Armchair biodiversity discovery during COVID

With the COVID-19 pandemic, many of us are working at home, instead of in the field. In spite of this, I think that I have made a taxonomic discovery that is important for phorid classification.

My friend, Lonny Coote, recently wrote to me, saying he would help me with some entomological questions if I “describe[d] a second species of Cootiphora”, a genus I described and named for him in 1993. Since then, the genus Cootiphora has languished in obscurity. I have seen only three specimens: the holotype from Tinalandia, Ecuador, a second specimen from La Selva Biological Station, Costa Rica, and a third specimen from Bastrop State Park, Texas (near Austin).

The genus Cootiphora is known only from these three female specimens. They are extremely distinctive because of their cockroach-like, limuloid body form, their reduced, triangular wing rudiment, and their pointed, parasitoid-type oviscape.

The female Cootiphora specimen from Bastrop, TX

The holotype was collected (as the label states) on “the clubhouse windows”. Tinalandia was (is?) a golf course with some nice forest on the property that makes it a popular place with naturalists. It was on my first trip to the tropics, accompanied by Lonny Coote, John Acorn, and Jane O’Donnell, that I visited this resort and received the kind hospitality of Tina herself. One day, army ants (Eciton burchellii) invaded the clubhouse, and the windows were rife with flies associated with the ants. I had a great time collecting numerous phorids on that day, much to the chagrin of Lonny, who saw one of the wasps that he worked on during the first day of that trip, and not a single one thereafter. Amongst those phorids was the little female fly that I later described and named for him. I always assumed that it was associated with the army ants, but of course had no evidence of this.

Coote’s demand that I “describe a second species of Cootiphora” prompted me to look at the specimens I had gathered from Bastrop, Texas, just for fun. In doing so I noticed another peculiarity about the Bastrop fauna- I had described a new species of an unusual tropical genus known only from males from this park. Finding these flies, which I named Macrocerides (Tauricornis) taurocephalus, the name referring to the bull-horn like antennae of the males of all species in this subgenus. In my Macrocerides paper, I described two new species, one tropical (M. (T.) borkenti) and one nearctic (M.(T.) taurocephalus). I had never put this association together in my mind, but it seemed provocative that I would find these two tropical flies, one known only from males, and one only from females, at this site in Texas.

There was one thing left to check: I looked through the prepared, mounted, labelled material from that day in the clubhouse at Tinalandia, and found, to my delight, a single male specimen of M.(T.) borkenti.

Of course, this is still speculation, but if true, I would consider M.(T.) borkenti to be a synonym of Cootiphora angustata (the type species of Cootiphora); at least the subgeneric name Tauricornis to be a synonym of Cootiphora; Macrocerides (Cootiphora) angustata.

But wait, there’s more! Among the material I treated in my Macrocerides review were two further species, M. abaristalis Borgmeier from Costa Rica and M. attophilus Disney from Argentina. Both of these might be further species of Cootiphora, but most interesting is the latter, which was collected in association with leaf-cutter ants. This is significant, because one of the most distinctive things about the Bastrop State Park site back in Austin, Texas, is the large number of Apocephalus wallerae specimens also in the Malaise trap samples. This species is a parasitoid of Atta texana, whose colonies are common in the area (according to Alex Wild of the University of Texas, Austin, who kindly corresponded with me about this). Thus we have another hint of where to look for further Cootiphora specimens- in association with this leaf-cutter ant.

               Finally, I have some serious reservations about the monophyly of genus Macrocerides, discussed in my review of the genus. There is little to link the type species, M. curtifrons Borgmeier, with the species of subgenus Tauricornis, other than the moderately modified antennae. If further research indicates that these species are not closely related to M. curtifrons, we will be free to use Cootiphora as a full genus name again, something that I think would please my friend. At any rate, I think I have fulfilled his request: I speculatively have “found” not one, but two more species of what would be subgenus Cootiphora!

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)

Flyobsession back online

After transferring from WordPress.com to my own site, flyobsession is back! Look forward to more stories and photos of our two-winged friends.

Pictured here is a male Adelopteromyia, a common phorid fly from the ZADBI project in Costa Rica (photo by Inna Strazhnik). The females of this genus are brachypterous (have reduced wings) and are found in army ant colonies.

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ps. This image looks much better full size- click on it to see.

Diptera blitz continues

Uber-collector Wayne Mathis

Uber-collector Wayne Mathis

The 30 participants in the Zurqui all Diptera biodiversity inventory in Costa Rica have been pushing back our veil of ignorance about the fly fauna of tropical cloud forests. So far, we have spent two full days at the site (Zurqui de Moravia, 20 minutes north of San Jose), and one full day at our partner institution INBio. New species and even new genera of flies are being found everywhere we look. Some examples:

Collaborators Dr. Greg Curler and Sergio Ibanez have found a strange new type of psychodid (moth fly) that holds its wings unlike any other they have ever seen. Most moth flies hold their wings either flat over the body, or roof like over the body; this one inverts its wings over the body so that they make a trough. Significance? Who knows, but it is amazing that such a fundamentally different body form has been found. Last night, Curler hoisted some light traps up into the canopy of the forest to see what other strange things he could discover. Stay tuned for what he found.

Collaborator Dr. Wayne Mathis has been collecting for several days and has pushed our family list to 69 with this collection of diastatids (sorry, no common name), therevids, and anthomyzids, as well as boosting the list of shore flies (Ephydridae) to 26 species.

Netting bats at night has been particularly productive for inventorying bat flies. Dr. Carl Dick has added several genera and species to our list that would never have been included if we had only relied on Malaise traps and other standard insect collecting methods.

In a small raid of army ants, Anna Holden I collected some bizarre short winged phorid flies running around with the ants. These flies, genus Acontistoptera, have fully winged males that transport them in-flight among army ant colonies. We know this because the flightless females turn up in our Malaise trap samples along with their more mobile mates.

acont

A tiny Acontistoptera female

Yesterday, we spent the day in the lab at our partner institution INBio, where our collaborators could interact with our Costa Rican staff and examine some of the ZADBI material in the collection. Collaborator Dr. Jeff Skevington looked at our 34 specimens of pipunculids (big headed flies) and pronounced almost every one of them a different species! Little duplication means that there are many more new things still to find.

 

This is just a taste of what has transpired over the last three days; I’ll get more reports from our scientists today. Lots of ideas have been exchanged, plans made for the future, and scientific papers outlined for publication. Two days still to go!

Getting lucky

Collecting flies while they are mating is a surefire way to establish conspecific identity. Many female phorids are extremely different in appearance than the males (search this site for lots of examples), and linking the two together can be almost impossible, based on morphology.
< Borgmeieriphora in copula

Borgmeieriphora in copula oblique

In this case, we have a pair of Borgmeieriphora collected in a Malaise trap by Wendy Porras in Costa Rica. The females are wingless, reduced creatures, which however have a sharpened, parasitoid type ovipositor. They lived in army ant colonies, but are rarely collected. The only large series of specimens known is a group that I caught over such ants at La Selva Biological Station many years ago. The males were flying over the ants, carrying the females, as many phorids do. Since then, males have shown up frequently in trap samples, but until now females almost never. This new capture was surely a huge stroke of luck, because most mating pairs separate before they die in the alcohol.

Something “new ” for Central America

When in Brazil a few years ago, my team of Giar-Ann Kung, Wendy Porras, and I found the spiny, brachypterous (short-winged) females of phorid genus Pheidolomyia. These flies, which live in the nests of the ant genus Pheidole, were only known from Brazil. Once back home in Costa Rica, however, Wendy quickly found them there, too. This demonstrates two important principles of dipterology: 1) Wendy is a great collector, and 2) our knowledge of the distribution of tropical flies is extremely fragmentary.

A female Pheidolomyia from Costa Rica - photo by Inna Strazhnik

A female Pheidolomyia from Costa Rica – photo by Inna Strazhnik

After I remarked on yet another interesting “South American” phorid fly showing up at La Selva Biological Station during the ALAS (Arthropods of La Selva) project, the ant ecologist Jack Longino agreed with me, only partly joking that “if you collect long enough at La Selva, you get the entire Neotropical fauna!” Now, that’s a hypothesis that would be fun to test.