden, as a closely-related but somewhat selectively bred alternative to S. oleraceus. I have also taken trouble to establish another weedy daisy more associated with the tropics, namely Bidens pilosa (https://en.wikipedia.org/wiki/Bidens_pilosa and https://www.jstor.org/stable/4252362 and https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7099298/), as an annual volunteer in my garden. However, both attempts failed: in the case of Cichorium because the plants require too much care and in the case of B. pilosa because the shoots turn out to be too rich in kidney-irritating oxalates (of which S. oleraceus seems blissfully free).

Sonchus oleraceus remains so underappreciated in human diets that, in my decades of gratefully consuming it and recommending it to others in health-conscious circles, I have never seen it eaten by even one other person.

However, in my personal experience: my everyday environment has presented me with a choice of two species of weedy daisies - one a nominal thistle and the other the prototype for lettuce - originally indigenous to the same parts of Eurasia and gone cosmopolitan in the modern world. The one that has prevailing in my diet, so far, is the one that has not been domesticated but could hardly have been improved even if it had.

Everyone knows that the roots of most plants have an intimate mutualism with fungi, and that the invisible mycorrhizae formed in this way help plants to extract nutrients from soils (https://en.wikipedia.org/wiki/Mycorrhiza and https://mycorrhizas.info/ecm.html).

However, how many realise that some of these associations are more powerful than others, and that the most powerful mycorrhizae were naturally absent from regions where herbivores are naturally most abundant?

For example, one of the puzzles of global biogeography is that South Africa was virtually devoid of 'supermycorrhizae' common elsewhere in the world.

How did this come about?

Most mycorrhizal fungi have microscopic hyphae. However, ectotrophic mycorrhizae - or ectomycorrhizae for short - can be macroscopic (https://en.wikipedia.org/wiki/Ectomycorrhiza) and are particularly powerful metabolically. Whereas in other mycorrhizae the fungal cells merely infiltrate the roots, in ectomycorrhizae they form a mycelium actually replacing root tissue. Furthermore, hyphae originating at the roots reach up above the soil surface, breaking down litter such as fallen twigs by oxidising them.

It is from these 'supermycorrhizae' that most wild mushrooms and truffles (both edible and toxic) grow as reproductive structures (https://www.sciencedirect.com/science/article/abs/pii/S0065229618300880?via%3Dihub). This means that even those naturalists ignorant about microbiology have seen, and may have eaten, ectomycorrhizal fungi (https://link.springer.com/chapter/10.1007/978-3-319-53064-2_7 and https://academic.oup.com/femsre/article/31/4/388/2398987 and https://en.wikipedia.org/wiki/Truffle).

That ectomycorrhizae are effectively 'supermicorrhizae' is evident in the fact that many of the fastest-growing trees useful to humans, such as pines, oaks, poplars, eucalypts, dipterocarps and casuarinas (https://www.academia.edu/27848967/Ectomycorrhizas_in_plant_communities), depend on these associations. Most of the timber and paper we use comes from plantations nourished via ectomycorrhizal mutualisms.

Africa includes vast areas of miombo woodland (https://en.wikipedia.org/wiki/Miombo), dominated by ectomycorrhizal trees (https://nph.onlinelibrary.wiley.com/doi/abs/10.1111/j.1469-8137.1982.tb03398.x). However, the only species penetrating South Africa are two species of Brachystegia (restricted to a few marginal locations, http://www.soutpansberg.com/brachystegia/), one species of Afzelia (sparsely distributed in the north of the country, https://en.wikipedia.org/wiki/Afzelia_quanzensis), and perhaps Manilkara.

Although most of South Africa lies at the same latitudes as lands dominated by ectomycorrhizal trees such as pines, oaks and eucalypts, it naturally lacked ecological counterparts for these trees. Ectomycorrhizae were even absent from South African species of the genera Salix and Myrica, which are ectomycorrhizal elsewhere.

The lack of ectomycorrhizae in South Africa is consistent with the odd treelessness (http://pza.sanbi.org/vegetation/fynbos-biome and http://pza.sanbi.org/vegetation/nama-karoo-biome) of this country under mediterranean-type and semi-arid climates, where the dominant plants are heath-like shrubs belonging to families (e.g. Asteraceae) and genera (e.g. Erica) unassociated with this type of mutualism.

By contrast, ectomycorrhizae are common in similar climates in Australia (https://nph.onlinelibrary.wiley.com/doi/abs/10.1111/j.1469-8137.1980.tb00768.x), even where trees are scarce. For example, in mesic heathlands ectomycorrhizal Myrtaceae include small shrubs (e.g. see https://florabase.dpaw.wa.gov.au/browse/profile/5365). And in semi-arid scrub some of the acacias in Australia are ectomycorrhizal.

Part of the explanation may be the abundance - even by African standards - of indigenous herbivores in South Africa.

Worldwide, vegetation dominated by ectomycorrhizal plants tends to have sparse populations of herbivores, an obvious example being the boreal forest - where spruces, firs, larches, alders, willows and birches are all ectomycorrhizal.

There is likely to be a fundamental incompatibility between the nutrient-recycling regime associated with faeces and urine, and that associated with fungal decay. The former emphasises bacteria and anaerobic decomposition conserving organic matter in the soil, whereas the latter emphasises macroscopic 'white rot' and aerobic decomposition in which organic matter is wasted.

And a previously overlooked link in any explanation of the biogeographical anomaly is as follows:

South Africa is one of the few lands on Earth where fungus-culturing termites (Isoptera: Termitidae: Macrotermitinae: Odontotermes, https://www.inaturalist.org/observations/38179116 and https://www.inaturalist.org/observations/66913664) penetrate temperate climates. These insects feed litter, including coarse faeces of herbivores, to underground cultures of the fungus Termitomyces. The fungal culture produces small food-bodies eaten by the termites as well as unusually large mushrooms eaten by humans, and the rapidly recycled nutrients act as fertiliser.

Fungus-culturing termites, by removing surface litter, effectively extend the realm of herbivory at the expense of the sort of decomposition that would favour 'supermycorrhizae'. Whereas ectomycorrhizal fungi promote trees, the fungi mutualistic with macrotermitines are equally powerful in promoting a different form of recycling of nutrients, which favours grasses and herbivores.

In summary:

South Africa was - until the anthropogenic establishment of trees introduced from other continents - unsuitable for ectomycorrhizae. And part of the reason may be that it was the only land on Earth combining temperate climates with a regime of consumption of plant material in which diverse large herbivores, together with various termites operating beyond the usual roles of such insects, precluded the nutritional regime essential to ectomycorrhizae.

Africa, South America and Australia are all southern continents (https://en.wikipedia.org/wiki/Continent#/media/File:Continental_models-Australia.gif and https://www.vecteezy.com/vector-art/532993-world-map), but they differ biogeographically.

Africa broadly straddles the equator and has always been connected to Eurasia. South America was formerly isolated, but is now connected via North America. Australia remains an isolated southern landmass, an island continent.

Africa has a balance of wet and dry environments, South America has extensive wetness, and Australia has extensive drought (https://en.wikipedia.org/wiki/Earth_rainfall_climatology#/media/File:MeanMonthlyP.gif).

The comparative biogeography of various families of animals, plants, fungi, and algae on the southern continents is of obvious interest. Here I investigate the caesalps (including both https://en.wikipedia.org/wiki/Caesalpinioideae and https://en.wikipedia.org/wiki/Detarioideae), a major category of mainly woody and mainly tropical plants capable of dominating vegetation - but with seemingly unpredictable ecological patterns.

I realise that, among the legumes, the pea (Fabaceae) and mimosa (Mimosaceae) families are the clearly-defined ones and 'Caesalpiniaceae' refers to what is really a heterogeneous assemblage of 'other legumes' (https://www.sciencedirect.com/topics/agricultural-and-biological-sciences/caesalpinioideae and https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5558824/ and https://nph.onlinelibrary.wiley.com/doi/10.1111/nph.14474). However, I think we can learn something interesting about biogeography by categorising the non-pea and non-mimosa legumes as caesalps.

With this caveat in mind, caesalps are surprisingly different in their incidence among our three continents. By incidence I mean the size of the plants and the proportion they occupy in the vegetation.

In a series of four Posts, I will sketch the respective floras of caesalps from this ecological viewpoint. I will then try to explain the overall pattern: major in Africa but minor in Australia, with South America intermediate.

All legumes tend to have nutritional niches in which the supply of the protein-forming element, nitrogen, is supplemented from sources other than the soil. What sets caesalps apart from other legumes is that they are the least likely of the three families to form nodules on their roots, in which bacteria symbiotically fix atmospheric nitrogen.

Of the three southern continents, it is Australia that has the least incidence of caesalps.

About 18 genera are indigenous (https://keys.lucidcentral.org/keys/v3/FFPA/key/FFPA/Media/Html/Caesalpiniaceae.htm), but most of these have failed to penetrate the most extensive vegetation types in Australia, and those that have done so tend to consist of small plants despite the drought-tolerance of trees on this continent (https://search.informit.org/doi/abs/10.3316/ielapa.360289688263431). Furthermore, it is the cosmopolitan genera of caesalps that tend to be prominent in Australia (Senna, Cassia, Chamaecrista, Bauhinia, Caesalpinia).

The few genera of caesalps restricted to Australia (e.g. Petalostylis https://herbalistics.com.au/product/petalostylis-labicheoides-butterfly-bush-seed/, Labichea http://anpsa.org.au/l-lance.html, Barklya https://bie.ala.org.au/species/https://id.biodiversity.org.au/node/apni/2894458 and http://www.plantsoftheworldonline.org/taxon/urn:lsid:ipni.org:names:21784-1) are ecologically unimportant in the sense of being small plants making minor contributions to the vegetation.

Several genera are restricted to tropical rainforest, which covers only a small proportion of this continent. Most of these (e.g. Cynometra, Crudia, Maniltoa, Sindora, Storckiella) occur in tropical Asia and have only marginally penetrated Australia. However, it is noteworthy that Cassia grows to the size of a tree adjacent to rainforest in Australia.

The only genera of caesalps which qualify as dominating any vegetation in Australia are Senna (e.g. https://www.researchgate.net/figure/Senna-artemisioides-n-coriacea-low-sparse-shrubland-Floristic-group-56-Mapping-class_fig3_275964125 and https://www.anbg.gov.au/photo/vegetation/other-shrublands.html) and perhaps Lysiphyllum (https://en.wikipedia.org/wiki/Lysiphyllum_cunninghamii and https://www.territorynativeplants.com.au/bauhinia-cunninghamii-bean-tree and https://bie.ala.org.au/species/https://id.biodiversity.org.au/node/apni/2914442). However, Senna in Australia forms shrubs rather than trees, and possesses nitrogen-fixing nodules; and plants of Lysiphyllum are small for trees.

The genus Erythrophleum (https://www.flickr.com/photos/13639096@N06/3832663517 and https://florabase.dpaw.wa.gov.au/browse/profile/3662 and https://www.territorynativeplants.com.au/erythrophleum-chlorostachys-ironwood) occurs mainly in Africa, and the genus Peltophorum (https://en.wikipedia.org/wiki/Peltophorum_pterocarpum) occurs mainly in South America. Both have a minor incidence in tropical Australia.

The only genus recorded to be ectomycorrhizal is Intsia (https://www.academia.edu/27848967/Ectomycorrhizas_in_plant_communities and https://www.inaturalist.org/observations/68132536), which has an extremely restricted incidence in rainforest in Australia (https://en.wikipedia.org/wiki/Intsia).

In summary, the incidence of caesalps in Australia is slight in several ways. Those species growing as large plants make only a slight contribution to the canopy where they occur. Most of the tree species have penetrated Australia only slightly. And most of the species deeply penetrating the continent are slight in the sense of being shrubs, dominant over only small areas, and replaced successionally by trees belonging to families other than caesalps.

To be continued...

Having examined caesalps in Australia, let us turn to South America, where tree-dominated vegetation is most extensively rainforest (see https://www.nature.com/articles/s41598-019-50323-9) but also includes large areas of woodland and savanna.

Caesalps have a far greater incidence In South America than in Australia. However, they are not dominant in the vegetation, with a few exceptions such as Mora (https://en.wikipedia.org/wiki/Mora_(plant)) and Dicymbe (http://www.grupoecologiatropical.com/wp-content/uploads/2016/10/2008-Mycorrhiza-paper-McGuireal.pdf).

Shrubby/weedy genera are common, e.g. Caesalpinia, Senna (which includes trees in South America, https://en.wikipedia.org/wiki/Senna_(plant)), Erythrostemon and Chamaecrista.

Many genera form trees, e.g. Libidibia (https://www.inaturalist.org/observations/95870454), Schizolobium (https://en.wikipedia.org/wiki/Schizolobium_parahyba), Copaifera (https://en.wikipedia.org/wiki/Copaifera), Hymenaea (https://en.wikipedia.org/wiki/Hymenaea), Eperua (https://en.wikipedia.org/wiki/Eperua) and Pterogyne (https://en.wikipedia.org/wiki/Pterogyne). In the case of several species of Tachigali, the trees can reach up to 40 metres.

Caesalps in the form of trees are present in many vegetation types. However, as in Australia, those types of woodland or savanna that are dominated by legumes tend to favour Mimosaceae (e.g. see https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0220151).

Some examples of genera according to the various ecosystems are as follows:

• northernmost rainforests: Mora, Dicymbe (which can be ectomycorrhizal and dominate vegetation), Tachigali, Dialium (https://en.wikipedia.org/wiki/Dialium_guianense)
• amazon forest: Macrolobium (https://en.wikipedia.org/wiki/Macrolobium), Tachigali, Dicymbe, Schizolobium amazonicum, Campsiandra (https://tropical.theferns.info/viewtropical.php?id=Campsiandra+angustifolia and https://en.wikipedia.org/wiki/Campsiandra and https://www.inaturalist.org/observations/96654430)
• caatinga: Bauhinia, Cenostigma (e.g. https://species.wikimedia.org/wiki/Cenostigma_pyramidale)
• cerrado: Apuleia, Copaifera, Dimorphandra (https://en.wikipedia.org/wiki/Dimorphandra_mollis), Hymenaea, Tachigali (https://archive.org/details/cbarchive_121081_asynopsisfothegenustachigalile2008/page/n9/mode/2up?view=theater)
• chaco: Denisophytum, Copaifera, Cynometra, Hymenaea, Gleditsia (https://www.inaturalist.org/taxa/48497-Gleditsia-amorphoides and https://www.inaturalist.org/observations/99311523 and https://www.inaturalist.org/observations/99370608), Libidibia (https://www.inaturalist.org/observations/99483178)
• Atlantic forest: Paubrasilia (https://en.wikipedia.org/wiki/Paubrasilia), Plathymenia (https://en.wikipedia.org/wiki/Plathymenia and https://www.inaturalist.org/observations/99397857), Schizolobium (https://www.inaturalist.org/observations/97173996 and https://www.inaturalist.org/observations/96300503), Bauhinia forficata
• atacama: Tara spinosa (https://en.wikipedia.org/wiki/Tara_spinosa and https://www.inaturalist.org/observations/49650269)

Ectomycorrizal mutualisms have been recorded in Dicymbe, Eperua and Gleditsia (https://www.academia.edu/27848967/Ectomycorrhizas_in_plant_communities and https://mycorrhizas.info/ecm.html). Fixation of nitrogen in nodules has been recorded in Dimorphandra and Tachigali, the latter being remarkable because this nutritional mode is not usually associated with genera capable of growing into such large trees. The following gives further information on the nutritional modes: https://www.jstor.org/stable/2588772.

To be continued...

Having covered Australia and South America, let us turn to Africa, where caesalps dominate some of the most extensive types of vegetation, and characterise important ecosystems (https://en.wikipedia.org/wiki/Angolan_miombo_woodlands and https://en.wikipedia.org/wiki/Central_Zambezian_miombo_woodlands and https://en.wikipedia.org/wiki/Southern_miombo_woodlands and https://en.wikipedia.org/wiki/Zambezian_and_mopane_woodlands and https://en.wikipedia.org/wiki/Eastern_miombo_woodlands).

Much of tropical Africa is covered by woodlands and savannas dominated collectively by the caesalps Brachystegia (https://en.wikipedia.org/wiki/Brachystegia), Julbernardia (https://en.wikipedia.org/wiki/Julbernardia) and Isoberlinia (https://en.wikipedia.org/wiki/Isoberlinia), all of which are ectomycorrhizal.

Extensive savanna in tropical southern Africa is dominated by Colophospermum (https://www.wikiwand.com/en/Zambezian_and_mopane_woodlands and https://en.wikipedia.org/wiki/Mopane), which is not ectomycorrhizal.

Minor areas are dominated by the small trees Baikiaea (https://www.wikiwand.com/en/Baikiaea_plurijuga and https://en.wikipedia.org/wiki/Baikiaea) and Burkea (https://www.inaturalist.org/observations/84055435 and https://www.inaturalist.org/observations/84280090 and https://www.inaturalist.org/observations/40353399 and https://en.wikipedia.org/wiki/Burkea_(plant)).

Tropical rainforest is in places dominated by Gilbertiodendron (https://en.wikipedia.org/wiki/Gilbertiodendron), Julbernardia and Cryptosepalum (https://en.wikipedia.org/wiki/Cryptosepalum_exfoliatum), all of which are ectomycorrhizal, and possibly also Baikiaea (https://en.wikipedia.org/wiki/Baikiaea_insignis), Cynometra and Tetraberlinia (https://en.wikipedia.org/wiki/Tetraberlinia_tubmaniana).

Tropical rainforest elsewhere contains Anthonotha (https://en.wikipedia.org/wiki/Anthonotha), Berlinia (https://en.wikipedia.org/wiki/Berlinia), Microberlinia (https://en.wikipedia.org/wiki/Microberlinia_bisulcata), Aphanocalyx (https://en.wikipedia.org/wiki/Aphanocalyx), Paramacrolobium (https://tropical.theferns.info/viewtropical.php?id=Paramacrolobium+coeruleum), Didelotia and Afzelia (https://en.wikipedia.org/wiki/Afzelia_africana), all of which are also ectomycorrhizal (https://www.academia.edu/27848967/Ectomycorrhizas_in_plant_communities and https://mycorrhizas.info/ecm.html) despite failing to dominate the canopy.

Dialium occurs as a minor component of equatorial rainforest in Africa (e.g. https://en.wikipedia.org/wiki/Dialium_corbisieri), as it does in South America.

As on the other southern continents, there are shrubby/weedy caesalps in the genera Cassia, Senna, Chamaecrista, Piliostigma and Bauhinia (https://en.wikipedia.org/wiki/List_of_Southern_African_indigenous_trees_and_woody_lianes#Caesalpiniaceae). In addition there is Pterolobium (https://en.wikipedia.org/wiki/Pterolobium and https://en.wikipedia.org/wiki/Pterolobium_stellatum), which does not occur on the other southern continents.

Caesalps in the form of large shrubs or small trees occur in semi-deserts in Africa, e.g. Parkinsonia (shared with South America), Ceratonia (https://en.wikipedia.org/wiki/Ceratonia) and Dialium (https://en.wikipedia.org/wiki/Dialium_guineense and https://www.inaturalist.org/observations/27754490 and https://en.wikipedia.org/wiki/Dialium_orientale).

Guibourtia in the form of trees (https://en.wikipedia.org/wiki/Guibourtia_coleosperma and https://www.inaturalist.org/observations/33591043) extends to extratropical southern Africa, as a minor part of the vegetation, as do small trees of Schotia (https://en.wikipedia.org/wiki/Schotia) and Peltophorum. However, the only ectomycorrhizal caesalp that extends considerably into South Africa is Afzelia quanzensis.

To be continued...

In this series of four posts I have summarised the incidence of caesalps according to the size of individual plants and their appropriation of the canopy, averaged over the entirety of each continent.

This crude estimation suffices to show that the southern continents differ by two orders of magnitude in the incidence of legumes other than peas and mimosas: about tenfold greater in South America than in Australia, and about tenfold greater again in Africa than in South America.

This variation is particularly surprising in view of the fact that various genera (e.g. Hymenaea, Cynometra, Copaifera, Erythrophleum, Bauhinia, Piliostigma, Crudia, Senna, Parkinsonia, Dialium) are indigenous to more than one of these continents.

The incidence of ectomycorrhizal (https://mycorrhizas.info/ecm.html) caesalps varies in the same direction and by similar orders of magnitude, suggesting that the prevalence of caesalps in Africa is linked to a mode of supplementation targeting not only nitrogen but also other nutrient elements.

The intercontinental differences are not merely the result of the drought of Australia and the wetness of South America. Instead, they apply even within categories of climate and arborescence.

Taking tropical savannas for example: in Australia the dominant trees tend to be ectomycorrhizal Myrtaceae (eucalypts), with caesalps relegated to a minor incidence and lacking ectomycorrhizae. By contrast, in Africa the dominant trees tend to be ectomycorrhizal caesalps, of which Brachystegia is representative.

So, if the intercontinental variation ultimately reflects environmental differences, what could these be?

Well, Africa is the continent on which there is the greatest incidence of large herbivores, which has implications for the cycling of crucial nutrients such as phosphorus and zinc (https://onlinelibrary.wiley.com/doi/pdf/10.1111/jbi.12100 and https://onlinelibrary.wiley.com/doi/abs/10.1046/j.1365-2699.2000.00436.x).

It also differs from South America and Australia in the occurrence of fungus-culturing termites (see https://www.inaturalist.org/journal/milewski/59182-comparisons-of-termites-and-termite-eating-animals-in-africa-and-australia-part-1#), which are the most powerful recyclers of nutrients among the social insects.

Furthermore, the three continents form a series w.r.t. the histories of anthropogenic influence on the cycling of nutrients. The human species has farmed in tropical Africa for most of the last ten thousand years, whereas there was no farming in Australia until Europeans settled. As in various other aspects both ecological and biotic, South America is intermediate.

Miombo woodland and other caesalp-dominated vegetation in Africa has been affected continually by proboscideans. The bush elephant (Loxodonta africana) and other large herbivores were hardly affected by the spate of megafaunal extinctions which wiped out proboscideans from the Americas by ten thousand years ago. Although practised in tropical South America for at least five thousand years, farming seems not to have affected the main types of woodland and savanna (llanos, caatinga, chaco, cerrado), leaving these vegetation types largely unaffected by both shifting cultivation (https://en.wikipedia.org/wiki/Shifting_cultivation) and all herbivores heavier than 100 kg.

Because woodlands dominated by ectomycorrhizal caesalps usually occur on relatively nutrient-poor soils, the role of fungus-culturing termites (particularly Macrotermes, https://en.wikipedia.org/wiki/Macrotermes) may be crucial. These termites concentrate trace elements in their mounds (https://zslpublications.onlinelibrary.wiley.com/doi/10.1111/j.1469-7998.2008.00544.x), which can form patches of sufficient fertility to grow crops. This advantage sets Africa apart from the other two continents and perhaps helps to explain the antiquity and extent of shifting cultivation and the adaptation of caesalps to an anthropogenic regime on a large scale.

Miombo woodland and its equivalent north of the equator are so resilient from slash-and-burn cultivation (https://www.jstor.org/stable/4620417) that these vegetation types can be seen as, in some sense, anthropogenic (https://en.wikipedia.org/wiki/Chitemene#:~:text=Chitemene%20(also%20spelled%20citemene)%2C,agriculture%20practiced%20throughout%20northern%20Zambia.). Disturbance by humans and large herbivores may be integral to the dominance of caesalps.

What looks like pristine woodland may actually be the 'climax' of a successional cycle in which the trees are intermittently broken down by physical means, and their eventual regeneration is facilitated by the patchy suppression of wildfires carried out by farmers and gregarious herbivores.

Among leguminous woody plants, mimosas (Mimosaceae) are far more likely than caesalps (see https://www.inaturalist.org/journal/milewski/59528-caesalps-on-southern-continents-part-1#new_comment) to be favourite foods of large herbivores. Accordingly, it is mimosas, not caesalps, that tend to possess large spines.

However, the genus Gleditsia (https://en.wikipedia.org/wiki/Gleditsia) is exceptional among caesalps (see https://www.inaturalist.org/journal/milewski/59528-caesalps-on-southern-continents-part-1#) in the degree to which it looks and acts like a member of the mimosas (http://www.nzdl.org/cgi-bin/library?e=d-00000-00---off-0hdl--00-0----0-10-0---0---0direct-10---4-------0-1l--11-en-50---20-help---00-0-1-00-0-0-11----0-0-&a=d&c=hdl&cl=CL1.12&d=HASH011ec19a37bb817d72319187.33).

Gleditsia has extremely large spines (https://en.wikipedia.org/wiki/Honey_locust and (https://www.britannica.com/plant/honey-locust-tree-genus and https://www.feedipedia.org/content/honey-locust-gleditsia-triacanthos-thorns-madrid-spain and https://canr.udel.edu/udbg/?plant=gleditsia-triacanthos)
and https://www.etsy.com/au/listing/1065477392/huge-honey-locust-thorns-real-acacia?ref=pla_sameshop_listing_top-1 and https://www.etsy.com/au/listing/777216641/huge-honey-locust-thorns-real-acacia?ref=pla_sameshop_listing_top-2&cns=1), as well as small leaflets, flowers lacking obvious petals, and fruits functionally similar to those of mimosas such as that widespread 'acacia', Vachellia nilotica (https://en.wikipedia.org/wiki/Vachellia_nilotica).

Implicit in this convergence with mimosas is that nutritional modes will have likewise converged: we can expect Gleditsia to be adapted to exploit nutrients recycled in urine and faeces, while at the same time fixing atmospheric nitrogen.

It comes as a surprise, therefore, that Gleditsia not only lacks nitrogen-fixing nodules but is listed as possessing ectomycorrhizae (see https://www.inaturalist.org/journal/milewski/59485-the-mystery-of-the-missing-mycorrhizae#). These macroscopic fungal root-extensions are rare among mimosas (https://mycorrhizas.info/ecm.html) and antithetical to the nutritional strategy typical of mimosas.

Because the largest mycorrhizal fungi tend to be associated with the roots of plants unattractive to large herbivores, and to promote nutritional regimes antagonistic to herbivory, Gleditsia presents a puzzle. Is its nutritional strategy really similar to those of caesalps which are neither attractive as food for herbivores, nor spinescent (https://mycorrhizas.info/ecm.html)?

If it really is ectomycorrhizal, Gleditsia is unique in combining some of the largest stem-protecting spines and some of the largest root-assisting fungi.

I cannot resolve this question because I have yet to find the original paper reporting ectomycorrhizae in Gleditsia. However, I can further explain spinescence in the context of ectomycorrhizal plants generally.

Spines, thorns and prickles vary greatly in their size, the part of the plant from which they are derived, and their adaptive functions. The large spines of many species of 'acacias' (now genus Vachellia) are derived from stipules (e.g. http://elephantseyegarden.blogspot.com/2010/06/fever-tree-umbrella-thorn-mimosa-port.html).

The largest spines derived from stipules and stems of dicotyledonous plants serve to control the foraging of the largest herbivores. This control should not be confused with deterrence, because plants with large spines are likely to be promoted by regimes of intense herbivory that recycle nutrients via faeces and urine while retarding competing plants. The function of these large spines is not to prevent loss of foliage as much as to protect the crucial regenerative organs/tissues, such as shoots and cambium, in the interests of plant and herbivore alike.

Spines have been established to occur in several genera of ectomycorrhizal plants, but they are of a different kind, compatible with a nutritional regime marginalising large herbivores.

The boreal conifer Picea (https://www.shutterstock.com/nb/video/clip-17336719-closer-look-picea-pungens-blue-spruce-thorny), the Australian pea Gastrolobium (https://www.ukwildflowers.com/Web_pages/gastrolobium_spinosum_prickly_poison.htm), and several Mediterranean and Californian species of oaks Quercus (https://naturenet.net/blogs/2009/01/15/crimson/) are ectomycorrhizal and possess leaf-spines similar to that familiar plant, holly (https://en.wikipedia.org/wiki/Ilex_aquifolium).

Leaf-spinescent plants are basically different from stem- or stipule-spinescent plants in that the leaves are too sclerophyllous (rigidly fibrous and lignified) to be particularly attractive to herbivores in the first place. They generally occur in ecosystems poor in large herbivores, where the main consumer of plant matter is episodic wildfires (https://www.academia.edu/7565234/Ecology_of_Australia_the_effects_of_nutrient_poor_soils_and_intense_fires). The spines can be thought of as an extension of the unpalatability of the leaves themselves, rather than a means of collaboration with herbivores as seem in spinescent 'acacias'.

Spinescence in ectomycorrhizal plants can therefore be summarised as follows:

In general, ectomycorrhizal plants lack spines, which is unsurprising because the nutritional regimes to which they are adapted are unsuitable for large herbivores. Those ectomycorrhizal species possessing spines tend to have a form of spinescence (i.e. leaf-spinescence) associated with limited herbivory. The stem-spinescence of Gleditsia is so unusual among ectomycorrhizal plants, and the association of Gleditsia with large herbivores is so obvious, that it calls into question the very claim that Gleditsia possesses ectomycorrhizae.

Does anyone doubt that, in general, the colouration of giraffes is a form of camouflage?

The flags which I have described (see https://www.inaturalist.org/journal/milewski/48447-conspicuous-features-of-colouration-in-giraffes#) hardly negate this generalisation, because they are subsidiary features.

However, one subspecies of giraffe has overall colouration so conspicuously pale that it does seem to mean a partial negation (https://africa.cgtn.com/2018/11/22/niger-to-move-protected-giraffes-as-habitat-shrinks/ and https://www.dreamstime.com/stock-photo-salivating-giraffe-its-dripping-saliva-epic-image70986808).

I refer to Giraffa camelopardalis peralta (https://en.wikipedia.org/wiki/West_African_giraffe and https://www.researchgate.net/profile/Bertrand-Chardonnet/publication/296408231_Antelope_Survey_Update_n9_November_2004_IUCN_SCC_Antelope_Specialist_Group_Report_Special_Issue_West_and_Central_Africa/links/56d5466608ae2cd682b9a145/Antelope-Survey-Update-n9-November-2004-IUCN-SCC-Antelope-Specialist-Group-Report-Special-Issue-West-and-Central-Africa.pdf#page=36 and http://travel2unlimited.com/niger-koure-giraffes/) of the western Sahel.

The particular pallor of Giraffa camelopardalis peralta is owing mainly to the breadth of the whitish 'matrix' among the blotches (see https://www.inaturalist.org/observations/91931958 and https://www.inaturalist.org/observations/67258492).

This differs from any pale overall effect in the form of giraffe inhabiting the edge of the Namib desert in southern Africa, which is owing mainly to fading of the blotches themselves and does not particularly affect the head or legs (Giraffa giraffa angolensis, see https://app.nimia.com/fpvideo/746972087/746972087-giraffe-walking-namibia and https://giraffeconservation.org/programmes/nw-namibia/).

The following photos of the most pallid individuals show how the camouflage effect has been compromised in Giraffa camelopardalis peralta in its current habitat. Although a corollary of the pallor is that the auricular and pedal flags are reduced, the caudal flag is if anything enhanced, because the tail-tassel remains black in all individuals.

https://www.inaturalist.org/observations/89357382
https://www.inaturalist.org/observations/66647598
https://www.inaturalist.org/observations/66511242
https://www.inaturalist.org/observations/66511269
https://www.inaturalist.org/observations/91633228
https://www.inaturalist.org/observations/84033823
https://www.inaturalist.org/observations/66511261
https://www.inaturalist.org/observations/66510605
https://www.inaturalist.org/observations/66510563
https://www.bornfree.org.uk/articles/conservation-update-gcf

Can it not be said that G. c. peralta has, in evolutionary terms, switched from inconspicuous colouration to conspicuous colouration? The mechanism has been a quantitative shift (mainly an encroachment of the pale matrix relative to the blotching), but the effect seems qualitative.

If so, this seems convergent with the conspicuous pallor of three other gregarious ruminants of the southern fringes of the Sahara, as exemplified by the following views of Oryx dammah:

https://www.shutterstock.com/nb/image-photo/herd-scimitar-horned-oryx-dammah-walking-1114267055 and https://www.shutterstock.com/nb/image-photo/herd-scimitar-horned-oryx-dammah-walking-1114267097.

Oryx dammah (https://www.inaturalist.org/observations/80301962), Addax nasomaculatus and Nanger dama ruficollis (https://www.hatadaranch.com/dama) all inhabited the vicinity of the Sahara, with O. dammah coexisting in the Sahel with giraffes (including G. c. peralta), A. nasomaculatus penetrating the Sahara itself, and N. d. ruficollis living in the eastern Sahel (where the local form of giraffe was not as pallid). In converging on a pattern of conspicuous overall pallor, these species set an unique pattern among the arid-adapted ungulates of the world.

I realise that:

• arid-adapted reptiles, birds, and small mammals tend to be pallid as a form of crypsis in the pale environments of deserts, particularly sandy deserts;
• there is seasonal variation in the colouration of A. nasomaculatus and possibly also the other two bovids referred to here; and
• hippotragins and gazelles tend to be adaptively conspicuous even in savannas, which can be partly explained by the futility of trying to hide as gregarious animals in the open.

However, ruminants in the southern African deserts and semi-deserts (https://www.istockphoto.com/photo/gemsbok-oryx-in-namib-desert-gm146923746-14197980 and https://www.agefotostock.com/age/en/details-photo/springbok-antidorcas-marsupialis-adults-walking-on-sand-namib-desert-in-namibia/YS1-1722745) lack the relevant pattern of colouration. In the degree of their pallor, our three bovids seem to have adapted in convergent ways to the particular conditions of the Sahara and its southern edges.

Seen in this context, does the pallor of G. c. peralta not seem to be part of a regional faunistic pattern?

The following illustrate the pallor of our three bovids, and the adaptive convergence that this represents.

Oryx dammah:
https://www.shutterstock.com/nb/image-photo/huge-herd-scimitarhorned-oryx-sahara-wildlife-1988644823
https://www.shutterstock.com/nb/image-photo/herd-scimitar-horned-oryx-dammah-walking-1114267079
https://www.inaturalist.org/observations/74899599
https://www.shutterstock.com/nb/image-photo/antelope-scimitar-horn-oryx-leucoryx-due-1605546127 https://www.shutterstock.com/nb/image-photo/antelope-scimitar-horn-oryx-leucoryx-due-1606769167 https://www.shutterstock.com/nb/image-photo/scimitar-oryx-aka-sahara-endangered-animal-1093058519 https://www.shutterstock.com/nb/image-photo/very-rare-scimitarhorned-oryx-dammah-extinct-1932362348
https://www.shutterstock.com/nb/image-photo/scimitar-oryx-aka-sahara-endangered-animal-1093058531
https://www.shutterstock.com/nb/image-photo/scimitar-oryx-aka-sahara-endangered-animal-1219348936
https://wildlifeconservation101.files.wordpress.com/2012/04/images-2.jpeg

Addax nasomaculatus:
https://www.dreamstime.com/stock-photo-beautiful-addax-desert-eilat-image72805856
https://www.dreamstime.com/addax-walking-jebil-national-park-tunisia-image126127489
https://creatures-of-the-world.fandom.com/wiki/Addax?file=Adult_addax.jpg https://pixabay.com/photos/addax-negev-desert-desert-dweller-4764789/
https://www.istockphoto.com/photo/a-critically-endangered-addax-also-known-as-the-screwhorn-or-white-antelope-stops-to-gm1141454086-305811175
https://www.dreamstime.com/royalty-free-stock-photos-antelope-addax-image19237788

Nanger dama:
https://naturerules1.fandom.com/wiki/Dama_Gazelle?file=3e2b7d460fe86e7d50d1bfec49723c12.jpg
https://www.iucn.org/ssc-groups/mammals/mammals-a-e/antelope/resources http://lh3.ggpht.com/_1wtadqGaaPs/TCiTe14fKOI/AAAAAAAAGm4/SeKuL3yo1c4/s1600-h/tmp1127_thumb4.jpg
https://www.texasdivide.com/dama-gazelle

to be continued...

To understand the pallor in Giraffa camelopardalis peralta (see my last Post), it may also be useful to review the masculine darkness that occurs in most forms of giraffe.

In full maturity, male giraffes become much larger-bodied than adult females: they can weigh up to 1.5 tonnes as opposed to the average 0.8 tonnes of their mothers. They also darken and produce odours from skin glands (https://zslpublications.onlinelibrary.wiley.com/doi/abs/10.1111/j.1469-7998.2012.00904.x and https://theses.ncl.ac.uk/jspui/handle/10443/1461 and https://www.facebook.com/jumbojunction/photos/a.1554448954625642/6189916674412157/).

Darkness and smelliness are individually variable (https://www.sciencedirect.com/science/article/abs/pii/S0003347219302453). However, the pattern of sexual dimorphism in giraffes is such that males combine:

• indeterminate development (in contrast to the determinate development of females), with
• shorter lifespan than that of females (https://zslpublications.onlinelibrary.wiley.com/doi/abs/10.1111/j.1469-7998.2012.00904.x).

Estes (1991, page 203) confirms that "Rival bulls recognise and react to one another from a long way off, demonstrating the capability of long-distance communication".

The changes in males at the peak of masculinity detract from their camouflage. However, this poses no particular puzzle, because:

• it is normal for males, particularly in sexually dimorphic species, to trade off security for sexual advantage,
• masculine self-advertisement needs not compromise the camouflage of females and juveniles, because giraffes are only loosely gregarious and dark males tend to be solitary (https://www.sciencedirect.com/science/article/abs/pii/S0003347219302453), and
• males may be less vulnerable than females to predation by day, and dark figures (although conspicuous in bright light) remain inconspicuous by night.

It is noteworthy that the various flags (see https://www.inaturalist.org/journal/milewski/48447-conspicuous-features-of-colouration-in-giraffes#) tend to remain despite the overall darkening in mature males.

Another noteworthy finding is that, in maturing males of Giraffa tippelskirchi tippelskirchi, the whole head remains pale in some individuals (e.g. https://www.istockphoto.com/photo/masai-giraffe-gm1296042780-389584472 and https://www.istockphoto.com/photo/giraffe-in-east-tsavo-park-in-kenya-gm655519816-119240823 and https://stock.adobe.com/sk/search?filters%5Bcontent_type%3Aphoto%5D=1&filters%5Bcontent_type%3Aillustration%5D=1&filters%5Bcontent_type%3Azip_vector%5D=1&filters%5Bcontent_type%3Avideo%5D=0&filters%5Bcontent_type%3Atemplate%5D=0&filters%5Bcontent_type%3A3d%5D=0&filters%5Bcontent_type%3Aaudio%5D=0&filters%5Binclude_stock_enterprise%5D=0&filters%5Bis_editorial%5D=0&filters%5Bfree_collection%5D=0&filters%5Bcontent_type%3Aimage%5D=1&k=%22masai+giraffe%22&order=relevance&price%5B%24%5D=1&safe_search=1&limit=100&search_page=4&search_type=pagination&get_facets=0&asset_id=442967946 https://stock.adobe.com/sk/search?filters%5Bcontent_type%3Aphoto%5D=1&filters%5Bcontent_type%3Aillustration%5D=1&filters%5Bcontent_type%3Azip_vector%5D=1&filters%5Bcontent_type%3Avideo%5D=0&filters%5Bcontent_type%3Atemplate%5D=0&filters%5Bcontent_type%3A3d%5D=0&filters%5Bcontent_type%3Aaudio%5D=0&filters%5Binclude_stock_enterprise%5D=0&filters%5Bis_editorial%5D=0&filters%5Bfree_collection%5D=0&filters%5Bcontent_type%3Aimage%5D=1&k=%22masai+giraffe%22&order=relevance&price%5B%24%5D=1&safe_search=1&limit=100&search_page=4&search_type=pagination&get_facets=0&asset_id=442967951).

The following illustrate various degrees/extents of masculine darkness in various species and subspecies of giraffes:

Giraffa giraffa giraffa

https://www.tripadvisor.in/LocationPhotoDirectLink-g312616-d7122939-i314711382-Moholoholo_Mountain_View_Bush_camp-Hoedspruit_Limpopo_Province.html
http://www.timaverybirding.com/photos/albums/2013/08-09_africa/20130901/safari/giraffe_b.jpg
https://es.123rf.com/photo_21431497_giraffe-bull.html
https://www.facebook.com/mafojanisafaris-100408458676624/photos/pcb.143965800987556/143965734320896/
https://www.africahunting.com/threads/dark-giraffe-bull-hunt-south-africa.55821/
https://www.facebook.com/mafojanisafaris-100408458676624/photos/pcb.143965800987556/143965737654229/
https://www.shutterstock.com/image-photo/big-dark-male-giraffe-swaziland-nature-1719565081
https://www.dreamstime.com/stock-photo-giraffe-browsing-high-leaves-bush-image50867157
https://www.inaturalist.org/observations/25833033
https://www.alamy.com/vertical-portrait-of-a-walking-male-giraffe-with-gymnogene-flying-above-its-head-in-kruger-park-south-africa-image369493417.html
https://www.alamy.com/vertical-portrait-of-lighter-and-darker-giraffe-walking-passed-each-other-in-kruger-park-south-africa-image369044615.html
https://www.inaturalist.org/observations/58934167
https://es.123rf.com/photo_14344382_old-giraffe-standing-next-to-dead-tree.html and https://es.123rf.com/photo_14344348_old-giraffe-standing-next-to-dead-tree.html
https://www.dreamstime.com/male-adult-giraffe-dark-pattern-walking-dry-winter-bush-kruger-south-africa-vertical-portrait-ox-peckers-his-back-image194142317 and https://www.alamy.com/vertical-portrait-of-a-walking-giraffe-with-ox-peckers-on-his-back-in-kruger-park-south-africa-image369083259.html
https://www.inaturalist.org/observations/10160335
https://www.youtube.com/watch?v=5CxgiE512aA

Giraffa giraffa angolensis

https://www.alamy.com/a-large-giraffe-giraffa-camelopardalis-peralta-namibia-image244651145.html
https://www.inaturalist.org/observations/5647366
https://www.inaturalist.org/observations/19426797
https://www.inaturalist.org/observations/56245174

Giraffa tippelskirchi tippelskirchi

https://www.inaturalist.org/observations/44386793
https://www.flickr.com/photos/davidbygott/4037387278
scroll in http://travelwithkally.blogspot.com/2015/10/serengeti-national-park.html
https://www.shutterstock.com/nb/image-photo/masai-giraffe-giraffa-camelopardalis-tippelskirchii-spelled-1669283635
https://www.alamy.com/a-giraffe-family-with-five-members-including-young-calves-standing-on-the-savanna-image360612331.html
https://www.offset.com/photos/giraffe-is-walking-between-the-bush-on-safari-in-kenya-912635
https://www.inaturalist.org/observations/54487214
https://www.inaturalist.org/observations/35913219
https://www.alamy.com/adult-giraffe-grumeti-tanzania-image3531887.html and https://www.alamy.com/adult-giraffe-grumeti-tanzania-image3531361.html
https://www.robertharding.com/preview/832-387758/masai-giraffes-giraffa-camelopardalis-tippelskirchi-animal-pair-mating/

Giraffa tippelskirchi thornicrofti

https://www.livescience.com/19686-aging-male-giraffes-black-spots.html
https://www.dreamstime.com/vast-open-plains-south-luangwa-giraffe-baboon-warthog-zambia-southern-africa-baboons-african-natural-image106164904

Giraffa reticulata
https://www.dreamstime.com/royalty-free-stock-photos-reticulated-giraffe-image26813158
https://www.dreamstime.com/full-body-portrait-reticulated-giraffe-giraffa-camelopardalis-reticulata-walking-northern-kenya-savannah-landscape-africa-image131865776
https://www.dreamstime.com/stock-images-male-reticulated-giraffe-zoo-waiting-food-miami-south-florida-image38695804

Giraffa camelopardalis camelopardalis

https://www.dreamstime.com/reticulated-giraffe-reticulated-giraffe-standing-sun-nice-day-image108993437
https://www.dreamstime.com/stock-photo-giraffes-mysore-zoo-i-clicked-photo-my-visit-baby-giraffe-was-eating-food-box-known-s-unique-image46191110

Giraffa camelopardalis rothschildi

https://www.flickr.com/photos/wildlifepictures/49439204907
https://www.istockphoto.com/photo/female-giraffe-with-a-baby-in-the-savannah-gm669752120-122872959
https://www.alamy.com/stock-photo-mature-male-rothschilds-giraffe-giraffe-camelopardis-rothschildi-at-47964879.html
https://www.sciencephoto.com/media/441396/view/giraffe-in-uganda
https://www.alamy.com/stock-photo-mature-male-rothschilds-giraffe-giraffe-camelopardis-rothschildi-at-47964865.html
https://www.inaturalist.org/observations/38456853
https://www.alamy.com/stock-photo-rothschilds-giraffe-rothschild-giraffe-baringo-giraffe-ugandan-giraffe-47925641.html
https://www.alamy.com/stock-photo-rothschilds-giraffe-rothschild-giraffe-baringo-giraffe-ugandan-giraffe-47925639.html
https://www.alamy.com/stock-photo-rothschilds-giraffe-rothschild-giraffe-baringo-giraffe-ugandan-giraffe-47925649.html
https://www.alamy.com/stock-photo-rothschild-giraffe-giraffa-camelopardalis-rothschildi-herd-walking-125631277.html
https://www.inaturalist.org/observations/190129545

If it is true that masculine darkness is poorly-developed in G. reticulata, this is in line with the observation that this is the species of giraffe possessing the greatest uniformity in various aspects of its colouration.

We need more information for G. c. peralta. The few photos available suggest that the blotches do darken in mature males (see e.g. https://www.inaturalist.org/observations/28558145). However, this is not enough to give the figure a dark aspect overall, because the matrix seems to remain as pale as in females and immature males.

Masculine darkness in giraffes does not necessarily contradict the idea that the conspicuous pallor of G. c. peralta of the Sahel/Sahara represents a basic adaptive shift among giraffes. It is this pallor, rather than masculine darkness, that is an important exception to any generalisation that the colouration of giraffes functions as camouflage.