Even with modern improvements such as lightning-fast auto-focusing and motor drives capable of firing the camera shutter ten times a second, plus digital memory cards that can store 40 times as many images as the old-style individual 36-frame films, photographing aculeates in flight remains a complicated affair. They are not the fastest invertebrate fliers, with some Diptera and Odonata travelling much quicker. One Australian dragonfly reportedly has been 'clocked' at 36mph whereas 15-20mph would be a maximum for most bees or wasps. Also, speeds vary depending on individual species, with larger ones tending to fly faster due to their greater mass, and on what they happen to be doing. For example, the rate of progress when foraging is usually relatively low, and for obvious reasons - extra weight and wind resistance - headway is slower when returning to a nest than when leaving it.
Our largest aculeate, the Hornet (Vespa crabro), uses between 117 and 247 wingbeats a second, which certainly makes catching its movement on an image a fairly daunting prospect. For smaller species the challenge is greater even if they are going a bit slower, and as a result flight for these is not attempted very often even in the studio, where much flight photography is effected. This method involves using such equipment as high-speed flash units, laser beams, microprocessors and sometimes artificial 'flight tunnels' measuring a metre or more. The paraphernalia can be fairly considerable both in size and in cost. Indeed, the celebrated nature photographer Stephen Dalton, the first person in Britain to take outstanding photographs of bees in flight over 30 years ago, claimed his equipment looked "a bit like a power station, with wires everywhere".
The studio approach has never had any appeal to me, since many of the resulting images tend to show a wasp or bee in flight, but not much more. From a purely aesthetic point of view, with the wings completely frozen at high speed, say 1/25,000th second, there is no genuine sensation of movement actually occurring. But forgetting this, and acknowledging that the technical marvel that is flight is indeed shown in all its glory in many studio images, there is still a major drawback. Put simply, nothing is ever being carried in the legs or mandibles of any wasp pictured in a studio, or in the mandibles or on the scopae of any bee. In other words, there is little characteristic behaviour on offer. Additionally, photographs taken in studios, and especially inside flight tunnels, quite often seem to have a backdrop which in colouration is not entirely natural.
To photograph an aculeate doing something in flight besides merely moving one has to leave the studio and go into the field, which even with modern equipment presents its own set of problems that have tested my inventiveness and patience any number of times over the last few years. The effort has been worthwhile though, since systematically watching and photographing any aculeate ferrying loads of one sort or another to a nest is rewarding in terms of the information it can provide about natural behaviour as well as the excitement generated on the sadly rare occasions when the subject is in focus and in a central position on an image.
The first rule is to capture any female stocking a nest in a suitable position, in order to identify her before attempting action photography. Taking images, whether it is of interesting behaviour or not, is of questionable value without knowing what the species is. None of the aculeates I have photographed (or tried to photograph) in flight at nests have been too difficult to determine. They are Ancistrocerus nigricornis, Odynerus spinipes, Psenulus pallipes, Crabro scutellatus, Ectemnius cavifrons, Ectemnius cephalotes, Rhopalum clavipes, Ancistrocerus trifasciatus, Crossocerus annulipes, Crossocerus megacephalus, Crossocerus podagricus, Crossocerus megacephalus, Crossocerus tarsatus, Trypoxylon clavicerum, Mellinus arvensis, Cerceris arenaria, Cerceris quinquefasciata, Cerceris rybyensis, Philanthus triangulum, Colletes hederae, Andrena cineraria, Andrena fulva, Andrena vaga, Dasypoda hirtipes, Megachile ligniseca, Anthophora plumipes, Ceratina cyanea, Chelostoma campanularum, the Red Mason Bee (Osmia bicornis), Osmia caerulescens, Osmia leaiana, Heriades truncorum, Hylaeus communis and Bombus lapidarius plus the cleptoparasites Nomada fucata and Nomada signata.
Nomada bees are among the slowest fliers in the family, and often fly close to the ground, both of which tendencies are helpful, though their small size is not a boon. Some bees gathering pollen can end up providing an in-flight shot almost by accident, because they flit from bloom to bloom. This particularly applies to Colletes succinctus and Lasioglossum prasinum, which use Heather as their pollen source. The flowers are tiny, so constant movement is required. For anyone wishing to try and obtain images of bees in flight, flowers are the best bet and bumblebees are among the easiest options. Bombus muscorum is one of the species I have managed to photograph in this way. Some wasps are good on flowers too as they take nectar or look for prey. The former include Ancistrocerus gazella and Ancistrocerus nigricornis, with Canadian Goldenrod a classic plant, and Argogorytes mystaceus, while Cerceris rybyensis is regularly seen around Common Ragwort looking for bee prey.
My standard equipment used to be a Canon 1D Mark II digital camera with a 100mm or 180mm macro lens, 12mm extension ring and Canon Macro Twinlite MT-24EX flash unit set manually on 1/32nd power, backed up by a Canon EOS 30D camera. Now the lenses are the same, but there are two of the flash units and two Canon EOS 7D cameras, which take much bigger images allowing a bit more distance between the subject and the lens. Even so, the field of view is still pretty small, around 36mm x 24mm.
On ISO 320 at f16 and 1/250th second seven or eight frames can be taken before the flash has to recharge, which takes only a couple of seconds. 1/250th is too slow to freeze many aculeates in flight but it is an acceptable compromise, allowing a good burst of well-lit images at an aperture affording respectable depth of field, i.e. ensuring that as much as possible of the subject is in focus. Manfrotto tripods are used where circumstances permit and cable releases are the norm on the cameras to prevent crowding the insects and allow full vision of what is occurring
Fossorial species that fly with prey or pollen are among the toughest to try and photograph near the nest, even when they are fairly large such as Crabro species, Cerceris species, some Andrena species or the Hairy-legged Mining Bee (Dasypoda hirtipes). Placing the camera on the ground pre-focussed on a point above the burrow rarely works since the bees do not all fly straight in - Andrena cineraria is a fair-sized one that usually does - and the wasps rarely fly in at the same angle. Moreover, providing the prey is not too large they tend to enter at pretty high speeds. Crabro scutellatus is the fastest I have seen, but Cerceris arenaria is not far behind.
One species that does not go at a breathtaking pace is Odynerus spinipes, which carries weevil larvae into its nest via a chimney with a diameter of 4mm or so. This is approached at a fairly consistent angle and since the speed is not too great there are definite opportunities and I photographed a female in flight, and in flight with prey, in 2006. In theory wasps ferrying full-grown weevils or almost mature instars should be easier to photograph but this is far from the case with Cerceris arenaria. However, lying on the ground and relying as much on luck as skill, I managed to photograph a female carrying a weevil in 2004. The wasp was well to the left of the image that resulted, a typical but not disastrous outcome - central positioning is a real bonus in any such photo but modern digital images are so large that even after cropping, a respectable portion large enough for an A4 print may be left.
Size of prey can be a crucial consideration but even though some of the solitary bee prey caught by Cerceris rybyensis are pretty substantial I have never managed to obtain an image of the process, just a female on her own in 2007. My only successes with Cerceris quinquefasciata in flight occurred that same year when a female caught presumed Sitona lineatus weevils, which were too large to allow her to go straight into her burrow at speed, instead forcing her to land gently close by and quickly walk in. In the same vein, the Beekiller Wasp (Philanthus triangulum) is not always too demanding since the nests sometimes are in banks rather than on level ground and the species does not fly at breakneck speed, in part surely because the Honey bee (Apis mellifera) prey quite frequently are almost as large as their captors. I managed several images of this species in 2009 but all were with the camera handheld rather than on a tripod owing to the nests being in a shallow bank of unstable sand above a fairly sheer drop of 20 feet. The same applied to Mellinus arvensis at the site. This wasp, which catches various types of fly, is one that is shown in flight more often than most in publications perhaps because it seems not to fly so fast as many others.
As a point of interest, Philanthus triangulum was one of a dozen species of wasp, mostly fossorial and also including Mimesa equestris and Lindenius pygmaeus, successfully photographed in flight with prey in the late 1950s by Gunter Olberg, probably the finest photographer of wasps there has ever been and a splendid naturalist to boot. Despite this, aerial nesters on the whole offer much better opportunities for in-flight photography than those using the soil since suitable places to build nests can be provided readily in a relatively controlled though still wild environment - the garden.
My first attempts at this were in 2005 at a friend's cottage in the Cotswolds where a number of Anthophora plumipes and Red Mason Bees nested in the walls. The former in particular is a large species and one that doesn't fly excessively fast, allowing enough scope for some reasonable results using only an early digital camera, a Canon EOS 10D, with its on-board flash and handheld to boot. Tripods make life much easier though. Setting them up and leaving them with one or more cameras carefully focused manually on the likely path an aculeate may use is feasible on private property but unthinkable in locations open to the public. My first successful use of a tripod for a small species was in 2007 on logs in my garden at Reigate. The wasp was Rhopalum clavipes, measuring 7mm, and while the results left room for improvement in terms of sharpness they were acceptable.
With recording for the book Wasps of Surrey just about complete and no more images required for that publication, I decided to make 2010 a year in which to concentrate on in-flight images and the results were reasonably satisfactory. I purchased three boxes, one made by the Oxford Bee Company and two by the German company Schwegler. All three provided fairly large, flat faces, which made it relatively easy to place photographic equipment on the same plane as the nest entrances, an essential element of the venture since even if a wasp or bee that is heading straight for a nest doesn't necessarily approach at right angles the whole way, at some point, usually the last moment, she is almost certain to be square on. Whenever possible I placed a piece of fairly pale wood parallel with the flight path a few centimetres behind the nest entrance as a means of preventing a black background, which can be distinctly off-putting.
The species using the Oxford Bee Company box were, predictably, the Red Mason Bee and, less expected, the Eumenid wasp Ancistrocerus nigricornis. Through May and into June several female Red Mason Bees nested, completing nine burrows, but just one Ancistrocerus nigricornis was involved, with four burrows stocked over five days containing an average of six micromoth caterpillars in each of three cells. On the face of it, given the number of times the female bees and the wasp had to bring materials into their nests the opportunities for obtaining images were prolific. However, the fact that none of the subjects necessarily flew to the correct burrow complicated matters - the Red Mason Bees in particular sometimes seemed clueless about where they were going, including twice entering the wasp's current burrow and ending up in a noisy dispute with the resident. Ancistrocerus nigricornis also did not go straight to the correct burrow every time.
Using two cameras mounted on tripods on either side of the bee box and sitting close by with cable releases for the shutters I took getting on for 500 images of Red Mason Bees and Ancistrocerus nigricornis in flight with pollen, prey or moistened earth (usually sand), and in action at the nests. Neither species flew in particularly fast, which helped, but even with the cameras set back a little, giving a field of view of around 36mm x 24mm, they still passed in front of the cameras in the blink of an eye.
The vast majority of the results, obtained over a fortnight or so, were of little account, especially as the angle of approach was often wrong and my judgment of when to fire the cameras was not always 100% either. An obvious problem is that unless the wings happen to be above the abdomen there is a strong likelihood that a crucial aspect, be it face, pollen brush or prey, may be obscured. There were enough satisfactory results though to encourage me to continue with species using the other two boxes from mid-June onwards. One interesting observation was that Ancistrocerus nigricornis closed the nest with a much smoother finish than the Red Mason Bees, whose work was relatively uneven. At those boxes there was room for just one tripod. Heriades truncorum was the main focus of attention and provided plenty of good results from hundreds of attempts, though neither of the females approached the nest in a straight line more than a fraction of the time, often landing a few centimetres away and walking to the burrow. The approach with resin in the mandibles tended to be more direct than with pollen on the scopa, but could not be guaranteed. The field of view was 32mm x 21mm, meaning that the bee, which measures 8mm or so in length, could occupy a maximum of just one quarter of the space available when the photographs were taken. This was less than ideal but another necessary compromise.
Psenulus pallipes and Trypoxylon clavicerum, which also nested in the cut reeds from July onwards, are no larger than Heriades truncorum but proved slightly easier to photograph, since both usually went in at right angles to the nest at something less than full pace. Only one Psenulus pallipes was involved, completing two burrows - I ascertained that the second nest was being stocked by the same female by briefly blocking the entrance soon after she commenced operations on it, whereupon without hesitation she went unerringly to the first nest site. In full flow she took one prey item about every five minutes and as a bonus every so often she hovered momentarily in front of her nest before going in. Trypoxylon clavicerum was, or were since four burrows were completed, less speedy, averaging one spider caught every 20 minutes. The spiders, which were not from the same species, were all fully legged, with no amputation having occurred. Like Ancistrocerus nigricornis, but unlike all the other wasps I have seen flying to a nest, the Trypoxylon clavicerum used her mandibles to help grasp her prey en route and on one occasion she had the spider entirely in the mandibles when approaching the burrow.
By late August the boxes were no longer being stocked by any species but several beech logs I had positioned vertically close by in 2008 were. The same principles applied in taking photographs. The best opportunities were presented by Crossocerus annulipes and Ectemnius cavifrons. The former is small, only 6mm-7mm in length, and usually flies in with its bug prey much too fast to be caught properly on camera at a mere 1/250th second. However, a stroke of luck saw one of the six or so females of the species using the wood construct her nest, rather oddly, in the top of one of the logs rather than in the side. This meant that she was unable to fly in at right angles and had to slow down ever so slightly and adjust her flight path when close to the nest, giving a fraction of a second to take an image. Since she was catching up to 18 bugs a minute at the height of her activity she provided me with every chance but it still proved extremely difficult.
As with so many other of the subjects of this venture, the Crossocerus annulipes could come in from any angle with her prey, and the fact that the two Ectemnius cavifrons that nested showed consistency in this regard was a real plus. My experiences when photographing Ectemnius cephalotes with prey in 2005 and watching other species such as Ectemnius continuus as well as Ectemnius cavifrons indicated that at the speeds available to me members of the genus generally fly too fast to allow taking images without blurring. However, when getting close to the surface of the wood they usually lessen the pace slightly, sometimes lowering the abdomen to an angle of as much as 45 degrees to prepare for entrance. This knowledge provided me with an edge. The Ectemnius cephalotes nest was right by a public footpath so using a tripod was tricky but fortunately I managed to obtain several images, one of which was used on the cover of Wasps of Surrey.
This knowledge stood me in good stead with Ectemnius cavifrons in 2010, and with that species and Ectemnius cephalotes again in 2015. This was particularly gratifying given the brightness of the wasps' markings and of many of the prey, especially hoverflies. In passing, capturing images of either species leaving a nest is much harder work than may be imagined. Without the encumbrance of prey, they exit very fast and hardly ever at right angles to the nest. The same comment holds good for the other species photographed during 2010, especially Heriades truncorum, which sometimes flew straight out though usually upwards or downwards, and often walked for a centimetre or two before launching herself into space.
Be that as it may, the results with Ectemnius cavifrons justified the effort and, if anything, fired me to try even harder in future years. The result was that in 2011, assisted by placing significantly more tubes, stems and pieces of wood for the species to use, I managed to photograph eight species not dealt with adequately, if at all, before. Two of these were ones that had provided no joy in 2010, Crossocerus megacephalus and Pemphredon lugubris. Obtaining images of the former flying towards its nest in a block of beech wood carrying prey including Soldier flies was gratifying and locating a Pemphredon lugubris that unlike most of them tended to fly straight into her nest was also a bonus.
The species which were completely new to the nest boxes were the wasps Ancistrocerus trifasciatus and Symmorphus bifasciatus, and the bees Osmia caerulescerns, Hylaeus communis and Stelis breviuscula. Ancistrocerus trifasciatus nested in cardboard tubes alongside Ancistrocerus nigricornis and the behaviour was pretty well identical. Symmorphus bifasciatus nested elsewhere in the garden but allowed flight images when she came to the drilled brick in the Schwegler box to take mud for her nest from a Red Mason Bee nest presumably ransacked by a woodpecker. The technique for both wasps was exactly the same as for Ancistrocerus nigricornis in 2010.
The bees were rather more problematical. Two Osmia caerulescerns, which nested in the cut stems, were irregular in their activity, with often very long gaps between their returning carrying either pollen or leaves and some days when neither bee appeared. Satisfactory images were obtained though. Hylaeus communis nested in the brick and stems but this is a small species, only 7mm or so, and they did not fly straight into the nest every time. The fact that there were two broods helped reduce that problem. The toughest was Stelis breviuscula, since this is a parasite of Heriades truncorum and as such does not have a nest but instead trawls around looking for a suitable one to enter and lay her egg in. My method was to focus on a spot where there were two host nests and hope the parasite would fly into view briefly to allow a photograph to be taken. Almost miraculously this did indeed happen, something of a coup as well a pleasant surprise.
A late-nesting Rhopalum clavipes gave me the chance to improve on my images taken in 2007, though sadly not with her ferrying prey into the log. The other new species photographed were social wasps, Vespula germanica and Vespula vulgaris. Both spent time hunting among the heather in a border near the house and the results were respectable.
From 2012 to 2016 a lot more dead wood was added to my collection and various species were added too, including Osmia leaiana, Megachile ligniseca and Chelostoma campanularum. The Megachile ligniseca was a bonus since leaf-cutter bees are very photogenic in action and this species is a good size. In the soil, Crossocerus tarsatus presented some problems owing to its small size. In 2015 more than 600 nests were completed in the garden by over 20 species of solitary bee and wasp and in 2016 the figure rose to more than 700. With luck this plethora will continue, allowing even more photography.
Images © Jeremy Early. All rights reserved.
In 2013 I published My Side of the Fence - the Natural History of a Surrey Garden. Details may be found, and orders placed, via this hyperlink My Side of the Fence. In November 2015 Surrey Wildlife Trust published the atlas Soldierflies, their allies and Conopidae of Surrey, jointly written by David Baldock and me. Details are on this web page: Atlas.