Far From Dead: The Surprising Biodiversity of Bare Ground

Occasionally, biodiversity might just be thriving beneath our feet.

bare hard ground at sunset
The deflation hollow which is home to so much diversity

The resilient Karoo landscape

During my explorations of the Wolwekraal Nature Reserve, in the arid Karoo region of South Africa, a seemingly desolate stretch of land consistently caught my attention. This area, characterised by hard, ancient sediment, served as a shortcut from the jeep tracks to a wild colony of honeybees nesting in an aardvark (antbear) burrow. The flora surrounding this bare ground is a stark reminder of the Karoo’s remarkable resilience, showcasing a rich tapestry of species that thrive in one of the world’s extreme climates. Over centuries, the vegetation here has adapted to withstand dramatic temperature fluctuations, from severe winter frosts to scorching summer heat at times exceeding 50°C, often enduring prolonged periods of drought.

Yet, amid this tenacity, certain areas of the landscape remain barren and hardened. Recently, I buried a wild hare—a tragic victim of roadkill—in an aardvark dugout; a deep, empty cavity, in an area of hard ground I could never have dug myself. My sister, with her characteristic humour, had remarked, “Everyone needs an aardvark.” As I reflected on the hare’s untimely death, I was once again captivated by the number of solitary bees nesting in holes on the inner edges of these burrows. Like the wild honeybees in these landscapes, many species are dependent on the :aardvark for their nest sites; a reminder of nature’s interconnectedness. All around the dugout vibrant yellow swathes of Gazania lichtensteinii were in flower, their annual beauty enhanced by early winter rains.

Top row left to right, solitary bees living on the edge of this aardvark dugout: Samba female, Colletes female, and a Patellapis female.

Misconceptions of bare ground

The seemingly lifeless stretch of ground, a wind-scoured deflation hollow, was located close to this dugout. Deflation hollows form where vegetation is lost allowing wind to remove sandy topsoil and expose a hard subsoil comprising desert dust cemented with calcium carbonate. They are often associated with stone age human settlements of hunter-gatherers and herders. At this particular deflation hollow, there are various stone tools made from chert including a stone arrowhead. Standing on this hardened ground I was struck by a common misconception: that bare earth signifies death. Often dismissed in environmental assessments, this apparently barren, hard ground was, in fact, teeming with life and intrigue. Initially mistaking the sounds I was hearing for a drone congregation area – where honeybee males dart through the sky waiting for a queen – I quickly realised that the sound was emanating from the ground. A closer look revealed a fascinating gathering of male Tetraloniella cf. brevikeraia bees. These short-horned longhorn solitary bees were eagerly vying for a chance to mate with a female as she emerged from her underground nest. Although solitary by nature—females work alone in building nests and provisioning food—these bees form dense aggregations in favourable environments. The apparent barrenness of the ground belied its role as a prime breeding ground, and I counted an astonishing 114 nests in the vicinity.

The author documenting the diversity on the deflation hollow; it is so unassumingly rich in life (photo: Collette Hurt)

The evolutionary journey of bees

The evolutionary journey of bees, stretching back around 100 million years, began with solitary, predatory mud-dauber wasps, coinciding with the rise of flowering plants. Today, bees exhibit remarkable diversity, particularly in arid regions such as the Karoo where solitary species thrive. They range in size from a mere 2mm to 39mm and come in various forms, from densely hairy to smooth and shiny, often adorned with striking colours, masks, stripes and patterns. Most species of solitary bees prefer to nest in the ground, often using plant materials or resin to line their nests. On this hard bare ground, the thriving community of Tetraloniella served as a vivid testament to the vibrant life hidden beneath the surface.

Dynamics on the deflation hollow

The deflation hollow measured 24 m by 13 m, with nests concentrated in a mere twelve square meters. The spacing of the nests varied, with some holes nearly touching while others were separated by more than 20 cm. The solitary male bees, have one primary role: to mate. Males are remarkably specialised, with their entire existence revolving around this singular task. To prepare for mating in the earlier hours of the day with temperatures in the lower teens, the males press their bodies against the sun-warmed sand. In September, when daytime temperatures can reach 28°C, the ground can heat up to a hot 44°C. Males dart close to the surface in circular and zigzag patterns, seeking the radiated warmth that boosts their speed. I counted 26 males on this day, but their numbers waned with each subsequent day. I could only imagine how busy this aggregation had been at the beginning of mating season. Many of the males were covered in bright yellow gazania pollen from hasty sips of nectar essential for their energy needs.

A small section of the Tetraloniella nest aggregation

Top row: Shortly after 9 a.m., the first male Tetraloniella bees make their appearance on the deflation hollow, braving air temperatures below 15°C. Seeking warmth, they press their bodies against the sun-soaked sand, where ground temperatures already reach the mid-20s. Middle + bottom row: Males congregating around the nest holes, waiting for the females to emerge

Coexisting species: The diversity of life

In addition to the Tetraloniella bees, there were other bees thriving in this environment. Among them were various species of leafcutter, dauber and mason bees (Megachilidae) that make their nests in pre-existing burrows. Initially chased by the males, they started inspecting holes abandoned after the females emerged. These bees possess unique nesting techniques and have the broadest range of nesting behaviours. Unlike honeybees that collect pollen on their hind legs, the Megachilidae collect pollen on tufts of red hairs on the underside of their abdomen, known as a scopa. When this is fully packed with pollen it is a bright patch of colour.

Leafcutter bees construct thimble-like cells lined with leaves or petals to protect their young from moisture and predators. For their nests they were using both leaves of krimpsiektebos (Lessertia annularis) as well as gazania petals. Both of these plants contain extremely bitter compounds that possibly serve to deter parasites and provide beneficial antimicrobial properties. Another of the females (Hoplitis sp.) used chewed, reddish-pink plant pulp together with a mouth secretion to line her burrow. The source of the plant was not established but was similar in colour to bellbush (Hermannia grandiflora) flowering nearby. Similarly diverse are the materials used for plugging the entrances, with some bees choosing a combination of leaf pieces and mud, while others used mud and stones. The collected pollens for the provision of larvae with food came from plants distinct from those used for nesting materials, possibly from honeybush (Melobium candicans) or the brightfig (Rushia bijliae), both in flower and in range of the nest site and on which the bees were sighted. Each female completed one burrow per day. 

Megachilid bees with their diverse nesting techniques, utilising burrows abandoned by Tetraloniella bees

Further careful study of the ground revealed a Camponotus rufopilosis (balbyter) ant carrying a dead conspecific. With mandibles featuring 5 to 7 teeth, these ants defend themselves by spraying formic acid when threatened. Amidst the male Tetraloniella bees, this ant dropped its prey and assumed a defensive stance. Larger, robust dauber bees (Megachile nasicornis) were also sighted, distinguished by their different, deeper  sounds, and by their striking, singular patches of black and white coloration. They were also on the lookout for abandoned Tetraloniella nests to use for their own reproduction.

Balbyter ants (Camponotus rufopilosis) and the larger, more robust, dauber bees (Megachile nasicornis) on the deflation hollow

While documenting these interactions, a brown-and-white striped fly (probably in the genus Parisus) scraped her abdomen along the ground and then hovered above several bee nest holes. This Bombyliidae fly is known to parasitise a range of insects including bees. Hovering above a nest, the fly deposited 33 eggs in less than a minute. To lay eggs in this way, some Bombyliidae have a chamber near the ovipositor filled with sand which they stick to the egg, giving it enough weight to shoot deeper into the host nest and helping to prevent the egg from drying out too much. This observation might represent a new host record, and underscores the intricate relationships between host and parasite.

Top row left to right: A Bombyliid fly filling her ovipositor with sand granules, then hovering over a nest burrow to lay eggs. Bottom row left to right: A male Tetraloniella bee investigating the fly; the fly shooting an egg into the burrow (spot the egg!)

A climax

The climax of my observations came when a chaotic scrum formed around a single nest hole, where male bees gathered in a frenzied attempt to mate with the emerging virgin female. In this state where the males were fixated on the hole and seemed vulnerable to predation, I wondered if the female released a pheromone to signal her arrival. As mating commenced, the male produced a sound known as “piping,” a result of the vibrations created by the wing muscles. The male, mounted on the female, used his antennae to possibly release a volatile pheromone, engaging in a behaviour known as antennal fanning. He fans his antennae near hers without direct contact. Research indicates that a courtship pheromone may exist in bees, which is believed to induce receptiveness in the female. Clasped tightly to her, other males attempted to dislodge him, displaying a complex mating struggle. Uniquely, the female grasped a red stone during copulation, remaining mostly still amidst the chaos around her. After more than 3 minutes of mating the primary male was dislodged, and the female executed an intense spinning motion to escape, eventually flying off to establish a new nest elsewhere. (To watch the incredible mating video, click here.)

Top: A scrum of males around the hole where the female is about to emerge. Top right and middle: a tussle ensues as one male battles another for the opportunity to mate with the female. Bottom: the mating pair.

I was unable to establish the mode of excavation of the nests, or whether this species use pre-existing nest cavities as the Megachilidae do, though did note that no turrets or soil mounds were present. Similarly, I could not locate the males at night, though these are thought to form sleeping clusters hanging from a branch.

A rich tapestry of creatures

While at the study site, I encountered numerous other creatures. Among them were a rock agama, Namaqua sand lizards, cryptic Sphingonotis grasshoppers, beetles, robber flies (Asilidae), and a wingless female mutillid wasp (velvet ant). This ant-mimicking wasp was intriguing as she used her abdomen to push aside stones to enter the nest of a solitary bee—they are parasites of the larvae of ground-nesting bees. I also heard barking geckos and, with much patience, managed to photograph one in its burrow. Overhead many kinds of birds flew by, including two pale chanting goshawks. Beyond this deflation hollow, I discovered an extraordinary mud nest in the shallow of a stone; a Chalicodoma mason bee builds her nest in a hollow on a rock, sealed over by sand cemented with a secretion from her mouth.

The hidden life beneath our feet

This study illuminated a critical lesson: even the most unassuming, barren stretches of land may be far from lifeless. They may harbour intricate ecosystems, teeming with life that defies initial perceptions. The conservation of these natural nesting habitats is crucial for the survival of solitary bees and other species. Therefore, these often-overlooked spaces must be included in environmental impact assessments (EIAs). Bare ground is too rich in life to be dismissed; recognising such ecosystems is essential for maintaining biodiversity and ecological resilience, while still allowing for erosion control and restoration efforts such as reseeding rehabilitation and replanting on damaged lands or vacant erven as a means to enhance ecosystem health. Remember, biodiversity might be thriving unseen beneath our feet.

From top to bottom: Wingless female mutillid wasp (in contrast, the males can fly) entering a nest burrow of a solitary bee; Sphingonotis grasshopper and a frantic surface beetle; leafcutter bee and a pale chanting goshawk; Chalicodoma bee and a barking gecko

ACKNOWLEDGEMENTS:

Dr Sue Milton-Dean of Wolwekraal Nature Reserve

Dr Connal Eardley and Dr Michael Kuhlmann for their help with bee identifications

Dr John Mark Midgely for his assistance with fly identification and behaviour

Prof Ben-Erik van Wyk for his ID of Lessertia annularis and his extensive knowledge of plant compounds

Dr Geoff Tribe and Collette Hurt for their assistance with other species and stone artefacts

REFERENCES:

Batra, S.W.T (1984) Solitary Bees, Scientific American Vol. 250, No. 2, pp. 120-127. Scientific American, a division of Nature America, Inc., 8 pgs

Fuchs, M., Kandel, A.W., Conard, N.J., Walker, S.J. and Felix-Henningsen, P. (2008).

Geoarchaeological and Chronostratigraphical Investigations of Open-Air Sites in the Geelbek Dunes, South Africa. Geoarchaeology.

Gess, S.K. and Gess, F.W. (2014). Wasps and bees in southern Africa, SANBI Biodiversity Series 24.

Michener, C.D. (2007). The Bees of the World, The Johns Hopkins University Press.

Packer, L. (2023). Bees of the World. A Guide to Every Family, Princeton University Press.

Romani, R., Isidoro, N., Riolo, P., and Bin, F. (2003). Antennal glands in male bees: structures for sexual communication by pheromones? Apidologie 34 (2003) 603-610.

Skaife, S.H. (1963). Strange Adventures among the insects, National Boekhandel.

Slingsby, P. (2017). Ants of Southern Africa. The ant book for all, Slingsby Maps.

skink in wild honeybee nest

The nest environment as more than just a dwelling for the honeybee

Burrowing mammals actively change their environment, and provide valuable microhabitats used by other species. These burrows offer other animals vital thermal refuges from temperature extremes and environmental fluctuations particularly in hot arid and semi-arid habitats.

wild honeybee nest in aardvark burrow

The architects of the burrow

All structures have stories to tell. At Wolwekraal Nature Reserve in the arid succulent Karoo region of South Africa, three wild honeybee nests were located in aardvark burrows. Aardvarks (Orycteropus afer) are extensive burrowers in sub-Saharan ecosystems, actively modifying their environment in the construction of shelters and in excavating termitaria for food. This, in turn, generates nest sites and unique habitats that support a variety of other species, amongst them also cavity-dwelling honeybees and the numerous other creatures that co-exist in a wild nest. This article looks at the ecology of the honeybee nest site, and the dependence on the honeybee for numerous creatures as a source of food.

Honeybees are cavity dwellers

Nest 1, an aardvark burrow with a colony under a sandy rim

Honeybees are cavity dwellers. At Wolwekraal trees are scarce and rock cavities are a rare sight. Dwellings, much like the food supply, are transient. Wolwekraal is situated in the  Karoo Basin which once formed a deep, inland sea. Through the erosion of the mountains to its south and of the landscape towards the north, a cycle of sedimentation began. Over hundreds of millions of years layers of sediment were deposited some 3km deep, and later tilted to form the seemingly endless horizons north of the village of Prince Albert. This area supports many different and unique plants, adapting to extreme climatic conditions together with their pollinators, including the honeybee. This is also where the aardvark roams. 

wild honeybee colony in aardvark burrow
Nest 2, a wild honeybee colony in an aardvark burrow

It has been suggested that the limiting factor on the number of wild honeybee colonies in any area is determined by the worst month when there is little forage. In this desert region, not only is forage challenging and limited in the hot summer months with very little other than the Vachellia karroo in flower, but also the availability of nest sites. The three wild honeybee nests located at Wolwekraal all lie along a single north flowing minor drainage line west of the Dorps river in aardvark burrows situated on deep alluvial sediments. Aardvarks avoid areas of solid or rocky ground, preferring a sandy, loose soil optimal for digging. Aardvarks are skilled burrowers, and their tunnels provide crucial thermal refuges in the extreme temperatures of the Karoo, where summer highs often exceed 40°C and rainfall is scarce. These burrows help maintain a more stable climate, with internal temperatures ranging from 25°C to 32°C, offering a respite for many species. For honeybees, whose ideal brood temperature is around 35°C, these burrows provide an ideal nesting environment. 35°C is also an ideal temperature for the moulding of wax comb by honeybees.

Nest 3, a small colony situated between two aardvark diggings that are connected

The importance of shade

One of the key benefits of these aardvark burrows is the shade they provide. The relentless heat of the Karoo can make the ground scorching—often 10°C to 20°C hotter than the air.  Even the sparse shade from the branches of the Vachellia karroo trees offer some relief from the heat. It is therefore not surprising to see a vast diversity of other creatures sharing the dwellings which honeybees inhabit. 

Co-habitants of the nest

The scavenging Tortoise Darkling Beetle (Epiphysa flavicollis) at the edge of the honeybee colony

Termites were the first co-habitants noted in the documented nest sites. Aardvarks, in excavating these cavities, were, of course burrowing for termites upon which they feed. Over the current study period of 12 weeks, only twice did it appear as though one of the three honeybee colonies had been attacked by termites. The skirmish with the guard bees was short-lived and within a day dead termites and honeybees were removed from the nest by worker bees and dropped at distances of 20m – 50m from the colony. In environments where food is limited, it does not take long for the dead to be predated on. Immediately Pheidole and other ants arrive on the scene to carry off the bodies, as do beetles such as the Eurychora sp., commonly known as mouldy beetles. Several mouldy beetles have been documented living side-by-side with honeybees in two of the three aardvark burrows. If bees detect incursions by chemical means, it would appear as if these beetles, by covering themselves with soil and nest debris and by rubbing soil particles on each other—as documented at one of the nests—would possibly use this form of camouflage to be able to move around the periphery of the nest largely unhindered by the honeybees. Soil and vegetative debris is adhered by bristles and waxy filaments on these beetles’ backs. Mouldy beetles are detritivores, feeding on dead plant and animal litter. Such detritus is an important component of food reserves in desert regions. As such, honeybee nests offer a constant concentration of detritus to support this population. Other beetles documented in these nests include the tortoise darkling beetle (Epiphysa flavicollis) which scavenge plant and animal material at night and live for many years. 

Mouldy beetles cover themselves with soil particles from their environment
mouldy beetle
A mouldy beetle feeding on organic matter close to the colony

The role of wax moths and other detritivores

Other detritus feeders are the larvae of the greater and lesser wax moth, Galleria mellonella and Achroia grisella respectively. These larvae are mostly tolerated by wild honeybees as they play an important role in maintaining nest hygiene. They help by cleaning out sections of old wax comb and are frequently found feeding on leaf litter, beeswax particles, and pollen present in the nest debris. 

On the Cape Peninsula an abandoned honeybee nest in a cavity situated under rock was documented. In the nest were numerous wax moths and the deserted wax comb was writhing with wax moth larvae. In addition, two shrews made their home in the nest, benefiting from the plentiful food source provided by both the larvae and the moths. Once the larvae have cleaned the comb and both they and the shrews have departed, the nest becomes ready for re-occupation, illustrating the ingenious cycle of nature. The role of the wax moth is thought to be that of a scavenger, helping therefore to control potential diseases much like hyenas do in their ecosystems. Interestingly, the moths and the larvae continue to play a role even while the bees occupy the nest.

Shrews on the Cape Peninsula feeding on wax moth larvae

A diverse community

Skinks, too, are found feeding on wax moth in honeybee nests. In addition, the variegated skink (Trachylepis variegata) was seen feeding on small invertebrates like ants and flies. Flies abound at every honeybee nest site. Rondanioestrus apivorous, a bee tachinid, lays eggs on bees in the air as they fly into their nest. The egg almost immediately hatches into a larva which feeds on the soft tissue of the honeybee until it is ready to pupate. The bee conopid, Physocephala fascipennis with its distinctive hourglass abdomen does the same. In addition, the Blue-green bottle fly of the Lucilla sp. is also a regular visitor to the nest as are the robber flies of the family Asilidae.

A robber fly having caught a honeybee
The variegated skink feeds on small invertebrates like ants and flies, and is mostly tolerated by the honeybees

When a small number of Banded bee pirate wasps (Palarus latifrons) were documented at two of the three nest sites, the wasp, upon catching a returning forager flying into the entrance of the burrow, would fly off in the direction of its nest followed by a scurry of small jackal flies. These kleptoparasitic milichiid flies (Desmometopa spp.) feed on :haemolymph rich in nutrients, and they typically steal food from other insects. They are thought to be attracted by a chemical pheromone given off by the injured or stressed honeybee, and they need to be fast to keep up with the flight of the much larger predatory wasp. Banded bee pirates are most active when temperatures are between 24C and 40C. Their diet consists largely of honeybees. The wasps compete with each other for prey, each vying to catch a honeybee, darting back and forth across the nest entrance and diving at each other. The captured honeybee is stung and paralysed by the female wasp and taken to its burrow where she lays an egg on the bee. The egg hatches into a larva and consumes the bee. Other wasps, such as the Bembix wasp, have also been noted flying higher above the nest site. Bembix hunt their prey on the wing, often catching forager bees flying off or returning to the nest. Bembix typically prey on flies of various families, however not exclusively, as seen at the wild honeybee nests and where honeybees are foraging on the sweet thorn (V. karroo).

Top row: Banded bee pirate wasp carrying off a honeybee caught on the fly. Bottom row: numerous fly species wait at the nest entrance to lay eggs on returning foragers

Ants are so important in the ecology of a wild honeybee nest and are ever present in the nest. They are constantly contributing to nest hygiene by carrying away dead bees and other organic matter including seeds blown into the burrows. Seeds are also carried out by the bees themselves, thus both ants and honeybees are vital to seed dispersal, with ants playing a much larger role. While ants are tiny, they have a really large ecological impact. In the wild certain species may be tolerated by honeybees and we have documented ant nests side by side with wild honeybee nests, separated by a good portion of propolis. As long as there is a healthy balance, ants and bees, which are related, both being of the order Hymenoptera, coexist in close proximity, as do the termites and the honeybees at the nest site at Wolwekraal.

A balbyter ant (Camponotus rufopilosis) carrying a dead honeybee into its nest

Spiders, too, find their place within these burrows. Some spiders tunnel into the walls, and Pholcid spiders (daddy-longlegs, cellar spiders) hang upside-down in their haphazard webs. They rely more on the irregular structure of the web to trap honeybees returning to the nest, rather than it having adhesive properties. Upon detecting struggling prey these spiders swiftly wrap their prey with silk-like material, either eating it immediately or storing it for later. Jumping spiders (Salticidae) do not spin webs, but are free-running and actively stalk any prey they detect, and are often found on the soil surface. They are known to have the sharpest vision of all spiders, along with tactile chemoreceptors on their pedipalps. They mostly specialise in preying on ants which are usually plentiful, but generally prey on a wide range of insects and other spiders, and prey may sometimes include a honeybee.

Honeybee followed by a jumping spider
Honeybee followed by a jumping spider
Wolf spider with many babies on its back
Wolf spider with many babies on her back in a tunnel in the burrow wall

Observing nature’s complexity

The diversity in these extreme environments is surprising. So often one is reminded of the extraordinary privilege of being able to still wander in such untouched spaces; or in areas that were once farmed but have been allowed to bounce back again boasting bygone beauty. This phenomenon is particularly evident in the case of wild honeybee colonies established within burrows, where the bees thrive abundantly and do not perceive humans as threats to their resources or existence. Instead, observers are presented with the unique opportunity to engage in passive observation, immersing themselves in an alternative ecosystem characterised by intricate ecological interconnections that often transcend human comprehension. It is as if one enters a state of mindfulness previously unexplored.

A mouldy beetle (centre) and daddy-longlegs (bottom right) in a wild honeybee nest

SELECTED REFERENCES

Begg, J. (2005). A Ramble through the Rocks of the Swartberg, Prince Albert (Kweekvallei), 88-96.

Dippenaar-Schoeman A.S., Van den Berg M.A. & Van den Berg A.M. (2001). Salticid spiders in macadamia orchards in the Mpumalanga Lowveld of South Africa (Arachnida: Araneae). African Plant Protection 7(1): 47–51.

Gess, S.K.& Gess, F.W. (2014). Wasps and bees in southern Africa. SANBI Biodiversity Series 24.

Haddad, C.R. & Dippenaar-Schoeman A.S. (2002). The influence of mound structure on the diversity of spiders (Araneae) inhabiting the abandoned mounds of the snouted harvest termite Trinervitermes trinervoides (Sjöstedt). Journal of Arachnology, August 2002.

Heiduk, A., Brake, I., Shuttleworth, A. and Johnson, S.D. (2023) “‘Bleeding’ flowers of Ceropegia gerrardii (Apocynaceae-Asclepiadoideae) mimic wounded insects to attract kleptoparasitic fly pollinators,” New Phytologist.

Henschel, J. R. (2021).Long-Term Population Dynamics of Namib Desert Tenebrionid Beetles Reveal Complex Relationships to Pulse-Reserve Conditions. Insects 2021 Sep; 12(9): 804.

Hepburn, H.R. (1986). Honeybees and Wax, An Experimental Natural History. Springer Verlag, 205 pgs.

Knöthig, J. (2005). Biology of the Aardvark (Orycteropus afer). Diplomarbeit, Fakultät für Biowissenschaften der Ruprecht-Karls-Universität Heidelberg.

Milton, S. J., Short, S., & Dean, W. R. J., 2022. Decline in whistling rat (Parotomys brantsii) density: Possible response to climate change in the Karoo, South Africa. African Journal of Ecology, 60, 969–979.

Pike DA, Mitchell JC, 2013. Burrow-dwelling ecosystem engineers provide thermal refugia throughout the landscape. Animal Conservation, 16, 694–703.

Tribe, G. (2009). Creatures inside the hive, Village Life No 34, Autumn 2009.

Tribe, G. (2021). Honey Mountain. Pinewood Studios, Cape Town. 216 pgs.

Weyer, N. M. (2018). Physiological flexibility of free-living Aardvarks (Orycteropus afer) in response to environmental fluctuations. Faculty of Health Sciences, University of the Witwatersrand, Johannesburg.

Leaf cutter bee on deflation hollow

About Bees, Naturally.

Should we keep bees?

Arid to semi-arid environments are surprisingly rich in bee diversity. When most people think of bees, they often picture the honeybee (Apis mellifera). However, the reality is far more complex. Globally, there are over 23,000 bee species, the majority of which are solitary rather than social, as seen in honeybee colonies. In these solitary species, each female constructs her own nest, gathers pollen and nectar to provision her young, and lays her eggs before her brief life of sometimes just 6 to 8 weeks comes to an end. Each season often hosts its own unique bee species. This contrasts with honeybees, which remain active throughout the year in most regions of South Africa. Many of the solitary bees have co-evolved together with the flora of each region. This intricate tapestry of adaptation and survival has unfolded over millennia, leading to a remarkable diversity that thrives even in the harsh climates of areas like the Karoo. It’s astonishing to think that such diversity exists in seemingly desolate landscapes. 

As I observe the bees at Wolwekraal Nature Reserve in the arid Karoo, I often feel a sense of urgency; there simply aren’t enough hours in the day to document the incredible ecology and behaviour of solitary bees currently emerging with the spring flowers. In SA we have over 1300 species of solitary bees, but many remain to be both discovered and described. So little is known about them. 

Wolwekraal is a paradise for bees, but bees don’t thrive in isolation. Their presence also transforms Wolwekraal into a haven for all life that relies on them. As examples of what exists, for the past three weeks I have been engrossed in a bare patch of hardened ground surrounded by vygies, daisies, and other typical Karoo flora. Currently annuals like Gazania lichtensteinii are flowering in swathes of yellow. At first glance this sandy deflation hollow appears desolate. But I have been incredulous at the life on it! It is home to a nest aggregation of solitary bees. I counted 114 nests in just twelve square metres. The males hatch first, engaging in frantic competition to mate with the newly emerging females. Once mated, she flies off in search of a new nest site and begins her foraging for pollen and nectar to start reproducing and to ensure the survival of future generations. 

The abandoned nests are then repurposed by other solitary bees, such as those in the Osmiine tribe, which includes leaf-cutting, mason, and resin bees. These female bees utilise diverse nesting techniques, often lining their burrows with remarkable reddish-pink pulp made from chewed plant material. Others use leaves and petals to construct thimble-like chambers. After foraging for nectar and pollen on flowers close by, each burrow is meticulously sealed with a mixture of sand, tiny stones or leaves, and a secretion from a special gland. Some even place stones on top for additional protection. Once her work is complete, the female dies, leaving her young to develop underground until they hatch in a year, if conditions are right. 

Leaf cutter bee returning with leaf material to her nest in the ground

However, the nests are not always safe from other threats. Wingless velvet ants (a kind of wasp) are able to push out the stones and lay eggs within. Moreover, bee-flies can parasitise open nests, further complicating the lives of solitary bees. Amidst this all, balbyter ants carry off any organic debris, frantic surface beetles and sand lizards scurry by, grasshoppers leap across the terrain, and robber flies, birds and barking geckos are all in the neighbourhood.

Beyond this deflation hollow, I discovered an extraordinary mud nest in the shallow of a stone. A bee known as Chalicodoma builds her nest in a hollow in a rock sealed over by sand cemented with a secretion from her mouth. And just a short distance away, along the ancient riverbank of the Dorps, exists another active bee nesting site with a multitude of bee nests; here root and burrow fossils create extraordinary patterns between nest entrance holes. This is merely a glimpse into the myriad stories waiting to be uncovered at Wolwekraal.

Many aspects in nature remain undocumented. For example, the fly I filmed parasitising the solitary bee nests may represent a new host record. Some solitary bee species are highly specialised, acting as the sole pollinators for specific flower species. This fragile interplay highlights the delicate nature of plant-pollinator relationships. Our landscapes are vastly under-documented, and with many species facing extinction due to habitat loss and human encroachment, we must act quickly to better understand and protect these vital ecosystems.

Wild honeybees are equally fascinating, and again so little is known about them. In most places honeybees have become a commodity and are typically associated with hives (and honey), and traded as pollination units. In South Africa, fortunately, more than 90% of honeybee colonies remain wild residing in natural habitats. Contrary to common belief, we have no shortage of bees. This natural abundance fosters adaptation in micro-habitats, and to changing environments and climates. In both our solitary bees and wild honeybees, SA has a huge natural pool of excellent pollinators. In Europe, most wild honeybee populations have been lost, with the majority of western European nations seeing almost total domestication and hybridisation of their once indigenous honeybee species. As a result, many colonies succumb annually to disease and other stressors. In contrast, wild honeybees in their natural cavities create habitats conducive to survival. Their use of abundant, local, site-specific plant resins, mostly in the form of propolis, and their interactions with many other creatures, organisms and microbes within the nest, is what keeps their environment healthy. 

Human activity can disrupt this delicate balance. Introducing hives to a new area can drastically alter the local ecosystem. A single hive can house up to 60,000 bees, all of which require substantial amounts of forage to thrive. The ecological consequences can be severe, especially when unaware of the existing species. Besides the risk of disease and microbial shifts, and the introduction of different strains of honeybees into an area, honeybees can outcompete solitary bees for nectar and pollen, further stressing already vulnerable populations. Wild honeybees living naturally in an area are equally affected by such increases in honeybee numbers. During periods of extreme heat or drought, wild honeybees can enter a state of dormancy, relying on stored honey to survive. It’s been suggested that the number of wild honeybee colonies in an area is limited by the worst month for forage availability. 

Nature possesses a remarkable intelligence and a fragile balance that we must strive to protect. We should be cautious not to unintentionally contribute to ecological loss through changes like hive introductions in finely tuned environments. Instead, by planting indigenous flowers and veld types and allowing some areas of our gardens to remain wild, we can invite nature in and help create vibrant ecosystems. Just as Wolwekraal serves as a unique ‘laboratory’ for observing nature, our gardens can also become spaces for wild bees and other creatures, revealing the wonders of biodiversity season after season, and allowing us all to become bee-keepers of a different kind. Who knows what bees we may yet discover. 

Honeybee collecting nectar from a milkweed plant in South Africa

Honeybees and Gomphocarpus filiformis (lammerlat): A source of propolis in the Great Karoo

Karoo landscape with Lammerlat (Gomphocarpus filiformis)

Wild honeybees and propolis

The wild honeybees of Southern Africa are renowned for their propensity in the use of propolis which enhances their survival in the wild. The vast amount of propolis collected and deposited in and around a wild colony attests to its value to the honeybees. But where do the bees collect the waxes, resins and plant exudates that form the basis of propolis, and in such large amounts? Individual honeybees are occasionally seen scraping waxes from buds of Protea species or resin oozing from a wound on the trunks of Vachellia karroo, but most sources of propolis are largely unknown. Although the topography and hence plant species composition may vary greatly between regions, the honeybees appear to have little difficulty in locating a source.

Recorded here is an observation of honeybees collecting the exudate from an oviposition puncture caused by a tephritid fruitfly whose larvae feed on the seeds of a plant widespread in the Karoo.

Extensive use of propolis in a wild honeybee nest
The extensive use of propolis in a wild honeybee nest.

Lammerlat (Gomphocarpus filiformis)

Gomphocarpus filiformis, a plant commonly known as lammerlat (an Afrikaans word, lammer = lambs, lat = stick, often used as a stick to chivy lambs when herding), is widespread across Namibia and the western arid interior of South Africa primarily in Desert, Nama-Karoo and Succulent Karoo habitats. Honeybees were first documented on lammerlat early in December 2023 at KRAAL, a garden in the Prince Albert municipality, where they were collecting resins on horn-like seed pods. The horns indicate that it belongs to the Asclepiadaceae into which family the stapeliads are prominent. There is a fruitfly species which lays its eggs in the stapelia pods where the fly larvae consume the seeds as they develop. A limited survey showed that 98% of the seeds on several Hoodia gordonii plants were entirely consumed (G. Tribe, unpub.). Was this happening on the horn-like seed pods of lammerlat?

Gomphocarpus filiformis (lammerlat), is found primarily in arid habitats.
Gomphocarpus filiformis (lammerlat), is found primarily in arid habitats.
A honeybee collecting resins on a seedpod of Gomphocarpus filiformis (lammerlat).
A honeybee collecting resins on a seedpod of Gomphocarpus filiformis (lammerlat).

Adaptive significance of plant resins and cardiac glycosides in defense and medicinal contexts

Resins are a defensive mechanism of plants and have a physical function (expelling an organism or encasing it) and a chemical antibiotic function (especially for fungi but also insects). For especially mammalian predators, there are heart poisons called cardiac glycosides (also in Erythrina seeds), hydrogen cyanide, etc. Yet porcupines feed without consequence on a wide variety of poisonous bulbs that are deadly to other animals. Nomadic peoples eat locusts or saturniid caterpillars by first removing the guts for they may contain poisonous plant material. Fireflies make cardiac glycosides and they have evolved warning colouration. Toads also possess such compounds. In very small doses these cardiac glycosides may stimulate the heart and are thus used in the treatment of congestive heart failure and arrhythmia, but at higher doses can cause havoc and death.

When injured, lammerlat oozes a milky sap-like resin, a latex, as a defensive response against pathogens. This sap contains cardiac glycosides that are likewise toxic to herbivorous animals. But some insects, like the monarch butterfly and locusts, incorporate it as a defence mechanism and thus become toxic to their predators (S. Milton-Dean, pers. com., Dec. 2023). Thus a defence strategy evolved by a plant to avoid being eaten can be co-opted by another organism so that it too can avoid being eaten. Most early medicines used by humans involved the same principle in plant-based medication when taken at the correct dosage rate.

Honeybee collecting milkweed resins.
When injured, lammerlat oozes a milky sap-like resin. Here a honeybee is collecting the resins.

Defensive resin response of Lammerlat seed pods to Tephritid flies

When its seed pods are punctured, lammerlat immediately exudes a resin to seal the pod and protect the seeds within. The puncture is usually a sign that eggs have been laid by the tephritid fly. Seed pods were collected and examined. The larvae pupate in the pod after completing feeding. At Wolwekraal Nature Reserve (113ha) on the outskirts of Prince Albert are several populations of lammerlat where honeybees were immediately observed on the lammerlat collecting resin, but there were also milkweed bugs Spilostethos, muscid flies, and allodapine bees at various locations on the plant. The allodapines collected mostly nectar from the flowers together with a male Coelioxys kleptoparasite bee.

Then a fly which initially looked very wasp-like was spotted on the seed pods. Sue Milton-Dean who offers guided walks on a trail through the nature reserve had studied tephritid flies in Pteronia seeds. One of the conclusions of this study was that flowering and number of seeds containing tephritid fly embryos, were positively correlated with annual rainfall. Sue had a keen eye for these flies. This tephritid fly which was later identified as most probably Dacus bistrigulatus, was laying an egg into the seed pods. We collected a number of seed pods with visible resin and later dissected them under a microscope. Inside each seed pod a larva was found which Sue recognised as a tephritid larva. In seed pods where all of the seeds had been eaten, the biggest mature larvae were found prior to pupation. 

A tephritid fly laying an egg into the seed pod.
A tephritid fly laying an egg into the seed pod.
Tephritid fly larva consumes the seeds as it develops.
The fly larva consumes the seeds as it develops.

Role of resins in honeybee nest construction

The bees collecting the exudate possibly originated from the two wild honeybee nests located in aardvark burrows on the reserve. At these nests honeybees were seen preparing the surface of the burrow for the application of propolis, enabling the colony to either increase the number and/or length of combs, thereby expanding the size of the colony. Once the surface had been cleaned by worker bees, resins were brought in by the resin-collecting foragers to be used as a sealant. Many of the resins used within a honeybee nest are the main components in propolis. These resins often have beeswax added to it to be more pliable, but can also consist of pure plant exudates in outer structures where it adheres tightly to the substrate, even if the substrate is partly loose.

A honeybee collecting lammerlat resins.
A honeybee collecting lammerlat resins.
The resins of Vachellia karroo being collected by a resin-collecting honeybee.
The resins of Vachellia karroo being collected by a resin-collecting honeybee.

Once the propolis is adhered to the substrate, comb building begins and the colony is able to expand to the outer edges of the adhered propolis. Elsewhere in the nest architecture, propolis may incorporate sand grains or plant material to strengthen it. Its thickness depends on where the propolis is being applied and its purpose. Resins play an important and essential role in honeybee nests, far beyond being a sealant. Being defensive chemical exudates of plants, propolis has many anti-bacterial and anti-fungal properties and is aromatic and health-giving with its essential oils. Lammerlat resin as propolis with its cardiac-glycosidic properties could perhaps be beneficial for honeybees in ways that are not yet known, possibly also as a defence mechanism against predators of honeybees, or ants, wax moth larvae and the like. An interesting aspect to consider would be whether the use of propolis for human consumption containing lammerlat resins could be toxic if taken in high doses.

Ecosystem dynamics of Lammerlat

Recently on one of Sue’s nature walks at Wolwekraal, both Rufous-eared warblers and the Karoo eremomelas were seen eating tephritid larvae on the lammerlat. It has also been found that the larvae of these flies are themselves parasitised by a small wasp. The ecological cycle around this arid shrubland species is both complex and remarkable.

A Rufous-eared warbler ((Malcorus pectoralis) photographed here on a Vachellia karroo tree.
A Rufous-eared warbler ((Malcorus pectoralis) photographed here on a Vachellia karroo tree.

SELECTED REFERENCES:

  1. Bruyns, P.V. (2002). The Stapeliads of Southern Africa and Madagascar Volume 1 pg 57.Umdaus Press, 329 pgs
  2. Ellis J.D. Jr and Hepburn, H.R. (2003). A note on mapping propolis deposits in Cape honeybee (Apis mellifera capensis) colonies. African Entomology 11(1): 122-124.
  3. Ghisalberti, E.L. (1979) Propolis: A review. Bee World 60(2):59-84.
  4. Iannuzzi, J. (1983). Propolis: The most mysterious hive element. American Bee Journal 123(8): 573-575.
  5. Iannuzzi, J. (1983). Propolis: The most mysterious hive element Part II – Conclusion. American Bee Journal 123(9): 631-633.
  6. Milton, S.J. (1995), Effects of rain, sheep and tephritid flies on seed production of two arid Karoo shrubs in South Africa, Journal of Applied Ecology 32, 137-144.
  7. Tribe G.D., Tautz J, Sternberg K and Cullinan, J. (2017). Firewalls in bee nests – survival value of propolis walls of wild Cape honeybee (Apis mellifera capensis). Sci. Nat. 104:29. DOI 10.1007/s00114-017-1449-5
  8. Whiteman, N. (2023), Most Delicious Poison, Little, Brown Spark.

The Historic Heuningklip sites at Kleinmond (Overberg) and Herbertsdale (Southern Cape)

“… we passed a large rock, which on account of its harbouring bees, has obtained the name Honingklip…”

There are two historic Heuningklip sites in the Western Cape – one near :Herbertsdale (inland from Mossel Bay) and the other along the coast near Kleinmond in the Overberg. Whenever ‘Heuning’ (‘honey’) is used as a name for various landmarks, it invariably indicates that wild honeybee colonies inhabit the site. They are usually found in deep inaccessible recesses in the rock, and are a permanent presence from year to year. However, names such as ‘Heuningvlei’ indicate good pasturage for domestic stock.

Kleinmond Heuningklip.

Heuningklip with its wild honeybee nests
Fig.1. The Heuningklip consists of a broken granite outcrop standing alone in the valley below the Perdeberg on the northern outskirts of Kleinmond.

There are almost no historical records about the Heuningklip near Kleinmond and this is probably because it was not on a major route travelled by the early pioneers. Nor was there a spring nearby to further enhance its attraction as an outspan venue, typically a place to unharness the oxen from the wagons and camp, although the nearby Isaacs stream flows from the mountains to the sea. Yet, from its position as an isolated chunk of broken granite, it remains an easily identifiable landmark [Fig. 1] and its name has remained as a feature on maps for hundreds of years. The Hottentot Chainouqua clan that originally occupied the Overberg was no doubt familiar with the presence of the nests in the rocks and possibly had raided them from time to time. The Witteberg peak (499m) of the Perdeberg Mountains lies immediately to the north of the valley in which the isolated Heuningklip occurs [Fig. 2]. On the western border of Kleinmond is the mouth of the Palmiet River (Houtema = ‘snake river’) which was not easily forded in the days of ox wagons. Today the Heuningklip remains in private hands and is not accessible to the general public.

'Heuningklip' literally means 'honey rock' and often indicates a wild honeybee nest
Fig.2. The southern aspect of the Heuningklip surrounded by Rockwood trees.

About 5km to the west is the Harold Porter Botanic Garden where a honeybee colony has been present from at least before 1980 until today – 45 years later. The exposed colony is high up under a small ledge on a smooth rock face opposite the waterfall [Fig.3a+b] but in an adjacent property. Here it is in an almost continuous shadow, dark and damp most of the year and the colony waxes and wanes to some extent but never appears to get any larger.

Four honeybee colonies were found in the crevices and cavities in the rocks that comprise the Heuningklip [Figs 4+5]. All the entrances to the nests were fully covered with propolis, leaving holes through which the foragers had access to the interior. Several beehives had been placed at the foot of the Heuningklip.

Indigenous vegetation

Some indigenous vegetation still remains; the oldest and largest trees were :Heeria argentea [Fig.7]. Also present were Pterocelastris tricuspidatus trees [Fig. 8], a cabbage tree Cussonia spicata and two clumps of the succulent Cotyledon orbiculata, (pigs’ ears) amongst other vegetation.

Herbertsdale Heuningklip.

This historical site, also recorded by the botanist Carl Peter Thunberg in 1794 when he camped there and described the area, was then on the main wagon road from Knysna [‘place of ferns’] to Cape Town. Besides the regular wagon traffic of goods needed by the early pioneers, sawn timber was transported from the indigenous forests of the Tsitsikamma [‘place where the water begins’] to Cape Town. The need for timber was critical because the forests around the Cape Peninsula such as those at Houtbaai [‘Wood bay’] and Rondebosch [‘circular thicket’] had quickly been felled from the earliest settlement in 1652. By 1811 there was little or no wood near Cape Town and slave-labour was required to walk 10 to 12 miles in order to carry home a couple of bundles of firewood borne on the ends of a piece of bamboo laid across their shoulders. Once the forests around Table Mountain had been exhausted new sources of timber had to be found, especially for wagon making. From about 1787 the forests along the Rivier Zonderend [‘Kannakamkanna’] mountains were exploited, followed by those of Swellendam and Grootvadersbosch. Next to be plundered were those of the Outeniquas and Tsitsikamma. Already by 1776 a woodcutter’s post had been established at the Swartrivier near the present day town of George where Commandant Mulder in charge of 16 men cut and transported timber for the Dutch East India Company. Herbertsdale (34° 01’S 21° 46’E) is situated east of the Gourits River [named after the Gouriquas or ‘cattle people’] in the Langtou Valley 56km north-west of Mossel Bay on the farm Hemelsrood.

Fig.1. The mound or ‘klip’ of conglomerate comprising the historic Heuningklip, with neighbouring farmer, Jannie Leonard, at the base, alongside the Heuningklip River which shares its name.

The Nouga [‘blue stone/ground’, from !noa , ‘blue’ and !ga, ‘back/ridge’] River which flows southwards from the northern boundary of Gondwana Private Game Reserve through the farm ‘Nauga’ enters the Gouritz River at Outeniquas Drift. Thunberg gave the location of the Heuningklip as 10km east of Outeniquasdrift and Helsdrif [from ‘helling’ = steep descent] on the Gourits River. He recorded that:

 “… we passed a large rock, which on account of its harbouring bees, has obtained the name Honingklip and reached a farmhouse, situated near the Attaquas Kloof [Hottentot tribe location in the 17th century]”. 

A small river also called the Heuningklip flowed at its foot. Thunberg found wild olive trees growing at the Heuningklipkloof that were large enough for the making of chairs but the area at the time of his visit was dry and unattractive.

Searching for Heuningklip

On the Herbertsdale map [3421BB: 1:50 000] there are three farms named Heuningklip along the Heuningklip Road and a Honigklips Kloof to the north of Gondwana Private Nature Reserve. Several attempts were made to locate this site along the Heuningklip Road, especially by calculating the distance of 10km from the Gouritz River and from that of an old milestone distance marker still in situ. With the aid of Jannie Leonard, the owner of the farm Honigklipkloof, the area opposite his farm house was scoured. There was the Heuningklip River, the kloof [ravine] with an additional spring, and furrows moulded out of natural clay that led the water from the mountainside to a pool from which the water flowed through a pipe under the Heuningklip Road to a dam beside the farmhouse. But the entire face of the outcrop was of loose conglomerate which afforded no cavities at all – and therefore no possibility of the presence of honeybee nests.

The Heuningklip stream at the foot of the Heuningklip encourages the growth of trees which cover most of its face.
Fig.2. The Heuningklip stream at the foot of the Heuningklip encourages the growth of trees which cover most of its face.

During a later visit to the area, it was decided to take the early pioneers’ description verbatim and to look for an outcrop that could possibly be described as a ‘large stone’ or klip. This led to the discovery further on, opposite the entrance gates to the farm “Elandsdans”, approximately 100m in from the Heuningklip Road of what could be appropriated as a huge ‘stone’ [Fig. 1] which was covered for the most part by large trees growing at its base [Fig.2]. This was at the historic milestone marking the 7-mile distance on the Heuningklip Road.

With permission to access the property from the owner of Elandsdans, Jannie Leonard of the adjacent farm, ‘Honigklipkloof’, joined me to reconnoitre the area. The huge mound that was the historic Heuningklip was again comprised of loose conglomerate [Fig. 3], half of which over the aeons had been sheared off by the Heuningklip River at its base through which we ‘waded’ to reach the cliff itself. What was different here was that there were horizontal bands of yellowish, powdery sandstone laid down in layers at intervals [Fig. 4]. It appeared that in the long past, that stones brought down by an ancient river were deposited in a wide area [about 17 kilometres long] stretching from the Du Plessis Pass (where the :Aloe ferox covered hill is composed of conglomerate) to the end of the Heuningklip Road. This appears to have been followed by a deposit of a layer of sandstone which varied from 15 to 45cm in width [gemaFig. 5] and then another layer of conglomerate on top of that deposited over many years. Conglomerate is a sedimentary rock deposited either by fast-flowing rivers or wave action and the sand and pebbles are held together [cemented] by silica, calcite or iron oxide. It was within this sandstone that several cavities suitable for bees to nest were found –but no bees [Fig. 6].

All the cavities were dank and wet, some with their entrances covered in black mould [Fig. 7]. In places there was a continual drip from the top of the ‘mound’. One piece of discarded honeycomb was discovered at the base of the cliff [Fig. 8] but it was not possible to discover from whence it had been extracted. Baboon droppings were found in several places but honey gathering by labourers cannot be discounted either. The last two years had been exceedingly wet and the huge indigenous trees, such as the wild olive (Olea europaea africana), growing at the base of the cliff and placing the cliff face in shadow, were possibly well over 100 years old [Fig. 9]. Kystervaring (Gleichenia polypodioides) grew on the floor below the cliff and fungi adhered to the trunks of trees [Fig. 10a+b] while stinkhorn mushrooms (Aseroë rubra) ‘sprouted’ from the moist earth [Fig. 11]. 

 This site fits all the criteria of the “Heuningklip” as described by Thunberg who acknowledged that this name was much older than when he recorded it in 1794. In the region are several rock art sites, of which two are reported to have depictions of honeybees but they are well guarded and no photographs are allowed. It is feasible that some of these paintings depict the nearby Heuningklip bees.

Caught in the headlights on the return journey to Herbertsdale were three magnificent bushbucks (Tragelaphus scriptus). This area is an ideal habitat for them as the appropriately named farms ‘Grootbosch’, ‘Buffelskloof’ and ‘Renosterbos’ attest. The Gondwana Private Nature Reserve is adjacent to the Heuningklip site and has been restocked with animal species which in the past had been hunted to local extinction.

Blocks

Fruits of the Rockwood (Kliphout) tree, Heeria argentea.

Rockwood

The Rockwood (Kliphout) tree, Heeria argentea (previously Rhus thunbergii) was the dominant tree species encircling the Heuningklip. It is a monotypic species in which this is the only species in this genus and belongs to the mango family Anacardiaceae. It has a corky bark which can withstand fires, and thus rocky outcrops represent fire refugia for the tree. It is endemic to the Western Cape and occurs amongst rocks and boulders on dry arid mountain slopes from Kleinmond into the Cederberg Mountains. The fruit is dispersed by the Namaqua rock mouse (Aethomys namaquensis) which hides the fruits in rock cracks after eating the outer layer, allowing the seeds to germinate. Its species name refers to the leaves which are dark green above with fine, silvery-white hairs below. Although not on the threatened species list, the tree has been widely used in the past for fire-wood, tannin from its bark, and was especially valuable for wagon brake-blocks, as was that also of the waboom, Protea nitida. The fruits are reported to be consumed by dassies (Rock Hyraxes).

Aloe ferox

William Burchell, an English explorer, naturalist, traveller, artist, and author, recounting in 1814 in his field journals from his South African expedition, notes that not even a bridle path existed through the Tsitsikamma forest to Plettenberg Bay – Burchell’s map indicating that his route led past the ‘Honigklip’ from Mossel Bay and where they crossed the Gouritz River at Outeniqua Drift. The predominance of Aloe ferox covering the hills between Duiwehoks River at Heidelberg eastwards over a distance of 65 km was already being tapped for ‘aloe gum’. This aloe is of course a great source of nectar and pollen for the honeybees during winter. The pollen and nectar produced by Aloe ferox would have been able to sustain a considerable population of honeybee colonies.

Additional historical information on the Gouritz River

Honeybees appear in the historical record of the Gouritz River in a most bizarre way. This concerns the last man to be publicly hanged on the town square of George in the southern Cape on 28 April 1856. We know that the cliff faces along the Gouritz River were long recorded as harbouring honeybee colonies due to the notorious criminal Bloubaard Swanepoel , a stock farmer at Rietfontein in Attaquaskloof. He would employ labourers to rob honey from the honeybee colonies found on the cliff faces of the Gouritz River. After stealing their honey he would push them off the cliff into a maalgat (whirlpool) in the Gouritz River below where their bodies were never found. However, he was only convicted for the murders of several persons who had bought cattle from him and whom he followed on their return journeys to kill them and then return home with ‘his’ cattle. 

Historically, the first official expedition to the Gouritz River was by Ensign Hieronymus Cruse [Croese] who traded for cattle and sheep with the Hessequa near Swellendam and the Gouriquas near Mossel Bay in 1666. He again came by sea to Mossel Bay in 1667 without wagons. The following year Cruse travelled by sea to Mossel Bay where he landed and made another extensive trading tour along the coast back to Cape Town, but on neither of these trips were wagons used.

 An expedition led by Ensign Isaacq Schrijver consisting of 27 men and several Khoi guides with two wagons loaded with supplies and goods to barter departed the Cape on 4 January 1689. They were to make contact with the Inquas who lived beyond the Attaquas near the present day town of Aberdeen and who had sent an emissary to the Castle to see the Commandant Simon van der Stel. Both the Outeniqua and Inquas were recorded as growers of ‘dagga’ [Khoi dachabCannabis sativa]. The Hessequa kraal under the chief Goukou was visited on the banks of the Goukou River and following along the base of the Langeberg, Schrijver reached the Gouritz River on 22 January. From here he turned inland, and with the aid of a Gouriqua guide, they followed an elephant pass that led them through the Attaqua Kloof. The region was covered in dense brushwood which had to be cut and burned to make a path for the wagons. Eventually they entered the Little Karoo; traversing the Kammanassie [‘washing water/river’] Mountains they entered the Great Karoo. Burchell recounted in 1814 that not even a bridle path existed through the Tsitsikamma forest to Plettenberg Bay. But by then the wagon trail pioneered by Isaacq Schrijver was well developed.

The author:

Geoff Tribe at a wild honeybee nest

Dr Geoff Tribe is a Specialist Researcher – Entomology, Plant Protection Research Institute (retired), has done research on dung beetles, honeybees, solitary bees, forest entomology, slugs & isopods.

Acknowledgement:

James & Noeleen Wessels of Honeystone Cottages, Kleinmond, and Jannie & Amelia Leonard of Heuningklipkloof, Herbertsdale, are thanked for their assistance in accessing the historic Heuningklip sites and for their hospitality.

References:

Binckes, R. (2013). The Great Trek uncut. 30° South Publishers + Helion & Company, 560 pgs.

Burman, J. (1988). Towards the far Horizon. Human & Rousseau, 168 pgs.

Du Preez, F.M. (n.d.) A History of Bees and Beekeeping in South Africa. Self-published, 201pgs.

Loos, J. (2004). Echoes of Slavery. David Philip, 168 pgs.

Nienaber, G.S. and Raper, P.E. (1983). Hottentot (Khoekhoen) Place Names. Butterworths, 243 pgs.

Raper, P.E. (1989). South African Place Names. Jonathan Ball Publishers, 421 pgs.

Ross, G. (2002). The Romance of Cape Mountain Passes. David Philip,224 pgs.

Schoeman, K. (2011). Cape Lives of the Eighteenth Century: Outeniqualand (1768-1794). Protea Book House, 676 pgs.

Skead, C.J. (2011). Historical incidence of the larger land mammals in the broader Western and Northern Cape. Second edition (eds: Boshoff AF, Kerley GIH and Lloyd PH). Nelson Mandela Metropolitan University, 519 pgs.

Stewart R. and Whitehead, M. (2022). Burchell’s African Odyssey; revealing the return journey 1812-1815. Struik Nature, 248 pgs.

White, J.D.M. and Midgley, J. (2017). Dispersal of semi-fleshy fruits to rock crevices by a rock-restricted rodent. South African Journal of Science 113:11-12. dx.doi.org/10.17159/sajs.2017/20170159

The Karoo is a tough place to be a bee!

Surviving the Karoo: resilience in a harsh environment

Being a bee on the Karoo plains is challenging. The Karoo is a boom and bust environment with short periods of spectacular productivity and long periods of drought and famine. Plants survive the droughts as seeds in the soil, or as long-lived small shrubs that are able to reduce their water needs to the minimum by discarding leaves, storing water and desisting from flowering and grow for months or years as need be. Nomadism is/was the preferred option for many birds and larger mammals, whereas many reptiles and invertebrates wait out the tough times in underground burrows reducing their activities to the minimum to save energy and water. Populations of small mammals follow boom and bust cycles in unison with the weather with numbers dwindling in drought and growing exponentially in the good times.

A small Apis mellifera capensis swarm two days after rain.

A forager feeding on nectar of ou rambos (Tetraena retrofracta)

Honeybees as cavity dwellers

Honeybee colonies in more stable environments may persist for many years in hollow trees and cavities in rocks. On the Karoo plains there are no trees and rarely any rock cavities. Accommodation, in common with food supply, is ephemeral and honeybees need to be mobile and opportunistic. From the North African deserts comes the 2000 year old biblical tale of the bees that colonised a lion carcass¹. In the Karoo the deep dens and foraging excavations of aardvark offer a somewhat more hygienic place to build a nest – but underground accommodation is temporary and risky.

Apis mellifera capensis nest site on Wolwekraal Nature Reserve near Prince Albert in October 2023 after drought-breaking rains stimulated mass flowering.

Beneath the surface: termites, aardvark and honeybees

Aardvark feed on Harvester Termites which build their nests 1-2 m below the soil surface. The long-snouted aardvark digs down to the nest and uses its sticky tongue to extract its food. Aardvarks eat a portion of the termite workers at night but seldom if ever wipe out the colony. The termites immediately start to repair the damage by filling the aardvark foraging hole with soil and their droppings. Aardvarks also dig deeper, wider burrows or dens in patches of deep soil and use these as a daytime retreat where they sleep. After a few months they move on to find a new source of food, and the abandoned den is quickly occupied by porcupines, bat eared foxes, mongooses, meercats, shell duck or honeybees.

A honeybee dragging out a harvester termite with a dead honeybee attached to it.

Dead worker bee dropped outside the nest with a dead soldier termite that never lets go.

Dead honeybee and harvester termite with a curious ant.

The downsides of making a nest below ground in a termite colony are threefold: flooding during heavy rain, attacks by the termites wishing to reclaim their territory, and unwanted interest from potential predators of termites, particularly Aardvark, Bat-eared Fox, Yellow Mongoose and Meercat. Dealing with these challenges is time consuming and costly for bees. After flooding, the workers clean the mud out of their nest by carrying mudballs out one by one and dumping them above ground. Bees injured or killed by soldier termites need to be carried out of the nest by fellow workers, and unwanted visitors such as mongooses chased away. All this defence and maintenance work reduces the workforce and eats into the time needed to gather the nectar and pollen resources needed to build combs and grow the swarm.

 

Dead honeybees and soldier termites in the aardvark burrow.

This is probably why the small dark Cape Honey bee (Apis mellifera capensis) with its multiple false-queens, small colonies and mobile lifestyle is much better at exploiting patchy and unpredictable food resources in the Karoo than the larger and more productive yellow race² (Apis mellifera scutellata) from the summer rainfall region of southern Africa.

 

Camera trap footage from Wolwekraal Nature Reserve taken at the wild honeybee nest site:

Yellow mongoose near bee nest at aardvark burrow.

Bees emerging from the subterranean nest in an aardvark burrow.

Aardvark looking into foraging hole occupied by bees.

Yellow mongoose at bee nest.

Yellow mongoose at bee nest.

A nectar foraging ant at the nest site.

Footnotes

  1. Lyle’s Golden Syrup tin boasted the now famous logo depicting Samson’s ‘lion and the bees’ (from the Old Testament, Judges 14:8). It was registered as Lyle’s trademark. Just as the bees produce honey inside the lion’s carcass, rich sweet syrup pours forth from the well-loved tin. Apparently the subject was raised at a beekeeper’s meeting in Pretoria – asking why this should be so? One farmer answered that when animals were slaughtered on the farm, honeybees would come and collect the moisture from the carcass (G. Tribe, personal communication, October 25, 2023).
  2. They are the same species, but from different geographical races of that same species. There are many such geographical races throughout the distribution of Apis mellifera that naturally inhabits Africa, much of Europe, and some of the Middle East.

The author at work:

Dr Sue Milton-Dean has immense experience in plant ecology and veld restoration and dynamics. Sue offers spectacular and highly informative and interesting walks on the Wolwekraal Nature Reserve in Prince Albert. She takes one on a deep-dive into the ecology of the Karoo biome, looking also at its geology, botany, natural and cultural history. For more information visit her website at Wolwekraal Nature Reserve*.

*Wolwekraal Nature Reserve

The vastness of the Tanqua Karoo with Dave in the foreground

Nature of the Interaction dynamics in the Tanqua Karoo

Fig. 1. The succulent Karoo scrub on the farm with the shearing shed in the far distance.


Fig. 2. Honeybee nest in a deserted aardvark burrow which was pestered by banded bee pirates.

Fig. 3. Honeybee nest in a crevice in a shale outcrop on the farm.


Fig. 4. The succulent Tylecodon paniculatus which grows on the surrounding hills.


Fig. 5. Longhorn beetle larvae in the stem of a rotting Tylecodon paniculatus.


Amphibians

Fig. 6. The Karoo toad, Bufo gariepensis.


Fig. 7. A string of Bufo gariepensis eggs.


Fig. 8. The aggregating tadpoles of Bufo gariepensis.


Fig. 9. The aquatic Platana, Xenopus laevis.


Fig. 10. The Cape honeybee, Apis mellifera capensis, the dominant sub-species on the farm.


Fig. 11. Apis mellifera scutellata visiting the flowers of Eberlanzia ferox.


Fig. 12. The emerald fruit beetle Rhabdotis semipunctata on Vachellia karroo flowers.


Fig. 13. Succulent Tylecodon wallichii plants in flower.


Fig. 14. The carrion flower, Hoodia pilifera var. pilifera which attracts pollinating flies.


Fig. 15. Chewed remains of the roots of a Euphorbia rhombifolia plant uprooted by baboons.


Fig. 16. Sundowners in front of the ‘skeerhok’.


Fig. 17. Asilid robber with a honeybee captured on the Tanqua farm.


Zig-zag emperor moth larvae, Gonimbrasia tyrrhea, on Vachellia karroo.


Brunsvigia bosmaniae.


Haemanthus coccineus growing out of a slope.


The authors: Geoff Tribe, A. David Marais & Karin Sternberg