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Stone-Hard Nests in the Karoo: Observing Megachile taraxia

A female Megachile taraxia exiting the mud nest she is constructing on Wolwekraal.

It was spring on Wolwekraal Nature Reserve in the arid Karoo, and the bare patches of ground were alive with solitary bee activity. Besides the busy work of Tetraloniella brevikeraia—the short-horned long-horned bees—other solitary bees were active, too.

The :deflation hollow, its surface cemented by fine dust particles, was bordered by scattered stones and sparse vegetation. Several of the larger stones provided ideal nesting sites for another solitary bee, Megachile taraxia. Unlike the better-known mason bees (Osmia), which usually build in cavities, this species constructs exposed domes of mud on rock surfaces, hence its description as a mud-nesting or “dauber” bee. Until recently, M. taraxia was known only from the male and recorded at just two localities, one in Namaqualand and the other in Gauteng. Finding it here, and finding both males and females, seemed an unusual and important turn up for the books.

Stones on the edge of a deflation hollow at Wolwekraal, providing firm foundations for Megachile taraxia mud nests.

I first noticed a female sitting on a prominent stone. At a glance I thought she was resting; she had been circling me for a while with a deep buzz. Only when she alighted did I see a small wet patch. On closer inspection, I realised I was looking at a mud nest. Without that faint spot of moisture, it might have seemed no more than a chance splash of mud.

Female Megachile taraxia completing the outer seal of her mud nest using saliva.

Reading through the literature on how such mud nests are built, I came upon the works of Jean-Henri Fabre (1913) in his celebrated Souvenirs Entomologiques. He applied the name “mason bees” to those that build their cells with materials similar to our own building supplies. “It is masonry,” he wrote, “but made by a rustic workman, better used to dried clay than to hewn stone.” In his essays he describes the mud-dome nests of Chalicodoma bees. While Megachile taraxia belongs to the subgenus Pseudomegachile, Fabre’s evocative accounts of Chalicodoma nest building, with its mixing of clay, sand, and saliva into a stone-hard mortar, help to illuminate how enduring these dauber bees’ constructions can be. He writes that a calcareous clay is mixed with a little sand and kneaded with the mason’s own saliva (a mix of long-chain hydrocarbons, mainly hentriacontene and tritriacontene, produced by the labial glands). Damp spots that might make the work easier and spare her saliva, are disdained by this dauber bee, which refuses fresh earth for building, just as our builders refuse old plaster and lime. Such materials, when soaked with humidity, would not hold properly. What is needed is a dry powder that readily absorbs the disgorged saliva and, with its albuminous principles, forms a kind of Roman cement that hardens quickly, something like what we obtain with quicklime and egg white. The glandular mixture is used not only as mortar but also spread over the exterior of the nest to render it hydrophobic. The mud dome dries as rapidly as our hydraulic cements, becoming almost as hard as stone. A strong knife blade is needed to cut it. In its final form the nest scarcely recalls the original work, yet it lasts through the year without notable injury. The dome remains much as it was at the start, so solid is the masonry; only the round holes, corresponding to chambers inhabited by the larvae of the past generation, mark its surface. Such dwellings, needing only minor repairs, save much time and effort. Mason bees reuse them, and build new nests only when old ones fail.

Various stones on Wolwekraal showing Megachile taraxia mud nests, their circular openings indicating completed development and emergence. The original mud nest where the first M. taraxia female was discovered is shown top left.

Ten months after my first sighting of the female M. taraxia on her mud nest, and with the early onset of warm spring weather, the first holes appeared in the mud dome nests on Wolwekraal, marking the emergence of the next generation. Within weeks, one hole became four, and soon I began noticing many more nests I had previously overlooked, camouflaged until the perfect round openings revealed them.

Megachile taraxia males and females both utilise mud nests for resting. Top: Female resting in a nest cell (left) with close-up view (right). Bottom: Male resting in mud nest (left) with close-up detail (right).

In one abandoned hole I found a male asleep. On other occasions, I discovered females sleeping inside nests they were still constructing and provisioning. I spent several hours a day watching a female at work. She arrived with her crop full of nectar and her scopa bright yellow with pollen. Head first, she entered the mud cell to disgorge nectar. Once emptied, she backed out, turned, then reversed in to deposit pollen, brushing it from her abdomen with her hind legs. She made several trips to build up a paste-like mass of nectar and pollen, shaping it into a ball like playdough. When the provisioning was complete, she laid an egg and sealed the cell. Only then did she begin work on the next, each cell fully finished before starting another. The rock beneath her nest proved a firm and lasting foundation.

Megachile taraxia female provisioning behaviour sequence. The female arrives with her crop full of nectar and scopa laden with bright yellow pollen, enters the mud cell head-first to regurgitate nectar, then backs out, turns around, and reverses into the cell to deposit pollen.

At another location within Wolwekraal, I discovered more mud nest sites and watched both male and female M. taraxia visiting Blepharis mitrata flowers within a 50-meter radius of these nests. This dense, hard-leaved plant is locally called “scorpion bush” for its formidable spines, or “shooting seed” for its explosive seed pods when wet, and appears to be a preferred foraging resource. The males, who emerge from their mud cells first, use a smart strategy: they wait at the flowers where females come to collect pollen, which is the essential protein food for their developing larvae.

Female arriving at Blepharis mitrata to gather nectar and pollen.

By positioning themselves on or near the flowers, males dramatically improve their chances of meeting a potential mate. Between flower visits, I watched males sunbathing on rocks, stones, and patches of bare sand to warm up, often returning to the same favourite spots. Following these males actually helped me locate more nest sites, as they regularly checked mud nests, apparently monitoring for newly emerging females.

Megachile taraxia foraging and mate-seeking behavior. Left: Female with head deep in Blepharis mitrata flower collecting nectar and pollen. Right: Male basking for thermal energy while waiting for females.

Mating typically occurs either at flowers or near nest sites. Male solitary bees have developed remarkable features for finding females, including modified eyes, antennae, and legs that work like sophisticated detection equipment. Some species have even evolved enlarged brain regions that boost their vision or sense of smell, depending on whether they track down their partners by sight or scent.

Watching these bees at work left me with a sense of awe for their resilience and ingenuity. In the middle of the Karoo, on bare stones near a wind-swept deflation hollow and exposed to temperature extremes, a species so seldom recorded constructed homes almost as enduring as the stone they were built on and carried on its quiet cycle of life. To stumble upon Megachile taraxia here, in numbers and with nests so well hidden, felt like uncovering a treasure. It’s one that reminds us how much there is still to discover, and how much depends on noticing the small, stone-hard wonders at our feet.

Male Megachile taraxia emerging from a mud cell after resting overnight.

A glimpse inside the mud nest: the brood cells are lined with glandular secretions, though little is known about this aspect of bee nesting.

Field Notes from Wolwekraal:

While documenting the story of Megachile taraxia, I often found myself lying so still and so quietly that it felt as though I had become invisible to the creatures around me, or perhaps absorbed into the very terroir of the place. On several occasions, giant tortoises walked past within only a few metres, undisturbed by my presence. Once, a grey mongoose passed less than a metre away, its nearness betrayed only by the scattering of small stones shifting beneath its paws. These encounters made me feel part of the ecology of Wolwekraal, allowing me to observe not only the focal species but also other elements of the ecosystem.

A giant leopard tortoise on Wolwekraal Nature Reserve, Stigmochelys pardalis pardalis.

References:

Buchmann, S. (2023). What a Bee Knows: Exploring the Thoughts, Memories, and Personalities of Bees. Island Press.

Danforth, B. N., Hinckley, R. L., & Neff, J. L. (2019). The Solitary Bees: Biology, Evolution, Conservation. Princeton University Press.

Fabre, J. -H. (1913). Insect Life: Souvenirs of a Naturalist. MacMillan and Co., Ltd.

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.