New paper: The whimsical long-tongue fly and its favourite colour.

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The flowers on one of these plants conceal drops of sticky nectar. The other is a cheating orchid, presenting empty flowers and false promises. Can you tell which is which? Even if you knew which one carried nectar, how can you tell the difference between them? The two plants might look a bit different to human high-res optics, but now try blurring your eyes. Pretty similar, huh?

What about this pair?

Screenshot 2018-10-30 15.09.50If it’s difficult for our brains and eyes to discern the difference between the flower with the reward and the one that’s falsely advertising, then what hope does a nectar-hunting fly with low resolution compound eyes and a smear of a central nervous system have?

Specifically, I’m talking about this fly…

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If this fly looks embarrassed, its because it has orchid pollen stuck to its face.

Until now, you probably thought lion, or elephant, or rhino were the most impressive animals roaming the grasslands of southern Africa. Well you’re wrong, and it’s ok to change your mind after seeing the majestic long-proboscid fly of South Africa. There are several species of these magnificent beasts, and this one is named Prosoeca ganglbaueri.

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That giant proboscis hanging from its face is a tool crafted by evolution for sucking nectar from the bottom of long flower tubes, and it can grow as long as 5 cm (which is longer than the fly’s own body length). Unlike butterflies who coil their proboscises, the long-proboscid flies simply hinge the instrument down, tucking it away underneath their bodies to trail out behind them. And this species isn’t even the most extreme: proboscises in Moegistorhynchus longirostris get up to 8 cm!

Sometimes handling that long instrument can be a challenge…

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In some areas of South Africa, P. ganglbaueri is the only creature capable of extracting nectar from flowers with very long floral tubes, and because of this it has become the exclusive pollinator for 20 species of plant. Altogether, the long-proboscid flies as a group bear the great responsibility as the only pollinator for approximately 130 species of plant, making them a truly important creature for the ongoing survival of many South African plants.

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Figure 1 from Whitehead et al. (2018): Prosoeca ganglbaueri feeding from a variety of nectar sources. (a) Zaluzianskya microsiphon, (b) Scabiosa columbaria, (c) Agapanthus campanulatus, (d) Dianthus basuticus.

An interesting fact about flowers that are pollinated by long-proboscid flies, is that most of them are pink, or white, or some variation in between (with one blue exception). This strong colour preference is a critical feature directing the evolution of the cheating orchid flowers introduced earlier. For a deceptive orchid to attract this fly, the orchids’ flower colour must match the flies’ colour preference, or the mimicry simply won’t work.

In my recent paper, we asked whether the colour preference of flies was something that they learned, like we learn to associate that perfect golden-brown hue of fried food with a mouth-watering culinary experience, or if it was instead a more hardwired innate response, like a moth drawn to a lamp. The answer is important for understanding ultimately what is driving the evolution of false advertisement signals in mimic orchids. So, for example, if flies had an innate bias to pink or white, then cheating orchid flowers would evolve to match that bias, in the same way that any good advertisements are designed to appeal to the fundamental desires of its audience. On the other hand, if flies learned to associate nectar reward with certain colours, their preference should be determined by the colour of their local nectar diet. Under the learned scenario, orchids should be evolving to match local flowers’ colours, not any intrinsic bias of the fly.

To test this, I took advantage of just how easy it is to bamboozle these flies. With a home-made artificial flower, painted to match the pink and white flowers visited by the fly, anyone can fool a fly into attempting to feed. So I mounted a pink and a white model to my “interview stick”, and travelled across the rugged Drakensberg Mountains to interview various populations of flies. In each location, I recorded whether the local flies preferred probing the pink or white model flower, as well as the colour and species of flower that the flies were using for nectar there.


The results were clear. Flies used to feeding mainly on pink flowers preferred the pink model. Flies that fed mainly on white flowers preferred the white model. And flies that fed on both pink, white, and violet flowers, showed no clear preference between pink and white.

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Figure 3 from Whitehead et al (2018): Pink-white preference for flies at seven sites. The x-axis shows colour preference, with pink on the right, white on the left. Measured preference at seven sites is represented, with the colour of local nectar sources depicted in the small pie charts.

This tells us that the flies are very flexible in their preferences, and the strong implication is that these flies are learning to associate colour and reward. A further result showed that as the variation of colours flies fed from increased, this made them less choosy in the pink-white preference choice. So the bottom line is that the colour of their local nectar-buffet strongly controls a fly’s colour preference.

What does this mean for orchid cheats? Well, the colour of nectar cheats is all important, and what matters most for the success of a deceptive orchid is the colour composition of the surrounding nectar-rich floral community.

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Post-script:
Still wondering about which flowers in the opening images were cheats, and which had nectar?

In both cases the deceptive orchid is on the left. The first image features Disa nivea (left), and Zaluzianskya microsiphon (right), the second features Disa pulchra (left) and Watsonia lepida (right).

Reference:

Whitehead MR, Gaskett AC, Johnson SD. (in press) Floral community predicts pollinators’ color preference: implications for Batesian floral mimicry. Behavioral Ecology 

Bumping into old floral friends, and pollination with a hug.

Rare plants nurseries are like second hand bookshops. It’s always so tempting to browse on the off chance you find that little treasure. I recently visited a charming rare plants nursery in Mt Macedon (boutique-y town outside Melbourne, Australia) where I discovered these for sale:

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Hello old friend! (Hesperantha coccinea)

The last time I saw this elegant iris, it was flowering on stream banks 10,000 km away in the Drakensberg Mountain range in South Africa. There in its natural habitat, it is pollinated in some areas by a very special butterfly: the Mountain Pride (Aeropetes tulbhagia). In other places, it is pollinated by the amazing long-tongue fly (Prosoeca ganglbaueri). The two forms are a wonderful example of “pollination ecotypes”, where different populations are undergoing adaptation to their unique pollinators. The fly-serviced ones are a pink hue with narrow petals, while the butterfly-pollinated ones are much redder with broader petals.

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Hesperantha coccinea at home in South Africa with its pollinator (Prosoeca ganglbaueri).

Fast forward two weeks, and I’m home walking the dog in my quite unremarkable Melbourne suburb, when who should I see?

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Hello old friend! (Diascia sp.)

It’s winter here, with very little in flower, but these brilliant little pink blooms volunteering themselves from underneath a fence in suburban Melbourne really made my day. The last time I saw a Diascia, it was growing amongst the boulders on creek beds and on cliffs in the Drakensberg Mountains. These are Diascia, or “twinspur” and its this common name that alludes to their fascinating pollination story.

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Hug-pollination by oil-collecting bee (Rediviva sp.) in Diascia.

Diascia have two spurs on the back of the flower, which is distinct from the usual arrangement of a single nectar-spur. The difference is that these flowers don’t reward pollinators with sugary secretions, instead they provide oil to specialised oil-collecting bees in the genus Rediviva. The bees use this oil to line their nests and provision their young. In order to collect the nectar, they must reach deep into the twin spurs with their lanky forelimbs, and comb it out. In so doing, they effectively hug the reproductive parts of the Diascia flower and effect pollination.

In Spring, I plan to take some cuttings from this little Diascia. Keeping species with special personal significance is a deeply satisfying part of cultivating plants. A plant can be kept like a souvenir or memento marking a time in one’s life, just like a photo or trinket. But plants have an advantage over these inanimate reminders. Because biological reproduction requires the physical donation of part of the mother’s cells to the daughter cells, my keepsake plant can be viewed as a physical part of the plant that appears in my fond memory. If I could see in four dimensions, I could literally look down the line of cell-divisions all the way back to where the Hesperantha in the nursery physically intersects as the same individual with the Hesperantha I observed flowering in the Autumn sun of the Drakensberg Mountains in South Africa.

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The Drakensberg Mountains, South Africa, Autumn 2014.

 

Plant pollinator interactions in the South African flora

The slides from my recent departmental seminar at the ANU are below.

The first half of the talk concentrates on plant-pollinator interactions, floral guilds and floral evolution. The second half is a slideshow of vistas, creatures and plants I encountered in my work.

My bruised human ego

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This is the best photo I got of a group of baboons who gave me quite an experience the other day.

On a sandy fynbos trail, I rounded a corner obscured by vegetation and came abruptly face to face with a troupe of seven of these creatures. The closest member was only 3 metres from me. All of them were stopped, standing or sitting,  looking at me as I did the same. My first reaction was one of awe, these creatures are impressively muscular and intimidating up close. One of them, a very large male, was wearing a radio collar. My second instinct was to take advantage of the photo opportunity, but my camera was in my backpack.

My only close experience with monkeys comes from Indonesian macaques, and extrapolating from the damage these ones wreak on tourists’ belongings I was not keen to get the baboons interested in anything I owned. I was also aware that some baboon troupes in the Cape have a reputation for raiding. Bins, bags, picnics, cars, houses are all fair game. They have overcome their fear of humans and are now a famous nuisance requiring full time management.

My bag therefore remained zipped and in place on my back. I raised my arms and hissed, to try and persuade them off the trail. One of the leaders began to advance on me, and the others stood up to follow suit.

Finally, I was forced to concede the path to the baboons. I back-stepped into the bush beside the track, allowing them a 2 metre thoroughfare which they calmly took in an orderly and nonchalant fashion. Only after passing me, when their backs were exposed, did they pick up speed into a quick trot for a dozen metres to put some distance between us.

 

Red Hill fynbos track

Kleinplaas Dam fynbos track

 

A most engaging mantid

Recently, I was fortunate enough to spend eight days in Ndumo Game Reserve, where for several hours a day I remained perched above clusters of large flowers smelling rather like a long-drop toilet. Tagging along as help on a study of Stapelia gigantea promised to be a chance to see a new South African biome and the wonderful creatures that come along with it.

The carrion flower (Stapelia gigantea, bottom right) in rural Zululand aloe country.

Driving to and from the field site every day we would encounter giraffe, wildebeest, nyala, impala and warthogs going about their daily activities. At dusk we’d sit in a bird hide, count waterfowl and watch crocodiles cruise on by. Our nights were serenaded by the wailing bush baby, the guttural grunting of wildebeest, the booming-bass of hippos and occasionally the manic whinny of a hyena, while the porch light drew in a bewildering buffet of invertebrate curiosity.

Croc on dusk, silently sweeping past the bird hide.

But perhaps the most endearing animal found all trip was one of the most captivating mantids I have ever seen. She is a cryptically coloured Hymenopodidae, belonging to the same subfamily as the spectacular orchid mantis. Unlike other mantids I have encountered she is very easy to handle and shows no desire to flee the hand or captivity. She is a voracious feeder and any moth or fly introduced to her enclosure scarcely lasts 5 minutes before straying too close to her raptorial forelimbs. On her second night in our field accommodation she had already laid a small ootheca.

She also displays a charming and unusual shadow boxing routine complete with weaving, jabs and feints.

Edit: I have since learned that she belongs to genus Oxypilus, a group of mantids called “Boxer mantis”, for reasons made obvious in this video. (Thanks Mantidboy for the ID).

The above video was shot with a Canon 500D, Canon 100mm f/2.8 macro in an improvised stove-top studio. A piece of white paper provided the background, the camera was stabilized on a bag of rice. This left my hands free to experiment with the lighting, provided by a cheap head torch. 

Mount Gilboa’s meadows.

This has been my sometimes workplace for the last two weeks:

The slopes of Mt. Gilboa. Watsonia densiflora in the foreground.

The slopes of Mt. Gilboa. Watsonia densiflora in the foreground.

To catch pollinators in action you need fine weather. On those days when the skies are clear and there’s little more than a gentle breeze in the air, Mt Gilboa is an exciting place to be. Gleaming green Malachite sunbirds chase one another between aloes, eagles and vultures wheel overhead, a startled bush buck bounds down the slope and out of view.

On these days the flowering veld is humming with the noise and motion of uncountable beetles, bees, flies and wasps, flitting, buzzing, mating and feeding. Protea heads crawl with furry monkey beetles, massive grasshoppers zoom by on the wing and bees of varied colour, shape and size forage diligently.

The flowering veld

The flowering veld

I come here to collect long tongue flies. As you prowl among the Watsonia inflorescences you first hear the telltale loud buzz, then look for the hovering fly probing a flower with its long proboscis.

Philoliche aethiopica foraging on Watsonia densiflora

Philoliche aethiopica is a specialist forager on Watsonia densiflora. This fly’s thorax is completely covered in pollen.

Netting the flies is not too difficult—they are lazy fliers. Keeping them alive in my flight-cage back closer to sea level has proved to be the big challenge. With the season wrapping up for this site, I’m unfortunately looking at the possibility of coming away with little more than just jars of dead flies.

Watsonia lepida, common veld iris and long tongue fly host plant.

Watsonia lepida, common veld iris and long tongue fly host plant.

Despite the setback there are other research avenues to pursue as the Summer field season unfolds. The luxury of a long field season is one factor that makes this veld such a productive place to study pollination.

Test post: Captive fly video

Currently in South Africa, my time right now is largely being spent on catching flies, planning to catch more flies and working out how to keep them alive and happy in captivity. The poor little video below is a quick capture of what I wish all my captive flies would do—buzz around and visit flowers like they’re just hanging out back in the veld they came from.

More on the fly project to come in the near future.

 

I hope to use this space in future to update on research progress, life in South Africa and occasionally sound off on things of a biology, botany, entomology and overall scientific nature.

 

Thanks for looking.