Unrecognized species of Bembidiina in the USA and Canada

I’ve been mentioning in this blog several unrecognized species which we have been finding in the fauna of Bembidiina in America north of México. By “unrecognized” I mean both undescribed species (those without a name), and species that have already been described, but that are currently and incorrectly considered in the literature to be a synonym of another species.

For my amusement, I decided to tally up how many species we now know about that are unrecognized in the literature, and I present the following list. The number of unrecognized species in the genus or subgenus or species group is listed in parentheses. A “+” after the number means we have hints of additional species, and ++ means that there are surely many more species, but we haven’t worked as much on that group yet and so we don’t understand how many more there are. I have not included the genus Amerizus, as Dave Kavanaugh is working on that group and is more familiar with it.

This list will continue to grow as we focus on various groups, but for now it gives a sense that we already know of a lot of undocumented species.  For context, I should mentioned that there are about 250 recognized species in Canada and the continental USA.

Unrecognized species of  Bembidiina (excluding Amerizus) in Canada and the continental USA

  • Lionepha (2+)
  • Bembidion (40++)
    Subgenera of Bembidion:

    • Lindrochthus (1+)
    • Furcacampa (2+)
    • Notaphus (5++)
    • Trepanedoris (2++)
    • Ocydromus (1)
    • Peryphus (2)
    • “Nearctic Clade” (5+)
    • Hydriomicrus (1)
    • Plataphus (16+)
      • breve group (4+)
      • arcticum group (1)
      • kuprianovi group (2)
      • gebleri group (2+)
      • prasinum (i.e., rusticum/sulcipenne/hyperboraeorum) group (1+)
      • curtulatum group (4)
      • planiusculum group (2)
    • Hydrium (1)
    • Trechonepha (1)
    • Liocosmius (3+)

This totals to 42 unrecognized species.  Immediately evident is the unrecognized diversity within the subgenus Plataphus; at least 16 species unrecognized, and there are likely more.  This high number is partly because we have done a lot of DNA sequencing on that group.  It would not surprise me if subgenus Notaphus might rival that once we study them more.  Subgenus Trepanedoris might not hold as many new ones, but it clearly has quite a few. Subgenus Hirmoplataphus is very complex, and may have many undetected species.

Update: see my follow-up post about showing the geographic distribution of these species.

Posted in Revising Bembidiina, Taxonomic Process | 1 Comment

The legacy of a taxonomist

I have recently been looking through the notes of Kenneth W. Cooper, a renaissance entomologist who wore many hats, one of which was as someone passionate about the same beetles I love, the genus Bembidion.  Kenneth and I began corresponding about these beetles when I was a graduate student, and in later years we discovered a mutual interest in the subgenus Liocosmius, and began a collaboration on that group.  Kenneth gave me his notes in 2005, so that I might use them to finish our joint project.  Kenneth died in 2007, at age 94.

Kenneth’s notes are a treasure trove of detailed information, and a perusal of them led me to ponder the legacy of taxonomists, and our struggles to make those legacies last.

Kenneth Cooper (right) and me.  Riverside, California, January 2005.

Kenneth Cooper (right) and me. Riverside, California, January 2005.

Those of us who are explorers in the land of biodiversity are continually gathering new data, even when we don’t intend to.   As we follow the backwaters and trails laid down by our predecessors,  and then strike out through the bush on our own, we are continually noting patterns in biodiversity’s tapestry.  Through the years, we construct in our brains a detailed map of whatever group we are fascinated in.  For me it is Bembidion and other carabids; for you it might be a clade of spiders, or a group of fungi.

Much of that map of life, including the patterns of morphological and ecological variation, is housed in the delicate folds of tissue inside our skulls.  We also occasionally write down into our notebooks (paper in the past, now digital) observations about the variation in one subgenus, or an unusual series of specimens from a recently sampled locality, or the details of some morphological structure.  Even more rarely some subset of these discoveries gets synthesized into a paper that is published, for the rest of the world to see.  As frequently, our knowledge dies with us.

The nature of the taxonomy is such that it is perhaps particularly prone to such losses.  The pattern recognition engines and data storage in our brains makes us all natural taxonomists; we are continually building maps of the world around us, although of course most humans don’t build maps of small beetles.  When taxonomists go into the field or museum, they are continually adding to this map, including in areas they had not meant to explore.  When I go into the field and look for a particular Bembidion, I will find other Bembidion, notice new species, observe previously unknown microhabitat associations, and see geographic patterns I hadn’t noticed before.  I store that information in my brain (and notebooks), with the intention that the newly discovered territory in my map will be published, someday, but, if you please, after I finish the last ten or fifty discoveries I serendipitously uncovered as I explored.

It is both one of the joys and burdens of taxonomy that we have this vast territory to map (at least in groups like insects), and discoveries are so easy to come by that we are continually, often unintentionally,  uncovering nuggets worthy of record.  In other fields of  science one might plan an experiment, execute it, and gather the data, with the rhythm of the work leading to natural breakpoints for writeup and publication.  This is less the case with taxonomy, where we are bombarded, willy-nilly, by the revealing of small new puzzle pieces in our enormous maps.

Kenneth’s notes illustrate the burden. I have papers and notebooks of his that would fill two filing-cabinet drawers.  About a quarter of this is on the subgenus Liocosmius, a group of delicate, pretty Bembidion in western North America.

An undescribed species of Bembidion (Liocosmius) from California

An undescribed species of Bembidion (Liocosmius) from California

Kenneth had become interested in this group in the 1960s, and his notes indicate an ever-expanding fascination with them.  He delved into the detailed structure of their genitalia, and made excellent sketches of his dissections.


He made hundreds of slide mounts of genitalic preps, and filled notebooks with standardized drawings of every specimen.


He took copious measurements, and made table after table containing data.


He was puzzled over the variation in the number of setae on the eltyra of these beetles, and started to explore the literature on fluctuating asymmetry. He filled pages with speculation about species boundaries as he tackled this difficult group.  He marveled at  Laboulbeniales (a group of very cool fungi – more about those later) on the beetles’ bodies, and carefully collected data on those, too.

His notes are a treasure map, but only he could unlock some of the secrets: these notes were meant for his eyes, not mine.  The parts I can interpret show a very careful, thoughtful, thorough mind; the drawings are beautiful. However, there is so much information he gathered, and so many paths that Kenneth’s exploration led him, that he never managed to rein it all in and publish.

Kenneth was not alone in this by any means.  The shelves, filing cabinets, and museum drawers of any taxonomist of his generation (or mine, for that matter) are filled with knowledge of which the world at large is unaware.  I remember visiting Henry Dybas (a world expert in featherwing beetles) in 1980, not long before his death, and heard many tales and saw many drawings of amazingly cool things about ptiliids.  Some of that knowledge died with Henry, and some is now only in his notebooks, or in the brains of the very few other people who have seen his notebooks.  I am sure that when I die many things I have learned will not be passed via paper, bits, or human brains, and will have to be discovered anew by future generations.

Also at play here is the nature of those who choose to be taxonomists.  I suspect we in general are people who like everything neat and tidy, organized, and complete.  It is rare that we sense that we have fully understood a group, and so we are perhaps prone to wait for that never-to-be-achieved enlightenment before we publish, and yet we continue to accumulate knowledge.

One of the goals of my current revision of North American Bembidiina is to participate in the development of tools to capture some of this information before it is lost, via quantum publication.  But it will require more than just tools – it will require a rethinking of how taxonomists view their work.  How we can use the pattern-recognition engines and memory banks of our brains to our best advantage, in the context of the administrative structure of modern science, is something I struggle with; a blog post on this is in my near future.

Posted in Academia, Taxonomic Process | Tagged , , , | 6 Comments

Sheep Creek in the Cascades of Oregon

Here’s a lovely creek in the Cascades of Oregon.   This part of Sheep Creek is at 795m elevation.  It has a rich bembidiine fauna living in the gravel patches along the shoreline.  In addition to two species of Lionepha (L. casta and L. sequoiae), there are six species of Bembidion.  Five of the species belonging to the Plataphus Complex of Bembidion: B. turbatum, B. curtulatum, B. haruspex, B. nigrocoeruleum, and an undescribed species related to B. sierricolaB. iridescens, a widespread species in the Pacific Northwest, also lives here.


Posted in Collecting Site of the Day | Tagged , , | Leave a comment

Programming Zephyr


My brother Wayne is visiting, and we are making very good progress on Zephyr. Programming is especially appealing when the outside world is covered in quiet snow.

David (left), Wayne (right)

David (left), Wayne (right)

Zephyr is a package for Mesquite that links Mesquite to phylogeny inference programs.  For example, one can have a matrix of DNA sequences in Mesquite, and ask that GARLI finds the maximum likelihood tree.  A dialog box from Zephyr pops up with GARLI settings, you choose them, and then Zephyr starts up GARLI.  As GARLI is running, Mesquite/Zephyr is monitoring its progress, and showing you the trees live in a tree window in Mesquite.  Once GARLI is done, Mesquite harvests the trees or trees and stores them in the current project.  

Below is a view of Mesquite with GARLI running in the background (in the black terminal window at right), and the latest GARLI tree shown in the tree window.  As GARLI runs, one can watch that tree being continually adjusted. (Touch on the image below to see a larger version of it.)

Screen Shot 2013-12-11 at 10.41.18 AM

We’ve been using Zephyr for a few years in our own work, and are now putting the effort in to making it usable by others.  So far it enables one to run GARLI, RAxML, TNT, and PAUP from within Mesquite, although the options for TNT and PAUP are limited.

We’re very pleased with the progress we’ve made, and hopefully will have something available for folks in the spring sometime.

(By the way, the image on the Zephyr banner is a closeup of the elytron of Bembidion zephyrum, one of my favorite beetles.  It lives on the sandy beaches of the Pacific Ocean in western North America.)

Posted in Mesquite, Software Development | 2 Comments

Dreaming of Saint Helena

Saint Helena is a small island in the Atlantic Ocean that is a very long way from any continent. It is approximately 1850 km from Africa, and about 3290 km from South America.  It is only 16 km by 8 km in size, and the highest peak is 818 m above sea level. It is the place where Napoleon was exiled, and where he lived the last five and a half years of his life.  It currently has a population of about 5,000 people, and is a British Overseas Territory.

And it is a place with an endemic radiation of extremely odd bembidiine carabids. I have dreamed of going there to collect some for DNA studies, to find out where they belong in the bembidiine evolutionary tree.  They are currently classified as genera separate from Bembidion.

Here is where Saint Helena lies:

I’ve looked into traveling to Saint Helena, and it is not a trivial undertaking.  There is no airport in Saint Helena (although one is under construction with a scheduled opening in 2016), and the typical route is to fly to Capetown, South Africa, and from there take a Royal Mail Ship to the island, a sea journey of five days (one way).

The flora and fauna of Saint Helena has many endemics, including all but one of the bembidiines.  In addition to the widespread Bembidion mixtum, the bembidiine fauna consists of only 12 described species, but these are fantastically diverse.  Here are three of the species:


From left to right, Pseudophilochthus nubigena, P. rufosuffusus, and Apteromimus platyderoides,
all to the same scale. Scale bar is 1 mm.

Some of these simply don’t look like bembidiines.  The oddest one isn’t shown here – it is Endosomatium megalops, whose huge head is vastly out of proportion to the rest of its body.

Unfortunately, some or all of these may now be extinct.  The introduction of  invasive plants and animals, including goats and Homo sapiens, has destroyed much of the habitat in which these beetles lived – the special forests on the island.  Howard Mendel, as part of his month-long survey in 2005-2006, sought these beetles, and only found a single specimen of an undescribed species.  As Mendel, Ashmole, and Ashmole note in their survey report (available here), “we failed to find any of the twelve known species in these three endemic genera. They are of ancient lineage and closely associated with the cabbage tree / Tree Fern cloud forest on the Peaks. Their survival is of grave concern and in serious doubt. ”

This has made me rethink going to Saint Helena, in part because I think more work needs to be done re-establishing the beetles’  forest habitat before any more collecting is attempted.  Instead, I have focused my attention on seeing if I can extract and sequence DNA from some of the pinned specimens already in museums.   To that end, I borrowed specimens of three species from the Royal Museum for Central Africa in Tervuren, Belgium. I’ve extracted DNA from one of each of these.  The DNA from one of them, a Pseudophilochthus nubigena, looks as if it will be of high-enough quality to allow us to obtain sequences using next-generation sequencing.  And, in fact, we have PCR products for 500 bases of 28S and 450 bases of the wingless gene.

In advance of the next-generation sequencing, we will be sending the PCR products off to be sequenced, and hope to have results within a week.  We should have the first information then about where at least that one species falls.

And so, I will make the following prediction: the Saint Helena species are simply highly derived Bembidion, and should not be treated as separate genera.  They form a single, endemic clade (to test this, sequences from more than one species will be required).  My first guess is that they belong to the Bembidion Series (Maddison 2012), perhaps near the Antiperyphanes Complex of South America or the Ananotaphus Complex of Australia, New Zealand, and Hawaii.  My second guess is that they are in the Philochthus Complex.  These guesses are based more on gestalt than a careful character analyses;  the dominance of the Bembidion Series in the southern hemisphere fauna also sways me.

Posted in Tree of Life | Tagged , , , , | 7 Comments

The Group Photo

One of the required products of a weekend field trip is a group photo.

On our Big Loop Trip this summer, we joined up with the Essig Museum group for one weekend in the southern Sierras, and had a very fun time with them.  Just before Kip, John, and I departed for New Mexico, we took a group photo.  Here it is. and I would claim that, while nice, it is relatively boring.  We are all standing up nice and straight, which is good, but it just doesn’t have much compelling about it.


Much more interesting is the progression toward that point.  Instead of a group photo, why not a group stop-motion animation, frame by frame?  It might start with this picture


and then one of the players might move to the side, making room for other folks,


and there can be a dash as one of the photographers (Joyce Gross) runs into view after setting up her camera


and then the other photographer (me) steps in


and we are almost ready


and now that we have established our boring poses, the default group photo:


If I hadn’t had the camera going earlier, I wouldn’t have realized how much I prefer one of the less static images.  The one I like the best is one of the middle ones, after we were all in view, but with just enough dynamism in it to be interesting, and to give a sense of spontaneity. This one:


Much better.  We look much more lively.  By maybe all of us should be running around?  We should have taken a picture while we were all pretending to be scurrying insects.

Posted in Fieldwork, Musings | Leave a comment

My favorite phylogeny

In a previous post, I showed my second-favorite phylogeny. Well, here’s my favorite one, posted on the wall outside my lab.

bembTreeNot that you can really see any details in this picture, but you get the idea:  it’s big.  It has about 600 species of Bembidion and related genera.  That’s also a maximum likelihood bootstrap tree based on seven genes, and you can see that it is pretty well resolved – there are very few polytomies.  This tree gives me so much pleasure because it tells me so much about the history of this group of beetles, and because there are so many things that make sense in light of it.

I’ll share some details about the tree later, but for now I would direct your attention to the caption under the tree.  I didn’t add that caption; it mysteriously appeared one day, like a gift-wrapped present under a sparkling tree.  I really, really like it.  Here’s what it says:


I have no idea who wrote it.

It would be very amusing to have that as the caption in the journal article when the phylogeny gets published.

Posted in fun stuff, Tree of Life | 2 Comments

Art.Science: Shedding a Scenic Tear

Last week I had what I hope will be a life-changing experience.


Art by (from left to right) Matt Norris, Mindy Lighthipe, Marjorie Moore and Cornelia Hesse-Honegger;
layout by Tuan Tang.

This fall I had the honor of being a juror for an exhibit of insect-inspired art, called ECLOSION, hosted by  Art.Science.Gallery. in Austin, Texas.  The event was organized by Hayley Gillespie (Art.Science.Gallery. founder) and Barrett Klein, both biologists/artists.   I’ve known Barrett for quite a few years, ever since he was a graduate student in one of my classes in Arizona; his brilliant melding of art and science was one of the reasons I was so pleased to help out with ECLOSION.  (If you haven’t seen it yet, you must look at the Damselflies of Texas field guide which Barrett illustrated – it’s gorgeous.)

[Note: all images in this post are courtesy of Art.Science.Gallery.]


A portion of ECLOSION at Art.Science.Gallery.

The opening of ECLOSION coincided with the 61st annual meeting of the Entomological Society of America (ESA) in Austin. I have been going to these meetings intermittently since my first one in 1981 in San Diego.  I have always enjoying catching up with old friends, and seeing what cool research folks were doing, especially that being done by graduate students and junior faculty members. But the ESA meeting last week was special for me because of two talks in a “Broader Impacts” symposium.  The first was by Hayley on “Using art to enhance your science” and the second was by Barrett called “Entomoart: the power of esthetically communicating entomology”.

I had a very strong emotional reaction to these two talks, and at one point I am sure the smallest tear might have been visible in my eyes to anyone who looked closely. I sat there, as Hayley was talking, thinking, “What have I done with my life? Why am I not doing my artwork anymore?”  I suppose in part the answer is the hectic pace of my biological work, which seems to leave little time for me to re-enter the world of art.  But as I was sitting in the audience I realized that I must return to my art, and I have vowed to do so.  This is just too important to me not to.  Hayley’s and Barrett’s talks also made me realize how important art can be not only to inspire scientists in their work and inspire people to be scientists, but also to inspire artists to appreciate the wonders of science and the natural world.

Re-energized, the next day I took the bus over to Art.Science.Gallery., and saw the exhibit in person.  It was fantastic.  There were some gorgeous pieces there, in both two and three dimensions. Some of these were art that conveyed scientific knowledge about insects, others presented in a dramatic fashion messages about conservation biology or invasive species, and others celebrated the striking form and beauty of insects.

Although there were many pieces that appealed to me, I’ll mention only a few.

Barrett Klein’s “Biodiversity” is a pie-chart showing the relative diversity of various groups of organisms, with a representative of that group used to “label” that slice of the pie:

Biodiversity by Barrett Klein

Biodiversity by Barrett Klein

I really liked the sparseness of “Differential Grasshopper 3 (Melanoplus differentialis)”, created by Inked Animal by painting parts of dead grasshoppers with inks and pressing those against the paper; it is thus like classical Japanese fish prints but done with insects.

Differential Grasshopper 3 (Melanoplus differentialis) by Inked Animal

Differential Grasshopper 3 (Melanoplus differentialis)
by Inked Animal

Pen Brady’s beautifully composed images of animals, drawn in a style reminiscent of northwestern First Nations cultures, were very inspiring:

Returns the 5th of Fibonacci by Pen Brady

Returns the 5th of Fibonacci by Pen Brady

This bronze bowl by Jessee Smith, inspired by galleries made by bark beetles, captivated me by its design, texture, and weight; it was so comforting to look at and touch:

Galleries by Jessee Smith

Galleries by Jessee Smith

In addition to static art, the exhibit featured video art, including this beautiful one by Barrett:

The importance of melding art and science came to me not only through Hayley and Barrett’s talk and the ECLOSION exhibit, but also through a chance encounter with a former student, Bruce Noll.  Bruce took my Systematic Entomology class when I was but a wee junior faculty member, 19 years ago.  Bruce is a poet who does dramatic interpretations of Walt Whitman, and who is fascinated by insects.  After all of these years it was delightful to see Bruce, with his quiet and gentle passion.  Bruce intermingles entomology or phylogenetics into his poems, which appear regularly in American Entomologist.  I realized, as I was talking to Bruce at the airport on my way home, that he too was part of this message I was receiving in Austin to return to my artistic roots.

Posted in Musings, Scientific Illustrations | Tagged , | 4 Comments

(Almost) my favorite phylogeny

I recently reported on my favorite beetle illustration.  In the same collection as that illustration was a favorite phylogeny of mine:


This drawing was also done by my daughter Julia at age 3 or 4; perhaps she was depicting what her father did at work.  (I think I need to add smiling hearts to my published phylogenies from now on. And happy faces on the beetles.)

The title of this post has the word “almost” in it because, as wonderful as this phylogeny is, my favorite has got to be our growing phylogenetic tree of the subtribe Bembidiina – 600 species and counting.

Posted in fun stuff | 3 Comments