milewski's Journal

Coaton W G H (1937)

(writing in progress)

See https://www.gettyimages.com.au/detail/photo/giant-termite-nest-in-the-okavango-delta-savannah-royalty-free-image/638725834?phrase=termite+nest&adppopup=true and https://www.gettyimages.com.au/detail/news-photo/termite-mount-in-front-of-a-marula-tree-in-the-gomoti-news-photo/1200229952?adppopup=true.

My references are McCarthy et al. (1991, https://www.sciencedirect.com/science/article/abs/pii/088329279190071V)

https://www.sciencedirect.com/science/article/abs/pii/000925419090065F

https://journals.co.za/doi/abs/10.10520/AJA052550590_272

https://onlinelibrary.wiley.com/doi/abs/10.1002/esp.1008

https://onlinelibrary.wiley.com/doi/abs/10.1111/j.1365-3091.1991.tb00362.x

https://www.tandfonline.com/doi/abs/10.1080/00359199809520384

https://journals.co.za/doi/abs/10.10520/EJC-1abed244bb

https://www.sciencedirect.com/science/article/abs/pii/003707389390078J

https://onlinelibrary.wiley.com/doi/abs/10.1111/j.1365-3091.1992.tb02153.x

https://onlinelibrary.wiley.com/doi/abs/10.1046/j.1365-2028.1998.89-89089.x

https://www.sciencedirect.com/science/article/abs/pii/002216949490216X

https://www.sciencedirect.com/science/article/abs/pii/S0022169405003501

What I have learned from these studies:

The Okavango is not a delta, but an alluvial fan

The channels keep changing, as the accumulation of organic matter blocks them

Peat fires deplete this organkc matter

Most (96%) of the water evaporates, instead of sinking into the earth

Thus there is a pan-like effect, in which a considerable quantity of salt builds up in the Okavango

Islands are formed partly by termite mounds, via a mechanism of precipitation of solutes

Evaporation and precipitation are aided by the transpiration of trees

The centres of islands tend to be so saline that they are bare of vegetation

I infer that:

• the accumulation of peat presumably acts as a bank for iodine, over the century of accumulation of peat in each channel, and
• the concentration of calcium carbonate on islands, particularly based on termite mounds, would also act as a bank for iodine, in the form of iodate.

Iodine flows in, in extreme dilution, in the water of the Okavango River. It is somewhat concentrated by evaporation, and then greatly concentrated by evaporation from islands, aided by

• transpiration from trees,
• precipitation of calcium carbonate, and
• binding to peat.

Iodine is thus stored in non-volatile forms, over time.
The peat fires that deplete the organic matter are mild, with combustion so slow that minimal smoke is produced. Thisnpossibly minimkses volatilisation, conserving much of the iodine.

Because the watershed of the Okavango River is flat and sandy, silt is scarce in the Okavango. Thus, remarkably, the main land-building material for the construction of islands is actually precipitated solutes, e.g. calcium.

The groundwater under the islands is actually saline.

I infer that the islands of the Okavango act almost like the pans of the semi-arid Kalahari, farther afield.

In both cases,

• fairly fresh water surrounds the evaporational surface, which concentrates salt and calcium from the water,
• ungulates are likely to seek nutrients by geophagy, perhaps at the periphery of the saline centre of the evaporative surface,
• groundwater under the evaporative surface is likely to be saline, in contrast to the surrounding water at the same horizontal level.

Basically, both pans in the Kalahari and islands in the Okavango can concentrate iodine from the surrounding fresh waters, and store it in calcium-rich material at tne surface, where iodate is likely to be stabilised. In the case of the islands of the Okavango, leaching of IO3 by rainfall is likely to be prevented on termite mounds.

Peat layers aggrade by about 5 cm per year, to a cumulative depth of about 5 m, before the channel blocks and shifts elsewhere. This takes about one century.

"Water in the Okavango swamps is of the carbonate-rich variety typical of continental regiions" (McCarthy and Metcalfe, 1990).

Islands have precipitation of calcium carbonate from evaporation of the waters of the Okavango River. This precipitate is concentrated by termites.

In the capillary fringe area of islands, calcium precipitates out first (edges of islands), whereas sodium reaches the centre of the island in solution, precipitating out there to form a trona crust (https://en.wikipedia.org/wiki/Trona).

Island centres:
"During the rainy season, surface crusts are dissolved and leached down into the soil...the rainfall rarely contributes to the water table, and the rainfall and its dissolved salts are merely held in the profile...During the dry season, the salts are returned to the surface" (McCarthy and Metcalfe, 1990).

McCarthy and Metcalfe (1990): the observed concentrations of calcium in the Okavango are calculated to have required 22,000 years, to the nearest order of magnitude. Thksnis likely to err on the conservative side, because rates of evaporation are greater now than in the past.

My commentary:

• I infer that these deposits are about as old as those in heuweltjies in the Western Cape province of South Africa.
• Peat may possibly act as a trap and bank for selenium, as well as iodine.

Salt tends to accumulate in the centres of the islands, to be eventually leached to the saline groundwater table, or redissolved, leading to the recovery of vegetation on the saline surfaces.

The result:
Although certain zones do become saline, there are many opportunities for geophagy of iodate and calcium, without excess salinity.

McCarthy does point out, in certain of his papers, that the calcium precipitated on islands is brought up (and concentrated?) by termites (mounds?).

Notes from McCarthy, McIver and Verhagen (1991):

I infer that the whole island fringe acts like a termite mound, in the sense that it is

• elevated,
• colonised by trees,
• replete with calcium carbonate,
• unleached,
• perpetuated by the precipitation of solutes, by virtue of transpiration, and
• ?fire-free.

The centres of the islands act like land, in which bare ground develops because of salinity.

The fringe of the island is raised by the precipitation of silica and calcium originating in the waters of the Okavango River. The centre of each island is at a lower altitude than not only the fringe of the island, but also the water level of the surrounding marsh/swamp. This is because, in contrast to the fringe lf the island, the centre is actually deflated. Obviously, the level of the groundwater beneath the centre of the island is likewise lower than the water level of the surrounding marsh/swamp. This is true for all of the islands, regardless if the stage in the flood-cycle. The saline water in the centre of the island may actually sink into the briney watertable there, because saline water is denser (heavier) than fresh water.

(writing in progress)

@tonyrebelo @beetledude @botswanabugs @jtermiteo @r_r_r_ @hamishrobertson

The aim of this Post is to provide an interpretive review of several half-century old, but still best-of-their-kind, papers about the African members of an ecologically extremely important family of insects, viz. Isoptera: Hodotermitidae.

HODOTERMES

My first reference is Coaton W G H and Sheasby J L (1975, https://pascal-francis.inist.fr/vibad/index.php?action=getRecordDetail&idt=PASCAL7650284276).

Hodotermes is restricted to subSaharan Africa, and contains two spp.

Hodotermes occurs in southern and East Africa, as far north as Eritrea. However, H. mossambicus occurs only as far north as Ethiopia. Hodotermes erythreensis replaces H. mossambicus in Eritrea, Somaliland, and possibly Somalia and easternmost Ethiopia (https://termites.myspecies.info/content/hodotermes-0).

My commentary:
Biogeographically, H. mossambicus is remarkable in paralleling a pattern in the ungulates, in which

• many genera and spp. occur in southern and East Africa, but not West Africa,
• there is a broad disjunction between southern and East Africa, and
• the Horn of Africa tends to have a distinctive fauna.

Hodotermes mossambicus occurs mainly where mean annual rainfall is <750 mm, except for East Africa, where it may extend to 800 mm. It occurs widely in Namibia (50-600 mm) and Botswana (200->600 mm), including the southwestern Kalahari (Kgalagadi Transfrontier Park, https://en.wikipedia.org/wiki/Kgalagadi_Transfrontier_Park).

Hodotermes mossambicus is absent from most of the Namib, but present in the Namib in Kaokoland and at Walvis Bay. In the Kalahari, it is restricted to the finer-grained substrates, and perhaps absent from the coarsest, nutrient-poorest sands. However, it is present (see Coaton 1963, https://koedoe.co.za/index.php/koedoe/article/view/811 and https://koedoe.co.za/index.php/koedoe/article/view/811/918) in sparse populations on dunes/sandflats.

My commentary:

• Hodotermes mossambicus occurs mainly where rainfall is not enough to leach nutrients from the soil, thus inhabiting dry/eutrophic ecosystems (thorn savanna and karoo).
• However, it occurs in mesic climates in e.g. KwaZulu-Natal, Mpumalanga, much of Zimbabwe (albeit not miombo and the eastern highlands, where this termite is absent), and the Serengeti ecosystem, where soils are relatively nutrient-rich.
• In extending to mesic climates in East Africa, H. mossambicus parallels the ungulate superspecies Madoqua damarensis/cavendishi/thomasi.
• The outlying occurrence of H. mossambicus in southern Malawi (https://www.inaturalist.org/observations/99337392 and https://www.inaturalist.org/observations/40692683) may not have been known to Coaton and Sheasby (1975).
• It reaches the winter-rainfall area in Namaqualand (https://en.wikipedia.org/wiki/Namaqualand), and the small enclaves of karoo vegetation at Worcester (https://en.wikipedia.org/wiki/Worcester,_South_Africa) and Montagu (https://en.wikipedia.org/wiki/Montagu,_South_Africa), where it broadly overlaps with Microhodotermes (see below).

Hodotermes mossambicus is somewhat tolerant of salinity (also see Popov et al.). In Zimbabwe, it is absent from granitic substrates, except where saline.

My commentary:
Hodotermes mossambicus seems to prefer base-saturated substrates.

Each colony has several/many hives (diameter <60 cm), at densities of 85-96 hives per hectare. The hives are 1.8-12.2 m deep. Only a few of the hives in each colony contain reproductives. Storage chambers for food are extraneous to the hives, and are mainly <30 cm deep. The hives within each colony are linked by tunnels, 2.5-7.5 cm in diameter.

My commentary:
The subterranean location of the hives, with no clue at the surface, protects H. mossambicus from predation by large myrmecophages (https://en.wikipedia.org/wiki/Myrmecophagy). This helps to balance the exposure of the workers as they scurry on the surface (including in cold weather) while foraging.

The diet is mainly, but not exclusively, grass.

My commentary:
In contrast to the normal conception of a termite, Hodotermes mossambicus acts as a grazer of mainly dry 'sweet' grass. It drinks at depth, as opposed to the horizontal commuting performed by grazing ungulates.

MICROHODOTERMES

My second reference is Coaton W G H and Sheasby J L (1974) National survey of the Isoptera of southern Africa. 6. Microhodotermes Sjoestedt (Hodotermitidae). Cimbebasia Series A, vol. 3, no. 6, pp. 47-59.

Like Hodotermes, this genus is approximately restricted to non-leaching climates (mean annual rainfall <750 mm). Whereas Hodotermes is associated with summer rainfall, Microhodotermes is associated with winter rainfall.

My commentary:
The threshold value of 750 mm of rainfall seems to correspond to Paarl Mountain, where I observed M. viator (https://www.inaturalist.org/posts/80176-thought-provoking-observations-from-a-brief-visit-to-the-karoo-desert-national-botanical-garden-and-paarl-mountain-botanic-garden-october-2001#).

Microhodotermes occurs under winter rainfall not only in southern Africa, but also in North Africa, where Hodotermes is absent. The North African spp. are:

• Microhodotermes maroccanus (Morocco), and
• Microhodotermes wasmanni (Tunisia, Libya, and Egypt).

My commentary:
Note that this genus, and the whole family, is absent from the Sahel (https://en.wikipedia.org/wiki/Sahel). This parallels various ungulates and myrmecophages such as Equus, Alcelaphus, Connochaetes, Raphicerus campestris, and Sylvicapra grimmia, which are typical of semi-arid climates in southern Africa, but have no wild counterparts in the Sahel in West and west-central Africa.

Microhodotermes viator

• collects faeces of ungulates to some extent, as food,
• inhabits a single hive per colony, in contrast to H. mossambicus,
• builds a low, broadly conical mound, up to 1.5 m diameter, above each hive, the latter being buried just below the general level of the ground and about 0.5 m below the apex of the mound,
• forages over a radius of up to 45 m around the (buried) hive, by emerging from small holes at various distances from the hive, and
• sometimes makes little piles of sticks and other litter at the entrance of these holes.

My commentary:
There are regions of winter rainfall (mediterranean-type climates) in Europe, the Levant, California, Chile, Western Australia, and South Australia. However, none of these has any termite ecologically similar to Microhodotermes. Microhodotermes viator does not extend to the 'karoid' vegetation (https://www.inaturalist.org/observations/33095087 and https://www.inaturalist.org/observations/112265311) of the southwest Kalahari (it does not extend north of the Orange River (https://en.wikipedia.org/wiki/Orange_River) at the longitudes of the Kalahari). Please note that these authors do not mention any relationship between H. viator and heuweltjies (https://en.wikipedia.org/wiki/Heuweltjie).

ECOLOGICAL IMPORTANCE

In Zululand, the mean annual production of grass is at least 3 tonnes (dry matter) per hectare. Hodotermes mossambicus is capable, in its own right, of harvesting most of this production.

My commentary:
This implies suppression of wildfire, and conservation of nutrient elements, e.g. nitrogen, sulphur, and selenium, that are easily volatilised and depleted by combustion.

Coaton and Sheasby (1975) state:
"Coaton (1958) reported that extensive areas in Zululand carried saturated populations of H. mossambicus which were consuming grass at an estimated rate of 1-3 tons per morgen (1.06-3.17 metric tons per hectare) per year." Some populations consumed the entire crop of hay on a given site, where production of hay is about 3 tons per morgen (3.17 metric tons per hectare).

"Veld hay was produced at a rate of just on 3 metric tons/hectare over stretches of Lowveld and Zululand Thornveld in the Game Corridor between the Hluhluwe and the Umfolosi Game Reserves, an area not grazed by livestock in which H. mossambicus is present, although not in pest proportions...an extreme case is cited of an ungrazed area in Zululand, which for years had been completely denuded by the end of winter, yielding 3 metric tons of baled hay per hectare per annum after saturated harvester termite infestation had been exterminated; at the other end of the scale, a far lighter infestation in an ungrazed test enclosure in the central Orange Free State has been recorded as removing only about 100 kg per hectare per year out of a total hay production of just over one metric ton".

My commentary:
I infer that H. mossambicus is fully a part of the habitat of wild ungulates in Zululand, even though this area is on the moist side, climatically. The map in Coaton and Sheasby (1975) shows the species to be present in Ndumo. uMkhuze, and St Lucia reserves. This termite shares a guild with the fauna of ungulate grazers of Zululand, which are noteworthy in that they extend from tropical to subtropical latitudes.

"On numerous occasions foraging workers of H. mossambicus have been observed to be cutting down lush green grass, the supplies gleaned being taken directly underground or left lying on the surface around the foraging ports, for subsequent collection...Thriving colonies have also been recorded as doing extensive damage to growing small-grain, legume and fodder crops where the diet must have consisted almost exclusively of green plant material."

Coaton and Sheasby (1975) describe a situation in which the entire diet of a colony for years was lawn grass, viz. Cenchrus clandestinus (https://en.wikipedia.org/wiki/Cenchrus_clandestinus) in a green state.

"From the observations presented above it is clear that under natural conditions in the field colonies of H. mossambicus can thrive on a practically undiluted diet of green vegetation" (appropriately cured, of course, after cutting).

My commentary:
This is a clear reference to

• foraging by day in winter, and by night in summer,
• damage to agricultural crops, and
• consumption of not only dry but also green grass, the latter being in some cases 100% of the diet, and/or 100% of the green grass present on a given site.

My third reference is Coaton and Sheasby (1972, https://books.google.com.au/books/about/Preliminary_Report_on_a_Survey_of_the_Te.html?id=2DhDAAAAYAAJ&redir_esc=y).

Referring to Namibia, these authors state:
"Including the vast Northern Kalahari region, which is rather lightly infested by harvester termites except on the more consolidated soils of river flood-plains, omurambas, vleis, and dune valleys, it is very conservatively estimated that H. mossambicus consumes approximately 25% of the aggregate hay yield of South West Africa during every year of average rainfall. In years of drought, when grass regeneration is poor, this figure would be doubled or even trebled. Apart from lowering the stock-carrying capacity of the territory by at least one-quarter these termites are facilitating soil erosion and desert encroachment through their removal of the protective grass cover."

The following seems to have no presence on the Web, even as a mere citation:
Grube S (2000) Soil clumps - indicators of foraging activity by Hodotermes mossambicus (Hagen) (Isoptera, Hodotermitidae) in northern Namibia. Cimbebasia 16: 269-270.

This author states that dumping of soil at the surface, by Hodotermes mossambicus, is not correlated with foraging activity. Foraging activity peaked in the rainy season, whereas the dumping of soil peaked in winter.

My commentary:
Are these termites feeding fertiliser to the grass and herbivores? Why is it that no author mentions the faeces of Hodotermitidae, which - in my experience - appear dark and nutrient-rich, like valuable fertilizer?

Lee and Wood (1971, https://books.google.com.au/books/about/Termites_and_Soils.html?id=Hj5DAAAAYAAJ&redir_esc=y), on page 180, state in a worldwide context: "It is only in South Africa that the depredations of termites in pastures have been considered sufficiently injurious to merit the adoption of extensive control measures."

However, even in central Asia, the hodotermitid Acanthotermes ahngerianus is a pasture pest, where grass grows. Bare areas around the nests may occupy 20% of the surface. (Also see the second comment below.)

My commentary:
I take this as evidence of the role of herbivory - with ungulates and termites working in concert - in producing low, palatable, fire-free vegetation in South Africa. I note that no termite on Earth is known to cut and maintain hedges or lawns. The fact that H. mossambicus eradicates lawns (patchily), rather than promoting or maintaining them, is crucial for understanding how the same species can be despised as a pasture pest by pastoralists, and valued as a natural member of the grazing guild by wildlife managers who understand the longer-term, more holistic relationships among herbivory, fire, and nutrient-cycling.

@gonsaro @r-a-p @kfinn @jeremygilmore @santiagoramos @enricotosto96 @nicoolejnik
@paradoxornithidae @botswanabugs @tonyrebelo

The Patagonian mara (Dolichotis patagonum, https://academic.oup.com/mspecies/article/doi/10.2307/0.652.1/2600781?login=false) is a relatively cursorial caviid rodent (https://en.wikipedia.org/wiki/Caviidae), with a body mass of about 10 kg.

Please see https://www.researchgate.net/figure/Photo-of-a-mara-Dolichotis-patagonum-with-some-adaptations-for-running-being-highlighted_fig3_333504431 and https://go.gale.com/ps/i.do?id=GALE%7CA609539093&sid=googleScholar&v=2.1&it=r&linkaccess=abs&issn=03279383&p=IFME&sw=w&userGroupName=anon%7E9e205385&aty=open+web+entry.

This may be the only rodent on Earth that

• stots (https://en.wikipedia.org/wiki/Stotting) by means of stiff-legged, bouncing locomotion, and
• possesses a bleeze (defined as a dark/pale pattern so bold that it makes the whole figure conspicuous, even at distance, and even when stationary).

The combination of stotting and displaying a large-scale conspicuous pattern on the hindquarters is associated with ruminants, from Ourebia ourebi (about 15 kg, https://www.dreamstime.com/oribioribi-sprinting-across-savannah-slightly-longer-shutters-speed-to-capture-motion-oribi-sprinting-across-image143323292) to Alcelaphus caama (about 130 kg, https://www.flickr.com/photos/neeravbhatt/6191239661/).

Among animals featuring this combination, the Patagonian mara is possibly

• the only rodent, and
• the smallest of all mammals.

(Please also see https://www.argentinat.org/journal/milewski/64011-flags-as-features-of-adaptive-colouration-in-lepus-part-2-other-species-of-semi-arid-north-america#.)

The following show that the bleeze of the Patagonian mara

• is situated on the hindquarters, from the base of the tail to each haunch,
• has a mainly horizontal orientation,
• shows dark/pale contrast, independent of sheen/anti-sheen effects, and
• subsumes but is not contributed to by the tail, which is short and bare.

https://www.inaturalist.org/observations/152145094

https://www.inaturalist.org/observations/149507669

https://www.inaturalist.org/observations/74038522

https://www.inaturalist.org/observations/143733867

The following shows that, in some individuals, there is a whitish patch near each knee (at the junction of flank and hindleg). In posteriolateral view, this extends the bleeze in an anterior direction (https://www.inaturalist.org/observations/51901419).

The following show that the bleeze of the Patagonian mara is conspicuous even when the standing figure is viewed in full profile (https://www.inaturalist.org/observations/116824024 and https://www.inaturalist.org/observations/35690755).

The following suggests that the white band of the bleeze can be flared/erected (https://www.inaturalist.org/observations/131837234).

The following shows the appearance of the bleeze when the animal flees (https://www.inaturalist.org/observations/73664556 and https://www.inaturalist.org/observations/147930616).

The following shows that the bleeze is concealed by sitting, which is a frequently used posture in the Patagonian mara (https://www.inaturalist.org/observations/126838562 and https://www.inaturalist.org/observations/109675491).

The following show the stotting gait of the Patagonian mara:

https://www.debrecensun.hu/local/2023/03/24/the-patagonian-mara-of-the-debrecen-zoo-got-a-partner/

https://www.inaturalist.org/observations/106457844

https://www.alamy.com/patagonian-mara-cavy-dolichotis-patagonum-stotting-valdes-peninsula-chubut-patagonia-argentina-image263028463.html

https://www.mindenpictures.com/stock-photo-patagonian-mara-cavy-dolichotis-patagonum-stotting-valdes-peninsula-naturephotography-image90672455.html

https://www.gettyimages.ie/detail/photo/rabbit-in-patagonia-argentina-royalty-free-image/1331275428?phrase=patagonian+hare&adppopup=true

https://www.alamyimages.fr/cavi-de-patagonie-dolichotis-patagonum-peninsule-de-valdes-site-du-patrimoine-mondial-de-l-unesco-province-de-chubut-patagonie-argentine-image396012030.html?imageid=67D29AC8-2DDD-4730-BA46-8F64B855A9A5&p=1360155&pn=4&searchId=fdf407efb395ee4e1da5818b4f0aac6e&searchtype=0

https://www.alamy.com/stock-photo-patagonian-mara-cavy-dolichotis-patagonum-stotting-valdes-peninsula-86749556.html?imageid=518140A7-DE66-4D56-AC01-B56AB482092B&p=269381&pn=1&searchId=589d9d24beda2dcae20b7ee79d560a38&searchtype=0

The Patagonian mara has long, slender legs for a rodent, and might be described as a 'cavy on high heels'. However, it remains digitigrade (https://en.wikipedia.org/wiki/Digitigrade), not unguligrade (https://en.wikipedia.org/wiki/Ungulate).

The following show the walking gait of the Patagonian mara:

https://www.inaturalist.org/observations/164868054

https://www.inaturalist.org/observations/104542547

https://www.inaturalist.org/observations/62059798

https://www.inaturalist.org/observations/38340582

https://www.shutterstock.com/image-photo/patagonian-mara-dolichotis-patagonum-rodent-genus-1207058716

https://www.shutterstock.com/image-photo/patagonian-mara-dolichotis-patagonum-599052566

https://www.alamyimages.fr/photo-image-mara-de-patagonie-81921558.html?imageid=A36AB68D-A326-4319-8811-EE760D18CF22&p=46571&pn=1&searchId=3f16e8ad5731abf977a4bcd41d46be22&searchtype=0

The following show the similarity in walking gait between the mara and a like-size African bambi, Raphicerus campestris (about 10 kg):

https://www.shutterstock.com/image-photo/patagonia-mara-portrait-while-running-grass-399321796

https://www.alamy.com/stock-photo-steenbok-raphicerus-campestris-walking-south-africa-krueger-national-76132028.html

DISCUSSION

Bambis (ruminants with body mass 15 kg or less) occur in South America. However, all of these depend on cover, and have inconspicuous colouration. The niche for a bambi in open vegetation has been usurped by a rodent, on this continent.

With the possible exception of one species/subspecies of Ourebia (which is marginal to the body mass criterion for bambis, anyway), no bambi on Earth possesses a bleeze.

(Please see https://colombia.inaturalist.org/journal/milewski/70050-the-three-main-types-of-oribi-ourebia-at-a-glance.)

The steenbok possesses a conspicuous white patch on the buttocks. However this qualifies as a flag rather than a bleeze, because it is

• located relatively ventrally, where it is normally out of sight,
• hidden by slight hunching of the sacrum (https://www.inaturalist.org/observations/59156961), and
• only raised and flared in alarm (https://www.inaturalist.org/journal/milewski/70293-the-bambis-part-9-bleezes-flags-and-semets-in-the-bovid-genus-raphicerus#).

Furthermore, the steenbok does not stot.

Aspects of the Patagonian mara that are evolutionarily convergent with bovid bambis include

• extremely large eyes, adapted for good vision by day,
• hue-differentiation in the ground-colour
• monogamy, and
• precociality at birth.

However, there is negligible convergence with bovid bambis in

• number of offspring per birth,
• dentition,
• anatomy and function of the stomach,
• cud-chewing (the Patagonian mara is instead coprophagous),
• diminution of the faecal pellets (compare https://www.inaturalist.org/observations/114536300 with https://www.inaturalist.org/observations/76872272 and https://www.inaturalist.org/observations/106239384 and https://www.inaturalist.org/observations/96703224),
• form of the claws, and
• form and mobility of the ear pinnae.

in view of the limited convergence between the Patagonian mara and ungulates, its possession of a well-developed bleeze is surprising. In displaying this conspicuous pattern, particularly by stotting, it outdoes its bovid counterparts - despite being otherwise only superficially ungulate-like.

Everyone knows that the only part of the world that bears any faunistic resemblance to Africa is tropical Asia, particularly the Indian subcontinent.

However, what may be poorly appreciated by most naturalists is how this relates to the ecologically important fungus-culturing termites, Macrotermes and Odontotermes.

Both of these genera occur in both Africa and Asia. However, the main differences are that the largest above-ground structures

• are built by Macrotermes in Africa, vs by Odontotermes in Asia, and
• are far smaller in Asia than in Africa.

Lee and Wood (1971), on page 172, states:
"Large termite mounds (species not identified) are common on grey soils near Soekamandi (https://mapcarta.com/15598374) in Java."

Mounds, large enough to be sites for the cultivation of dryland crops, are scattered through rice paddies on the flat lowlands of Thailand.

In Sri Lanka, mounds of Odontotermes redemanni (https://en.wikipedia.org/wiki/Odontotermes_redemanni#:~:text=Odontotermes%20redemanni%2C%20is%20a%20species,of%20sugarcane%2C%20tea%20and%20coconut.) are large enough for the cultivation of crops.

In central Congo, there are large fossil mounds (now abandoned) of termites, dating from a previous dry period. These nave densities of 4-7/ha, and occupy 30% of the ground surface. The outer layer of these mounds is fairly fertile. However, the central mass consists of subsoil, which has proven to be infertile for cultivated crops when levelled. In this area, horticulture has produced large patches of sterile ground.

Hesse (1955) noted that, in many areas of East Africa, the mounds of termites are used as earth-licks (i.e. for geophagy), by Bos and wild ungulates. (My commentary: in my experience, this includes the relatively small mounds of Odontotermes.)

In iNaturalist, the coverage of termites is poor.

This is unfortunate, because these insects are ecologically important.

In this Post, I compensate somewhat for this deficiency by reviewing the literature on the biogeography of certain termites in southern Africa, in an interpretive way.

The references are

• Coaton and Sheasby (1972, https://books.google.com.au/books/about/Preliminary_Report_on_a_Survey_of_the_Te.html?id=2DhDAAAAYAAJ&redir_esc=y),
• Ruelle et al. (1975, https://pascal-francis.inist.fr/vibad/index.php?action=getRecordDetail&idt=PASCAL7536010953),
• Marais and Irish (1989, https://scholar.google.com.au/scholar?hl=en&as_sdt=0%2C5&q=Marais+and+Irish+1989&btnG=),
• Turner (2000, https://scholar.google.com.au/scholar?hl=en&as_sdt=0%2C5&q=Turner+2000+macrotermes&btnG=), and
• Tinley (1985, https://researchspace.csir.co.za/dspace/handle/10204/2353 and https://books.google.com.au/books/about/Coastal_Dunes_of_South_Africa.html?id=WuUHAQAAIAAJ&redir_esc=y and https://www.si.edu/object/siris_sil_312931 and https://www.abebooks.com/first-edition/Coastal-Dunes-South-Africa-Tinley-K/16862566834/bd).

Let us begin with the astonishing fact that termites (presumably Macrotermes) have been recorded burrowing as deep as 84 m. This has important implications ecologically, particularly for the retrieval of leached nutrients and the boosting of productivity in whole ecosystems.

Marais and Irish (1989) state "we have observed active termite tunnels at the following depths in South West African caves: 25 m (Pofaddergat), 27 m (Ghaub Cave), 47 m (Dante's Cave), 84 m (Arnhemgrot)"

The following provides information in the location of these caves: https://en.wikipedia.org/wiki/Caves_of_Namibia.

Ghaub Cave https://africantourer.com/attraction/ghaub-cave and https://www.booknamibia.com/listings/ghaub-cave-ghaub-nature-reserve-farm

Dante Cave https://www.ncei.noaa.gov/access/metadata/landing-page/bin/iso?id=noaa-cave-14268

Arnhem Cave: 85 km east-southeast of Windhoek https://www.namibweb.com/arnhemcave.htm and https://www.arnhemcaves.com/

I infer that the species penetrating underground to a depth of 84 m is Macrotermes natalensis.

The subfamily Macrotermitinae of the family Termitidae consists of fungus-culturing termites. This means that they function more like large herbivores than like mere invertebrate agents of decomposition.

Macrotermes is an extremely important genus of termites. However, few naturalists have a grasp of it, even at the generic level - partly owing to confusion with Odontotermes. One of the reasons why the biogeography of Macrotermes remains so obscure is that the size and shape of the mounds varies greatly among, and within, species.

For example:
The mounds of Macrotermes natalensis (https://www.inaturalist.org/taxa/431392-Macrotermes-natalensis)

• vary from low and conical to massive and domed, with diameter up to 30 m,
• tend to be relatively low at high altitudes, and
• tend to be open at the top in Uganda (https://en.wikipedia.org/wiki/Uganda), but sealed at the top in Ivory Coast (https://en.wikipedia.org/wiki/Ivory_Coast).

Macrotermes natalensis

• is the only species in its genus to penetrate south of 28 degrees South,
• reaches part of the Highveld (https://en.wikipedia.org/wiki/Highveld), as far south as 29 degrees South in Free State province in South Africa, - marginally penetrates the Nama Karoo (https://en.wikipedia.org/wiki/Nama_Karoo), and
• reaches nearly as far south as 33 degrees South in Eastern Cape province of South Africa.

Tinley (1985) refers to the occurrence of large mounds of Macrotermes natalensis in the Eastern Cape, at the southerly limit of the genus. The large mounds occur in a strip, hugging the coast from Transkei (https://en.wikipedia.org/wiki/Transkei) to near East London (https://en.wikipedia.org/wiki/East_London,_South_Africa). The soils in these landforms are red loams with clay subsoil.

Near East London, the mounds of M. natalensis have diameters of about 10 m and heights up to 2.5 m. They are covered in bush-clumps of Phoenix reclinata (https://www.inaturalist.org/taxa/166729-Phoenix-reclinata) and Phyllanthus reticulatus (https://www.inaturalist.org/taxa/340305-Phyllanthus-reticulatus), exempt from wildfire.

Macrotermes penetrates rainforest in tropical Africa, but not in southern Africa.

Macrotermes subhyalinus (https://www.inaturalist.org/observations?taxon_id=673979) builds mounds at least 6 m high in Angola and northeastern Africa, but no higher than 4.2 m in Kaokoland (https://en.wikipedia.org/wiki/Kaokoland), where it penetrates the pro-Namib.

Macrotermes bellicosus (https://www.inaturalist.org/taxa/346641-Macrotermes-bellicosus) in Uganda transports earth into its mounds at rates of 0.4-0.9 cubic metres per hectare per year. This produces mounds averaging 4-8 cubic metres per hectare.

Some spp. of Macrotermes (e.g. vitrialatus and falciger) harvest plant matter, including living plants which they cut down, in the open. In this way they resemble the hodotermitids Hodotermes (https://www.inaturalist.org/taxa/558313-Hodotermes) and Microhodotermes (https://www.inaturalist.org/taxa/568761-Microhodotermes-viator). Adding to this similarity is the fact that M. vitrialatus - like the ecologically powerful Hodotermes mossambicus (https://www.inaturalist.org/taxa/558312-Hodotermes-mossambicus) - does not build mounds.

Other spp. of Macrotermes forage on the ground surface (mainly for litter and faeces, including those of Bos and Loxodonta) under the cover of mud-runnels. They construct these temporary shelters ad-hoc, as they go along. Examples are M. natalensis, M. michaelseni, and M. subhyalinus.

Let us focus on Namibia, where the relationship of large-bodied, fungus-culturing termites to aridity and temperature is relatively easy to describe.

The biogeography of Macrotermes and Odontotermes in Namibia, proceeding from north to south, is as follows:

• Macrotermes is widespread (together with Odontotermes) north of Windhoek, except for the arid western regions,
• Macrotermes reaches the pro-Namib only in the northernmost area (Kaokoland), where M. subhyalinus builds structures up to 4.2 m high,
• Macrotermes michaelseni is widespread in the north of the country (north of 22 degrees South), building spires rather than mounds; these spires are tallest in the extreme north, where they reach up to 6 m high;
• Macrotermes vitrialatus occurs in places in northern Namibia;
• Macrotermes natalensis (which is particularly widespread in Africa) is the main species on the sandy substrates of the east-central part of the country, where it co-occurs with Odontotermes (https://www.inaturalist.org/observations?place_id=113055&taxon_id=492047&view=species),
• Macrotermes is absent from the area west and south of Windhoek, and the southwestern Kalahari, where Odontotermes remains present by virtue of its greater latitudinal tolerance, and
• this leaves about 40% of Namibia, in the west and particularly the south, devoid of fungus-culturing termites; here, the ecologically most powerful termites - none of which builds above-ground structures comparable with those of Macrotermes and Odontotermes - are Psammotermes (https://www.inaturalist.org/observations?place_id=any&taxon_id=643360&view=species), Hodotermes, Microhodotermes, and Trinervitermes (please see https://www.inaturalist.org/journal/milewski/80318-scarcity-of-termitaria-of-the-grass-harvesting-termite-trinervitermes-in-namibia-and-implications-for-the-aardwolf-proteles-cristatus#).

@tonyrebelo @jeremygilmore @botswanabugs @paradoxornithidae @matthewinabinett

In 2001, I was told the following by Elisabeth Straube, who had lived on a farm in the Khomas Hochland of Namibia, 135 km southwest of Windhoek (https://www.hakos-astrofarm.com/en/anreise/).

EQUUS HARTMANNAE

Over the years, Elisabeth had hand-reared various species of wild and domestic animals.

A neighbouring farm had a large population of Equus hartmannae (https://www.inaturalist.org/observations?taxon_id=129691). Intruders illegally killed a mother, orphaning its female infant.

When Elisabeth took this infant into her care, it was already in a thin and weak state. The milk of domestic Bos proved unsuitable, resulting in diarhhoea.

Despite time being against the survival of the infant, Elisabeth kept experimenting to find a solution. Based on advice from an elderly woman in the area, she followed local lore in giving the infant 4-5 tablespoons of dry, black tea-leaves. This did indeed stop the diarrhoea, buying some time.

Other farmers tipped off Elisabeth about the sweetness of equine milk compared to ruminant milk, so she resorted to klim milk (https://en.wikipedia.org/wiki/Klim_(powdered_milk)), and added fructose (fruit sugar). This she dispensed to the infant up to 14 times per day, in a wine bottle, solving the problem of surrogate lactation. Later, this infant would lick sugar from the sugar-pot on the tea table.

As the infant grew, it soon became jealous. It bit car tyres, and tried to bite humans. It would enter the house and jump between the chairs occupied by Elisabeth and others, as if to separate them. When still an infant (with long pelage on the belly), it tried to bite Elisabeth's husband.

This led to the infant/juvenile being banished to a property near Okahandja (https://en.wikipedia.org/wiki/Okahandja), at the age of about six months.

Several months later, Elisabeth visited this individual, which was locked in a kraal. The zebra recognised Elisabeth's voice before seeing her, and cried out immediately. It then accepted Elisabeth in reunion, despite the elapsed interval. Elisabeth was told by experienced locals that E. hartmannae will remain emotionally attached to a foster-mother for the rest of its life - even if this was a man, not a woman.

I infer that this reflects both the strength of maternal imprintation in Equus and the particular social structure of E. hartmannae.

ANTIDORCAS MARSUPIALIS

Antidorcas marsupialis has a reputation - somewhat paradoxical in view of its common name - of being unwilling to jump over fences, preferring to crawl under them.

However, Elisabeth observed the carcase of an individual of A.marsupialis that had tried to jump a fence. It had failed to clear the top wires, so that its hind hooves got caught. The animal died - in the way more familiar for Strepsiceros strepsiceros - hanging upside down with the tips of its fore hooves just above the ground (for a photo of a similar casualty, please scroll in https://pronamib.org/).

MADOQUA DAMARENSIS and RAPHICERUS CAMPESTRIS

Elisabeth hand-reared an individual of Madoqua damarensis. This animal covered its faeces, as does the domestic cat (Felis catus). It dug a hole with its hooves, then squatted over the hole to defaecate, then used its hooves again, to scrape earth over the faeces.

Elisabeth also hand-reared an individual of Raphicerus campestris steinhardti. She found that this species did not behave in a similar way.

STREPSICEROS STREPSICEROS

Elisabeth told me that, during a lean period, she observed that Strepsiceros strepsiceros zambesiensis on the farm were weak, and seemed incapable of jumping fences in the usual way. Several individuals were found to have failed to clear the top wires, and died hanging on the fence from their hind hooves. Under these conditions, Elisabeth found that the meat of culled individuals was dark red and, somewhat paradoxically, jelly-like yet not softened by cooking.

During the same period, when foliage was scarce, she observed that S. s. zambesiensis resorted to eating the spines and prickles - which are normally avoided by delicate movements of the lips and tongue during foraging - of shrubs such as acacias.

Elisabeth noticed this when she tried to prepare the tongues of the carcases for the table. The tongues - although appearing fairly normal on the surface - proved to be unfit for consumption, because numerous spines were lodged inside them.

These lodged spines and spine-tips included

• large and small spines,
• straight spines and prickles (short, curved thorns such as those of Senegalia mellifera (https://www.inaturalist.org/taxa/527007-Senegalia-mellifera), and
• both Vachellia (pale, straight spines) and Prosopis glandulosa (https://www.inaturalist.org/taxa/58160-Prosopis-glandulosa).

This observation indicates that

• when starving, S. s. zambesiensis can endure the pain of a riddling of the tongue with embedded spines, in an attempt to survive on the leafless twigs, and
• this does not result in so much inflammation that the tongue becomes obviously swollen.

SUS SCROFA

Elisabeth also hand-reared an individual of the domestic pig (Sus scrofa), and found it to be so intelligent and responsive that it made a particularly rewarding pet.

I find this surprising, for the following reasons.

Most mammals subjected to domestication have become decephalised in the process (https://www.inaturalist.org/journal/milewski/67392-decephalisation-in-domestication-part-1#), but this is particularly so for Sus.

Furthermore, there is little about the niche of the wild ancestor that would suggest an evolutionary pressure for intelligence exceeding that general for artiodactyls. If anything, one would expect minimal intelligence in Sus relative to other artiodactyls, because of the extreme fecundity of this genus.

Homo and Sus are in various ways biologically similar enough (omnivory, relatively pale muscle-tissue, sundry physiological processes) for Sus to be a suitable model in laboratory work in medicine. However, Homo and Sus could hardly differ more in pace of life (particularly in rates of growth and reproduction). This has been the basis for an extremely successful 'mutualism' in domestication, in which the gulf between Homo and Sus has been further widened by selective breeding in which fecundity has increased, and braininess has decreased, in Sus.

Thus, to find that, despite the conceptual framework outlined above, the domestic pig remains more intelligent than most artiodactyls, is deeply puzzling.

@tonyrebelo @jeremygilmore @botswanabugs @paradoxornithidae @ish_crew @matthewinabinett @chris_whitehouse @karoopixie

On 14 August 2001, I visited Kosierskraal Game Farm, southwest of Bredasdorp (https://en.wikipedia.org/wiki/Bredasdorp) in Western Cape province, South Africa. The precise location is shown in https://www.inaturalist.org/observations/101930627.

My guide was the owner, Mick J D'alton.

The location is on the Agulhas Plain (http://www.maphill.com/south-africa/western-cape/swellendam/bredasdorp/maps/physical-map/), extending slightly on to the lower slopes of the 432 m-high hill Soetmuisberg (https://mapcarta.com/19068450 and https://za.geoview.info/soetmuisberg,3361386), which forms a northern boundary to the Agulhas Plain.

The area of the property was 750 ha, of which 250 ha bad been cleared of natural vegetation. 550 ha was game-fenced, of which 250 ha was still natural fynbos (Elim Flats dwarf fynbos, still in good condition, although rather grassy). An additional 250 ha, also game-fenced was being leased on an adjacent property.

There was an additional area (200 ha) of natural vegetation - belonging to a third landowner - on a slope adjacent to Kosierskraal, where Leucadendron platyspermum (https://www.inaturalist.org/taxa/589081-Leucadendron-platyspermum) was being commercially picked.

The total area for wild ungulates at Kosierskraal was 800 ha, of which about 500 ha retained fynbos vegetation.

The substrates are mainly sandstone and gravel. Ferricrete (https://en.wikipedia.org/wiki/Ferricrete) was visible as a heap where earthworks had occurred.

VEGETATION

I visited a population of Protea pudens (https://www.inaturalist.org/taxa/592534-Protea-pudens). This threatened species regenerates germinatively. The population in question had been subject to foraging by the common eland, the black wildebeest, and even the springbok (which requires short vegetation).

I also visited an area of Elim Flats dwarf fynbos, last burnt about 7 years before, that lacked proteas and was characterised mainly by Restionaceae.

I noted Erica regia (dark pink) https://www.inaturalist.org/taxa/570046-Erica-regia

I also noted Erica cerinthoides, scattered but conspicuous, in fynbos still short, having been burnt on 23 December 1999 (I observed Nectarinia famosa in this area, https://www.inaturalist.org/taxa/13300-Nectarinia-famosa).

Mick D'alton pointed out that

• Asparagus africanus or a lookalike (https://www.inaturalist.org/observations?place_id=48632&taxon_id=60271&view=species), although spinescent, was untouched by any herbivores, and
• Carpobrotus acinaciformis, although indigenous, acted as a weed at Kosierskraal.

Parts of the formerly cultivated lands were now covered with tussock grassland of Eragrostis (?curvula, https://www.inaturalist.org/taxa/76850-Eragrostis-curvula), which I assume to be a fairly palatable grass, although not lawn-forming.

I observed the introduced species, Eucalyptus luehmanniana (https://www.inaturalist.org/taxa/883158-Eucalyptus-luehmanniana), killed by fire, but with seedlings apparent.

UNGULATES

Bontebok (Damaliscus pygargus pygargus)

Although fully indigenous to Kosierskraal, the bontebok fared poorly here. A population of 45 individuals declined to 15, and never recovered.

Black wildebeest (Connochaetes gnou)

Body size:
The largest male individual probably weighed >150 kg; its carcase weight was recorded as 80 kg. However, mean carcase weights here averaged: males 75 kg, females 60-65 kg.

I observed excavations made by the black wildebeest in fynbos vegetation, where the substrate was sandy.

Common eland (Taurotragus oryx oryx, https://www.inaturalist.org/observations/115898959)

Body size:
A large male individual about 8 years old, killed in 2001, weighed 800 kg, with a carcase weight of 360 kg. This was considered average for this area. At 'Armskor' (https://www.saairforce.co.za/the-airforce/bases/8/air-force-base-overberg), elsewhere on the Agulhas Plain, carcase weight reached up to 400 kg.

Reintroduction at Kosierskraal began with 1 individual from Salmonsdam Nature Reserve (https://en.wikipedia.org/wiki/Salmonsdam_Nature_Reserve), followed by 5 from De Hoop Nature Reserve(https://en.wikipedia.org/wiki/De_Hoop_Nature_Reserve), totalling a founder population of 6. By 1990, these had increased to 75, with 25 having been culled. Remarkably, the population increased from 6 to 100 in one decade.

In the early years, some of the female individuals bred continuously, giving birth every 10 months. Births were mainly in September (spring) and March (autumn). Males remained with the females when the latter give birth.

Females stopped breeding at about 13 years old, which is the same as for the European bost (see below).

During my visit, I observed an individual of the common eland that was unusually dark. Mick D'alton explained that one of the females at Kosierskraal, then 12 years old and near the end of its reproductive life, was 'black' (melanistic), this being inherited by some of her progeny.

In the drought of 1991, 50 individuals out of 75 died.

More recently, the population had been reduced to 13. At the time of my visit, this had increased to 35.

Wildfire swept through Kosierskraal in December 1999, followed by good rain in early 2000. The common eland (like the black wildebeest) then spent most of the time in post-fire fynbos.

It is well-known that adult males of the common eland produce a clicking sound from the carpal joint when walking. However, I learned that adult females and juvenile males also produce this sound, albeit softly.

Mick D'alton told me of a clear record of the common eland here eating the young inflorescences of cultivated (and fertilised) Protea compacta (https://www.inaturalist.org/taxa/320042-Protea-compacta). The property was Waterford (https://za.geoview.info/waterford,3359604), near Hagel Kraal (https://mapcarta.com/19079848), owned by Pietman Cilliers but subsequently sold to Eskom (https://en.wikipedia.org/wiki/Eskom). Here, a population of 15 of the common eland had been kept together with stands of P. compacta, planted for commercial purposes. The stomach of a culled individual contained many young inflorescences of P. compacta. Pietman Cilliers also directly observed the common eland eating the young inflorescences wholesale.

Mick D'alton told me that, at Kosierskraal, the common eland ate Carpobrotus in summer. It also sometimes uprooted Carpobrotus with its horns.

I observed a small group of the common eland in a stand of Acacia saligna, which was tall but thinned-out, and half-broken in the case of saplings 2-3 m high. I asked Mick D'alton if he had seem the common eland breaking A. saligna with its horns, and be said no. However, he told me that branches are broken somehow in the process of the young, green pods being eaten. I.e. the common eland forages roughly on A. saligna, and its attentions do not seem to benefit this plant species, but it seems incapable of breaking down and destroying a stand of A. saligna.

I observed a group of the common eland on a patch of pasture, with Cynodon dactylon. This group included a creche of juveniles (close aggregation of juveniles is typical of this species).

In the experience of Mick D'alton, the common eland dislikes being approached in dense vegetation; it flees into open vegetation, where habituation to human proximity resumes.

Springbok (Antidorcas marsupialis)

This species, although not indigenous to this area, fared well enough on grassy pasture and burnt fynbos at Kosierskraal.

Mick D'alton told me that he had a serious problem with predation on the springbok by the caracal (Caracal caracal caracal, https://www.inaturalist.org/taxa/42042-Caracal-caracal and https://www.inaturalist.org/observations/80090747).

Grey rhebok (Pelea capreolus, https://www.inaturalist.org/taxa/42336-Pelea-capreolus)

The population of this species was only 3 individual at Kosierskraal at the time of my visit; it had been hard-hit by the caracal.

On the mantlepiece, Mick D'alton displayed a noteworthy specimen of the hooves, 10 cm long, presumably from an old female individual, which I infer to have been kept captive on soft ground. Apparently, all four feet were distorted in this way. This shows how rapidly the claws of this species, adapted to rocky, abrasive terrain, grow in compensation for natural wear.

Southern bushbuck (Tragelaphus sylvaticus sylvaticus)

This species was absent from Kosierskraal. However, Mick D'alton told me that it was fairly common in the thickets of introduced Acacia at Pearly Beach (https://en.wikipedia.org/wiki/Pearly_Beach).

Cape duiker (Sylvicapra grimmia grimmia)

This species was fairly common at Kosierskraal.

Cape grysbok (Raphicerus melanotis)

This species was fairly common at Kosierskraal at the time, but was said to have been more common in the past.

Steenbok (Raphicerus campestris campestris)

This species was scarce at Kosierskraal at the time of my visit. Mick D'alton explained that the habitat was suitable, and the steenbok continued to be seen frequently outside of the game paddock. His view was that competition for food by the springbok had locally usurped the niche of the steenbok.

Farmers hereabouts believed that the caracal had greatly depleted the small wild ruminants, which it prefers as prey over the domestic sheep (Ovis aries). However, it was also true that the domestic sheep had reduced the cover required by the small ruminants.

Fallow deer (Dama dama)

Mick D'alton told me that this species thrives in this area, going feral, and becoming elusive, secretive, and self-sufficient; it hides and forages in the local thickets of Acacia saligna. He emphasised that the fallow deer was as successful as the indigenous ruminants, given the availability of non-indigenous thickets.

Thickets of Australian spp. of Acacia had also favoured the southern bushbuck. I infer that any given patch of non-indigenous thicket was utilised by one or another of these ruminants, but not both.

Domestic goat (Capra hircus)

At Kosierskraal, this species

• ate the mature foliage of Acacia saligna, particularly if cut down,
• did not avidly eat the seedlings of A. saligna, which seem to be heavily defended chemically,
• did not seem to eat Carpobrotus, which increased in a paddock devoted to the domestic goat.

Mick D'alton also told me that, in another paddock inhabited by the domestic goat at a low-lying position topographically, Leucadendron spp. (including Leucadendron linifolium, https://www.inaturalist.org/taxa/589072-Leucadendron-linifolium) seemed to survive heavy browsing.

He also told me of an experiment to see if the domestic goat could control invasive stands of Acacia saligna. This was conducted on the farm Tripolanda Trust. However, the experiment failed, in the sense that, in the paddock concerned, the fynbos was damaged to the same extent and degree as the introduced acacia.

Domestic sheep (Ovis aries)

This species, although kept elsewhere on the Agulhas Plain, was absent at Kosierskraal.

European bost (Bos taurus)

This species fared well enough at Kosierskraal, normally reproducing every year up to 10 years old. Females produced their first infant in their third year, after mating during their second year.

Culling was applied once an age of 10 years was reached. After this, production was subeconomic.

At Kosierskraal, the European bost

• routinely ate fuzzy, low restios,
• ate Carpobrotus in summer (like the common eland),
• nibbled at juveniles of A. saligna; I watched an infant browsing a sapling of this species, about 1 m high, and
• preferred to eat the foliage of A. saligna one day after this was cut down, presumably because this reduces the secondary compounds.

MANAGEMENT OF UNGULATES

Lightning was the confirmed cause of wildfire hereabouts, in the summer of 1999/2000.

Mick D'alton, in an earlier conversation with me 16 years previously (in 1985) reported the following birthing times at Kosierskraal:

black wildebeest December (summer)
bontebok September (spring)
common eland mainly in spring, but year-round to some extent
springbok year-round with some concentration in autumn and spring

Nutrient deficiencies:

Copper-deficiency was known to be a regional problem for ungulates. Therefore, Mick D'alton put out a copper-based lick block.

However, none of the ungulates ever accepted any lick block.

Mick D'alton recorded no geophagy (https://en.wikipedia.org/wiki/Geophagia) here, but osteophagy (https://en.wikipedia.org/wiki/Osteophagy) was frequently recorded.

Drought:

A drought had occurred 10 years previously, in 1991. So little grass was available that the European bost could not survive. However, no food supplements whatsoever were given to any of the ungulate species.

Both the black wildebeest and the common eland lost condition, despite being given sufficient water. Both were foraging mainly in the same areas (with diets consisting partly of fynbos plants), as opposed to being segregated.

However, the black wildebeest survived the drought with few losses (only 2 of a population of 40, compared to 50 of 75 for the common eland, and 30 of 45 for the bontebok).

Most individuals of the common eland died in the drought, despite having access to Acacia saligna. Contrary to expectations based on indigenous status, the black wildebeest fared better during the drought than did the common eland.

Both species became susceptible to parasites under stress; the black wildebeest was not necessarily more resistant than the common eland to parasites.

The bontebok, too, fared poorly during this drought. It suffered from parasites.

OTHER ANIMALS

Termite-eating mammals:

The bat-eared fox (Otocyon megalotis) had recently become common hereabouts.

The aardwolf (Proteles cristatus, https://www.inaturalist.org/taxa/1306005-Proteles-cristatus) had been present about 8 years previously. Mick D'alton recorded three individuals (an adult plus two juveniles) as road-killed simultaneously on a road adjacent to his property. By the time of my visit, this species seemed to have disappeared from the area.

However, as I write, I note that there is a recent observation of a road-killed specimen in the immediate vicinity of Kosierskraal (https://www.inaturalist.org/observations/79498114).

Mick D'alton reported having previously observed a road-killed specimen of the aardwolf at Akkedisberg Pass (https://en.wikipedia.org/wiki/Akkedisberg_Pass).

Birds:

Ostrich (Struthio camelus)

The ostrich was absent from Kosierskraal. However, I note that this species is present today, in this immediate area (https://www.inaturalist.org/observations/33745752 and https://www.inaturalist.org/observations/26587578).

Fork-tailed drongo (Dicrurus adsimilis, https://www.inaturalist.org/taxa/8268-Dicrurus-adsimilis)

I watched this species attending a group of 4 of the common eland, standing on the ground next to the hooves, and flying up to land on the withers. Mick D'alton told me he had observed this species flying up between the legs to pick something off the skin, without perching. (The fork-tailed drongo seems to act partly as an oxpecker here, not just taking insects disturbed by the ungulate, but actually taking invertebrates associated with the animal, including directly off its body.) Then, as we watched, this happened in front of us.

Blue crane (Anthropoides paradiseus, https://www.inaturalist.org/taxa/144286-Anthropoides-paradiseus)

I observed this species on an extensive anthropogenic lawn, now dull green during a relatively dry spell, of Pennisetum clandestinum and Cynodon dactylon, maintained entirely by the European bost and the reintroduced ruminants.

Insects:

Snouted harvester termite (Trinervitermes trinervoides)

I observed large termitaria (1 m high) of this species of grass-harvesting termite in Elim Flats dwarf fynbos, including in the vicinity of Protea repens (https://www.inaturalist.org/taxa/355849-Protea-repens and https://www.inaturalist.org/observations/141358640). At the time, I wondered whether it harvests mainly Restionaceae in fynbos, and today I still do not know.

Mick D'alton told me that fires in Elim Flats dwarf fynbos are intense enough - despite the low stature of this type of fynbos - that the termitaria of T. trinervoides subsequently collapse. I noted that a collapsing, fragile termitarium consisted of sandy clay loam, soft and brittle.

He also mentioned that the domestic bost had a habit of rubbing on the termitaria, thus destroying them. Removal of this species of livestock from a paddock allowed the number of termitaria to increase.

I observed an old, abandoned termitarium of T. trinervoides. Excavating this, I found large, robust roots of Acacia saligna reaching up through it. The accompanying odour was strong, resembling cut onions.

DISCUSSION

I find scant reference to Kosierskraal on the Web today, suggesting that the game farm is now defunct, at least as a tourist-attraction.

However, it is now one of the sites for the quagga re-breeding project (https://www.quaggaproject.org/ and https://www.inaturalist.org/observations/113812038 and https://www.inaturalist.org/observations/115584188). This is confirmed by https://www.inaturalist.org/observations/101930627), despite the misspelling 'Coetzeekraal'.

The special interest of this region, for me, was the combination of fynbos (which is regarded as unpalatable and unproductive for large animals) and the incidence of considerable populations of wild ungulates in prehistoric times on the Agulhas Plain.

I was surprised that the ungulates at Kosierskraal showed no interest in supplementary nutrients, in the form of lick-blocks.

I was also surprised that

• the black wildebeest fared as well as it did, given that the Agulhas Plain is far from its recent distribution, and
• the bontebok fared poorly, despite this being its typical habitat.

The black wildebeest at Kosierskraal foraged mainly on previously cultivated land.

I infer the possibility that these two alcelaphin grazers competed with each other, to the detriment of the bontebok. I suspect that, if Mick D'alton had not introduced the black wildebeest, the bontebok might have fared well at Kosierskraal.

It is clear that the common eland fares well on the coastal plains of the southwestern Cape, even when its original seasonal movements are no longer possible. Furthermore, it grows to its full genetic potential here, in contrast to populations conserved in the Drakensberg (https://hikingthedrakensberg.blogspot.com/2017/09/wildlife-of-drakensberg-story-of-eland.html and https://en.wikipedia.org/wiki/Drakensberg), where stunted-looking individuals have been typical (https://www.inaturalist.org/observations/64785188 and https://www.inaturalist.org/observations/10007389 and https://www.inaturalist.org/observations/11332783 and https://www.inaturalist.org/observations/92906834).

In previous Posts, I have pointed out that fynbos (South Africa), although ostensibly the ecological counterpart of kwongan (Australia), is considerably more fertile.

One of the problems with intercontinental comparisons of this kind is that the biotas are so different that it is hard to disentangle phylogenetic constraints from ecological adaptations.

For example, the marsupial Notamacropus irma (https://www.inaturalist.org/taxa/1453435-Notamacropus-irma) metabolises, grows, and reproduces more slowly than its ostensible counterpart, the bambi Raphicerus melanotis (https://www.inaturalist.org/taxa/42376-Raphicerus-melanotis). Is this really because kwongan is nutrient-poorer, and more prone to wildfire, than fynbos, or is it merely because two fundamentally different clades of mammals are represented?

There are, however, a few genera of plants and animals are shared between these two continents, and these provide an opportunity for relatively straightforward comparison.

In the case of birds, the shared genera are mainly raptors and owls. This limits their relevance, because their position at the top of food-pyramids means that they have little interaction with soils and plants.

This reduces the relevant genera to Turnix (Turnicidae), Coturnix (Phasianidae), Zosterops (Zosteropidae), and Mirafra (Alaudidae).

https://www.inaturalist.org/taxa/7318-Mirafra-javanica

https://www.inaturalist.org/taxa/204543-Mirafra-apiata

@tonyrebelo @jeremygilmore @kevinatbrakputs @craigpeter @jrebman @pietermier @botaneek @mdevill @koosretief

On 21-22 May 2006, a colleague and I visited Kleinzee (https://en.wikipedia.org/wiki/Kleinzee) in Little Namaqualand (https://en.wikipedia.org/wiki/Namaqualand) in Northern Cape province of South Africa.

Today, I dusted off my field-notes from that visit.

Little Namaqualand falls within the succulent karoo biome (https://pza.sanbi.org/vegetation/succulent-karoo-biome).

Approaching Kleinzee from the northeast:

At the turnoff to Kleinzee from the Steinkopf-Port Nolloth road (https://en.wikipedia.org/wiki/R382_(South_Africa)), we found a vast vista of blanket-like vegetation consisting of low (0.5 m), succulent shrubs (https://www.inaturalist.org/observations/162520809), on flats, slopes, and drainage lines alike (https://upload.wikimedia.org/wikipedia/commons/a/a5/Houthoop%2C_Kleinzee%2C_Namaqualand%2C_Northern_Cape%2C_South_Africa_%2810465408374%29.jpg and https://www.flickr.com/photos/south-african-tourism/10993794574/in/photostream/ and https://www.flickr.com/photos/south-african-tourism/10993845063/in/photostream/ and similar to https://www.inaturalist.org/observations/11048822 and https://www.inaturalist.org/observations/120312638 and https://www.inaturalist.org/observations/58090550).

This landscape is completely exempt from wildfire, and the vegetation had not burned for at least half a century, for hundreds of thousands of hectares.

What struck me as noteworthy about this vegetation was that it was both shorter and denser than expected for a semi-desert.

Both heuweltjies (https://en.wikipedia.org/wiki/Heuweltjie) and the termitaria of Trinervitermes (https://en.wikipedia.org/wiki/Trinervitermes) were absent.

Houthoop Guest Farm (https://www.flickr.com/photos/south-african-tourism/10993785134/in/photostream/) got its name (English translation: wood pile) from an interesting circumstance in terms of the regional vegetation. Trees and even tall shrubs being absent as far as the eye can see, any incidence of wood much larger than gnarled twigs is exceptional in this region.

A major family in succulent karoo vegetation is Aizoaceae (https://en.wikipedia.org/wiki/Aizoaceae), the plants of which are usually small. However, the genus Stoeberia (https://www.inaturalist.org/observations?place_id=any&taxon_id=575290&view=species) - although not arborescent - is large and woody enough to be remarkably solid in the local context. Simply collecting the occasional dead plant of this genus, the landowner here eventually accumulated a pile (?several hundred kg) of wood, which became noteworthy enough to form the initial basis for a tourist-attraction.

Although I recall seeing this pile during our visit in 2006, I have found no photos of it on the Web, suggesting that it has not proven to be durable or renewable.

I know of nowhere else on Earth where the vegetation is

• perennial and technically woody (as opposed to herbaceous), and dense enough to appear to blanket the landscape, yet
• so low that dead wood is so hard to obtain that the collection of even of a few hundred kg is remarkable enough to become an anthropogenic landmark.

Herbivory/folivory in succulent karoo:

The main herbivores/folivores in this succulent karoo ecosystem northeast of Kleinzee at the time were Raphicerus campestris (https://www.inaturalist.org/observations/144416187) and, probably, Microhodotermes viator (https://www.inaturalist.org/taxa/568761-Microhodotermes-viator).

I observed the former (of which extremely dense populations have been reported near the coast hereabouts), but not the latter.

Two other indigenous species with similar diets, viz. Antidorcas marsupialis (https://www.inaturalist.org/observations/135399878) and Struthio camelus (https://www.inaturalist.org/observations/10816345 and https://www.inaturalist.org/observations?taxon_id=154483), were relatively scarce over much of the area owing to management by pastoralists.

(Driving south from Kleinzee subsequently, over a distance of about 50 km, I observed both A. marsupialis and S. camelus, but not R. campestris. This suggests competitive release, in which the bambi (body mass 10 kg) compensates for the scarcity of the gazelle (body mass 30 kg) and/or the large bird (body mass 80 kg).)

I encountered vigorous seasonal growth of Mesembryanthemum crystallinum (https://www.inaturalist.org/observations/163481201 and https://www.inaturalist.org/taxa/49319-Mesembryanthemum-crystallinum) on an old track.

I tried it as food, and found it remarkably palatable, raw. It was extremely succulent and tender, with a content of probably >95% water in the leaves and shoots. It lacked the taste I associate with oxalates in e.g. Tetragonia and domestic spinaches, and was only slightly salty and only slightly sour. I describe it, in my field notebook, as 'basically salad-flavoured green water, but very close indeed to a table-standard palatable wild plant'.

Farming of domestic sheep (Ovis aries):

The domestic sheep prefers grass as its staple diet, and cannot thrive in succulent karoo - in which the small shrubs virtually exclude grass - year-round.

Therefore, the pastoral practice in this region is for the livestock to be moved to Bushmanland (https://www.researchgate.net/figure/Northwestern-part-of-Northern-Cape-showing-the-area-that-we-consider-to-fall-within_fig1_336835392 and https://en.wikipedia.org/wiki/Bushmanland,_Northern_Cape), with its relatively grassy semi-desert vegetation (https://www.inaturalist.org/observations/150480278), for the summer, and then returned to Little Namaqualand for the winter.

It is noteworthy that the herbivorous termite in Bushmanland is not M. viator but instead Hodotermes mossambicus (https://www.inaturalist.org/taxa/558312-Hodotermes-mossambicus), which eats grass rather than shrubs.

Thus, the livestock spend winter in the habitat of M. viator, sharing a mainly dicotyledonous diet with one harvesting termite (https://www.inaturalist.org/observations/132288477 and https://www.inaturalist.org/observations/138127670), and spend summer in the habitat of H. mossambicus, sharing a mainly monocotyledonous diet with another harvesting termite (https://www.inaturalist.org/observations/150534825 and https://www.inaturalist.org/observations/139114838).

In Kleinzee:

At the time of our visit, Kleinzee was a mining town run by De Beers (https://en.wikipedia.org/wiki/De_Beers) - which subsequently abandoned the enterprise hereabouts.

Our official guide, Paul Kruger, told me that, at the stable on his property, M. viator had a noteworthy pattern of behaviour. This termite suddenly emerged from holes in the ground, just before rain, fervently cutting and burying the hay provided for Equus caballus.

I found it puzzling that M. viator was so eager to harvest grass, given that

• this species has a staple diet of small shrubs, and
• grasses are so uncommon in succulent karoo as to be hardly noticeable, even in good seasons.

Unusually wet season:

The normally dry Buffels River, which runs through Kleinzee, was in spate during our visit.

I was told that this river had flowed strongly three times in 2005, most recently in October; the current event brought that total to four times in one year.

Indeed, I was told that

• it had been drizzling about every 10 days in Kleinzee, since April 2006, and
• only a few days before we arrived, there had been a truly exceptional rain event in Little Namaqualand, with 130 mm recorded at Springbok (https://en.wikipedia.org/wiki/Springbok,_South_Africa), and 12 mm recorded at Kleinzee.

For comparison:
Mean annual precipitation is 16.7 cm at Springbok (part of the watershed for the Buffels River), and only 1.8 cm (plus frequent fog) at Kleinzee (https://tcktcktck.org/south-africa/northern-cape/kleinsee#:~:text=The%20district's%20yearly%20temperature%20is,%25%20of%20the%20time)%20annually.).

The situation at the time of our visit was exceptional, because drainage lines generally flow strongly only about once every 7-10 years in Little Namaqualand.

Based on a rule-of-thumb that depth of penetration is about tenfold the precipitation, 130 mm of rain can be expected to percolate >1 m deep in the substrate, but 12 mm cannot be expected to percolate much deeper than 10 cm.

The dominant small, succulent shrubs in succulent karoo have shallow roots, adapted to precipitation so light that the subsoil remains dry for years on end. It is by virtue of the fact that the precipitation, although minimal, occurs fairly frequently (on average, 36.5 days per year excluding the advection fog from the shore that occurs mainly in autumn and early winter), that perennial plants occur here, with enough density to seem to carpet an arid landscape.