The story behind "Latitudinal patterns in tachinid parasitoid diversity".

Burington, Z.L., Inclán‐Luna, D.J., Pollet, M. and Stireman, J.O., III (2020), Latitudinal patterns in tachinid parasitoid diversity (Diptera: Tachinidae): a review of the evidence. Insect Conserv Divers, 13: 419-431. https://doi.org/10.1111/icad.12416 

 

When I started my doctorate working with John Stireman of Wright State University, he and I already had discussions about the chapters in my dissertation. We agreed that there should be a broad biogeographic or ecological component, a narrower phylogenetic component, and a taxonomic revision. I am happy to say that the taxonomic portion passed review in 2020 and is currently in final edits. The ecological portion was published last March in Insect Conservation and Diversity. I long ago promised that I would blog all my publications, but I've obviously dropped the ball on that here. So this is the story behind the paper. 

In 2012, I started my doctoral program at Wright State in Dayton, Ohio. At the time, Stireman had been running malaise traps out in the woods adjacent to campus and in several other locations near the University. These flight intercept traps catch all sorts of things, but his main interest was to catch tachinid flies, of which he has a sizable personal collection. One of these locations was from Huffman Metropark; most of these data were published by his former student Diego Inclan. Stireman also had a large tachinid pan-trapping data-set from his doctoral research in Arizona. And, to top it all off, he had access to specimens from a large trapping project at Yanayacu Biological Station in Ecuador. These three regions, Ohio, Arizona, and Ecuador, are spread across 40 degrees of latitude from the temperate zone to the high tropics, which allowed a rough comparison of diversity between tropical and temperate tachinid communities.

The reason this was such an interesting idea, this latitudinal comparison, is that there is this historical idea that parasitoids are less species rich in tropical zones than in temperate regions, that they have a sort of "inverse diversity gradient", inverse to the normal strong gradient of lower to higher diversity as one moves towards the tropics. Dan Janzen wrote a article about this in the mid 1980s, where he used species diversity of parasitoid wasps across North American latitudes to suggest that these wasps have a mid latitudinal peak in species diversity, at around 30-25 degrees North Latitude, and that after that the diversity declines. And furthermore he, and other authors, suggested that tachinid flies would follow this pattern similarily or even more so.

Now, this particular hypothesis was a sort of creative impulse to write this chapter of my dissertation. But at the time of publication, and even ten years ago at the time I started writing this research, the "inverse latitudinal gradient" for tachinid flies was already a rejected hypothesis. The copious amount of new species coming forth from Janzen's own rearing project in Costa Rica made that abundantly clear. So, it was not my purpose to falsify that hypothesis with this paper, but it was an interesting place to start.

Over time I added several other datasets for comparison. Sonja Scheffer and Mathew Lewis at the University of Maryland allowed us to add their dataset for comparison. Karen Petersen, a former master's student under Stireman, had surveyed in another area at Yanayacu, so I added that small dataset. And Marc Pollet had a good sized survey of tachinid flies from Podocarpus-El Condor Reserve in Ecuador that he collected while trapping for Dolichopodidae. (Marc, Stireman, and Diego all joined me as authors on the final publication.) But by far the largest later addition was all the Tachinidae collected during the Zurqui All Diptera Biotic Inventory (ZADBI) project in Costa Rica, for which Stireman and I identified a large portion of the tachinid specimens. With the addition of that Costa Rica dot I also filled a latitudinal gap, so in the paper we had a general latitudinal spread of 40--30--10--0 degrees. The result was that, every time we thought we had the paper set for publication, it grew another dataset, so what should have taken a few years instead took most of a decade.

(As an aside, I'd like to reflect briefly on the ZADBI project. Since I wasn't a forefront author for those papers, I don't feel like it's right to do a full post about it. But I do want to comment on one point, that over half the tachinid species collected there were in the tribe Blondeliini, which is my taxonomic research focus. Blondeliine diversity is absolutely insane in the Neotropics, far beyond any other group, and the ZADBI project really bore that out.)

The unsurprising result was that we saw a general increase of tachinid species richness from the temperate to the tropical zone. But as I said this had already been established by taxonomists for decades. What is far more interesting to me, is the rarity of most species in all datasets. If you look at the accumulation curves in Figure 1, you can see that none, not ONE of those datasets reaches an asymptote. Even years into collecting, species new to the longer surveys were being recorded. And this is across the board, both large and small datasets, both datasets in the temperate region and in the tropics. In all cases, 55-68% of all species were collected only once (singlets or singletons) or twice (doublets or doubletons). This to me is absolutely astounding, that even in Ohio, more than half of all tachinid species can be counted as rare. And in Costa Rica, at Zurqui, more than two thirds of all species, 68 percent (!) were collected only once or twice. What this says to me, is that any accurate measure of tachind community richness requires a long period of collecting, perhaps decades of surveying. For tachinid flies, Preston's Veil seems wide.

Overall, I'd say I'm pleased with the paper. There's still that pesky gap at 20 degrees (Mexico) and it wasn't possible to make any strong conclusions due to the low number of datasets. There are more tachind survey datasets out there, but they're survey's of caterpillars that sometimes yielded parasitoids. Since caterpillar rearing is generally not a random selection process, it's not comparable with trap surveys. So, we could say that the metrics strongly suggest higher species richness in the tropics, but there is nothing in the paper that counts as a definitive test. My question is: If someone were to repeat the same survey at Zurqui, or Yanayacu, how much would those two datasets overlap? 90% species the same? 70%? We know nothing about the turnover of richness in these communities, how many species are transients or inhabitants, or even what "rare" means for Tachinidae. Does it mean "low number of individuals"? Does it mean "widely dispersed in the landscape", and thus rarely encountered? We know that these flies often hilltop, they gather at specific points in the landscape for mating, and we know they have the flight strength to travel long distances. After all, "tachinid" comes from the Greek word "táchos", meaning "speed". Maybe how we see landscapes, with our narrow surveys, means little to a speedy fly.

The hidden trials of the taxonomic impediment.

So. I'm coming back after a long hiatus because this paper dropped not long ago and I've got feelings. Here's the press release.

A quick summary: Sarah Meierotto of the University of Kentucky and several of her colleagues collaborated with the Area de Conservacion Guanacaste project to describe 18 new species of parasitoid wasps reared in Guanacaste province, Costa Rica. The kicker is they did this entirely from sequences of Cytochrome C Oxidase subunit 1 (CO1) mitochondrial DNA sequences, sometimes known as the "barcode gene". Aside from a single habitus photograph of each species, they offer no morphological treatment of any of these species, and their only diagnostic characters are DNA nucleotides at certain positions. This has resulted in a somewhat heated conversation among taxonomists about the validity and usefulness of these new species "descriptions".

Before I go into those criticisms, I should mention that this isn't the first time a taxonomist has described species from barcodes only. Andrew Brower published
ten new species of moths from barcodes only a decade ago, but his purpose was to preemptively criticize DNA-only descriptions of new species. Quoting his abstract, "the application of names to concepts does not corroborate or endorse the biological validity of those concepts."

Meierotto and her colleagues, in contrast, say this is a "revolutionary protocol" for species description. Are these descriptions available? According to Commisioner Thomas Pape they meet all the criteria for availability. Quibbling about whether listing of base pairs are symbols or words is not really helpful.

But, are they valid? That's a much more difficult and interesting (and hair-tearing) question. How did we get to this point?

The new species are described in two different braconid genera, Zelomorpha and Hemichoma. The genus Zelomorpha has more than 50 previously described species, while Hemichoma has less than ten. Both of these genera were revised in a 2006 dissertation by University of Kentucky student Carlos Eduardo Sarmiento-Monroy. As far as I can tell, none of these manuscript names were ever published. Dissertations are not considered available under the ICZN, so Dr. Sarmiento-Monroy would have needed to publish these names in a journal, book, or other publication to become available; he never did. This year, Meierotto published a taxonomic review of Zelomorpha and Hemichoma with coauthors including Michael Sharkey, who was both the dissertation advisor for Sarmiento-Monroy and is second author on the DNA-only paper. This taxonomic review includes a large number of "new combinations", but as far as I can tell it has no new species.

What I suspected happened is that Sarmiento-Monroy completed his dissertation and never got around to publishing the species names. Maybe he disappeared, or moved on to work on some other group, or decided he hated braconid wasps so much he never wanted anything to do with them again. The names were never published. However, all the tantalizing species descriptions, photos, diagnostic information, these all still exist in dissertation form. All of that was copyright to the author, meaning, anyone couldn't come along and publish it for him. So, Michael Sharkey passed the project along to his new graduate student, Sarah Meierotto. Or maybe Sharkey continued working on the group in the intervening time. Anyway. The morphological species concepts were well known, even some genetic concepts were well known, but all the descriptions, diagnoses, and photographs would have to be redone, from scratch.

Anyone who has ever done any taxonomic work before, I can hear your groans through cyberspace.

So, they got the idea to publish some new names using the simplest description, just the barcodes. They already had some of these, but Daniel Janzen has a ton more via the ACG caterpillar rearing and barcoding project. Meierotto and Sharkey knew which species in ACG referred to new species and previously described species, because they had all of Sarmiento-Monroy's barcodes. So they took Sarmiento-Monroy's species concepts that matched with those species in ACG, eliminated everything but the barcode, gave them all new names, took habitus photographs for each species, and voila. Taxonomy published. Or so I'm guessing.



This is all my imagination. I repeat, I don't know how this happened. But it seems likely, and I don't have a problem with it. There are all sorts of reasons for the taxonomic impediment: poor 100 year old descriptions, missing type specimens, a new genus named for every new species (thank you, Townsend). These are obvious, but there are many more hidden reasons for a lack of taxonomic progress. Often times there is a taxonomic scion, the last, greatest worker in a particular family. They remember everything, all the names, the relationships, all the hypotheses. And when they pass, they take all of that knowledge with them and leave behind no heirs. Often times a group isn't difficult by way of some taxonomic mistake, but just because it is naturally so. All the distinct species groups grade together at the boundaries, all the distinct species are apparently species complexes. And sometimes, all the knowledge is there, but it's behind a copyright wall and therefore cannot be published and acknowledged.


As a taxahacker, I only really care about what is helpful, about what actually progresses our understanding. So finding some way around the dissertation impediment appeals to me. Let copyright rot if it aids our work. However, the key word is "helpful". When I plan a natural history work, I think, who is this for? Who is this helping? Most of the time I consider my fellow and future researchers. They'll probably be using a stereomicroscope, similar to what I use or with worse optics. They'll probably not have a large research laboratory with genetic tools for identification. And they'll probably be people with only incidental interest in the group. Perhaps they work in field ecology, or pest management, or evolutionary biology. They will care little for single species descriptions, and appreciate in-depth revisions the most.


In sum, the average researcher that comes to your work is not going to have access to genetic sequencing equipment. They will have eyes and a microscope. We should aim to write for this level of technology. Any species description that omits morphological characters is effectively useless. Are Meierotto et al. species valid? The above train of literature would suggest yes. At least, they aren't junior synonyms of previously described species. But they aren't helpful, not to any reasonable student of braconid wasps. From Sarmiento-Monroy's dissertation I see that color patterns are necessary but insufficient for identifying Zelomorpha species. There are other necessary components, the shape of the head, the hairing and crenulations of the cuticle, the structure of the genitalia. Many of these aren't visible in a habitus photograph. I suppose I could refer the sequences in the paper back to the dissertation to find the diagnostic characters, but what a hassle!

Natural history science isn't a filing heuristic. Our job isn't to simply give all the things names, in whatever way possible, and organize them into the prettiest tree. The authors state that we must work ever faster with the biodiversity crisis upon us. To what end? I learned very little about Zelomorpha species from this article, nothing that wasn't already available through previous works or the ACG database. I learned that there are many species and have pretty colors. Replace "species" with "postage stamps" in that sentence; "all science is physics or stamp collecting." No, taxonomy is not Pokemon. Every insect species is a hypothesis of metapopulations. Every genus is a hypothesis of species relationships. Every piece of evidence presented, morphological, genetic, behavioral, either corroborates or negates these hypotheses. As Brower said, "the application of names to concepts does not corroborate or endorse the biological validity of those concepts." The names themselves do nothing. They are often satisfying or amusing, but they don't actually increase our knowledge. They're just a filing code. The problem with the paper isn't that the names are unavailable or invalid. The problem is that the species are poorly described. If this is the answer to the biodiversity crisis, then future generations will know nothing of Zelomorpha danjohnsoni, except that it lived in Costa Rica, fed on a species of noctuid caterpillars, and had black wings and an orange body. Oh, and its mtDNA sequence had A's and T's and C's and G's at particular positions. Hardly a useful fact if the species is extinct.

Yet, I understand authors' desires. There's a deep need to put a name on things, even if naming them doesn't actually help us understand them better. It's questionable science, but it's art I guess? So I'm torn.

Using Inkscape for Biological Illustration, Version 1.1.

I'm releasing an updated version of the UIBI book today, with the following changes.

-Minor editing and changes to figure names in all chapters.
-Minor updates to several chapters, including "Detailing" and "Special Structures" (formerly Setae and Hairs).
-Major rewrite of the "Shading" chapter.

I was planning to wait a few months before publishing a new version, but my previous ideas about hatching and stippling in Inkscape were so very very wrong. I have literally wasted tens of hours of my life (or more!) trying to make regular hatch and stipple patterns in Inkscape using work around methods, when the "Pattern" fill option works very well for both of these if you understand how to adjust the density of the pattern. I only discovered this today. Ugh. Fortunately, shading isn't a major part of my line art style. And everyone else can learn from my mistakes. There is also a new curved hatching method included, using the Calligraphy Tool.

UIBI Version 1.1

Using Inkscape for Biological Illustration

Years ago, I posted a short guide to illustrating insect genitalia in Inkscape. Now in the shadow of my dissertation writing, I've managed somehow to write a more complete booklet version.

It's free, just like Inkscape! I take you through my line drawing process step by step, from importing sketches, to laying down and modifying lines, to making special structures like setae, to finishing and exporting. I also include some more experimental techniques which may be of interest to other sorts of Inkscape users. Enjoy!

Download link: Using Inkscape for Biological Illustration (PDF)

The Fly Trap (Book Review)

     The Fly Trap is a modern creative novel: one part biography, a second part history of entomology, a third scientific explanation, with rapture at flies mixed throughout. I’ve been reading Moby-Dick recently, and the similarities are striking. There’s the same tendency to switch themes between paragraphs and chapters, to weave the scientific, historical, and biographical together, and to embiggen fact when necessary. But what draws the entomologist in is Fredrik Sjöberg’s replacement of Melville’s whales with flies and the hope that the author will do the subject of our work and life passions the justice they deserve. How often is any particular family of insects besides butterflies made the subject of literature?

     Sjöberg’s centerpiece is the two-winged fly family Syrphidae, more commonly known as “hover flies” or “flower flies”, names which refer to incredible flight capabilities and the tendency to be nectar feeders. They are commonly yellow and black mimics of bees and wasps. Unlike the animals they’re mimicking they have no stingers to deter predators, so they avoid predation by looking like something dangerous. Sjöberg peppers his chapters with vignettes about individual species, including everything from taxonomic and regional history, physical and ecological diagnoses, and personal anecdotes. My favorite is the sudden “invasion” of Eristalis smilis which overtook the Swedish countryside, contrasted with Doros, of which there are only occasional sightings and elaborate rumors.

     The other subjects are “islands”, whether those be Sjöberg’s home island of Runmarö or a tree stump in a recent clear-cut. “Islands are generalizations of a kind”, he writes. “And where there are no islands, we have to invent them. If only for the fun of it.” He cites the loneliness and isolation of islands both positive and negative. Islands are perfect ground for the cataloger, sometimes disparaged as “buttonologist”, who provides a complimentary and more detailed worldview for “mapmakers”. 

“But the person who makes maps can never include everything in his picture of reality, which remains a simplification no matter what scale he chooses. Both attempt to capture something and to preserve it.” 

I particularly enjoyed his description of the Fly Tree, an enormous, 500 year old black poplar that was an island ecosystem onto itself. These species descriptions and descriptions of “islands”, are the stepping stones on which Sjöberg’s stories rest. 

     Yes, stories. There are actually two stories here, two interwoven biographies. One is of the author’s work with hover flies. The other biography is of the heroic, larger than life Rene Malaise, who sits in sharp contrast to the author. Malaise was a great explorer, eponymous trap inventor, and collector abroad, especially in eastern Russia and Southeast Asia. Sjöberg tries to avoid all collecting and exploration beyond his small island in the Baltic Sea. He says of the tropics, “Tropical nights can build into tremendous explosions of downright Cambro-Silurian cacophony when a thunderstorm starts or cicadas celebrate their orgies in the treetops. They’re magnificent, but no more than that. The indescribable sound of the Madagascar nightjar is worth the entire trip, but in the end it is merely interesting and exciting and fun to tell people about later.” Of the Congo River basin, “What an adventure! What stories I would tell! About freedom! But it didn’t happen. I never managed to say much more than that the forests were vast and the river as broad as Kalmar Sound. And that I’d been there.” Yet he idolizes Malaise and his travels, to the point where he starts a collection of Malaise-related ephemeralia. This ends ironically with an expensive purchase of a painting once belonging to Malaise. The author, so adverse to crazed collecting, has become a buttonologist. But Sjöberg stays to his island, claming glorious isolation and “slowness” allow him an illusion of control over these impulses.

     One of The Fly Trap’s most overreaching themes is what Eliezer Yudkowsky calls “The Virtue of Narrowness”. Sjöberg’s collection only contains the 202 species of hoverflies (plus one) found thus far on Runmarö island. He feels he must justify his narrowness, so he writes that it’s purely for pleasure. No, it’s because he loves the D.H. Lawrence style of isolation provided by islands. No, it’s a sort of “buttonology”, a collecting disorder, which in his case is benign. No, it’s an attempt to slow down in our fast-paced world. He doesn’t beg the reader to accept his reasons for collecting and observing his island’s hover flies as scientific. Even when he claims his study allows him to “read nature’s language”, the result is for enjoyment. Maybe he feels he can’t explain the usefulness of this small study on his small island to broader natural history, not even to a lay reader, but I don’t think he needs to. The Virtue of Narrowness is the precision and accuracy of your knowledge. It’s enough to only explain hover flies on Runmarö, and Sjöberg knows it, but he still claims “hobby” because it’s not his “real” job.

     True, he does romanticize his narrowness whenever possible. But I enjoy some romanticized narrowness. In my favorite poem by the midwestern American Tom Montag, “The Farmer’s Manifesto”, the farmer says of his father, “He had no /ideas but the things which /his hands could touch, or /those his eyes could find /at great distance—a glint /of sun off farmhouse windows. /Or close at hand, beneath /his feet. What he could /catch as breath; wind would /carry. He knew those weeds.”  Romanticized or not, that sort of narrowness holds an incredible depth of knowledge, what Montag could only name as “strange /dark madness, some amazing avalanche /of wolves, lakes, stars, tongues” and the ability to “hear corn grow in summer; /can hide your face in /the curving surface of sky; /examine a potato in light /so special you know something /flies back at you”. This is the sort of knowledge that comes from doing the same thing repeatedly over a small stretch of world and small number of subjects until they become windows. What seems like buttonology is deep expertise.

     I don’t mean to say that The Fly Trap is perfection or free from cliche. It belongs firmly within a genre of creative natural history writing first made popular in the 19th century, a Euro-centric and primarily masculine genre written by men for men and boys. Women are largely incidental to the story and are mentioned mostly as love interests or as props. His wife features prominently at the end of the first chapter, but only as the nameless “girl who sat in the audience one evening”. Of professional meetings, he says, “Normally no women take part at all. And the few who do happen to show up are usually the better halves of the biggest crackpots, wives who could easily pass as personal assistance from a psychiatric open ward. Well, maybe that’s unfair. But the fact is that unattached women could hardly find a better hunting ground than entomological societies. Unusual men, no competition. Just a suggestion.” Does that mean the only reason for women to attend meetings is to pick up men? Even the preface quote ends with the condescending line, “Me, I just concern myself with flies — a much greater theme than men, though maybe not greater than women.” The only exceptions are the short biographical sketches of the incredible, possibly lesbian Esther Blenda Nordström, a writer, explorer, and ethnologist who briefly married Malaise and traveled with him to Asia. Unfortunately, her story was abandoned when Sjöberg realized Malaise hadn’t named a species after her, and therefore Malaise’s “love” for her couldn’t be verified (unlike for Ebba Soederhall). I could have read an entire book about Nordström and her travels. Fortunately she wrote several. Unfortunately, I don't read Swedish (but maybe you do).

     The Euro-centrism is more forgivable. The Fly Trap was originally written and released in Swedish. The intended audience was Swedes, the setting was (mostly) Sweden, and Sjöberg is Swedish himself. The English translation came ten years later, so it should be read as a Swedish novelslashbiographyslashcreative-nonfiction and shouldn’t be taken as worldly. Especially since Sjöberg repeatedly admits his own non-worldliness.

      I realize I haven’t said very much about flies in this review. Fact is, if you’re still reading this and you haven’t already read The Fly Trap, you probably already have some interest in flies and will be delighted as I was of the hover fly natural history in this book. There isn’t anything to criticize about those descriptions except to say that they’re wonderful and I wish there was more of them. I recommend The Fly Trap for entomologists and non-entomologists alike.

 Sjöberg, F. 2014. The Fly Trap [English translation, Thomas Teal], Pantheon Books, NY. Amazon

Zoobank is down (the future of taxonomic publishing).

ZooBank is currently down while I'm writing this, the "official registry of Zoological Nomenclature". This is the registry that all new electronically published names and nomenclatural acts must use before publication, and that all traditionally published nomenclatural acts SHOULD use before publication (but usually don't).

I don't know if this is just temporary down time, or this has been going for a while, but it's a definite problem. Especially with the way publishing is going.

I was just talking with Morgan Jackson about social media and taxonomic publications, because I woke up with a weird thought in my head this morning: what if I took the taxonomic portion of my dissertation, registered the new names with ZooBank, and published it as a PDF on my blog? Given the standards currently in the ICZN, and assuming I'm meticulous about referencing type specimens and depositories, etc., the new names would be totally available under the code! Any person can do this now, or at least they could if ZooBank was running.

This isn't only limited to nomenclatural acts originally published as PDFs. To quote Morgan,

"The Winnower is working to publish, assign DOIs and archive blog posts and reddit threads right now. There is very little standing in the way of someone publishing a new species name in an electronic place like reddit (with the proper [ZooBank] registration and everything) and having it become valid via Winnower sucking it up...As far as I know they haven't finalized their archiving with [CLOCKSS] yet, so they haven't met all of the Code requirements for digital publication, but last I talked to them it was in the works"

The Winnower, for those who aren't familiar, is an open access publishing site that uses an open access peer review system. They specialize mostly in commentary on publications (i.e., post publication peer review), but their targets include a wide variety of non-traditional publishing platforms like blogs and web forums. CLOCKSS is an archiving platform for electronic publications, which The Winnower is using to store publications as PDFs. What Morgan is suggesting is that a taxonomic work registered with ZooBank could be originally published on a blog, adopted by The Winnower, and archived with CLOCKSS; thus it would meet all electronic publication requirements of The Code despite not being available in it's original publishing context.

It seems convoluted, but the above scenario is totally workable under the current version of The Code. I can see both positive and negative elements of this. For one, the traditional taxonomic publishing method is incredibly ponderous, even with taxonomic journals such as Zookeys and Zootaxa. Publishing is further complicated by the general feeling in biology that taxonomic works are low priority under the categories "impact" and "significance". It also opens up low or zero cost ways for taxa-hackers to publish their work, and I'm all for that. (That new species of fungus gnat I've been sitting on, for example. Hmmm...)

Conversely, I see the recent trend in taxonomy for higher quality publications and the role Zootaxa, Zookeys, and other taxonomic journals have played in this. I would hate for taxonomic publishing to slide back into Townsend-esque quality or for taxonomic vandalism in the mode of a certain Australian snake hobbyist to become more common.

Someone will try one of the above methods eventually. Or whenever Zoobank is up and running again. At the time of finishing this, the registry website is available! But it still worries me, because electronic publication is only going to become more important in the next decade. If Zoobank is unreliable, then what of the future of animal taxonomy?


Thanks to Morgan Jackson (@bioinfocus) for help in fermenting these ideas.

Adopting Orphaned Taxa (TAD2015).

(For Taxonomist Appreciation Day 2015)

I know it’s been a while since my last blog post. I’m deep into research right now and about to defend my thesis proposal. I’m also working on a publication. Which means that my writing time is going elsewhere and not here.

My thesis research overall concerns a large and varied tribe of tachinid flies called Blondeliini (Blond-el-ee-ai-nai) or the blondeliines. The core of the work is the Blondelia group of genera, called such because it includes the type genus of the tribe, Blondelia. Females of the Blondelia group have a boat-like keel on the abdomen and a sharp piercing hook for poking holes in things, usually caterpillars.

The piercer isn’t an “ovipositor” in the homologous sense, because it doesn’t contain the tube that carries the egg into the host. Instead, the egg tube (the mostly membranous segments 8-10 of the abdomen) travels down the posterior groove in the piercer and into the hole the piercer has made in the host.

A female 'sword fly' of the genus Eucelatoria with hind legs removed. (1992: Mexico, Portillo de Reon.)

In one genus (Eucelatoria), the piercer can be half the body length! I’m not sure why these species have a piercer that long, but there’s some evidence they parasitize caterpillars hidden in rolls of leaves. Some Blondelia group species have spines on the ventral keel, and others have only bristles. Some males of the Blondelia group have hairy patches on their abdomens, and other closely related species are clean shaven. Host use varies; Costa Rican species of the Eucelatoria armigera complex are particular to one or a few species of noctuid moth caterpillar, while the polyphagous species Compsilura concinnata feeds on over 200 species of Lepidoptera.

With interesting oviposition behavior, morphology, and a large number of species (>135; not including all the many undescribed Neotropical species) the Blondelia group is an enticing project for a young taxonomist. Do not fall for this trap!

Orphaned taxa are those genera or families that are without a current expert or active worker. The Blondelia group, and Blondeliini in general, are a particularly frightening example. Abandonment can be for a number of reasons. In some cases the group isn’t charismatic enough, or is of minimal economic importance. In other cases high diversity and difficult diagnosis are deterrents. A history of poor descriptions and over-splitting genera may be to blame; for this final reason orphaned taxa often have a taxonomic impediment. The longer the period between experts, the greater the impediment to future research becomes.

In the case of the Blondelia group, our good friend Dr. Townsend is mostly to blame. He is responsible for naming nearly half of all valid blondeliine genera, and most of these with one species apiece. Add to this his notorious over-splitting, his mediocre descriptions, and his terrible, no good, very bad Manual of Myiology genus key, and very few people are courageous enough to venture forth.

However, not all blame can be placed on Townsend. Disregarding the history, blondeliines are a difficult group with many examples of morphological convergence. They are small, usually dark colored, and told apart mostly by arrangements of bristles. Color patterns often fool me.

Left: Sword fly male. Right: NOT sword fly male. Really similar, but really different. Can you see the difference? (Click for embiggen)

I thank Monty Wood for his great work on Blondeliini (1981), without which I would be lost. But this is a preliminary work of broad scope. Efforts focused on a single or a few genera have revealed the scale of the mess yet to be resolved.

Diego Inclan (a graduated MS from my lab) and Dr. John Stireman (my PI and advisor) provide a vexing example of this mess in their recent Zookeys paper. In his masters thesis, Diego found that some species considered part of the Neotropical blondeliine genus Erythromelana were clearly not. This lead to a convoluted taxonomic investigation. Below is a paragraph from the Zookeys paper as illustration.

“An example of the taxonomic instability of Neotropical tachinid genera is witnessed in the species Euptilodegeeria obumbrata (Wulp). This species was first classified in the former tachinid genus Hypostena by Wulp (1890; along with many other blondeliines), based on specimens collected in Guerrero, (southwest) Mexico. […] The species was moved by Townsend (1931) to the new genus Euptilodegeeria, moved again to the genus Erythromelana Townsend by Wood (1985) and recently excluded from Erythromelana and resurrected to its previous genus (Euptilodegeeria) by Inclán and Stireman (2013). Although the taxonomy of Tachinidae, particularly of the Blondeliini, is challenging due to the scarcity of clear synapomorphies, the confusion in the generic assignment of E. obumbrata was also due to the limited number of specimens evaluated, the lack of examination of male terminalia and the use of only males for the descriptions. In the present study, we use additional information from male and female terminalia to demonstrate that these “obumbrata” specimens, previously assigned to Hypostena, Euptilodegeeria and Erythromelana, actually belong to the genus Eucelatoria Townsend (1909), in which females possess a sharp piercer for internal oviposition in the host. We also argue that the former species Machairomasicera carinata described from a single female by Townsend (1919) in the monotypic genus Machairomasicera, and later synonymized with Eucelatoria by Wood (1985), belongs to this same species group of Eucelatoria, which we here define and characterize. In the end, taxa that were assigned to four different genera in fact belong to one species group of Eucelatoria, providing an example of the taxonomic confusion that plagues many groups of Neotropical tachinids.” [Emphasis mine]

Many similar issues remain in the genus Eucelatoria. The group may not even be monophyletic. I am not revising all the species in the Blondelia group for my dissertation—or even all the species in Eucelatoria—but the challenge feels insurmountable.

There are two ways to publish natural history research. One is to be cautious, to wait until all possible evidence is covered and carefully recorded, all the museums have been visited, and every last lead has been pursued. “I only have one more type to look at, and it’s been missing for 40 years. But I can’t publish until I find it.” The other is to rush wildly into publication with any new finding, getting the information out as quickly as possible. “Never mind the types in that European museum, I have the specimens here and there’s nothing (well) written in the literature to say I’m wrong!” 

Taxonomists, myself included, fall more on the cautious side. Townsend was an exception. We want all the bits of evidence before we publish our scientific opinions, whether that be new species, synonyms, homonyms, or redescriptions. Caution is great when you start with a clean slate. But in the face of a huge mess caution is paralyzing. How do I start? I’m looking at a great wreck of a building. Do I take the debris out piece by piece and slowly repair? Or do I knock it down, bulldozer the area, and pour a new foundation?

I have sat and looked and sat and looked and wondered at my specimens guessing and second guessing myself if what I am seeing is really separate species, or if they have been previously described. This back and forth mental motion is useless. I fear too much of wreaking havoc. But plenty havoc has already been wrought.

I think there is a middle ground. Stride boldly, but document everything. Don’t worry too much about creating new species synonyms or mis-associating males and females. Those issues can easily be fixed later. Otherwise you’ll spend the rest of your life waiting for that visit to that one university in Chile (when the type was long since moved to a museum elsewhere). At the same time, document everything and carefully record your findings. If you provide photographs, written description, genitalia drawings, and adequate references to collections and literature in your publications, someone else can build upon this firm basis and correct your mistakes.

An excellent example of walking this line is Dr. Lee Herman’s 2013 revision of the New World species of Oedichirus, a genus of rove beetles (Staphylinidae). Dr. Herman, a Curator Emeritus at the American Museum of Natural History, received the “J. O. Westwood Medal and Award for Insect Taxonomy” [PDF] for this publication. Rove beetles have a taxonomic history as equally tortured as tachinid flies. As in tachinids, associating males and females of the same species is difficult. At times only male or female specimens are available, and species were described sometimes based on the male and sometime based on the female. Furthermore, the majority of specimens available (including the types) are too old for modern techinques like DNA Barcoding. Herman could have waited until new specimens were available, but instead he pushes forward. In the methods section he writes, “hypotheses of male/female association proposed herein for the other species can be corroborated or refuted by DNA barcoding techniques using newly collected specimens.”

Remember that every “opinion” of natural history is a hypothesis subject to further testing. When our hypotheses are presented as expert opinion but rest on shoddy work they are an obstacle. When we refuse to present hypotheses for fear of being wrong they are also an obstacle. But when our hypotheses are presented boldly and rest on good work, even our mistakes are outweighed by the scientific contribution. I have discovered that it does no good to worry. Any well documented progress is good progress. Anything mopped up is better than the mess we have now.

CHT Townsend, Vandal of the Calypterates. Epilogue.

Part I; Part II; Part III; Part IV; Part V.

When I began this series in January, my intent was to show parallels between recent and historical conflicts in taxonomy and systematics. You may remember Raymond Hoser from the first part and "Call for Comments" post. He is an extreme example, perhaps more extreme than Townsend with his repeatedly forced rhetoric and lack of decorum. In the history of taxonomy, however, this is not unusual.

And lest you all get this opinion, let it be said that I, Z. L. 'Kai' Burington, do NOT hate Charles Henry Tyler Townsend.

I chose to exhibit him in a negative light, the sort of light that people working on tachinid flies see him in. This is only one half of the story. Contrast my take with Neal Evenhuis's treatment (Page 15 in this PDF). He calls Townsend a "man of wanderlust and mystery" and outlines his many accomplishments, including: world expertise on Diptera and calypterate flies in particular; instruction in pest control (Jamaica); co-ownership of a taxidermy and zoological specimen company; biology professor (Philippines); biological control of the cotton square weevil (Peru); discovery of the fly vector for both verruga and Oruya fever (Peru); doctorate from Washington University; honorary custodian of calypterate flies at the US National Museum; control of leaf cutting ants (Brazil) and other pests (Peru).

Evenhuis told my favorite story of Townsend today during his talk at the 8th International Congress of Dipterology, that he traveled across the Amazon in 53 days (not including stops), and arrived in Peru just in time for his 60th birthday; he popped a champagne bottle near the beach in Pacasmayo to celebrate. I can't help but admire his intense energy and fortitude.

And then I'm brought back to the reality. Townsend was an ego driven, abrasive man who died quite bitter about his recession from science. The story feels like a one-two punch of schadenfreude, but the aftertaste is more like Townsend's own bitterness. He did many great things, and he is usually remembered for his vandalous legacy and his nutty ideas. Often I see the parallels between Dr. Townsend and the late Dr. Lynn Margulis. Both had incredible expertise. Both did great work. And both had some ego-related nutty ideas which threatened their public face and careers and which leave a shadow on their contributions.

I would like to say that things have changed since Townsend. Unfortunately they have not. And I would like to blame all the personality circus acts of this saga on Townsend alone. But I cannot do that either. Coquillett refused to speak with Townsend to try to make things right. Walton, instead of contacting in private, aired his equally abrasive opinions in a public forum. Aldrich, instead of simply cutting off contact and refusing to play along, continued his angry letter sending to his former colleague. And the rest of the dipterist community didn't step forward in outcry against these antics until after the 1925 "Insider History", long after the damage was done. In short, the egos of all people involved were to blame.

Things have not changed. I am, as I said above, at ICD8 this week. This morning as I pondered these questions and yesterday's panel on the "Future of Diptera taxonomy and systematics", a colleague said this to me. He said, it's not the differences in methods, or morphology vs. molecular, or ages of the participants that are why these issues--these apparent clashes--continue. The reason they continue is the difference in personalities.

The reason why the Townsend saga got out of hand--the egos and personalities of the people involved--is the same reason taxonomy lacks unity in our current crisis.

Just before my PhD program started I was working on a short term project at a U.S. museum. At the end of the project one of the curators gave me a long and personal lecture. He said that the most important thing for my future was to be kind and generous to everyone, to promote unity, and to tamp down ego. Because, he said, selfishness and other personality flaws are to blame for our problems in taxonomy.

And the people who are most public, most obvious, the loudest, most outspoken people are often the most abrasive, unkind and ungenerous people. I cannot tell you the number of times I have yelled at my email in the past months upon finding yet another message from the ICZN-listserv. This forum, which was supposed to be for finding and giving help and discussing zoological nomenclature, has become a platform for various taxonomists (including Hoser but not limited to him) to argue and curse at each other publicly over their personal disputes. I have spoken with several people about this on Twitter. The general response is that it is going to happen and there's nothing I can do about it. The Taxacom listserv is much the same. Better to ignore it, I was told.

Yet THIS is the face we present the world and it is not a pretty face. It is the face of irrelevance. If we let people like Townsend and Hoser be what people see in taxonomy, if they see anything at all, how much longer will our science be considered science at all? How can we live up to the many challenges if there is no public unity? How will our field of work continue if there is no kindness and generosity to each other and to the next generation?

Please prove me wrong.

CHT Townsend, Vandal of the Calypterates. Part V.

IN THE PREVIOUS POST...Townsend's ego-driven quest to propagate genus names ad infinitum leads to a libelous article attacking John Merton Aldrich. The backlash of his former colleagues suggests he had worn out his welcome.

Townsend never returned to North America.

Yet, he continued his research much as he had previously, or potentially more fervently. His output between 1915 and 1925 was smaller than the earlier years, but his publication record from 1926 until 1942 (2 years before his death) nearly matches those high numbers. There was the "Synopse dos generos muscoideos da regiao humida tropical da America" (Synopsis of muscoid genera from the tropical rainforest region of America), published in 1927. The Synopsis contains a 100 page dicotomous key with 605 individual couplets, in Portuguese, and uses a system of abbreviation conceived by Townsend. It was undoubtedly as difficult to use then as it is now. Of course, no Townsend publication would be complete without at least one new genus, so he includes 87 pages of them (with several on each page).

New species description from "Synopse dos Generos" (1927). It is both in Portugese and Townsendian abbreviation. Perhaps it is thankful in this case that Townsend described a new genus for nearly every species, as all of his genera are well described in the Manual of Myiology in English (with comparable abbreviations).

And there was the long awaited Manual of Myiology, published in 12 large volumes between 1934 and 1942. If Townsend could be considered to have a Magnum Opus, this is it. It includes complete keys to families, tribes, and genera of "Oestromuscaria" (muscoid flies), descriptions of all genera, and notes on biology and morphology of the various tribes. Volumes 11-12 contained a strange digression from the earlier sections, including chapters on the Tertiary origin of the Moon from a near Oceanic continent, the origin of humans ("Hands cannot remain idle. Doubtless driftwood clubs and fistsized pebbles were their first implements."), the flight mechanics of a Cnephanamyia bot fly traveling at 400 yards per second, and very, very wrong ideas about gravity.

Cretaceous map of Pangaea (According to Townsend (1942)). Note the clearly marked "Moon" attached to Oceana, which, as Terry Wheeler pointed out, "explains those Australia-Moon sister groups."

This second to last item, bot flies traveling at Mach I, has it's own story in one of the most bizarre papers ever to be published in an entomological journal. Dr. Peter Adler mentions Townsend in his Insect Morphology course, and says only two things about him. One, that he has a very strange species concept, and two, that he clocked a bot fly traveling at 800 miles per hour. Long before I started working on tachinids I was already aware that Townsend was a strange fellow.

Townsend claimed to have observed this physical impossibility in Arizona at 12,000 ft, which he described originally in the April 1926 issue of Scientific Monthly. After recieving several comments, he wrote in response in his paper titled "On the Cphenemyia flight mechanism and the daylight day circuit of the Earth by flight" (1927) that by traveling at this speed (815 miles per hour) one could circuit the earth in less than a day, or see two days traveling east. "It is of extreme interest as affording a mark [466 mph] that should be reached within the next decade; while the more remote future holds the possibility of riding the tail of high noon or speeding on the wings of the morning halfway between the equator and either pole. It can not be denied that the double daylight-day westward circuit will attain great poularity before the single daylight day circuit is realized." 

Since the vibrating wings of a fly are very different than that of a bird or a fixed wing aircraft, he gives the fly flight mechanics its own name, the "Myiopter" groundplan. Townsend proceeds to describe this groundplan in great detail, but not before inserting some off color remarks.

"Regarding the speeds of Cephenemyia, the idea of a fly overtaking a bullet is a painful mental pill to swallow, as a friend has quaintly written me, yet these flies can probably do that to an old-fashioned musket ball. They could probably have kept up with the shells that the German big-bertha shot into Paris during the world war."

This (to use Townsend's own word) quaint idea was thoroughly debunked by Nobel laureate Irving Langmuir in 1938, who brought the speed down to a more believable but still appreciable 25 miles per hour.

In the same year, Townsend published his second paper on synonymy. The first, as you may remember, was published in Science Journal (1911) and was relatively optimistic. The 1927 "What constitutes synonymy?" paper is decidedly bitter and full of schadenfreude. I have transcribed the majority for your enjoyment:

"I have never for a moment considered [these genera] synonymous with Hilarella. Such synonymy is quite ridiculous. As to the rest of the world, no one competent to form an opinion had studied material, hence no opinion existed but rather a complete indifference. Nobody cared a snap whether these genera were synonyms or not. This forcibly illustrates what a power lies concealed in the weapon synonymy. A careful worker may erect valid genera and species. An ignorant or malicious person may publish an article stating that these valid genera and species are synonyms, and henceforth they bear the synonymic stigma. The genera public is not competent to judge of the merits of the case, and besides has troubles of its own. No one cares a snap about the matter unless he is making a special study of the group in question. The original author may publish a refutation of the synonymy. Nobody pays any attention to him, the public not being interested, and his refutation is quickly forgotten. Fifty years later, a competent worker reconizes these genera and species as valid and concludes that they have lain in the synonymy a half century. Is he technically correct in this view?
Synonymy has too long masqueraded as a court of permanent and infallible decisions. There is nothing final about synonymy[...]
The synonymic pronouncements of a single individual carry weight in exact ratio to his ability in the groups concerned. But the general public has no means of judging of his ability. If he sets himself up as a specialist and speaks with confident authority, the public accepts him at his own valuation. He is henceforth at liberty to inflict his personal opinions on a long-suffering public and to manufacture synonymy ad libitum. This is the easiest thing in the world to accomplish as long as the manufacturers escapes detection as a fraud. In fact, it may be termed systematic pastime. He is knocking everything on the head right and left as suits his fancy, while the public looks on unconcerned and practically uninterested. He is destroyed, not building, but no one cares except the original builder who notes the attempt to level to the ground his laboriously erected edifices. Yet they are not really leveled and their status is just as good as before until the synonymy in question is abundantly endorsed [...] This strong weapon synonymy is not to be left at the beck and call of every individual."

Upon hearing the above, my darling partner declared "Dear Sir: No one will ever recognize your true genius, even long after you are dead" and "You mad, Bro?". The imagery of synonymy as a "weapon", of the good taxonomists as the "original builders", of the synonymizers as "knocking everything on the head" and being "destroyed, not building", and that "no one gives a snap" shows Townsend at his low point. This was, after all, just two years past the "Insider History", and before he found a way to publish the Manual of Myiology.

In 1944, only two years after the final volume of the Manual was published, Townsend died in his home at Itaquaquecetuba. The total number of publications over his lifetime is in the hundreds, and the total count of species described is near 1500. He seemed to have burned every bridge with his former colleagues. He outlived his "bitter hatreds". Aldrich died in 1934, the "nation's greatest accumulator of dipterological information" (from Melander 1934). Coquillett had long since passed. The works of both were celebrated. 

The Townsend obituary published in Revista de Entomologia (1943) paints him in a positive light, as a great entomologist, biologist, linguist, author, farmer, hunter of beasts, and a member of numerous scientific societies. Yet, to taxonomists who work with tachinids, he is remembered most for his ego and vandalism.

Epilogue

CHT Townsend, Vandal of the Calypterates. Part IV.

IN THE LAST EPISODE...we examined the short but pointed publication war between Townsend and Walton. Now we return to Townsend's "second bitter hatred", that of John Merton Aldrich.

 In 1914, Townsend returned from Peru and became the honorary custodian of muscoid diptera at the U.S. National Museum. He had just received his doctorate from Washington University and was working at the Bureau of Entomology. The plan was to quickly finish his Manual of Myiology, but the first volume wouldn't be published till 1934, long after he had left the U.S. Instead, Townsend was caught up in a conflict with his perceived rival, Aldrich. This can be seen in his publication output, which dropped from over 30 papers on tachinids in 1915-16 to only five in 1917, and never returned to the previous numbers.

Aldrich, also an employee of the Bureau, had recently left his job at the University of Idaho and was working in Indiana. In 1915 he published a summary of his 25 years collecting tachinid flies, which followed closely to Coquillett's 1898 revision. Townsend was not pleased; the paper ignored his many families of "muscoid" flies and condensed them all into a single family, Tachinidae.

It is interesting, though entirely unintentional I'm sure, that Townsend's "On Proper Generic Concepts" follows directly after Aldrich's paper in the same 'Annals' volume. This is yet another attempt to rally for more restricted generic concepts. Here, Townsend begins by separating all the Muscoid taxonomists into two categories, the "specialists" and "generalists". Specialists, such as Rondani, Desvoidy, and the much admired Brauer & Bergenstramm, used restricted generic concepts. Despite the many potential flaws in their work due to ignorance of internal reproductive characters, Townsend finds their work excellent. Conversely, authors such as Macquart, Schiner, van der Wulp, Walker, Bigot, and the ever hated Coquillett are all generalists, who

"attempted to apply the same broad generic concepts to the Muscoidea that they applied to the rest of the Diptera. Without going into lengthy detail, it is enough to state that their mistakes are many and often overshadow the good contained in their results. Their misidentifications of species are extremely numerous. Their wholesale confusion of distinct generic forms was the natural result of no concise generic concepts. Almost throughout, their genera are mixed-genera. They may be said to have practically lacked muscoid generic concepts, for their generic rulings were largely arbitrary and so loose as to admit numberous foreign elements. The true explaination of all this is that they possessed only the most superficial knowledge of their subject."

Townsend goes on to discuss proper generic concepts, those that do not group by "transitional species." "Groups of generic stems [as of tips of a tree] that happen to be connected throughout by transitional species can not be treated as a single genus, on account of their diverse combinations of characters." The branches have not become decimated over time due to their young age, removing the "transitional stocks".

Table from Townsend (1915) showing the pros and cons of restricted and unrestricted generic concepts.

Figure from Townsend (1915) showing two "families" of tachinids with some species used for illustration of convergence and transitional species. Older, decimated stocks would have more easily delineated genera.

The above figures are interesting as evidence that Townsend's basic ideas were not poorly thought out. He was attempting methods to classify a very recent group of insects which had not been decimated (see Wonderful Life (Gould 1989)), and thus there were many apparently intermediate forms between what would otherwise be clearly recognizable genera. The problem isn't Townsend's justifications, it's the extremism of his taxonomic splitting and inflationism.

Though not explicitly named, Townsend probably considered Aldrich as a generalist, making a "great number of egregious blunders."

Between 1915 and 1924 we have little to mark the falling out between Aldrich and Townsend. In the earlier part of this 10 year period, it seems the two were communicating about their work on the genera Imitomyia and Masiphyia, and there is no direct evidence they were arguing behind the scenes. However, in 1918 Aldrich was moved from his post in Indiana to the Smithsonian, where he became the Curator of Diptera. Less than a year later, in March 1919, Townsend left D.C. for Peru and Ecuador, and later Brazil, to Iquaquecetuba, near Sao Paulo. There's some reason to suspect that Townsend left America because of professional conflicts with his "new boss", Aldrich. The "Inside History" makes it seem that he was fearful of Aldrich undoing his organizational work at the museum, as he quickly published notes on the collections soon after leaving.

What Townsend does next is a bit astounding. He published two personal correspondences from Aldrich in his 1925 History. Such a thing seems completely absurd to us now, a total breach of academic conduct and courtesy, yet, it is similar to the methods of Raymond Hoser (our contemporary, who inspired this series).

On May 8th, 1924, Aldrich sent this letter. The context is a paper he just published with colleague Webber on a tachinid species complex. He writes: "You will not like it, because we did not recognize enough genera to suit you. I am responsible for the generic arrangement, which cost me an immense amount of work and study."


Townsend replies: "Your remark is highly significant. Instead of wasting time in an attempt to extend generic limits arbitrarily where they do not naturally fall, it is far wiser to strike a generic arrangement that shall be fairly simple and easy to follow out. Restricted genera, concisely defined, attain the greatest simplicity of treatment possible."

Aldrich responds, on August 2nd:  

"It would be useless to undertake any general discussion of the limits of genera. I have, as I freely admit, much difficulty in determining them. You solve the problem by making a genus for almost every species, but you encounter precisely my difficulty when you start to group these genera into tribes. So you are no better off than I am, and I am trying to classify muscoids as nearly as possible on the same lines as other animals. I never did take any stock in your oft-repeated belief that muscoids require a different taxonomy."

And, of course, hits the nail on the head. These sorts of issues will arise, no matter what rank Townsend decides is appropriate. Better to limit taxonomic inflation than to let the field become grossly distended with monospecific genera. But of course Townsend sees this differently.

"The work of Aldrich is destructive rather than constructive. He is attempting to relegate to the synonymy as many of Townsend's restricted muscoid genera as possible, with the sole aim of vindicating his own original commitment to broad categories. It is a pity that he is so unreceptive to progressive ideas and holds so stubbornly to long-exploded concepts. He refuses absolutely to change his ideas in the light of new facts. It is evident that his work will suffer proportionately in consequence. He has a better eye than Coquillett had for muscoid characters, but he persists in ignoring important characters which Townsend has pointed out, partly from prejudice and partly from the difficulty of interpreting them."

Remember, he's writing in the third person, about a professional colleague, in a professional journal. 

The ego-train continues:

"The numerous dicta put forth by Aldrich would be interesting if true, but the trouble is that no dependence can be placed on them. They are simply the individual prejudiced opinions of a man who is unable to learn because he will not keep a receptive mind."

And:

"Townsend harbors no animosity toward any one, for life is too short to waste in animosities. He writes this himself, standing off as a detached and impartial observer, contemplating his own work as though it belonged to another, and exposing this inside history only in the interests of fair play and a square deal."

You get the idea. He has an ego as tall as the Washington Monument, and a lack of decorum to match.

The drawn out rant finally concludes:

"Such is a brief outline of the work on muscoid taxonomy in North America to date, involving also recent work in South America. Younger students are arising, from whom we may expect much. Let the keep an open mind, for a closed mind is a fatal fault in an investigator. Let them beware of prejudices and commit themselves only to a search for truth. They, will then not be faced by the alternative of retraction, or continuance on a mistaken course."

All of this was published, March 1925, in the Journal of the New York Entomological Society.

In 1926, the Society's publication committee received a letter signed by 23 American entomologists, including John Merton Aldrich. It was published as follows:

"To the Publication Committee, New York Entomological Society.
The undersigned wish to express their great surprise and regret that you should have published in your Journal the article by [Townsend] [...] This article is in substance a bitter and uncalled for attack upon [Aldrich], a man of high standing, who is greatly respected both as a man and an entomologist. Dr. Aldrich's criticisms of Dr. Townsend's work in his studies of the Muscoidea have always been justifiable and were an honest endeavor to reach the truth. No one could do as much on this group as Dr. Aldrich has done and criticize Townsend less. Therefore we earnestly desire that you make it known in the next issue of the Journal that you greatly regret the publication of this article and extend to Dr. Aldrich your sincere apology."

With which the Publication Committee also published a reply: "[The Committee] regrets any hard feeling has been aroused and all of use feel that Townsend went too far. In fact, it seems to us that he spoiled his won case, if he has one, by indulging in personalities."

Townsend, the purported "insider historian", never returned to North America.

Concluded in Part V.

Canfield - Field Notes on Science and Nature

Just what is the best way to record and organize my research notes? I've long been interested in answering this question. On the first day of my master's degree, I started a bound notebook (I love the squared, softcover Moleskine notebooks), and used it for caddisfly-exclusive notes. In particular, the notebook filled with sketches, observations, and thoughts pertaining to my work on the genus Cheumatopsyche. When I started my PhD program, I did the same for my tachinid research. I also keep a Grinnel-style triad of field journal, catalog, and species accounts for any field work and dragonfly observations. These are less often used, as I spend most of my time in the lab looking at dead specimens under the microscope.

I had been meaning to read Field Notes on Science and Nature for several years now, and finally just got around to it. The book is a mixed collection of biologists, anthropologists, and geologists, writing about their methods of taking notes in the field (whatever "the field" might be). The individual chapters are accompanied by photographs of the actual field notes, so you get both the text explanation of methods as well as a visual example. The primary methods of these researchers range from the above mentioned Grinnel system, to more informal collections of notes and drawings, to careful logs of stratigraphy, to the completely electronic recording system of insect taxonomist Piotr Nasrecki.

However, the overall feel is less that of a textbook on field work and more artbook-slash-nature journal. Most of the chapter authors supply prose accounts of exciting field observations, particularly those working with large mammals. And the journals in themselves are both art and historical artifacts; they carry information, but are also pleasant to look at.

One theme stretching through the work that seemed most important to me was that "the field" is not necessarily out in nature. It can be, in many people's research, simply in the presence of potentially living specimens. For me, viewing specimens at the microscope is "the field", and the notebook in which I record my observations is my "field journal". Another point many authors made was that observations should be recorded as soon as possible, in a permanent method which other people can use in the future. Who knows what piece of information may be useful?

I appreciated these and other suggestions on design and maintinance of field notes, including Jenny Keller's heuristic for drawing biological specimens in the chapter "Why Sketch?". I have been illustrating genitalia for some time now, but I have no formal art training, so some of her methods were completely unknown to me.

I recommend this book for anyone who does natural history research, because, even if you have already found your perfect method, you will appreciate the diversity of approaches to keeping notes in the field.

CHT Townsend, Vandal of the Calypterates. Part III.

When we left off, in 1908, Aldrich's review of Williston's Manual of North American Diptera marked the "second bitter hatred" against Townsend. We leave Aldrich for a time and focus now on the events marking the "third bitter hatred" of William R. Walton.

That same year, Townsend began his work on the reproductive system of tachinid flies. This would lead to his doctorate in 1914 at George Washington University. A preliminary summary of this research was published in 1911. At the same time, he was taking collecting trips to Peru, and becoming increasingly disturbed by a large number of his names being relegated to synonymy. This resulted in the paper "On Muscoid and Especially Tachinid Synonymy" (June 1911).

This particular paper is not recounted in Townsend's 1925 history, but it is interesting for reasons beyond the content and timing. One, it is the first of two papers he published on the topic of synonymy, the other in 1927. And second, the journal is Science, which shows you how much that journal has changed over the past century, and how far taxonomy has fallen. At that time taxonomy was still considered a worthy and important pursuit by the majority of scientists, and a worker as prolific as Townsend was a superstar.

Aside from the mixed criticism of Aldrich and Coquillett, and praise for Brauer and Bergenstaumm, there are some opinions that seem ironic in the context of his own work. He calls for the careful examination of types, and wishes for it to lead onto synonymy. He writes,

"The statement that I am going to make now will probably astonish some people, but I can truthfully say that I would be greatly pleased to see half the generic and specific names that have been proposed in the Muscoidea safely relegated to the synonymy where they could rest undisturbed and buried forever, with no hope of a resurrection, a goodly sprinkling of my own among the number; but such a considerable reduction of names is hardly possible of realization. Looking toward a consummation of final synonymy, however, I shall hope to accomplish in the next few years some portion of the work necessary to this end, during the course of which I here pledge my word that those generic and specific names of my own making will receive the same impartial treatment at my hands as all others. My one wish in this matter is to secure certainty before putting a name into the synonymy. The making of incorrect synonymy is a much more serious taxonomic offense than proposing further names for forms already named. In the latter case the forms can always be definitely referred to by means of the names that have been bestowed upon them, but in the former case serious confusion is certain to ensue."

Indeed, it is astonishing. Without context, the writer would seem to be inclined towards synonymy and stability of nomenclature, but as Townsend's history and future shows, he is anything but.

1911 also marks the death of Daniel W. Coquillett, the man who, if you recall, was the "first bitter hatred against Townsend". Freed of Coquillett's reaction, Townsend published his "Readjustment of muscoid names" (1912), in which he claims that "dipterological nomenclature is getting a severe shaking up, and the Muscoidea comes in for their share." It was a victory dance of sorts.

Unfortunately, and possibly unbeknownst to Townsend, William R. Walton had become friends with Coquillett shortly before his death. He became, as Townsend put it in his history, Coquillett's "staunch defender".

At the November 7th meeting of the Entomological Society of Washington, Walton presented a paper "The variation of structural characters used in the classification of some muscoidean flies" (published in the Proceedings (1913)). He argued against the use of "characteristic bristles" as the main source of generic differences within calypterate fly classification, which Townsend and others had used to great effect. These include the number of hairs in lines on the thorax and the abdomen, and the apparent hairiness of the eyes. While some of these hair-based characters are conserved across lineages, others vary within species or even populations. Numbers and size of hairs can also be linked with the amount of food a larva receives during it's development, or the sex of the individual. Thus, Walton gives 4 recommendations:

"a. The erection of a genus on a single example of either is folly and should not be permitted.

b. The proposal of a new species on a single specimen or series representing only one sex is inadvisable.

c. The creation of either a genus or species on solely chaetotactic characters without a careful study of ample material is unwise.

d. The variants of a species should be conserved under species name until good and sufficient evidence is adduced prove they are otherwise. The splitting of species in the genus Lucilia as practiced by Mr. Townsend is a negative example what is here meant."

These are actually very good recommendations, still so 100 years later but especially so in the early 20th century. It was common back then to describe new species (or genera!) based on a single specimen, or on a series of one sex. For example, the male of Eucelatoria gladiatrix was originally described as Proroglutea pilligera; the female was named Xiphomyia gladiatrix. Both were from single specimens (and both by Townsend). Careful taxonomy is what Walton is asking for, and Townsend was considered to be a prime offender. He closes,

"It seems possible that the studies of the internal anatomy of these flies upon which Mr. CHT Townsend is at present working may eventually prove useful as an index to group relations. But the mass of undigested facts, near facts, and conjecture with which he is at present deluging the devoted heads of his confreres will require an immense amount of elucidation, rearrangement, and generous elimination before becoming available for use. To conclude, there is great need of careful rearings of species belonging to homogeneous groups, from known parents, for the purpose of studying variation of structure, color, and size within the species and, failing which our knowledge of the true relations of the Muscoidean flies will never extend much beyond its present meager limits."

 Townsend was quick to respond, and fired off three papers on taxonomic theory before the end of 1913. The first of these, "Criticism and Muscoid taxonomy", is a direct rebuttal to Walton's paper. Townsend appreciated Walton's dissection of variable characters, but disagreed with Walton's recommendations, especially the first. Walton is declared "a champion of Mr. Coquillett's work on muscoid flies", and that "time will fully demonstrate whatever merit that work may possess, and no one's commendation can increase its merit one whit." "Beginners" and "new students" are clearly not suited to make these judgements, and should wait until they have enough experience before they do. Townsend, in his own opinion, is the expert. Any may come and join him, there is more than enough work to be done; "no one need harbor petty jealousy of another's work." Townsend concludes, 

"It is unwise and unseemly for a beginner in a difficult subject to ridicule good work done by his predecessors. Caustic comment has no legitimate place in taxonomic literature, and solves no problems. In the minds of all right-thinking persons such comment serves no other purpose than to reflect on the commentator. I bespeak a spirit of cordial cooperation on the part of my confreres. Such spirit will be both highly appreciated and warmly reciprocated."

That is, as long as his "confreres" fall in line with his cherished taxonomic opinions. All are welcome, but know I am the ultimate authority. This line was laid out in his next paper, "A new application of taxonomic principles", which is a rambling and convoluted account of his "typic-atypic" system. I honestly can't make any sense of it, except that it seems like he is proposing a taxonomic unit between genus and subtribe. Perhaps this is an early formulation of his "natural genus" concept (see Part I for a full explanation).

The final 1913 paper, "Notes on Exoristidae" is full of the same haughty language, same mixed criticism and judgement upon other workers seen in the above 1911 paper. He praises Walton for his attention to detail and illustrations, and equally condemns Coquillett, calling the former works "constructive" and the latter works "destructive". Townsend writes,

"What is needed in the Muscoidea and especially in the Exoristidae [i.e., the subfamily Exoristinae of Tachinidae] and more nearly allied families, is an intensive study of the numerous forms thoroughly and conscientiously carried through, without bias and with that keen adjustment of character values and natural appreciation of phylogenetic relations which stamp the master zoölogist. Each one of us must strive as best he can to attain this result."

Among the "destructive" poseurs and the "creative" "master zoölogists", it is clear from Townsend's writing where he thought himself to stand.

In 1914, Walton published a scathing rebuttal to the "Notes on Exoristidae" and Townsend's opinions of Coquillett in general. He titled it "On the Work of the Late Daniel W. Coquillett and Others", but it's clear from the first sentence that the whole and entire target is Townsend. What follows is a catalog of errors, from outright mistakes in descriptions of type specimens (legs cannot be both "not yellow" and "wholly yellow"), to imagined microscopic characters and overlooked obvious characters, to the large number of Townsend names that had been rightfully sunk into oblivion.  It also contains more than a few beautiful burns, including my favorite:

"It would seem that the possession of "that keen judgement of character values and natural appreciation of phylogenetic relations," cannot preserve even a "master zoologist" from palpable error when he does take sufficient care to see what is visible."

Ouch.

The whole work is delightful and worth a read (free at the JSTOR link above), especially the final words:

"But I conceive these criticisms would much better be said now, while the subject of them is present to explain this position, than in some distant future, when time shall have sealed his lips and stayed his busy pen forever. His fine command of English and evident scholarship will then avail him nothing, if some surviving, or perhaps yet unborn student rise up and brand his work destructive."

Thank you, Mr. Walton, for predicting that piece of irony. We are doing just that. Indeed, my current research hinges upon fixing at least part of Townsend's destructive mess.

As far as I am aware, no more shots were fired between Townsend and Walton in public.  The 1925 history reports, "Here was born a third hatred of Townsend which became very bitter until it was fortunately dispelled a few years later." He does not record anything more.

Meanwhile, the conflict between Townsend and his US National Museum supervisor John Merton Aldrich was intensifying.

Continued in Part IV.