Heriades truncorum is only 8mm long but does not fly excessively fast when loaded with pollen

Colourful female Tawny Mining Bees (Andrena fulva) are among the best subjects for in-flight photography near the nest Anthophora plumipes is a large species that sometimes approaches its nest, often in building walls, at a relatively slow speed The Red Mason Bee (Osmia rufa) is another striking species that nests around houses
A fluky shot of a Blue Carpenter Bee (Ceratina cyanea), a tiny species nesting in stems Nomada bees are cleptoparasites and many fly at a much slower speed than their hosts, quartering the ground for occupied burrows Colletes succinctus takes pollen only from Heather and since the flowers are small there is a lot of flitting to and fro
Male Bombus muscorum: Bumblebees are large enough, and fly between blooms enough, to make photographing them on the move relatively easy The narrow chimney at the nest entrance of Odynerus spinipes provides opportunities for catching them in flight Another lucky shot since Cerceris arenaria usually approach their nests at great pace
The size of the prey taken by Cerceris quinquefasciata in 2007 made it impossible for her to fly straight in to her nest in the usual fashion The Beekiller Wasp (Philanthus triangulum) is often unable to fly fast on account of the size of the Honeybee prey Mellinus arvensis females are probably the wasps most frequently photographed in flight
Rhopalum clavipes is only 7mm in size This female Ancistrocerus nigricornis, heading for her nest in a bee box with a micromoth caterpilalr, was a revelation in 2010 The same Ancistrocerus nigricornis with moistened sand to close one of the nest cells

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 wingbeatsThe set-up used for photographing Osmia rufa and Ancistrocerus nigricornis in 2010 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 movingOsmia rufa female heading for the bee box with a reasonably neutral background inserted 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 difficult to determine. They are Ancistrocerus nigricornis, Odynerus spinipes, Psenulus pallipes, Crabro scutellatus, Ectemnius cavifrons, Ectemnius cephalotes, Rhopalum clavipes, Crossocerus annulipes, Crossocerus megacephalus, Trypoxylon clavicerum, Mellinus arvensis, Cerceris arenaria, Cerceris quinquefasciata, Cerceris rybyensis, Philanthus triangulum, Colletes hederae, Andrena cineraria, Andrena fulva, Dasypoda hirtipes, Megachile ligniseca, Anthophora plumipes, Ceratina cyanea, Osmia rufa, Heriades truncorum and Bombus lapidarius plus the cleptoparasites Nomada fucata and Nomada signata.

Osmia rufa's bright colouring makes the species very photogenic

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 is a Canon 1D Mark II digital camera, which on motor drive takes eight frames a second, with a 100mm or 180mm macro lens, 12mm extension ring and Canon Macro Twinlite MT-24EX flash unit set manually on 1/32nd power. The close focus on the 100m lens offers a field of view of only 24mm x 16mm, and on ISO 200 at f16 at 1/250th secondHeriades truncorum were very obliging but they did not always fly straight towards the burrow 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. Even so, at the closest point only 5mm or so is in focus, which makes it all a bit of lottery. Back-up is provided by a Canon EOS 30D camera, which has 'on-board' flash. 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 isA rare if not unique image of Heriades truncorum with resin in a ball and lengthy tail that must have made flying tricky 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 sometimesPsenulus pallipes sometimes hovered briefly outside her burrow, perfect behaviour for attempting to photograph her are in banks rather than on level ground and the species does not fly at breakneck speed, in part surely because the Honeybee (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.

Crossocerus annulipes is tiny and usually flies in too fast to allow photography but this female gave me a chance because of where she positioned her nest

My first attempts at this were in 2005 at a friend's cottage in the Cotswolds where a number of Anthophora plumipes and Osmia rufa 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 planeAnother small species, the elongate Trypoxylon clavicerum luckily is not the fastest of fliers approaching the nest 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 (Osmia rufa) and, less expected, the Eumenid wasp Ancistrocerus nigricornis. Through May and into June several female Osmia rufa 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 Osmia rufa 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 byFemale Ectemnius cavifrons showing the helpful tendency to lower the when about to enter the burrow with cable releases for the shutters I took getting on for 500 images of Osmia rufa 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 Osmia rufa, 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. TheFemale Ectemnius cavifronsshowing using a flatter, faster trajectory 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 spiderFemale Crossocerus megacephalus flying very fast and sadly without prey 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 principal nester was Pemphredon lugubris, which in my experience only rarely flies straight into the burrow. Other species included Crossocerus megacephalus, which I managed to photograph a few times as she flew towards her burrow but in only average fashion owing to her speed. Unfortunately not once did she have prey. Better 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 cavifronsThe boxes which had led to so much photography in the summer looked rather different in December 2010 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, which was particularly gratifying given the brightness of the wasp's markings and of many of its hoverfly prey. Over the space of ten days in September 2010 I probably spent 20 hours watching and photographing the two females, using the equipment outlined above and one tripod. Unlike the smaller black species they could be seen coming towards the nests, which helped considerably in preventing my missing shots. In passing, capturing an image of either female leaving her nest was much harder work than may be imagined. Without the encumbrance of prey, both exited very fast and hardly ever at right angles to the nest. The picture below was the only success story here. 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.

Unencumbered by prey the female Ectemnius cavifrons left her burrow at high speed, usually at an angle rather than straight

As a postscript, the heavy snowfall of December 2010, pictured above right, left the boxes where so much of my photography just a few months earlier had taken place looking as if life within would be out of the question. It is part of the joy of studying nature, though, that species can cope with such conditions and emerge the following spring or summer as flying adults to continue the process.

Images © Jeremy Early. All rights reserved.