Tag Archives: Prince Albert

Banded Bee Pirates: Honeybee Hunters with a Sweet Tooth

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

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.

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

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).

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.

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

The resilient Karoo landscape

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

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

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

Misconceptions of bare ground

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

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

The evolutionary journey of bees

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

Dynamics on the deflation hollow

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

A small section of the Tetraloniella nest aggregation

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

Coexisting species: The diversity of life

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

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

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

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

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

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

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

A climax

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

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

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

A rich tapestry of creatures

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

The hidden life beneath our feet

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

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

ACKNOWLEDGEMENTS:

Dr Sue Milton-Dean of Wolwekraal Nature Reserve

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

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

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

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

REFERENCES:

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

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

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

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

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

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

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

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

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

Honeybee collecting nectar from a milkweed plant in South Africa

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

Karoo landscape with Lammerlat (Gomphocarpus filiformis)

Wild honeybees and propolis

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

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

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

Lammerlat (Gomphocarpus filiformis)

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

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

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

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

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

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

Defensive resin response of Lammerlat seed pods to Tephritid flies

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

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

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

Role of resins in honeybee nest construction

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

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

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

Ecosystem dynamics of Lammerlat

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

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

SELECTED REFERENCES:

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

The Karoo is a tough place to be a bee!

Surviving the Karoo: resilience in a harsh environment

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

A small Apis mellifera capensis swarm two days after rain.

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

Honeybees as cavity dwellers

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

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

Beneath the surface: termites, aardvark and honeybees

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

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

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

Dead honeybee and harvester termite with a curious ant.

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

 

Dead honeybees and soldier termites in the aardvark burrow.

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

 

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

Yellow mongoose near bee nest at aardvark burrow.

Bees emerging from the subterranean nest in an aardvark burrow.

Aardvark looking into foraging hole occupied by bees.

Yellow mongoose at bee nest.

Yellow mongoose at bee nest.

A nectar foraging ant at the nest site.

Footnotes

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

The author at work:

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

*Wolwekraal Nature Reserve