ZADBI: Zurqui All-Diptera Biodiversity Inventory

giant sciarid fly

From a 4 hectare (about 9 acre) patch of cloud forest near San José, Costa Rica, a team of dipterists (fly researchers) have identified over 4000 species of flies, and extrapolations from this collecting suggest that the true total is 7,000 – 8,000. This incredible diversity was the result of the ZADBI project, funded by the US National Science Foundation. A team of talented and dedicated Costa Rican biologists sampled at the site, called Zurquí de Moravia, for an entire year, using several Malaise traps, regular light trapping, hand collecting and other methods. The resulting catch was prepared (mounted, pinned, slide-mounted, labeled) and sent to international experts on various fly families for identification.  The result was an eye-popping number of species, many of them new to science (especially in the families of smaller flies).

Leading the way were the gall midges, or Cecidomyiidae, which were represented by at least 800 species, almost all of the probably undescribed. During the course of the project, cecid expert Dr. Mathias Jaschhof actually described some of the new species, including those that belonged in a new genus he named Zadbimyia in honor of the project (there were 19 of the 22 species of this genus alone at Zurquí).

Second most species rich were the phorids, whose numbers were so large that I am still reeling from the effort of trying to understand their diversity (with help from Maria Wong, Giar-Ann Kung, John Hash, and others). The 400 species we identified is surely just part of what is really there.

There were hundreds of species of other families of flies as well, including lots of strange and wonderful new things. It will be years before we can finish identifying everything, but the project gives us some important insights into tropical Diptera diversity. It is also a testament to how effective a large group of dipterists can be, identifying thousands of specimens of one of the least-known biotas in the world.

I hope to bring you many more details about the results from the project, but for now, here are links to the two recently published papers:

Borkent, A., B. V. Brown, P. H. Adler, D. S. Amorim, K. Barber, D. Bickel, S. Boucher, S. E. Brooks, J. Burger, Z. L. Burington, R. S. Capellari, D. N. R. Costa, J. M. Cumming, G. Curler, C. W. Dick, J. H. Epler, E. Fisher, S. D. Gaimari, J. Gelhaus, D. A. Grimaldi, J. Hash, M. Hauser, H. Hippa, S. Ibáñez-Bernal, M. Jaschhof, E. P. Kameneva, P. H. Kerr, V. Korneyev, C. A. Korytkowski, G. Kung, G. M. Kvifte, O. Lonsdale, S. A. Marshall, W. N. Mathis, V. Michelsen, S. Naglis, A. L. Norrbom, S. Paiero, T. Pape, A. Pereira-Colavite, M. Pollet, S. Rochefort, A. Rung, J. B. Runyon, J. Savage, V. C. Silva, B. J. Sinclair, J. H. Skevington, J. O. Stireman III1, J. Swann, P. Vilkamaa, T. Wheeler, T. Whitworth, M. Wong, D. M. Wood, N. E. Woodley, T. Yau, T. J. Zavortink, and M. A. Zumbado. 2018. Remarkable fly (Diptera) diversity in a patch of Costa Rican cloud forest: why inventory is a vital science. Zootaxa 4402: 53-90

 

Brown, B.V., A. Borkent , P.H. Adler , D.S. Amorim, K.Barber, D. Bickel, S. Boucher , S.E. Brooks, J. Burger, Z.L. Burington, R.S. Capellari, D.N.R.Costa, J.M. Cumming , G. Curler, C.W. Dick, J.H. Epler, E. Fisher, S.D. Gaimari, J. Gelhaus, D.A. Grimaldi, J. Hash, M. Hauser, H. Hippa, S. Ibáñez-Bernal, M. Jaschhof, E.P. Kameneva, P.H. Kerr, V. Korneyev, C.A. Korytkowski , G. Kung, G.M. Kvifte, O. Lonsdale, S.A. Marshall, W. Mathis, V. Michelsen, S. Naglis, A.L. Norrbom, S. Paiero, T. Pape, A. Pereira-Colavite, M. Pollet, S. Rochefort, A. Rung, J.B. Runyon, J. Savage, V.C. Silva, B.J. Sinclair, J.H. Skevington, J.O. Stireman III, J. Swann, F.C. Thompson, P. Vilkamaa, T. Wheeler, T. Whitworth, M. Wong, D.M. Wood, N. Woodley, T. Yau, T.J. Zavortink, & M.A. Zumbado. 2018. Comprehensive inventory of true flies (Diptera) at a tropical site. Communications Biology (Nature). 

 

tachinid fly, photo by W. Porras

 

 

 

 

 

More on greenbottle flies

Questions remain about differentiating greenbottle flies (Lucilia spp.) from muscid lookalikes (Neomyia) in iNaturalist photos. I wrote previously that the distribution of the tomentum (silver reflection caused by microtrichia) would help; below are some further photos to aid in this process.

Male Neomyia

Male Lucilia

Female Neomyia

Female Lucilia 

In each instance, you can see that the tomentum in Lucilia extends much farther dorsally on the frons than in Neomyia.

There was also the tougher question about separating Lucilia cuprina from L. sericata in photos. This is more difficult, because the main character used by the calliphorid expert, Terry Whitworth, is on the back of the head, usually out of sight in photos. There are supporting characters, however, on the dorsum (top) of the thorax. If you look at the humeral callus (“h”, below) and the notopleuron (the triangular area marked “n”), there are a  few tiny setulae along the posterior margins. In L. sericata, there are usually around 6, in L. cuprina 2-3. If you can see that kind of detail in an iNaturalist photo, then you can make a call; otherwise, I suggest leaving them as “Lucilia sp.”

sericata thorax

cuprina thorax

How big is Megaselia?

The phorid genus Megaselia Rondani is possibly one of the biggest genera of insects. I say “possibly” because we still don’t know if we should consider all the species currently classified as Megaselia as belonging to a single group. Recently molecular work by Sibylle Häggqvist seems to indicate that most of Megaselia is monophyletic (forms a natural group), but much more work needs to be done.

Just for fun, I made a graph of the described species of phorids, showing the number of species per genus.  This is a representation of how big Megaselia is within the Phoridae. Note that this is the number of DESCRIBED species, and does not take into account all the unknowns. 

For the phorid researchers: does this treemap surprise you? It did me, a little.  I thought Megaselia would be even more dominant. Presumably, once we get to do all the world’s fauna, Megaselia will end up being closer to half of the family.

Or maybe not.

 

Facing greenbottle flies

In the recent “City Nature Challenge” (an iNaturalist event co-hosted by our museum, LACM), the question was indirectly asked of me “how do you separate the common greenbottle flies,  classified in the calliphorid genus Lucilia, from the muscid lookalikes in the genus Neomyia that can also be green.” Especially, how would you tell them apart from the photos in iNaturalist, which are often not of stellar quality. My answer is to look them in the face. In both genera, the distribution of tomentum (microscopic extensions of the fly’s cuticle; not ‘hairs’ or setae, which are socketed) is different. The tomentose parts of the frons reflect light as silver or white, so this is often easy to see in photos.

You can see in Lucilia, on the left, that the tomentosity extends much farther up the frons, especially medially (in the middle) than in Neomyia, on the right, which has a dark, tomentosity- free section of the head.

You can see that the attached photo from iNat is a Lucila.

 

 

 

New fly book!

In spite of the importance and diversity of flies, there are not many books, especially popular books, devoted to them. Perhaps the reasons are obvious: they are are small, often inconspicuous, and sometimes annoying (or worse). Often, when I tell people I work on flies, they skeptically and sometimes disgustedly repeat the word “flies?” as if that is the most unthinkable group of insects to which one could devote their attention.

Certain people, the hardcore dipterists, however, truly love flies, and some few can convey that affection to the public. It takes a mixture of delight, quirkiness, and sometimes downright dark humor to unlock the secrets of this hidden world and make it available to the non-scientist. Few have tried, but one of the most successful to date is the British dipterist Erica McAlister, whose recent book “The Secret Lives of Flies” is required reading for all fly aficionados.

I am just returning home from the Entomological Society of America meetings, where I had the great pleasure of hanging out with some of the members of the “Diptera tribe” who came to deliver talks, exchange ideas, meet the newest crop of students, and discuss future plans. I got to spend some time with Erica, as well as Vladimir Blagoderov, Brian Wiegmann, David Yeates, and Riley Nelson, and many of the upcoming students. All are accomplished dipterists, yet only Erica has written a book that will be of wide interest to non-dipterists. Buy a copy- you won’t regret it!

Syrphid migration

Some flies migrate, just like some birds and butterflies, but it is rarely recorded. In fact, this might be the first record of a flower fly (or hover fly) migration for North America!

It took place on April 20th, 2017, at beautiful Montana de Oro State Park. I was walking with my dear friends Marianne and Gary Wallace  when Gary said “look at all those bees!” I looked, and realized that they were not bees, but flower flies, thousands of them. They were flying against the wind, in a northward direction, not stopping at all. I actually got some video of this migration; have a look.

 

[embedyt] https://www.youtube.com/watch?v=30u0dXZ0bjo[/embedyt]

 

The flies look like bullets going by. We didn’t have any collecting gear (duh, it’s a park), but we tried knocking some down, to no avail. They were moving fast and avoiding contact. I tried taking some stills, and this is my best (bad) photo:

Why would syrphids migrate? These flies feed on aphids as larvae, and aphids are found on tender new growth. California’s rainfall comes during a short time in the winter, and then things dry up. An hypothesis is that as the more southerly vegatation dies off from lack of water (and thus also the aphid supply dwindles) the flies move northward looking for green, still tender fields, well stocked with their aphid food.

This was something I never expected to see, but wanted to record so others could keep their eyes open for it in the future. I’ll be back next year looking for them at the same time of the year, hopefully to learn which species they are.

Urban phorid flies in Los Angeles, California, USA

Last week Emily Hartop and I published a new paper on the phorid fly fauna of Los Angeles, in the journal Urban Ecosytems (get the full text here). This is the culmination of years of work by all of the BioSCAN team, in getting the project up and running, sorting and identifying 40,000+ flies (mostly Megaselia, mostly the work of Emily), and looking for patterns in the community.

Lisa Gonzalez of the BioSCAN project and Joe Hogg, one of the site hosts, in Joe's backyard.

Lisa Gonzalez of the BioSCAN project and Joe Hogg, one of the site hosts, in Joe’s backyard.

The major findings for us is that the Los Angeles urban fauna consists of about 100 species (43 of which were new to science and described by us for the first time), but individual sites (mostly backyards) had from the mid-twenties in the area just west of the museum to 83 species in Glendale, right up against the mountains.

The commonest species by far is Megaselia agarici, a fungus-feeding species possibly introduced from elsewhere. It is sometimes a pest in commercial mushroom farms, but here they are extremely common in “wild” Agaricus mushrooms that we have found in the city and brought indoors for rearing the flies. This species is so prevalent, that it was found in every one of our 30 sites (as were a few other species), and in some sites it made up over 70% of the phorids captured. Many of the other commonest species in the study were also mushroom feeders, such that at least 40% of our collected phorids have this lifestyle.

In contrast, our most diverse site had many more species that do other things, especially those that parasitize ants (including the infamous “ant-decapitating flies” I have written about elsewhere). These ant-associated species are completely absent from the downtown area, almost certainly because the introduced Argentine ants have completely eradicated the other species of ants formerly found here. It is difficult to know how much damage was caused by the introduction of “argies” here, but it was a catastrophe for phorid flies.

Many other results are in the paper, and I urge all interested fly aficionados to read it. The next step will be to associate these diversity data with variables of climate and urbanization to see if we can find some correlations that make sense. We are far from finished working on L.A. phorids! But the outstanding remaining question about this phorid community we have here is whether it is representative of other urban phorid assemblages, here in California, here in North America, and throughout the world. Are all urban sites dominated by fungivores? Do they all have a superdominant species like M. agarici (or maybe even M. agarici itself)? We just don’t have any data to know the answers yet.

Additionally, one of our next studies will be to look at other groups of insects (including other families of flies) to see how they are distributed in the city. I expect that there will be some major differences among groups, attributable to small-scale differences among sites. One study (Avondet et al. 2003) in Ohio, looked at, among other things, the distribution of dumpsters to help predict the diversity of Drosophlia “fruit flies” (Drosophilidae, better referred to as vinegar flies or pomace flies) in the urban landscape! Probably the diversity of plants, flowers, amount of hardscape vs yard, mulch versus grass, and so on will greatly affect the number of species at a site, but our ability to predict this is still in the infancy stage. Stay tuned for more…

Cited article:
J.L. Avondet, R.B. Blair, D.J. Berg, and M.A. Ebbert. 2003. Drosophila (Diptera: Drosophilidae) response to changes in ecological parameters across an urban gradient. Environmental Entomology 32(2): 347- 358.

The white-tailed phorid, Megaselia albicaudata

It is amazing to me that some phorid flies are found in multiple environments around the world. Less surprising are the scavengers that live mostly in human-built structures or that need scraps of our civilization to thrive. Usually, these scavengers are well-known to us, through their large numbers and occasional nuisance status. But what about the”white-tailed phorid” (I just made that name up, based on the Latin name!), an obscure but seemingly ubiquitous fly?

There is no question that this 1.5 mm long fly is well-named: the male genitalia are a shocking white in comparison to the rest of the rather dingy brown body. It was described in 1910 from England, and is well-known from other parts of the world such as Finland, Israel, and China. It is also found in the two big fly inventories that I am part of: the Los Angeles County Natural History Museum’s urban biodiversity project called “BioSCAN” and the Costa Rican “ZADBI” project in a tropical cloud forest.

How can a species live in such disparate climates and habitats? Nobody knows, just as nobody knows what the larvae of this fly eat. I have looked at specimens from both North America and Costa Rica, and I am sure they are morphologically the same species, not two species masquerading as one doing different things in different places. It is one of thousands of Megaselia worldwide that we know nothing about, but discovering phorid lifestyles is part of our* phorid research effort. Obviously, we have a long way to go when such a widespread species remains a mystery.

*Note, when I say “our”, I mean the few people worldwide actively pursuing phorid research, including our group here at LACM, consisting of Emily Hartop, Lisa Gonzalez, and me.

Fly porn- what can we learn?

[embedyt] https://www.youtube.com/watch?v=-aNlk2rvtWE[/embedyt]

Take a look at this video of a pair of Phora I observed mating in Central California.

Apart from the humorous aspects of the “action”, including the hilarious way they kick each other to separate, there is actually some good science here.

Notice the movement of the male’s hind legs (how could you miss it!). Males of many species of Phora, and some other phorid genera, have tufts of setae (thick, hairlike processes) at the base of the hind femora. Presumably, the male uses them to contact the female during sperm transfer as a sort of courtship to persuade the female to use his sperm. Female flies often mate with multiple males, and can control to a remarkable degree whose sperm she uses. This is because the sperm from any mating is shunted to a storage organ called the spermatheca, where it is kept until needed. Females have been shown to have remarkable control over the sperm in a spermatheca, shunting unwanted or lesser quality sperm (as judged from mating stimuli received during copulation) to the back, or even expelling it. Thus, males not only have to copulate with females, but also have to convince them to use their sperm when they later lay an egg. Male signals, like the stimulation of hind femoral setae, are what convince her of the worthiness of the sperm.

Also of note is the position of the forelegs of the male, flat on top of the female’s scutum (“back”). Male Phora have thickened, flat foretarsi (the short segments at the end of the leg) that probably have specialized setae as well, but, well, nobody has really looked. It would be an interesting study for someone with spare time and access to a scanning electron micrograph.

The final aspect of the mating is the fastidious cleaning that the female subjects herself to. In insects, as in all animals, close contact with another individual always involves the possible transmission of parasites. For small flies, what we note most frequently, is the presence of mites, which can almost completely coat an unfortunate victim. This female spends a lot of time sitting on a leaf, exposed to predators (and photographers), while cleaning herself. It must be worth it, to keep away those mites!

This post is dedicated to my colleague Erica McAlister at the Natural History Museum in London, to give her something to read on a Sunday morning!

 

 

A new type of ant-decapitation in phorid flies!

Dohrn longi plateOkay, I literally have been waiting for years to write this story! Now it is published, and you can read about our amazing (at least to me) discovery.

Most phorid flies that decapitate their hosts do so by injecting an egg into the host ant; after hatching from the egg, a larvae feeds in the head, eventually causing the head to fall off, sometimes before the rest of the body stops moving.

In species of the Dohrniphora longirostrata group, however, the female flies are attracted to injured Odontomachus ants, which they decapitate themselves, and haul off the head to feed on its contents or to lay an egg. We have seen this in several countries, and on many occasions, so there is no doubt in my mind that this is a specific behavior to these flies. They are not attracted to injured ants of other types, only Odontomachus.

Here is some video of the action!

[embedyt] https://www.youtube.com/watch?v=GQbtXG48azU[/embedyt]

(video by Kate Lain)