Banded Bee Pirates: Honeybee Hunters with a Sweet Tooth

How These Predatory Wasps Use Honeybees to Fuel Their Survival and Reproduction.

Karin Sternberg

Survival and reproduction come with tough choices for many animals, and both parasitic and predatory wasps—like Palarus latifrons—are no exception. They rely on two key resources: hosts or prey for their offspring, and sugar to fuel their own bodies and activities. Since these resources are often found in different locations, wasps must carefully divide their time between hunting and feeding. With short lifespans and limited or unpredictable resources, every decision counts—should they pursue prey or stop to refuel? The choice can have a significant impact on their survival and reproductive success in the wild.

Among predatory wasps, Palarus latifrons exhibits particularly striking hunting strategies, preying on honeybees (Apis mellifera), capturing them mid-flight and immediately paralysing them. Subdued bees are then carried to the wasp’s nest, where several paralysed bees are stored in a chamber. A single egg is laid on one of them. The stored prey provides nourishment for the developing larva, until it reaches the pupal stage. After pupation, the fully developed adult wasp emerges from its cocoon.

Palarus latifrons is a medium-sized solitary wasp with a robust body. The female has distinctive black and yellow bands on its abdomen, whereas the male has a black and white banded pattern and is generally smaller than the female. The thorax is usually black and their broad faces have porcelain-like markings (lati- means broad, -frons means forehead or front in Latin). Palarus latifrons is indigenous to Sub-Saharan Africa, however, its distribution extends from the Cape to as far north as Ethiopia and the Arabian deserts of mainly the Arabian Gulf countries. These wasps have adapted well to desert conditions where their activity is high in the hot and dry summer months. Its predatory behaviour has earned it the common name “Banded Bee pirate” (BBP). While it is known to target managed hives, very little has been documented about its interactions with wild honeybee nests.

Hunting for their offspring

At Wolwekraal Nature Reserve on the edge of the Great Karoo—a region considered a desert with annual rainfall less than 200 mm—3 of the 5 wild honeybee nests in deserted aardvark burrows were visited by BBPs, both female and male. One of the nests had up to 8 BBPs hunting around midday. From December through February, maximum temperatures climb to the upper 30 and low 40°C range. Ground temperatures can reach over 50°C. The BBPs use the heat of the sand to gain thermal energy, often flattening their bodies to increase surface area and maximise heat absorption. When warmed up they are capable of hunting at optimal temperature.

One of the wild honeybee nests inside an abandoned aardvark burrow.

Female Banded Bee Pirate wasp soaking up the Karoo heat on the edge of the burrow entrance.

This wasp has a formidable hunting technique. As an agile flier, the BBP is able to crisscross hive entrances of managed bee colonies, while at wild nests located in burrows, wasps dart, in swooping, circular patterns over the nest opening, sometimes diving deep into the burrow and then flying sharply upwards attempting to intercept returning foragers. Notably, no direct attacks on bees clustered on the comb at wild nests were observed.

A returning forager to her nest faces two female BBP wasps, with a male BBP nearby watching a female.

The wasp captures and subdues its prey midair, primarily targeting bees returning from foraging. However, it has also been seen persistently taunting and luring bees out of hives, striking as soon as a bee steps away from the colony. With its long, slender body, the BBP can position itself advantageously to deliver a sting. Still, honeybees sometimes manage to overpower and sting a BBP to death. Where BBP population densities are high—such as at apiary sites in the Porterville area, where we counted an average of 50 BBP wasps per hive—they can severely impact bee colonies, leading to significant honeybee losses. In a study carried out in front of a hive in the Kalahari, researchers killed 131 BBP wasps within 75min.

Female BBP wasp captures a forager with pollen still on her legs at the nest entrance.

The presence of these wasps near hives and at wild nests, is thought to induce either a ‘moaning’ sound by the bees, or a distressed ‘piping’ sound. The distress signals as recorded at the Porterville apiary site were intense and acute. Here, foraging activities halted entirely and many of the bees formed a defensive line across the hive opening, preventing BBPs from entering. At a wild nest on Wolwekraal, short, singular piping sounds were heard in the presence of the BBPs. BBP numbers were meagre and not all foraging ceased, but fewer foragers were seen leaving the nest.

BBP wasps are found in high numbers at apiary sites in hot and dry regions, drawn to a concentration of managed hives. Notice the multitude of wasps flying in front of the defensive line of honeybees in the hives. In contrast, only a few BBPs are typically observed at wild honeybee nests, where colonies are more dispersed across the landscape.

SOUNDS FROM A HIVE:

In the presence of hordes of BBP wasps, these are the distress signals emanating from a hive. Listen for the high-pitched piping sounds.

SOUNDS FROM A WILD HONEYBEE NEST:

Sounds emanating from a wild nest in the presence of a few BBP wasps. Listen closely for the short piping sounds in the background.

BBP wasps are, however, not solely dependent on having to find honeybee nests and have been seen widely dispersed. An observational study in the Tanqua Karoo, recorded BBP wasps at flowering Vachellia karroo trees and water seeps. These seepages attract a diverse array of species, including wasps, solitary bees and honeybees, especially during the hot, dry summer months. At these seeps, BBP wasps were seen carrying off honeybees. Vachellia karroo in bloom serves as a key resource attracting numerous pollinators.

Water as a limiting resource

Water is the most limiting resource for all living creatures, a challenge intensified in desert regions where resources are scarce across both time and space. Wasps require water for survival, but they do not rely solely on seepages or riverbanks to obtain it. Research on water-deprived parasitic wasps (Pachycrepoideus vindemmiae) describes a water-intake strategy whereby the researchers observed increased feeding by the wasps on the water-rich :haemolymph of their hosts, :Drosophila suzukii larvae, for nourishment, in this case, to obtain water. This led to more host (larval) mortality, because the host-feeding by the wasps to sustain themselves with water (and not just to provide food for their offspring) killed the larvae. The paper notes, that these are interesting findings, not only because water has rarely been reported as a critical nutrient for adult wasps, but especially because using prey for the purpose of hydration does not appear to be a common strategy in nature. Or is it more common than thought?

Cape cobra, belly full—likely from a rodent—slips into an abandoned aardvark burrow, once home to a wild honeybee nest. Snakes obtain much of their water from the body fluids of their prey.

The Karoo’s extreme conditions demand remarkable adaptations from its inhabitants, with resident fauna deriving most of their body water from their food. The Cape cobra (Naja nivea) meets its water needs almost entirely through the moisture derived from the body fluids and tissues of its prey. Africa’s largest rodent, the Cape porcupine (Hystrix africaeaustralis), has developed a different strategy to cope with water scarcity—digging up and consuming dwarf succulents to access their moisture. Our observations of Cape porcupine diggings on Wolwekraal revealed an intense foraging pattern. It is also speculated that the eight-year drought in the Karoo from 2015 to 2023, forced porcupines to forage on aloe stems to access both moisture and food.

Besides water for hydration, nest cooling, humidity regulation, larval nourishment, and nest building in bees and wasps, every organism requires energy to survive. Honeybees rely on a steady intake of food to fuel their daily activities—from foraging and flying, to maintaining the nest and producing honey.

Fuelling their behaviour with an energy-rich resource

Foraging demands exceptional cognitive and flight abilities, requiring high metabolic and aerodynamic power. Nectar, a sugar-rich liquid produced by the glands—or nectaries—of flowering plants, provides the necessary energy. Foraging bees collect nectar using their long proboscis (or tongue) and store it in their honey stomach before returning to the nest. Nectar typically has a high water content, usually between 50% and 80%, depending on the plant species. Honeybees process this nectar through regurgitation, enzymatic fermentation, and evaporation to produce honey. Like bees, adult wasps, require quick and significant energy for flying and hunting, which they often obtain from flower nectar or other carbohydrate sources, such as honeydew from aphids, overripe or fermenting fruit, and tree sap.

BBP wasp poised at the burrow entrance, with the comb of a wild honeybee nest visible in the background.

At Wolwekraal, BBP wasps were observed not only capturing prey at wild honeybee nests, but also fulfilling their second key requirement for survival and reproduction: obtaining sugar for energy. While the behaviour of BBP wasps extracting nectar directly from honeybees has been documented in writing, unique footage captured the extraordinary moment, providing a visual record of this fascinating interaction that, to my knowledge, has never been captured before. By preying primarily on returning foragers, these predatory wasps secure nourishment for their larvae, and access to energy-rich nectar and water. The latter is particularly vital in arid regions where water becomes scarce during the peak of summer and flowering plants are few and far between.

The BBP wasp forces open the mouthparts of the captured bee to extract nectar.

The captured bee is not always carried away to the nest but is sometimes used only as a food source. After stinging the bee, the wasp waits for it to calm down before beginning a mouth-to-mouth feeding procedure. The BBP forces open the bee’s mouthparts and, by pressing down on the bee’s abdomen, forces out the crop contents, effectively robbing it of its collected nectar. After this, the BBP either leaves the dead bee to resume hunting or, once suitably positioned, flies back to its nest with the bee. The BBP digs tunnels in the ground in especially sandy, granular soils. Inside the nest, the wasp provisions a chamber within its burrow with up to five bees, laying an egg on the mass before sealing the chamber and proceeding to provision the next one. While only females were documented exhibiting this nectar-robbing behaviour at Wolwekraal, records indicate that male BBPs also engage in this behaviour.

Slow-motion footage (50% speed) capturing the extraordinary moment a female Banded Bee Pirate wasp forces open a honeybee’s mouthparts to extract nectar—a rarely seen interaction between predator and prey.

The path back to the nest is not always easy for the BBP and its prey. In nature, especially under extreme conditions, hazards abound and opportunities must be seized. At Wolwekraal, this duo racing through the sky is often followed by a scurry of small jackal flies, flying just below the honeybee. These kleptoparasitic milichiid flies (Desmometopa spp.) feed on nutrient-rich haemolymph and are known for stealing food from other insects. They are thought to be attracted by a chemical pheromone released by the stressed honeybee and must fly rapidly to keep pace with the much larger predatory BBP.

In the Tanqua Karoo, a small aggregation of six BBP nest entrances was observed within a 20 cm × 6 cm area. Kleptoparasite flies were spotted congregating at the entrances of the BBP nests. These flies were likely Craticulina seriata, known kleptoparasites of sand wasps. They lay eggs that hatch immediately, with the larvae feeding on the honeybee prey. The observations also revealed a BBP returning to its nest, carrying a drone—the much larger male honeybee.

Kleptoparasites, *Craticulina seriata*, at the entrance to a BBP nest. Photo by Geoff Tribe.

The role of male BBP wasps at wild honeybee nests remains unclear. Like honeybee drones, males lack a sting. They often chase female wasps, possibly mating near the nest, and sometimes mistakenly pursue honeybees. This results in chaotic, fast-paced activity above the nest entrance.

The need for a balanced ecology

Observing the ecological dynamics around :wild honeybee nests offers valuable insights. The presence of large numbers of BBP wasps at apiary sites raises serious concerns. These wasps, known for their mass attacks on managed honeybee colonies, exploit the high-density clustering of hives, particularly in arid environments where resources are limited. Their predatory behaviour is intensified in these artificial settings, leading to significant colony stress, as evidenced by the increased defensive responses and distress signals from the hives. While wasps serve a critical role in ecosystem regulation, their population dynamics are highly responsive to environmental conditions, sometimes tipping the balance in their favour. This highlights the importance of conserving wild honeybees in their natural habitats, where intrinsic ecological balance helps regulate predator-prey interactions without human interference.

REFERENCES

Al-Khalaf, A. (2021). Modeling the potential distribution of the predator of honey bees, Palarus latifrons, in the Arabian deserts using Maxent and GIS. Saudi Journal of Biological Sciences 28(2).

Arena, G. And Milton, S. J. (2025). Impact of drought-induced herbivory by Cape porcupine on Aloe claviflora on the Wolwekraal Nature Reserve, Prince Albert. Journal of Arid Environments 227 (2025) 105337.

Bernstein, C. (1996). Time sharing between host searching and food searching in solitary parasitoids: state dependent optimal strategies. Behavioral Ecology 7(2):189-194.

Bezerra Da Silva, C.S., Price, B.E. and Walton, V.M. (2019). Water-Deprived Parasitic Wasps (Pachycrepoideus vindemmiae) Kill More Pupae of a Pest (Drosophila suzukii) as a Water-Intake Strategy. Sci Rep 9, 3592.

Clauss, B. (1984). The status of the Banded Bee Pirate, Palarus latifrons, as a honeybee predator in southern Africa. Paper presented at the Third International Conference of Apiculture in tropical Climates, Nairobi.

Dean, W. R. J. And Milton, S. J. (2004). The Karoo. Ecological patterns and processes. Cambridge University Press. 374 pgs.

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 239(4).

Mally, C. W. (1908). Bee Pirates. Agricultural Journal of the Cape of Good Hope 33(2): 206-213.

Skaife, S. H.(1979). African Insect Life. Struik 279 pgs.

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

Wright, G., Nicolson, S. and Shafir, S. (2018). Nutritional Physiology and Ecology of Honey Bees. Annu. Rev. Entomol. 63:327-44.

Xylocopa io: The Rarely Seen Carpenter Bee

Karin Sternberg

Some Xylocopa have interesting adaptations, like Xylocopa io with its elongated middle legs.

A male *Xylocopa io* with a distinctively long middle leg, visible on the right of the bee.

Xylocopa bees are mostly large, robust bees. They are solitary bees, unlike honeybees that live in colonies. There are many species of carpenter bees and they are important pollinators. They excavate tunnels in dead branches which are protected from rain, and construct partitions between the individual cells, each of which is provisioned with a lump of nectar and pollen mixed together, on which a single egg is laid.

Recently while out hiking we spotted a male Xylocopa io high on a ridge between Prince Albert and the Swartberg Mountains in the Karoo. Not much is known about this species, and even less is known about the female X. io., but one interesting feature of the male is his two long middle legs.

A male *Xylocopa io* hovering, revealing its unique extra-long middle leg.

Insects are generally bilaterally symmetrical, so this male—with only one long middle leg—may have had a narrow escape from a bird, or something similar.

To attract females, Xylocopa males generally hover around a flowering shrub, then dart off, only to return seconds later to hover briefly again.

Males patrol a patch of flowers, which females visit while foraging, in order to mate with them. A patrolling male will aggressively chase away other males that intrude on his territory. The success of these encounters is enhanced by the permeation of a sex pheromone released by the male while patrolling his patch of flowers.

Male *Xylocopa caffra* on *Aspalathus capensis* (Cape capegorse).

Male *X. caffra* with less exposed membranes

Xylocopa caffra males make quick circuits over an area of several square meters. What is most fascinating is that the normally completely yellow male, when hovering around its patch of flowers, lowers its abdomen and exposes two of its interstitial membranes which appear as two black bands on the abdomen. This behavior is most likely a way for the male to release a sex pheromone, serving both as a species-specific chemical signal and as an attractant for females. When the male momentarily darts away, the membranes are less exposed compared to when he hovers.

Males of the carpenter bee Xylocopa hirsutissima have a different strategy: they fly to the top of the mountain where they spread mandibular gland secretions over the ventral surface of their abdomen while hovering in the air awaiting the female.

A rare sight: a male *Xylocopa io* with a strikingly long middle leg, a unique trait of this elusive carpenter bee.

In Xylocopa io, the male’s elongated middle legs are believed to be an adaptation for mating. During mating, the male launches into the air after the female, using his extended legs to grasp her along ridges below her three small simple eyes, known as ocelli. Interestingly, another species, Xylocopa flavrufa, exhibits a similar mating behaviour: the males also pursue females in flight, using their long middle legs for mating.

The parasitic fly, *Hyperichia bifasciata*, thought to mimic the carpenter bee *Xylocopa caffra*, .

Found in the same area as X. io, was a species of fly, Hyperichia bifasciata, known to mimic specifically Xylocopa caffra. The fly’s name, bifasciata, suggests that it may have two prominent markings or bands (the “bi-” prefix means “two” and “fasciata” refers to bands or stripes). There exists much individual variation in the colours of the bands on the thorax and abdomen, from whites to yellows. Hyperechia flies are notable for their parasitic relationship with carpenter bees. The larvae are predacious on those of Xylocopa. As adults, these flies feed on bees and wasps. H. bifasciata is not too picky and so will parasitize other Xylocopa species as well.

References:

Grünberg, K. 1907. Zur Kenntnis der Asiliden-Gattung Hyperechia Schin. (Dipt.). Deutsche Entomologische Zeitschrift: 515-524

Van Bruggen, A.C. A preliminary note on the genus Hyperechia

Schiner in Africa (Diptera: Asilidae). https://journals.co.za/doi/pdf/10.10520/AJA00128789_3712

Velthuis, H.H.W. and Camargo, J.M.F. de 1975. Further observations on the function of male territories in the carpenter bee Xylocopa (Neoxylocopa) hirsutissima Maidl (Anthophoridae, Hymenoptera. Netherlands Journal of Zoology 25(4): 516-528.

Watmough, R.H. 1974. Biology and behaviour of carpenter bees in southern Africa. The Journal of the Entomological Society of Southern Africa. 37(2): 261-281.

Far From Dead: The Surprising Biodiversity of Bare Ground

Occasionally, biodiversity might just be thriving beneath our feet.

Karin Sternberg

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.

Plant pulp in nesting hole of solitary bee

Megachilid bee emerging from nest burrow

Flower petal taken into hole by leaf cutting bee

Megachilid leaf cutter bee with pollen on scopa

Megachilid leaf cutter bee taking a leaf into a hole

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.

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.

Karin Sternberg

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

Aardvark burrow with a wild honeybee colony

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.

A small colony situated in an aardvark hole

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

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.

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

The role of ants

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.

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.

About Bees, Naturally.

Karin Sternberg

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.

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

Karin Sternberg

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.

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.

An Allodapine bee foraging on the lammerlat flower. — 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.

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.

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.

Honeybees preparing the surface and applying resins in a cavity dwelling situated in an aardvark burrow.

In a second cavity situated in an aardvark burrow honeybees prepare surface soils and apply propolis with lammerlat resins. — Honeybees preparing the surface and applying resins in a cavity dwelling situated in an aardvark burrow.

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.

SELECTED REFERENCES:

Bruyns, P.V. (2002). The Stapeliads of Southern Africa and Madagascar Volume 1 pg 57.Umdaus Press, 329 pgs
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.
Ghisalberti, E.L. (1979) Propolis: A review. Bee World 60(2):59-84.
Iannuzzi, J. (1983). Propolis: The most mysterious hive element. American Bee Journal 123(8): 573-575.
Iannuzzi, J. (1983). Propolis: The most mysterious hive element Part II – Conclusion. American Bee Journal 123(9): 631-633.
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.
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
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…”

Geoff Tribe

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.

Fig.3a. Decades old Apis mellifera capensis colony near the waterfall adjacent to the Harold Porter Botanical Gardens.

Wild honeybee colony — Fig.3a. Decades old Apis mellifera capensis colony near the waterfall adjacent to the Harold Porter Botanical Gardens.

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.

Fig. 4: Propolis covered entrance to the cavity housing the honeybee colony.

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.

Fig.7. Fruits of the Rockwood (Kliphout) tree, Heeria argentea.

Fruits of the Cherrywood (Kershout) tree, Pterocelastris tricuspidatus — Fig.7. Fruits of the Rockwood (Kliphout) tree, Heeria argentea.

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 17^th 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.

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].

Fig.3. The conglomerate structure of the mound that comprises the Heuningklip.

The horizontal yellow sandstone bands in which suitable cavities for honeybee nests were seen. — Fig.3. The conglomerate structure of the mound that comprises the Heuningklip.

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].

Dank, dark and wet cavities within the conglomerate that could have housed honeybee colonies in bygone years — Fig.7. Dank, dark and wet cavities within the conglomerate.

Piece of comb from one of the wild honeybee nests located below the cliff face. — Fig.7. Dank, dark and wet cavities within the conglomerate.

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.

Fig.9. Wild olive trees growing against the cliff face.

Substantial rainfall of the previous two years resulted in the profusion of mosses growing. — Fig.9. Wild olive trees growing against the cliff face.

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

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 dachab – Cannabis 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:

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!

Sue Dean and Karin Sternberg

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.

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.

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.

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.

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:

Footnotes

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).
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:

Nature of the Interaction dynamics in the Tanqua Karoo

Geoff Tribe, A. David Marais and Karin Sternberg

Environmental impact assessments

An Environmental Impact Assessment (EIA) is nowadays required before any development can proceed – yet how accurate are they? The year in question, the season, the size and duration of the survey (weeks/months/years) can influence the outcome of the EIA. What would constitute a minimum duration for such a survey? An ecological research project would require a minimum of 3 years, where 5 years is passable, but 10 years would give far better insight. EIAs are often inadequate in terms of genuine environmental protection, and many species are missed, particularly the geophytes and ephemerals, the invertebrates, frogs, birds and bats. Some EIAs are designed to facilitate and even legitimize development where it should not happen. They are mere snapshots in time and miss the far greater, fundamental ecological processes and responses. An erroneous decision could have far-reaching consequences for very sensitive and less well researched areas. The flora and fauna must be researched well, especially in areas that had not previously been well documented.

With sedentary plants and perennials which can be more readily located, identified and enumerated, a reliable assessment is more easily accomplished. Yet seeds from some plants can stay dormant for decades until favourable conditions cause them to germinate. This applies also to the erratic flowering of underground bulbous plants triggered usually by substantial rains. An example of this in relation to the account below is the dubbeltjiedoring, Tribulus terrestris, which prior to the flood was rare on the farm. Following the exceptional rains the unwelcome thorns appeared en masseespecially around the disused sheep pen and along the banks of the once dry streams, forming dense mats with single plants often covering an expanse of one metre or more in diameter and leaving literally thousands of the thorns to germinate when conditions are again favourable.

When it comes to mobile organisms such as birds, animals and insects which can migrate from regions when adverse conditions persist, re-colonization is necessary when conditions improve. This would necessitate ‘refuges’ for such organisms where, within the vast area affected, there are atypical areas that remain favourable due perhaps to underground water or seepages. Would such ‘refuges’ be identified during EIA? Re-colonization can be accomplished by migration or in the case of plants by seeds being blown or carried by animals back into the affected area. This would only be possible if the EIA properly takes into account the region and mitigation of damage and risk is done by judicious sparing of some zones for development.

The cyclical effect on nature through seasonal and climatic fluctuations determines and requires adaptive responses by different organisms. Yet it is surprising to see how readily reactions can occur in response to changes in the status quo. An example of this was the changes that occurred on a farm in the Tanqua Karoo about 35 km north-east of the town of Touwsrivier. Some interesting observations on this farm will be described after an account of recent climatic conditions.

Tanqua Karoo

The karroid vegetation of the Tanqua Karoo is composed of xerophytic, semi-desert shrubland with a large number of succulent-leaved species [Fig 1]. There is no surface water on the farm. A wind-pump supplies ground-water when required. From 20 years ago, average rainfalls were recorded on this farm. Although one year was never the same as the year before, the fluctuations in the fauna and flora were not drastic. However, a drought persisted for the preceding eight years during which time a marked deterioration of the veld was observed. Many succulent plants died and others failed to flower. Many insect species either disappeared or became rare. Four colonies of Cape honeybees (Apismellifera capensis) two of which nested in aardvark burrows [Fig. 2] and two in clefts in shale outcrops [Fig.3], absconded. The annual rainfall over the past decade averaged 102mm with a range of 34 to 226mm. With the exception of 2018, the preceding 5 years had rainfalls of less than 90mm per year. Generally, the rainfall during November and December was <10% of the annual rainfall but in 2021 it was 26mm (37%). It was completely different in December 2022.

The drought was broken by recurrent episodes of significant rainfall from November 2022 to May 2023 which were unprecedented for the last 20 years. The rainfall of 143mm in December resulted in a flood. Indeed, the Tanqua Karoo and adjacent regions experienced widespread floods. Dry river streams turned into raging torrents and the sand was washed down from the hills and deposited onto the plain below, leaving a wide river bed with solid shale bedrock. Over the six months of increased rainfall, the growth of the vygies in particular was particularly good. Whereas one could easily walk in the empty patches between the vygies, they now coalesced into an aggregation as they increased in growth. Vachellia karroo (‘soetdoring “ or karoodoring’) immediately responds to substantial rainfall . Vachellia karroo flowers develop only on the young growth of that season and so growth, dependent on rainfall, precedes flowering by 4 to 5 weeks. Within a short while these trees flowered twice in succession and became an attraction for all manner of insects including beetles, wasps, flies, caterpillars, butterflies, bugs, aphids, and solitary bees … but no honeybees were seen. Presumably the Cape honeybees that were observed collecting water at various seepages throughout the farm were visiting alternative sources which were more profitable.

Due to the floods, the Vachellia karroo and Searsia burchellii trees which were scattered along the rivulets had their roots exposed. The roots had been unable to penetrate the shale bedrock but instead had followed along the banks of the river, some being as long as perhaps 12 metres. In most cases it appeared that the combined mass of the roots of a tree was greater than that part above the soil. Another observation was that many of the larger succulent species such as Tylecodon paniculatus [Fig. 4] and Tylecodon wallichii had rotted and collapsed due to the perpetually rain- soaked soil. Surprisingly, larvae of an unidentified longhorn beetle (Cerambycidae) which are usually found in the solid wood of dead trees were found in the soggy, water-laden stems of a T. paniculatus [Fig.5].

Amphibians

Although five species of snakes and several lizard species occur on the farm, until this flooding no frogs had ever been seen. Yet over a period of at least six months the newly filled hollows in the impermeable shale of some of the riverbeds by seepages contained many hundreds of tadpoles. Two frog species were identified. The Karoo toad (Bufo gariepensis, also named Vandijkophrynus gariepensis)) [Fig.6] can, under favourable conditions, reproduce in large numbers because as many as 20 000 eggs can be deposited by a single female. These distinctive eggs are united into strings [Fig.7] by copious amounts of jelly-like oviducal secretion. The eggs hatch into small, dark tadpoles which concentrate into tight free-swimming clusters [Fig.8]. The adult Karoo toad may vary considerably in colouration but is cryptic in its prevailing surroundings.

The second species, the Common Platanna or African clawed toad (Xenopus laevis) [Fig.9] has webbed toes, and lacks a tympanum, tongue and movable eyelids as an adaptation for an aquatic life. Adults are both predators and scavengers. Their eggs are small, heavily pigmented and are enclosed in individual jelly capsules attached to submerged objects.

The question arises from whence these amphibians came? Because both species are indigenous over this vast and arid region, it can be surmised that they had buried themselves in the sandy banks of the dry streamlets in order to survive the adverse drought conditions. This phenomenon has been observed in the Namib Desert where frogs were found a metre deep in sand under a dried pool in dry-hibernation. Had the Tanqua floods not brought them to activity, they would not have been known to occur on the farm.

Honeybee races at hybrid zone

The Cape honeybee (Apis mellifera capensis) [Fig.10] is restricted to the winter rainfall region of southern Africa but is purported to have an interface with the Savannah bee (Apis mellifera scutellata) along the margins of this region which has been designated as the ‘hybrid zone’. Because the outer margins where the two honeybee races converge are mostly semi-desert, it has been postulated that the low numbers of wild colonies and the correspondingly reduced population pressure between the two sub-species, could result in co-existence but changes in the winter and summer rain could influence the success of a given species occupying the zone.

Several other factors may also influence the habitation and cohabitation of the bees. The area where the highest numbers of ovarioles occur in laying-workers of the Cape bee is regarded as the ‘heart’ of the capensis distribution. This lies in an inverted triangle that can be drawn from Stellenbosch in the west, to Swellendam in the east, and to Cape Agulhas in the south. The Cape bee has maintained its dominance in the winter rainfall regions for many thousands of years. The predominance of capensis is also observed when scutellata hives are brought into their region. Within a few years such hives become occupied by capensis because capensis workers find their way into scutellata colonies and become laying-workers and eventually take over these colonies by becoming pseudo-queens. This is due to the higher level of queen pheromone possessed by the Cape bees.

The ‘Capensis Calamity’

Cape honeybees taken to the aloe-flow on the Springbokvlaktes north of Pretoria were able to infiltrate scutellata colonies during the manipulations which take place when ‘making increase’. This is accomplished by taking advantage of the massive amounts of pollen available by dividing a hive and allowing queens to be produced in the queenless brood placed above the queen excluder. Such conditions are ideal for the acceptance of a Cape bee into the queenless partition and here to rapidly develop into a pseudo-queen. The upper partition rapidly fills up with capensis brood and then Cape honey bees emerge.

Despite this proliferation, it appears that capensis colonies do not survive for long in the summer rainfall region but steadily dwindle in numbers and eventually die out. Cape bees rarely are able to invade healthy widely scattered natural colonies of scutellata, but will readily invade commercial colonies because of their proximity in apiaries where regular manipulation takes place. There is much anecdotal evidence that honeybees are adapted to certain environments where they flourish, yet merely maintain themselves when taken to a different environment. For instance, honeybee colonies hived in the Delmas area and brought to Pretoria where they were average producers compared to resident colonies, excelled when taken back to the sunflowers around Delmas each year. Cape bees in Pretoria were seen in greater numbers to visit Cape plants like vygies compared to scutellata bees. After more than 30 years in scutellata territory, other than as invaders of manipulated commercial scutellata hives, the Cape bee has failed to become established in the summer rainfall region.

Shifting boundary

In May 2023 on one of the hills on the Tanqua farm where the vygie Eberlanzia ferox was flowering, some bright yellow honeybees were found with little pubescence on their abdomens which could only be scutellata [Fig.11]. Had they temporarily expanded into capensis territory due to the excessive (summer) rains and the resulting mass flowering of plants which could facilitate such an expansion? Although the identity of the bees has not been determined scientifically by dissection, having worked with both races for many years it is possible to superficially identify the two races with some certainty. Perhaps the history of the region should give an indication.

History of the Tanqua Karoo

The farm is situated in the Tanqua Karoo, the name probably being derived from “Sanqua” indicating that the San were the first inhabitants of the region. There is also a river bed named the Tanqua. The region is also known as the ’Onder Karoo’ and ‘Ceres Karoo’. The Khoi subsequently moved into this area with their livestock and the competition for pasture and water resources became intense. This was before the days of the wind pumps and the extraction of subterranean water. The Trekboers expanded into the interior in large numbers in the 18th century. In the 1800s this was a contested area which existed between Touwsrivier in the south, to the Hantam Mountains in the north and the Roggeveld escarpment inland. It was only possible to farm small stock in this area if a farmer was able to move across the boundaries between the winter and summer rainfall area in different seasons – which still is pertinent today. Farmers still take their sheep down to the Tanqua Karoo in winter to escape the extreme cold of the escarpment. The demarcation between winter and summer rainfall regions is not a precise line but rather a shifting corridor in which there might be little rainfall in both the winter and summer months. This all-season rainfall corridor coincides roughly with the line of the interior escarpment. Sheep and goats could survive in the winter rainfall region during winter but could not remain there when it dried out and the heat became intense. They would be moved further inland onto the higher escarpment in summer.

This region is subject to cycles of excessive rain followed by drought which resulted in competition for grazing amongst the Trekboers, Khoi and San hunters. This became an area of continual conflict for about a century. An example of this conflict is a report by Field Sergeant Charl Marais in September 1779. Although a comprehensive record does not exist for the climate, flora and fauna, some information is available about the rainfall in the region. Historically, the years 1700 to 1704 were years of poor rainfall but normal rainfall returned in 1705. Heavy rains in 1706 caused a great loss of livestock. July 1715 was exceptionally cold and wet and was associated also with an unknown cattle disease which led to high mortality. A severe drought in 1800 was followed by substantial rainfall in the Roggeveld in 1803 with many flowing streams. But during 1805 an extreme drought was experienced in the Tanqua Karoo. In contrast, there is a record that in one year the summer rains were so widespread that they swept through the Karoo to the sea.

Fauna and Flora

Visiting this farm regularly throughout the years unveiled the effect of climate and weather on the ecology of the area. Nothing stays the same from year to year. An insect species which is prolific one year may disappear totally for a number of years before reappearing again when conditions are suitable. A good example is the emerald fruit beetle (Rhabdotissemipunctata) which can be found cavorting on the flowers of Vachellia karroo. Prior to the long drought they had been prolific but disappeared until V. karroo flowered again due to the heavy rains in 2022/3 [Fig.12].

Botanists visiting the farm over a weekend were able to identify 98 species of plants – and many more have yet to be identified. Yet honeybees are rarely seen on flowers despite hours of hiking each day on the farm. However, they are desperate for water in the hot summer months and will visit artificial pools of water. Despite masses of vygies of various species flowering in spring, it is rare to see a single honeybee visiting them. It appears that the amount of forage in the worst month greatly influences the carrying capacity of wild colonies in an area. Honeybees have been observed visiting Haemanthus coccineus, Brunsvigia bosmaniae and various species of Asteraceae.

Many species of succulent plants appear to have niche pollinators, many of them solitary or sub-social bee species. Tylecodon wallichii produces an exceedingly long raceme which towers above the karroid scrub. It attracts a Xylocopidae (carpenter bee) species which flies from the one exposed flower stalk to the next which is clearly visible in the landscape as they project well above the karroid scrub [Fig.13]. There are also seven species of carrion flowers (Stapeliads) whose flowers produce a stench of either sweat, urine, faeces or a rotting cadaver by which they (deceptively) attract flies without any reward which pollinate them [Fig.14]. This makes at least a subset of plants independent of bees for pollination. The pollination of these plants is further amplified by the placing of all the pollen in a sac (pollinium) which the fertilising insect transfers. A similar strategy is employed by orchids, of which only one species has been found on the farm. The Gorteria diffusa uses a different strategy to attract insects. By expressing pigments on the petals, the spots which resemble insects, they attract the passing fly to visit the plant and it becomes a pollinator. Nevertheless, bees remain important for the pollination of some of the plants whether they flower in the summer or winter.

Occupation of the region by humans has undoubtedly affected the ecology of the region; especially with continual occupation and the introduction of livestock. The farm has not had stock on it for 20 years (except occasionally those of the neighbours which find an opening in the fence) but there are resident rhebok, duiker, steenbok and three pairs of klipspringers. Baboons regularly pass through and leave a tell-tale of the chewed roots of Euphorbia rhombifolia [Fig.15].

Game abounded in this region before firearms appeared. Colonel Robert Jacob Gordon in 1777 attested to the large number of lions in the Tanqua Karoo. Augusta de Mist (1803) also commented on the large numbers of lions, leopards and hyenas and was enthralled to see a flock of about 300 ostriches on the southern plain below the Rooiberg which forms the one boundary of the farm. The Bokkeveld in fact was named after the scattered herds of springbok which migrated from the interior into this territory at times. The only accommodation on the farm is an old ‘skeerhok’ (shearing shed) in which to sleep from which a beautiful view over the veld can be seen as the sun sets [Fig.16].

There is no doubt that weather and climate have a large effect on honeybees which expand in strength and number of colonies in favourable years. During unfavourable times, the honeybees decrease in numbers and possibly abscond. Judging from trap boxes placed out each year in the Swartland, drought can drastically curtail the numbers of migrating reproductive swarms. The banded bee pirate, Palarus latifrons, occurs on the farm and there are also generalist arthropod predators such as the Asilidae robber-fly [Fig.17]. However, although honey badgers have a wide distribution across Africa they have not been observed on the farm.

The interesting observations over approximately two decades need further study before the extensive development of facilities to generate electricity by harnessing wind or sunlight overwhelm the area. The Perdekraal Oos wind farm is located within 10 km of the farm where the observations have been made. Several other developments are envisaged in the Renewable Energy Development Zone (REDZ). The environmental impact studies for this development and other developments under consideration in the region reflect relatively short periods of study and do not provide detailed information on the waxing and waning of the numbers of bees and their nests, the interaction of the capensis and scutellata bees as well as other related pollinators. It is likely that periods of more summer rain have temporarily favoured scutellata bees while the capensis bees can be viewed as the predominant inhabitants of this region. It is also evident that there is an interaction between the bees and flora as well as fauna, including the provision of nests by aardvark burrows.

Cape Point Cape bee sanctuary

Because of the high numbers of Savannah bee hives that were being brought within the natural Cape honeybee distribution area, there was a fear of a threat to the existence of the Cape bee. It was decided to found a sanctuary for the Cape honeybee where they would be protected from hybridization. A cursory survey was carried out in the proposed sanctuary of Cape Point in1980. On the recommendation that no honeybees were present in the area, a decision was made to introduce colonies of Cape honeybees. These were fortunately located within the ‘heart’ of the Cape bee distribution at Fernkloof (Hermanus) and De Hoop Nature Reserve and approximately six colonies were introduced.

Yet the Cape bee has always occurred in Cape Point – one of the oldest maps of the Cape Peninsula having a location named Bynes, alludes to this. Over the past 8 years of research in locating and analysing wild colonies in Cape Point, 98 wild honeybee nests were recorded – and more still have yet to be located. Then how was the incorrect assessment of the total absence of honeybees in Cape Point arrived at? It appears that there was one visit on a specific day only and a cursory search of bees on flowers was made. At certain times of the year in Cape Point, masses of various plants can be in flower but no bees are seen visiting them. Honeybees gauge which flowers are most cost effective and will visit those despite other species in full bloom. Also, the carrying capacity of an area appears to be determined by the level of resources available in the month(s) of dearth. Thus the timing and duration of the survey becomes of utmost importance if it is not to become a mere snapshot in time!

The pressure on development of the land and the changes in climate should compel us to document our unspoilt areas in detail, to perform comprehensive EIAs and to monitor the impact of the developments for all our flora, fauna and landscape.

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

Geoff Tribe

Dave Marais

Karin Sternberg

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