Dr. Rob Dillon, Coordinator

Sunday, August 6, 2017

The Most Cryptic Freshwater Gastropod In The World

Through much of the 20th century, the #1 (and possibly only) expert in the biologically fascinating genus Fontigens was the legendary Mr. Leslie Hubricht (1908 – 2005).  Hubricht was trained as a repairman for adding machines and (later) for computers, but spent most of his life travelling around the United States in a van, digging through humus, rolling logs, and peering into holes [1].  He published over 150 scientific papers, primarily on land snails but also on the biota of caves.  Among these were 4 - 5 papers on spring or groundwater-dwelling hydrobiid snails, especially Fontigens [2].

So last month I shared an inspiring story about Lori Schroeder and her discovery of the tiny shells of a mysterious Fontigens species in deposits of storm water flotsam on the margins of several creeks at the Bernheim Forest in central Kentucky [3].  And I mentioned in passing that our good friend Bob Hershler, together with Leslie Hubricht and the cave biologist John Holsinger, had listed nine “recognized species” of Fontigens in their 1990 monograph [4].  Let me back up and expand that statement slightly.

Hershler and his colleagues actually listed ten species, nine of which they “recognized” and one of which was of “questionable status.”  That tenth species, mentioned in a single concluding paragraph on page 43, was Fontigens cryptica.

Leslie Hubricht described Fontigens cryptica in 1963 from under stones in a small spring along the Ohio River in southeast Indiana [5].  He seems to have had at least one living specimen in hand, because he described the animal as “translucent whitish, blind.”  But he immediately confessed, “verge unknown.”  Hershler reported in 1990 that he “was unable to find the snail during two recent visits to the type locality (and nearby localities), nor was it taken during an extensive survey of subterranean habitats in the region (Lewis 1983).”  Hence the uncertainty of its status.

Above I have scanned the 1.9 mm holotype figured by Hubricht in 1963 next to a photo of a 1.9 specimen collected last year by Lori Schroeder.

Notice also the reference to a "Lewis 1983" in the Hershler quote reproduced above.  My attention was called to the existence of this tenth, questionable-status Fontigens by none other than Dr. Julian J. Lewis himself, still very active in karst, cave, and groundwater research in Kentucky today.

By one of those strokes of fortune I have begun to take for granted in my long career, it materialized that in 2016 the Bernheim management engaged Lewis and Associates LLC to survey the subterranean fauna of its caves and springs, and that professional surveys of the entire property had been underway for several months prior to the date that Lori Schroeder first brought her Fontigens discovery to my attention.

So in April Lori offered me an electronic introduction to Dr. Julian (Jerry) Lewis, and we struck up a conversation.  And Jerry confirmed: 
“Bob Hershler and I looked for this species together at the type-locality at a spring in Clark County, Indiana and I've been there repeatedly with no luck.  The spring is high on a bluff overlooking the Ohio River and consists of a hole about the size of one's fist....not much habitat in which to search.  Subsequently I pulled shells of this species from a meter below the surface of the Blue River  -  from groundwater flowing through the hyporheic zone, in the company with a number of other non-cave subterranean species - using a Bou-Rouche sampling pumpwell. 
 I found the snails alive in the interstices of a gravels in a similar situation in a cave in Monroe County, Indiana (near Bloomington) using Karaman-Chappuis extraction. 
 So I suspect your snails are probably a groundwater species, likely living in the interstices of gravel and sand, and will require special sampling methods."
So no, Lewis and Associates LLC had not recovered any Fontigens whatsoever in their survey of the Bernheim property just recently concluded as of this spring [6].   Nor indeed was it Jerry’s expert opinion that we should expect to find any.  Jerry’s studies of the entire regional biota have led him to hypothesize the existence of a single subterranean zoogeographic province extending at least as far south as Mammoth Cave in central Kentucky and as far north as the Blue River drainage of Indiana.  But he has never seen a population of Fontigens cryptica in any cave stream he has explored, ever.  Gravel under a cave stream, yes.  But the little snail is no more an inhabitant of open flowing water under the ground than of open flowing water at the top.  Fontigens cryptica seems to be limited to a third, even more mysterious habitat: saturated interstitial spaces.

I concluded last month’s blog post with a series of three teaser questions: “From what dark recesses of central Kentucky knobland might Lori’s tiny little snails be emanating?  And what might be their identity?  And will Lori Schroeder surrender her quest?”  This month I have addressed questions #1 and #2.  Next month I will address #3.  And here’s your teaser.  No.


[1] He stood silently in the lobby outside AMU meetings in the 1970s and 1980s, wearing a stiff dark suit.  I wish I had gotten to know him.

[2]  Gerber, J. (2010)  Leslie Hubricht (1908 – 2005), His publications and new taxa.  American Malacological Bulletin 28:15-27.

[3] Lori Schroeder’s Tiny Snails [17July17]

[4] Hershler, R., J. R. Holsinger and L. Hubricht (1990)  A revision of the North American freshwater snail genus Fontigens (Prosobranchia: Hydrobiidae).  Smithsonian Contributions to Zoology 509: 1 – 49.

[5] Hubricht, L. (1963)  New species of Hydrobiidae.  Nautilus 76: 138 – 140.

[6] Only a single freshwater gastropod was listed among the 61 species documented by Lewis and Associates - Lymnaea humilis in a seep spring.

Monday, July 17, 2017

Lori Schroeder's Tiny Snails

Mrs. Lori Schroeder manages her husband’s dental practice in Bardstown, Kentucky.  She is also a skilled and dedicated malacologist, who in recent years has expanded her interests from seashells to land snails and into the freshwater fauna.  Back in 2009 she volunteered to lead a “Bioblitz” survey of the land snails of the Bernheim Arboretum and Research Forest south of Louisville [1], collaborating with our good friends Harry Lee, the late Henry McCullagh and Bill Frank [2].  And several years ago I was blessed to strike up a correspondence.

Organic litter by Harrison Creek
Lori’s favorite method of surveying land snails is to pick through samples of forest litter, such as are often deposited along the courses of the several creeks and streams running through the 15,000 acre Bernheim property.  So it is not surprising that she occasionally finds freshwater gastropods mixed with her land snail collections.  I received my first emails from her in 2013 regarding the pleurocerids and physids of Bernheim, which I was pleased to entertain.

And I was pleased to receive another email from Lori this past January.  She sent to my attention a jpeg photo of a small planorbid, requesting guidance on the difference between Menetus and Gyraulus.  And, as an afterthought in some follow-up correspondence, she mentioned, “I also have a tiny unidentified hydrobiid species from the same lot.”

My antennae raised immediately.  I replied, “What’s this I hear about a tiny unidentified hydrobiid?  I’d be curious to see a photo of that one as well.”  And on 1 February 2017, this is the jpeg Lori sent me:

My antennae raised even higher.  Clearly a little pile of Fontigens, that diminutive inhabitant of springs, spring runs, and caves scattered widely throughout the Eastern and Midwestern United States.  I reached to the shelf beside my desk and pulled out my well-thumbed copy of the monograph by our good friend Bob Hershler and colleagues [3].  And here is the reply I sent Lori, under the subject line “OK, now you’ve got my attention:”
Dear Lori, 
The jpegs attached to your message of earlier this afternoon appear to depict a hydrobiid population of the genus Fontigens.  My first guess would be the most common species, Fontigens nickliniana.  But upon closer examination, they might look more like published photos of Fontigens aldrichi, which I’ve never personally seen on the hoof.
The genus Fontigens was marvelously-well monographed by Bob Hershler and colleagues (1990).  And among the many surprises is that Hershler did not report any records of Fontigens of any species in Kentucky, period.  F. nickliniana is widespread in the Great Valley of Virginia, spreading up to Pennsylvania, and also in western Michigan, so one might assume that they range through Kentucky.  A lot of secondary references do indeed list F. nickliniana in your state.  But I myself have never seen a F. nickliniana population in Kentucky, and don’t know of any reliable record.
All that said, the flat nuclear whorl doesn’t really look like F. nickliniana.  Picture-matching to the other eight species recognized by Hershler, they look more like Fontigens aldrichi or Fontigens antroecetes.  But those species are not known outside of Missouri.
Bottom line.  Those shells were all dead-collected, am I right?  It would be great if you could find the living population, and send a preserved sample to Bob Hershler at the USNM.  Fontigens populations are often narrowly-restricted to springs and spring runs.  There’s a lovely, clean, clear, coldwater spring right upstream from your collecting site, am I correct?  Lots of watercress, I’ll bet?  You should find your big population of Fontigens at that spring head, covering the cress and every square inch of bottom.  Go look and tell me if I’m right.  Take a Nalgene bottle or something to collect in.
Standing by,
And here is Lori’s immediate reply: 
Hello Dr. Dillon, 
I've been down the hydrobiid road with Dr. Lee and Dr. Fred Thompson at the Florida Museum previously [4].  I failed miserably to locate any living specimens.  That being said, I am more than willing to give it a second go. 
 It subsequently materialized that my speculations about a spring head with “lots of watercress” and “Fontigens covering every square inch of bottom” were very, very wrong.  And the situation much more complicated than I initially imagined.

Lori had found her little pile of Fontigens shells sorting through debris deposited along Harrison Creek by a catastrophic flood that ravaged the area in January of 2016.  Harrison Creek washes down through a lovely valley of old fields flanked by mixed hardwoods.  Yes, there are (indeed) small springs and caves on the ridges surrounding, but most issue water only in wet weather.

A further complication was that Lori had actually found Fontigens shells in similar circumstances along three different streams on the Bernheim property during the course of her extensive studies of the area, all in dry debris deposited by floodwaters, none downstream from any spring or other obvious source of groundwater.

And on 3 February 2017 Lori sent me another set of more detailed jpegs, giving a standard shell length of just 2.1 mm.  Adult F. nickliniana reach 3 – 4 mm, in my experience.

Broadly speaking, there are two categories of Fontigens in North America, which (given my Presbyterian heritage) I think of as being either “black” or “white” [5].  The black species, such F. nickliniana, F. morrisoni and F. bottimeri, are usually somewhat larger and live above ground.  The white species, such as F. orolibas, F. tartarea and F. turritella, are usually smaller and subterranean.  The shell photos Lori sent us on Feb 3 struck me as depicting a white species – not something we might expect to find at the surface.

Undaunted, Lori and her husband Jeff set out on what she called a “full-on blitz attack,” ultimately extending over several months and involving Andrew Berry, Forest Manager at Bernheim.  And I cannot remember ever witnessing, first-hand or second, a more earnest or comprehensive survey of any patch of the earth for any mollusk population, of land or sea.  Lori and colleagues surveyed every spring or cave or hole in the ground actually, potentially, or historically upstream or in the vicinity of every location where any Fontigens specimen had ever been recovered from Bernheim.  They also installed mesh traps over the openings of several culverts draining into Harrison Creek.  And the results, to quote Lori directly, were “Nothing.  Nada.  Zilch.”

From what dark recesses of central Kentucky knobland might Lori’s tiny little snails be emanating? Indeed, what might be their identity?  And will Lori Schroeder surrender her quest?  Tune in next time! 


[1] The Bernheim Arboretum and Research Forest was originally developed in 1929 and endowed by the German philanthropist Isaac W. Bernheim “as a gift to the people of his new homeland.”  It is open to the public during regular daylight hours, with a small admission charge on the weekends.
Website: [html]

[2] The Kentucky land snail list is available at [Jaxshells-Kentucky].
Photos of many elements of the Bernheim freshwater gastropod fauna are also available from the jaxshells.org website, if you try the search button at the bottom of the front page.

[3] Hershler, R., J. R. Holsinger and L. Hubricht (1990)  A revision of the North American freshwater snail genus Fontigens (Prosobranchia: Hydrobiidae).  Smithsonian Contributions to Zoology 509: 1 – 49.

[4] Yes, it materialized that the 2016 specimens were not the first Fontigens Lori had found in her nine-year survey of Bernheim.  Several years previous she had corresponded briefly with Fred Thompson about the little snails, but Fred was not interested unless she could find live material, and the matter dropped.

[5]  For an illustration of the striking black/white dichotomy see:
  • Springsnails of The Blue Ridge [26July06]

Tuesday, June 27, 2017

The Freshwater Gastropods of The Ohio: An Interim Report

By popular demand!  Several of you have requested copies of the presentation I contributed at the Society for Freshwater Science meeting in Raleigh earlier this month.  A pdf download is available for the clicking below.

My collaborators on the work were Ryan Evans of KYDOW, Mark Pyron of Ball State, Tom Watters of Ohio State, Will Reeves of the USDA in Ft. Collins, Richard Kugblenu of SUNY-Albany, and Jeff Bailey & Mike Whitman of WVDEP.  Here’s the abstract: 
We report preliminary results from a survey of the freshwater gastropod fauna from the Ohio River basin above Paducah, including western Pennsylvania and most of West Virginia, Ohio, Kentucky, and Indiana.  Our database of 4,746 records was almost entirely drawn from state natural resource agencies (36%), museums (30%) and our own original collections (30%), almost all personally examined and verified.  We report 66 species and subspecies of freshwater gastropods [1], the most common of which are Physa acuta (959 incidences), Ferrissia rivularis (536), and Pleurocera semicarinata (all subspecies combined 528).  Nine species were collected in but a single population, four of which seem to be legitimately rare, the remainder peripheral.  The distribution of commonness and rarity appeared log-2 bimodal, with mean 3.84 (14.3 incidences) and standard deviation 2.79 (6.9 incidences).  Our Ohio River basin results are compared to similar databases previously assembled from the Atlantic drainages and from East Tennessee, and the 99 species of the combined 13-state fauna ranked by their incidence.
 I should emphasize that the results as I offered them in Raleigh were very preliminary.  Subsequent to the submission of the abstract above, our GIS analysis suggested that a couple subregions of our study area had been oversampled, resulting in a prune of our database back from 4,746 records to 4,570.  And I have just returned from two weeks in the field, adding quite a few sample sites from northern Tennessee [2] and central Kentucky, as well as a very large and impressive slug of data from Illinois [3].

Several of you have also asked me when the finished FWGO site might be expected to appear online.  That date, to be precise, is Not Quite Yet.  But we’ll keep you posted… 


[1] We recognize 6 subspecies of freshwater gastropods in the FWGO study area: the pair Campeloma decisum crassulum and C. decisum (ss), the pair Pleurocera simplex ebenum and P. simplex (ss), the pair Pleurocera canaliculata acuta and P. canaliculata (ss), the pair Pleurocera laqueata alveare and P. laqueata (ss), and the trio Pleurocera semicarinata livescens, P. semicarinata obovata, and P. semicarinata (ss).  Thus the total species we analyzed on slide #7 of our June presentation was 60.

[2] Yes, the Green River drainage includes a sliver of north central Tennessee.  The faunal similarity between Kentucky and Tennessee has been one of the biggest surprises of the FWGO survey.

[3] A tip of the hat to our good friend Kevin Cummings for his gracious hosting at the Illinois Natural History Survey.

Friday, May 26, 2017

Freshwater Snails and Passerine Birds

Editor’s Note – This is the fourth in a series of essays about the aerial dispersal of freshwater gastropods.  But no, you do not have to read the other three, which are entirely independent of this one.  My previous essays were posted 15Dec16, 11Jan17, and 24Apr17, if you’re curious [1].  But that material won’t be on the test.

So back in April of 2015 I received an intriguing email from our good friend Nora Foster, way up in Fairbanks, Alaska.  She wrote: 
"A colleague of mine has just brought me a whole sh-- load of tree swallow nests. It seems that the little birds select pond snails, presumably for food, there's some indication that it's a response to low calcium availability. I've only found one paper on this [2].  I wonder if you know of any other work done on freshwater snails being consumed by small birds. The shells are intact, by the way, I'm not sure if the swallows are spitting out the shells, or just what's going on."
From a swallow nest in Alaska
My quick answer was no, that I was not aware of any other work on the consumption of freshwater snails by small birds.  Stacks of papers have been published about ducks and other waterfowl, of course.  We’ve reviewed some of that work in recent months.  And there’s the Everglades kite, which is a raptor, feeding on Pomacea.  But no, Nora’s email of 4/15 was the first report I had ever heard that little dicky-birds might eat freshwater snails.

So Nora and I struck up a correspondence that extended a couple weeks.  Nora found one reference suggesting that momma swallows may bring freshwater snails back to their hatchlings and regurgitate them as food provisions [3].  Perhaps the empty shells we subsequently find in swallow nests were food offerings rejected by the hatchlings, or lost?  Not swallowed?  Sorry, I couldn't resist.
L. atkaensis?

In a sense this phenomenon would seem to be the complement of the situation we reviewed in December and April.  Waterfowl are very likely to ingest freshwater snails, but the snails are very unlikely to survive.  Passerine birds are certainly much less likely to ingest freshwater snails, but the likelihood of snail survival may be much greater.  Perhaps the products of the two sets of probabilities are comparable?

Now imagine a shot of a calendar flipping 17 pages.  This past September I was pleased to receive a cordial email from our good friend Bob Prezant, introducing me to Dr. T. J. Zenzal of the Migratory Bird Research Group at the University of Southern Mississippi.  Dr. Zenzal had a photo, a question, and an interesting story to tell.

Each spring migratory birds arrive on the southern Louisiana coast after long and perilous flights across the Gulf of Mexico [4].  Dr. Zenzal has worked many migration seasons and has captured (he estimates) over 20,000 birds.  Several years ago, while he was a graduate student, one of his labmates made the remarkable collection figured below from the body of an indigo bunting.

I responded to Bob and TJ that I was “about 80% sure” those snails are Lymnaea cubensis, with “maybe a 5% chance” that they could be fat, squat Lymnaea humilis.  (I’d have to count the cusps on their first marginal to rule out the humilis hypothesis.)  I also left 15% chance that those little snails might be referable to some Central or South American taxon, like viator/viatrix, neotropica, or cousini, and that any of those nominal species from South America really is specifically distinct from cubensis.  I confessed that I have zero experience with any element of the South or Central American fauna, however, and concluded with my (gratuitous) impressions that the South American lymnaeids are terribly oversplit, and that our understanding seems to be worsening with each new research paper published on the subject [5].

But the most interesting question (by far!) is not what those snails are, but how the heck did those snails get there?  Looks like 8-9 snails, from the photo.  Buried in the breast feathers, according to TJ’s note currently in press [6].  My mind was first called back to my 2015 correspondence with Nora Foster.  Apparently passerine birds do, at least sometimes, eat snails.  Possibly as a calcium provision?

And I also recalled W. J. Rees’ charming “presidential address” of 1965, on the aerial dispersal of the Mollusca [7].  Rees retold a story from one “Prof. G. E. Beyer” regarding upland plovers: 
“When the birds arrive in Louisiana, they invariably carry ten to thirty snails under their wings.”  “The occurrence was so regular and confirmed so often after many years, that I expected the habit to be generally known.” 
Prof. Beyer hypothesized that plovers deliberately carried snails under their wings as food provisions.  But the plover is a ground-nesting bird, and so (I suppose) a skeptic might suggest an alternative hypothesis that all those snails had climbed into all those plover arm pits accidentally, if they were nesting, which I don’t suppose they probably were. 

But an indigo bunting?  A male, by the way, who wouldn’t need augmented calcium supplies, who wouldn’t be feeding hatchlings?  Does that sound like a food provision?  I suggested this idea to TJ, and he replied, quite reasonably: 
“The Indigo Bunting is primarily a seed eater, however birds can show flexible foraging during migration - leaving it possible for this species to eat snails.  However, I am not sure the bird would willingly carry snails just to consume them later - especially if the bird made a trans-Gulf flight.”
Fair enough.  So for now, at least, I myself have filed the remarkable observation of eight Lymnaea hitchhiking on an indigo bunting breast in my very large folder labelled, “queerer than we can suppose.”

I’m not sure if it’s a blessing or a curse, but the attentions of practicing biologists are often called to “one-of” phenomena.  While such phenomena do not lend themselves to scientific inquiry, they often register extremely high values on the importance-meter.  The origin of life was such a one-of phenomenon.  And so are we all.


[1]  The three previous essays were:
  • Freshwater gastropods take to the air, 1991. [15Dec16]
  • A previously unrecognized symbiosis? [11Jan17]
  • Accelerating the snail’s pace, 2012. [24Apr17]
[2] St. Louis, VL and L Breebaart (1991)  Calcium supplements in the diet of nestling tree swallows near acid sensitive lakes.  Condor 93: 286-294.

[3]  I can’t open this link, but for the record:

[4] Many songbirds most certainly do make the 1,000 km crossing directly over the Gulf of Mexico.  But some apparently go around.  The possibility that any particular bird might have circumnavigated the Gulf, rather than crossing it, cannot be ruled out.

[5] The Lymnaeidae 2012: Fossarine football [7Aug12]

[6] Zenzal, TJ, Jr, EJ Lain, and JM Sellers (in press) An Indigo Bunting (Passerina cyanea) Transporting Snails During Spring Migration.  The Wilson Journal of Ornithology

[7] Rees, WJ (1965)  The aerial dispersal of mollusca.  Proc. Malac. Soc. Lond. 36: 269 - 282.

Monday, April 24, 2017

Accelerating The Snail's Pace, 2012

My faithful readership may recall a review I posted last December on the ever-intriguing phenomenon of passive molluscan transport entitled, “Freshwater gastropods take to the air, 1991 [1].”  That essay was focused entirely on unpublished material I developed for my (2000) book over 20 years ago.  What have we learned since then?

A lot, actually.  But before training our intellectual laser beams on recent research progress in the avian dispersal of freshwater gastropods in detail, I should spend a paragraph to acknowledge a pair of more general reviews contributed in the mid-2000s by Figuerola and Green [2].  The F&G reviews are especially useful to place our admittedly-narrow focus on mollusks into the larger perspective. Most research published on the broader subject of avian dispersal in recent decades has been directed toward protozoans and microcrustaceans, as well as algal and plant propagules.  The charming review of Rees [3] was skipped over entirely by F&G, although they did cite the works of Boag [4] and Malone [5].  I found it interesting that across the entire subdiscipline of avian dispersal, more attention seems to have been directed toward internal transport in the gut passage than to external hitchhiking.

So in 2012 I was pleased to find the PhD thesis of a promising young scientist from The Netherlands named Casper van Leeuwen delivered to my snail-mail inbox.  That beautifully-printed 175 page work, with the whimsical cover illustration reproduced above, turned out to be a treasure trove of research on the avian dispersal of freshwater gastropods.  Some of the material in its eight chapters had just been published elsewhere as of 2012, other material was still on editor’s desks around the world.  Casper’s entire thesis, as well as the five journal articles it ultimately yielded, are all available from his professional website, see note [6] below.

Casper opened his studies with a general introduction, and a meta-analysis of 81 previously-published papers on the gut transport of macroinvertebrates and seeds, updating and refining the work of Figuerola & Green.  He got down to business in Chapter 3.

Casper reported the results of a set of feeding experiments involving mallards and four small species of aquatic gastropods common in The Netherlands: the little planorbid Bathyomphalus contortus and the hydrobioids Hydrobia ulvae [7], Potamopyrgus antipodarium, and Bithynia leachii.  The birds were fed 100 – 300 living snails and their feces collected for 24 hours.  Casper’s observations confirmed those of Malone, zero survivorship for Bathyomphalus, Potamopyrgus, or Bithynia.  But Casper did recover approximately 21 viable Hydrobia individuals, of 6,600 fed to the ducks.  Given the vast flocks of ducks that must visit the coastal marshes of Europe, the vast populations of Hydrobia that inhabit those marshes, and the vast expanses of time ducks have been eating snails, flying elsewhere, and pooping along the way, even a 0.32% survival rate is not negligible, I suppose.

The ducks Casper used for his Chapter 3 studies were all held in individual pens for the 24 hours following snail ingestion.  So in Chapter 4, he reported the results of an experiment to examine the effects of subsequent locomotion. Six mallards were fed 300 Hydrobia ulvae each and their feces collected during one of the three treatments that followed: isolation, wading, or swimming.  The entire experiment was repeated four times, for 4 x 3 x 300 = 3,600 snails ingested.  Casper recorded a total of 29 (0.81%) Hydrobia surviving the experiment, marginally better than his Chapter 3 results, but still “too low to compare between treatments.”  My eyeball impression of his Table 4.S1 suggests to me, however, that swimming was more harsh on the gut contents than wading, and wading more harsh than isolation.  Which makes me wonder if both of Casper’s estimates of survivorship, 0.32% and 0.81%, might be biased above their natural values.

As unlikely as all these highly-unlikely events most certainly are, I really think transport of viable freshwater gastropods on the outside of birds is less unlikely than their transport on the inside.  So in Chapter 5, Casper reported results from several experiments designed to evaluate the likelihood that snails might adhere to the feet, feathers, and bills of ducks and survive their subsequent dehydration.  The most interesting of these experiments involved three small European pulmonates: Lymnaea peregra (aka “Radix balthica[8]), Gyraulus albus, and Anisus vortex.  Casper constructed cages with shallow removable trays in the bottom, into which he introduced a mixture of aquatic macrophytes and snails at four densities [9].  Individual mallards were placed in these cages for 60 minutes, totaling 48 ducks over 4 days.  Each duck was then released to walk through a 3 meter tunnel, washed and brushed, and given a final inspection.

The bottom line was that snails were transported out of the cages in 34 of the 48 trials – more of the little planorbids than the Lymnaea.  Casper reported that most snails were shed off the birds during their tunnel walk, but that some remained attached through brushing and washing to final inspection [10].

In his Chapter 6 Casper flew the barnyard for the field, picking up his genetic tool box as he flapped out the back gate.  The Donana National Park is a protected wetland region in southwest Spain, featuring marshes, dunes, shallow streams, and over 3,000 hydrologically-isolated ponds.  Roughly half of these ponds dry during an ordinary summer, and roughly half of those remaining are inhabited by populations of Physa acuta.

The nearest source for all these hundreds of isolated Physa populations is the Guadalquivir River, and especially a set of rice field connected to the river, 10 km east of the park.  Here’s a thought-experiment.  How could you distinguish vector-mediated dispersal from dispersal by sheetwater flooding in such a situation using a microsatellite survey?

Casper and his colleagues sampled the Physa populations at 21 sites, including 16 in the isolated ponds of Donana Park and 5 in the rice fields of the Guadalquivir River and estimated genetic variability using six previously-published microsatellite markers.  And as you might expect regardless of dispersal mechanism, the 16 park populations showed significant subdivision, while the 5 rice field sample sites did not.

But here’s the key.  Casper was able to distinguish two types of Physa ponds within the park, those that were often visited by large mammals (primarily cattle) and those that were rarely visited [11].  The high-cattle ponds showed less genetic subdivision than the low-cattle ponds.

The graph below shows a measure of genetic divergence plotted as a function of distance across all 16 populations [12] in the park.  These are all-pairwise distances, so the data are not independent.  But a Mantel test returned a significant correlation between geographic distance and genetic divergence in the low-cattle ponds, not in the high-cattle-ponds.

This observation has two implications.  Most importantly, it suggests that dispersal has been vector-mediated, since there is no reason to expect sheetwater dispersal to differentiate high-cattle ponds from low-cattle ponds.  It also suggests that at least sometimes, large mammals [13] transport Physa.

These results are reminiscent of a similar study Amy Wethington and I published back in 1995 involving 10 Physa acuta populations sampled here in the South Carolina lowcountry [14].  We documented a similar correlation between genetic divergence and simple overland distance across local “sea islands,” despite intervening brackish tidal creeks.  And extrapolating the results of Van Leeuwen and his colleagues about as far as you can go, they also remind me of my dissertation research on 25 populations of Pleurocera proxima in the Southern Appalachians, published way back in the dark ages [15].

The correlation between genetic divergence and simple geographic distance overland was strong for P. proxima too, over a 200 km study area extending into three states, completely independent of drainage system or mountain ranges.  Of course, levels of genetic divergence among pleurocerid populations are much more profound than among physids, and the duration of isolation must be far longer.  I continue to think my P. proxima populations evolved tens, if not hundreds of millions of years ago, possibly at the Appalachian orogeny, and have been diverging since [16].  But I also think this isolation has been punctuated by very rare, but nevertheless significant, airlifts of individual wildebeest into the bison herds [17].

As if all the research reviewed above weren’t enough to earn Casper a Ph.D. four times over, he actually finished his thesis with a Chapter 7 survey of ITS1 rDNA sequence divergence in Lymnaea (Galba) truncatula sampled over four continents.  But let’s defer that study to another day, shall we?  I feel as though the time has come to sum up.

That freshwater gastropods most certainly can be passively dispersed over extensive distances overland has been established for at least 50 years, indeed longer [18].  More recent generations of scientists have shifted our research focus away from the whether, and increasingly toward the how.  Birds can plausibly serve as vectors for the transport of viable freshwater snails over substantial distances, more likely on their outsides than their insides.  Genetic markers suggest that dispersal across terrestrial barriers, probably by birds but sometimes by other agents, has significantly influenced rates of interpopulation divergence in freshwater gastropods.  I, for one, have become increasingly intrigued by observations of their absence than their presence.


[1] Freshwater Gastropods Take To The Air [15Dec16]

[2] Figuerola, J. and A. J. Green (2002)  Dispersal of aquatic organisms by waterbirds: a review of past research and priorities for future studies.  Freshwater Biology 47: 483-494.  Green, A. J. and J. Figuerola (2005) Recent advances in the study of long-distance dispersal of aquatic invertebrates via birds.  Diversity and Distributions 11: 149 – 156.

[3] W. J. Rees (1965) The aerial dispersal of Mollusca.  Proc. Malac. Soc. Lond. 36: 269-282.

[4] Boag, D. A. (1986) Dispersal in pond snails: Potential role of waterfowl. Can. J. Zool. 64: 904-909.

[5] Malone, C. R. (1965a) Killdeer (Charadrius vociferus) as a means of dispersal for aquatic gastropods. Ecology 46: 551-552.  Malone C.R. (1965b) Dispersal of aquatic gastropods via the intestinal tract of water birds. Nautilus, 78: 135–139.

 [6]  Casper’s professional website is www.caspervanleeuwen.info/  From there pdf downloads are available for:
  • Van Leeuwen, C.H.A. (2012) Speeding up the snail´s pace: bird-mediated dispersal of aquatic organisms.  PhD dissertation: Radboud University Nijmegen, The Netherlands.
  • Van Leeuwen, C.H.A., G. van der Velde, J.M. van Groenendael and M. Klaassen (2012).  Gut travellers: internal dispersal of aquatic organisms by waterfowl.  Journal of Biogeography 39(11): 2031-2040.
  • Van Leeuwen, C.H.A., M.L. Tollenaar, and M. Klaassen (2012).  Vector activity and propagule size affect dispersal potential by vertebrates.  Oecologia 170(1): 101-109.
  • Van Leeuwen, C.H.A., G. Van der Velde, B. Van Lith, and M. Klaassen (2012).  Experimental quantification of long distance dispersal potential of aquatic snails in the gut of migratory birds.  PLoS ONE 7:e32292.
  • Van Leeuwen, C.H.A. and G. van der Velde (2012). Prerequisites for flying snails: external transport potential of aquatic snails by waterbirds. Freshwater Science 31(3): 963-972.
  • Van Leeuwen, C.H.A., N. Huig, G. van der Velde, T.A. van Alen, C.A.M. Wagemaker, C.D.H. Sherman, M. Klaassen and J. Figuerola (2013)  How did this snail get here? Multiple dispersal vectors inferred for an aquatic invasive species.  Freshwater Biology 58(1): 88-99.
[7] Hydrobia ulvae is not a freshwater snail.  But sometimes we bend the rules in this blog, a little bit.

[8] See footnote [6] here:
  • The Lymnaeidae 2012: A clue [9July12]
[9] Casper did not count the snails in these experiments.  The four densities were created using four different volumes of mixed weed, assuming uniform distribution of the three pulmonate populations (by ratio) within.

[10] It is difficult to estimate the actual numbers of snails transported from Casper’s graphs.

[11] Casper and his colleagues evaluated the intensity of large mammal visitation using the densities of “droppings” surrounding each pond.  This is the third time my essay has touched on the subject of poo.  A new record.

[12] Not all, actually.  Casper’s figure shows high/high comparisons and low/low comparisons only.  The high/low comparisons are not plotted.

[13] Including us humans.  The large-mammal hypothesis will be weaker if the high-cattle ponds also tend to be high-waterfowl ponds as well.  Perhaps the high-cattle ponds were consistently larger?  Data regarding the relative sizes of the sample ponds would have been welcome here.

[14] Dillon, R.T., and A.R. Wethington (1995) The biogeography of sea islands: Clues from the population genetics of the freshwater snail, Physa heterostropha. Systematic Biology 44:401-409.  [PDF]

[15] Dillon, R.T. (1984) Geographic distance, environmental difference, and divergence between isolated populations. Systematic Zoology 33:69-82.  [PDF]

[16] Dillon, R T. and J. D. Robinson (2009)  The snails the dinosaurs saw: Are the pleurocerid populations of the Older Appalachians a relict of the Paleozoic Era?  Journal of the North American Benthological Society 28: 1 - 11.  [PDF]  See:
  • The Snails The Dinosaurs Saw [16Mar09]
[17]  This is the “jetlagged wildebison” model I proposed last spring:
  • Mitochondrial Superheterogeneity: What it means [6Apr16]
 [18] Charles Darwin, Freshwater Malacologist [25Feb09]

Tuesday, April 4, 2017

SFS Raleigh 2017

This is just a quick note to alert all my colleagues in the Society for Freshwater Science that I am planning to attend this year’s meeting in Raleigh June 4 – 8.  Come visit me under the “Gastropoda” sign at the taxonomy fair, and bring all your vials of tiny beat-up Physa.  I’m also offering a first glimpse of our next big survey, The Freshwater Gastropods of The Ohio.  Evans, Pyron, Watters, Reeves, Kungblenu, Bailey, Whitman and I have logged over 4,500 records from an eight-state area, recording 64 species and subspecies.  Some other than Physa.  See you in Raleigh!

Tuesday, March 14, 2017

Fred Thompson, Elizabeth Mihalcik, and the Pleurocerids of Georgia

Science is a human enterprise, alas.  Last month [1] we reviewed the fitful progress of research into the systematic relationships of the Florida Pleuroceridae through the 1970s and 1980s, as a function of the personalities of two scientists, the late Dr. Fred G. Thompson and Dr. Steven M. Chambers.  In 1990 Steve Chambers published a malacological masterpiece, “The genus Elimia (= Goniobasis) in Florida and adjoining drainage basins [2],” correcting Clench & Turner’s 1956 seven-species model down to four.  He then shifted his professional interests out of the field.  Thompson, failing to cite the Chambers 1990 paper or indeed (almost) any of Chambers’ impressive body of research, published an update of his “Freshwater Snails of Florida” in 1999.  He was going in the opposite direction.

Thompson’s Second Edition [3] began with the Clench & Turner seven-species model but added an undescribed “spring Elimia” from Holmes Creek and three new species that “may occur in Florida” (Elimia taitiana, Choctawhatchee Elimia and Escambia Elimia), bringing his grand total up to 11 and foreshadowing things to come.  Simultaneously, Thompson was working with a committee of the AFS to publish a second edition of the Turgeon et al. “Common and Scientific Names of Mollusks,” which appeared in 1998 [4].  The Turgeon work was destined, in a perverse way, to become the Bible of American malacology.  Thus less than ten years after the publication of Steve Chambers’ landmark paper, the Clench & Turner model became gospel, and Chambers’ work forgotten.

Meanwhile, a new graduate student arrived at the University of Florida named Elizabeth L. Mihalcik.  She entered the laboratory of the noted herpetologist F. Wayne King, and was “introduced into the world of invertebrates through her mentor and friend, Dr. Fred G. Thompson.”  In 1996 she was awarded an EPA STAR graduate fellowship for her proposal entitled, “Analyzing morphological forms within southeastern river system species complexes.”
The Mihalcik [6] study area, showing P. floridensis
Elizabeth Mihalcik’s proposal originally focused on estimating gene frequencies at polymorphic enzyme-encoding loci in populations of what her mentor and friend referred to as “the Elimia curvicostata complex” of Central Georgia and the Florida drainages of Alabama.  And in fact, I did host Elizabeth at my lab in Charleston for several days in the late 1990s, and showed her everything I knew about allozyme electrophoresis.  But if she ever ran an allozyme gel independently, I never saw the data.

Elizabeth’s (1998) dissertation [5] focused primarily on shell morphometrics: 6 measures, 4 counts, spire angle and 8 ratios of the above in various combinations, blessedly naive of any inkling that any shell character might demonstrate any interpopulation variance for any reason whatsoever.  But the winds of change were blowing across systematic biology.  Her dissertation also included a little bit of mtDNA sequence data.

The first fully automated sequencing DNA machine had come onto the market in 1987, even as Kerry Mullis’ patent for PCR amplification was approved.  The Folmer et al primers to amplify the mitochondrial CO1 gene were published in 1994, and by the late-1990s what had been a trickle of DNA sequence data became a torrent.  Surely such data must be useful for systematic biology, if we could just settle on an algorithm to build our gene trees, yes?  The “phylogenetic species concept,” originally proposed by Cracraft in 1983, came into vogue.

So by the time Elizabeth published her dissertation with Fred Thompson in 2002 [6], her analysis was almost entirely based on mitochondrial CO1 sequence data.  She sequenced 25 individual snails from 24 Pleurocera populations scattered across Florida, Georgia and Alabama, with one individual P. catenaria from South Carolina thrown in to boot.  Her results (below) returned four clusters of roughly 5 - 7 individuals each, corresponding remarkably well to the four species that Steve Chambers documented in 1990.

Mihalcik and Thompson also discovered at least three apparent cases of mitochondrial superheterogeneity (mtSH above).  I have very nearly beat this subject to death on the FWGNA blog, but see note [7] below if you need your memory refreshed.  MtSH may indeed have some utility in identifying genuine biological species, but only if the phenomenon is recognized, which Mihalcik and Thompson did not [8].  So setting aside their sequence data for those three individual snails, the remaining snails demonstrate four clusters neatly corresponding to the species Chambers called floridensis, curvicostata, dickinsoni, and “boykiniana*.”

Alas, Mihalcik and Thompson saw substantially more than four species in their gene tree.  They interpreted their 25 mtDNA sequences as evidence of 15 species and subspecies of Pleurocera in Central Georgia and the Florida drainages of Alabama, as follows.  Starting with baseline curvicostata, a catenaria reference from South Carolina and two flava references from Alabama, M&T resurrected four old Isaac Lea names from synonymy: induta, mutabilis, viennaensis, and ucheensis.  They raised Goodrich’s subspecies timida to the full species level, and described two new subspecies underneath it: exul and nymphaea.  And they freshly described five new species: exusta, glarea, annae, buffyae, and darwini, the last three of which Elizabeth named for her dogs. 

When one steps back to consider that the authors did not even consider six of the seven Clench & Turner species as members of the “Elimia curvicostata complex” (albanyensis, athearni, clenchi, dickinsoni, floridensis and vanhyningiana), the Mihalcik and Thompson hypothesis constitutes a staggering multiplication of pleurocerid nomina [9] for the region.

I myself was not idly sitting by in Charleston, however.  Through most of the 1980s I confess that I became distracted by hard clam aquaculture, and by the 1990s Amy Wethington and I were beginning to work on Physa.  But I never forgot my first love, which was the pleurocerid snails.  And in 2002, just a few months prior to the work of Mihalcik & Thompson, an undergraduate student and I published “A survey of genetic variation at allozyme loci among Goniobasis populations inhabiting Atlantic drainages of the Carolinas” [10].

We estimated gene frequencies at 8 polymorphic loci in 12 populations, picking up in NW North Carolina where my dissertation left off and extending through South Carolina to touch eastern Georgia – 4 populations of P. proxima and 8 populations of P. catenaria, including the type locality 50 km W of Charleston.  Our most interesting result touched a very familiar theme on this blog [11], ecophenotypic plasticity in pleurocerid shell morphology.  Populations of P. catenaria bearing typical shells in the Piedmont of the Carolinas tended to lose their spiral cords and costae as they ranged into the coastal plain, independently developing the shell form associated with the subspecific nomen “dislocata[12].

In 2004 Thompson published a third edition of his “Freshwater Snails of Florida,” this update strictly online [13].  From seven species of pleurocerid snails in 1984 and eleven in 1999, Thompson’s fresh online edition now recognized 12 species in Florida alone, the “Choctawhatchee Elimia” formally identified as buffyae, the “Escambia Elimia” now annae, and a new “Rasp Elimia” waiting in the wings, not as yet described.  None of Steve Chambers’ papers whatsoever were cited in Thompson’s 2004 online version.  Even the passing mention of Chamber’s 1980 paper had now disappeared.

So ten years ago, this was the situation:  Two morphologically variable species, P. proxima and P. catenaria, were understood to range from southern Virginia through North Carolina and South Carolina to the Georgia border.  At the Georgia border P. catenaria disappeared, to be replaced by over 20 rare and endemic pleurocerid species, each occupying single subdrainages, or in some cases single springs.  Does that model seem biologically plausible to you?
P. ucheensis is dislocata, the remainder are P. catenaria
In 2011 John Robinson and I extended our allozyme survey across Georgia, linking at long last my intensive dissertation work in Virginia with my buddy Steve Chambers’ extensive dissertation work in Florida [14].  John and I sampled 14 populations, including populations identified by Mihalcik & Thompson as viennaensis, darwini, mutabilis, induta and timida nymphea, reaching down to the boykiniana and floridensis populations of Chambers.  We estimated genetic divergence at 11 polymorphic allozyme encoding loci, calibrating our observed values with three populations of P. proxima, the conspecific status of which is unchallenged.

We did not include any bona fide curvicostata or dickinsoni in our survey, because those two taxa seemed unambiguous to us.  But the bottom line was that all the Georgia populations we sampled were either floridensis (including induta and timida) or catenaria (including albanyensis, boykiniana, darwini, mutabiliis, and viennaensis).  There are exactly four species of pleurocerid snails in Central Georgia, Florida, and the Florida drainages of Alabama as hypothesized by Chambers: curvicostata, dickinsoni, floridensis and (*a minor correction) catenaria, all boasting extensive ranges.  And the most widespread of those species, Pleurocera catenaria (Say 1822) extends through five states, from Virginia through the Carolinas and Georgia to mimic P. floridensis at Ichetucknee Springs, bringing the discovery Steve Chambers first reported in 1978 [15] into a properly regional context.

I didn’t expect Dr. Thompson to embrace the four-species model in 2011 any more than he did in 1990.  His species concept was rooted in nineteenth-century typology, no different from such giants of American malacology as Henry Pilsbry and F. C. Baker.  Indeed, distinction of species by shell costations or spiral chords is not conceptually different from distinction by individual nucleotide substitutions, as do many of our colleagues in the present day.  But I confess, when I saw the species list published by P. D. Johnson and colleagues in 2013 [16], including as it did albanyensis, annae, athearni, boykiniana, buffyae, clenchi, darwini, exusta, glarea, inclinans, induta, mutabilis, taitiana, timida, ucheensis, vanhyningiana, and viennaensis, I was just a little bit disappointed.


[1] Fred Thompson, Steve Chambers, and the pleurocerids of Florida [15Feb17]

[2] Chambers, S. M. (1990) The genus Elimia (= Goniobasis) in Florida and adjoining drainage basins (Prosobranchia: Pleuroceridae)  Walkerana 4: 237 – 270.

[3] Thompson, F. G. (2000)  An identification manual for the freshwater snails of Florida.  Walkerana 10(23): 1 -96.

[4] Turgeon, D.D., J.F. Quinn, A.E. Bogan, E.V. Coan, F.G. Hochberg, W.G. Lyons, P.M. Mikkelson, R.J. Neves, C.F.E. Roper, G. Rosenberg, B. Roth, A. Scheltema, F.G. Thompson, M. Vecchione, and G.D. Williams (1998) Common and scientific names of aquatic invertebrates from the United States and Canada: Mollusks (second edition), American Fisheries Society Special Publication 26, Bethesda, Maryland, 526 pp.

[5] Mihalcik, E. L. (1998)  Elimia curvicostata species-complex within the river drainages of the southeastern United States: Morphology, DNA, and biogeography.  Ph.D. dissertation, University of Florida, 209 pp.

[6] Mihalcik, E. L. & F. G. Thompson (2002)  A taxonomic revision of the freshwater snails referred to as Elimia curvicostata, and related species.

[7] Pleurocerid populations (and a variety of other gastropod populations as well) often demonstrate “mitochondrial superheterogeneity,” where two or more members demonstrate 10% sequence divergence or greater for any single-copy mtDNA gene, not sex-linked.  For more see:
  • Mitochondrial superheterogeneity: What we know [15Mar16]
  • Mitochondrial superheterogeneity: What it means [6Apr16]
  • Mitochondrial superheterogeneity and speciation [3May16]
Mihalcik and Thompson only sampled single individuals from almost all [8] of their populations, so (strictly speaking) mtSH cannot be recognized.  But based on my personal experience with many datasets of this sort, the cases labelled flava-1, annae, and boykiniana all look like mtSH to me.

[8] The only population from which M&T sampled more than one individual was their reference “flava” from Alabama, the two individuals sequenced from which apparently demonstrating more than 20% sequence divergence.  They might have been the first researchers ever to discover mtSH in the Pleuroceridae, had their eyes been open.  Dillon & Frankis didn’t formally document the phenomenon until 2004 [PDF].

[9] Here’s something even more remarkable, if you stop to consider it.  The 15 populations M&T identified as being members of the “Elimia curvicostata complex” (explicitly excluding floridensis, dickinsoni, or boykiniana) apparently represented all four of the Chambers species (curvicostata, floridensis, dickinsoni, and boykiniana) about equally.   Shows you the value of shell morphology, doesn’t it? 

[10] Dillon, R. T. and A. J. Reed (2002)  A survey of genetic variation at allozyme loci among Goniobasis populations inhabiting Atlantic drainages of the Carolinas.  Malacologia 44: 23-31. [PDF]

[11]  Much more on cryptic phenotypic plasticity in pleurocerid shell morphology:
  • Goodbye Goniobasis, Farewell Elimia [23Mar11]
  • Pleurocera acuta is Pleurocera canaliculata [3June13]
  • Elimia livescens and Lithasia obovata are Pleurocera semicarinata [11July14]
[12] More about the pleurocerid subspecific nomen “dislocata” and its treatment by Turgeon [4] can be found at:
  • What subspecies are not [5Mar14]
[13] Thompson, F. G. (2004) on-line edition: An identification manual for the freshwater snails of Florida. Florida Museum of Natural History, Gainesville, FL [html].

[14] Dillon, R. T. and J. D. Robinson (2011)  The opposite of speciation: Population genetics of  Pleurocera (Gastropoda: Pleuroceridae) in central Georgia.  American Malacological Bulletin 29: 159-168.  [PDF]

[15] Chambers, S. M. (1978) An electrophoretically detected sibling species of “Goniobasis floridensis” (Mesogastropoda; Pleuroceridae).  Malacologia 17: 157 – 162.

[16] Johnson, P. D. et al. (2013) Conservation status of freshwater gastropods of Canada and the United States.  Fisheries 38: 247 – 282.  See:
  • Plagiarism, Paul Johnson, and the American Fisheries Society [9Sept13]

Wednesday, February 15, 2017

Fred Thompson, Steve Chambers, and the pleurocerids of Florida

Word has reached us of the death of Dr. Fred G. Thompson, who passed away December 27, 2016 at his home in Ocala [1].  Dr. Thompson was Curator of Malacology at the Florida Museum of Natural History in Gainesville for 40 years.  He was 82.

Dr. Thompson’s body of published work must run into the hundreds of titles – focusing primarily on the North American Hydrobiidae, but extending to include the Pleuroceridae and a broad range of terrestrial gastropod taxa as well, especially of Mexico and Latin America.  I understand that his complete necrology will soon be published in The Tentacle.

I had no personal relationship with the late Dr. Thompson.  I did attend his talks at the AMU (later the American Malacological Society) over a period of some thirty years, although I don’t recall his attending any of mine.  He did not respond to my letters or emails.  During my visit to the Florida Museum in July of 2006 he did not emerge from his office.

We did, of course, have many mutual colleagues.  The American community of freshwater gastropod workers is not a large one.  All I know about the personality and character of Dr. Thompson I have gathered from friends.  Good friends, like Steve Chambers.

Steve Chambers was an important influence on my young career.  He first met Dr Thompson on a field trip as a graduate student at the University of Florida in 1976, and became interested in the genetic relationships among Florida pleurocerid populations known at that time as Goniobasis, now Pleurocera.  Steve’s major adviser, Dr. Thomas Emmel, ran a big laboratory specializing in the promising new technique of allozyme electrophoresis.  Steve thought that the estimation of genetic divergence among populations of pleurocerid snails as a function of gene frequencies at multiple allozyme-encoding loci might elucidate evolutionary principles of great generality and importance [2].
The Clench & Turner model, as figured by Chambers [7]
The pleurocerid fauna of Florida is no less enigmatic than that of most other regions of the American southeast. Goodrich [3] recognized three species: catenaria (2 subspecies), clenchi and curvicostata, with boykiniana (3 subspecies) in nearby South Georgia.  Clench & Turner [4] took a big swing at this system, substituting floridensis for catenaria cancellata, raising catenaria vanhyningiana to the full species level, bringing one of the boykiniana subspecies down into Florida (albanyensis) and raising it to the full species level as well.  They also described two new species (athearni and dickinsoni) which, together with clenchi and curvicostata yielded seven species of Goniobasis in Florida.

It was the seven-species Clench & Turner model that Fred Thompson preferred for the Pleuroceridae of Florida, and the seven-species model is what Steve Chambers brought into his study design.  His (1977) dissertation [5] was an impressive effort to replicate the extremely influential Drosophila research that F. J. Ayala [6] had published in 1974.  Steve showed that, just as in Drosophila, populations of Goniobasis at increasing degrees of taxonomic divergence demonstrated increasing levels of genetic divergence at allozyme loci.  The paper based on his dissertation research was published in 1980 [7].

Perhaps unsurprisingly, however, Steve’s genetic data did not jive especially well with the Clench & Turner seven-species model.  Allozyme divergence suggested that vanhyningiana and clenchi be synonymized under floridensis and that albanyensis, athearni and the REF population (more about which anon) “are in the range of conspecifics,” although Chambers called for further study on that question.  Note that by the time he had published his dissertation in 1980 he had already synonymized clenchi (marked “F” in the figure above) under floridensis.

Meanwhile, on a small farm in Kansas, a boy was growing up [8].  Well, actually, he entered the graduate program at the University of Pennsylvania in the fall of 1977, on a scientific journey remarkably similar to that of Steve Chambers, just a couple steps behind.  My adviser was also running a big lab specializing in the promising new technique of allozyme electrophoresis, and I too was interested in the genetic relationships among populations of Goniobasis, mine in the southern Appalachians.  And it turned out that my adviser, Dr. George Davis, was also the Editor of the journal to which Steve Chambers sent his first two papers, the 1980 work based on his dissertation and an earlier one on that “REF” population I mentioned above.

Steve had discovered his REF population quite by accident.  He had gone to sample Ichetucknee Springs (the source of North Florida’s Ichetucknee River) expecting to find a typical population of Goniobasis floridensis, and that’s what he thought he had, until he got back to the laboratory.  But his gels showed two reproductively isolated populations, one of which matched floridensis genetically and the other of which was much more similar to his sample populations of Goniobasis athearni and G. albanyensis.  He published “An electrophoretically detected sibling species of Goniobasis floridensis” in Malacologia in 1978 [9].

Both Steve’s 1978 and 1980 papers were highly influential in my budding career.  I struck up a correspondence with him and he was happy to share techniques, recipes, and tips.  Imagine my surprise (and dismay) when my adviser showed me a manuscript he had received attacking Steve Chamber’s work gratuitously and viciously.

Fred Thompson had submitted a Letter to the Editor for publication in Malacologia.  The journal very rarely published letters in those days.  In fact, no letters whatsoever had been published in the previous ten years.  But Dr. Thompson was livid about research that had taken place at his own home institution, research that very carefully and thoroughly documented a model of evolutionary relationships among the Florida Goniobasis he did not share.

That letter was ultimately published in 1982, along with Steve’s reply [10], over the protestations of a farm boy from Kansas [8].  It included such whoppers as “Because the Ichetucknee population of athearni is distantly related genetically to floridensis, they cannot be sibling species.”  But the amount of genetic divergence has nothing to do with sibling species.  Ernst Mayr coined that term in 1963, and he defined it as “morphologically similar or identical populations which are reproductively isolated,” and genetic divergence does not enter into it [11].  In fact, whether the observed amount of genetic divergence might correlate with the taxonomic divergence suggested by systematic biologists was the very hypothesis that Steve was trying to test.

Well, by 1982 Steve had been gone from Gainesville for five years.  He told me later that he had been a finalist for the opening at the University of Michigan Museum of Zoology left vacant when Henry van der Schalie retired, but that Dr. Thompson had written an unsolicited letter to his colleagues in Ann Arbor, scuttling Steve’s chances for that job.

But Steve was able to land a job with the FWS Office of Endangered Species in Washington, which turned out to be a plum.  From Washington he was able to publish two high-visibility works on chromosomal evolution in gastropods [12].  He also became interested in the land snail fauna of the Galapagos, describing two new species of bulimulids in 1986 [13].  And with Christine Schonewald-Cox and other colleagues he edited a very influential book on the budding discipline of conservation genetics [14].

Meanwhile up north, the farm boy from Kansas [8] accepted an AAAS congressional fellowship and moved to Washington for the 1981-82 academic year, still right on the heels of his buddy Steve.  And we struck up a personal friendship that year which I wish had lasted longer.  Toward the end of my brief sojourn in Washington Steve remarked to me, and I’m paraphrasing here, “In the entire wide community of systematic biology, malacologists have the reputation of being the most petty and venal.”  He was broadening his professional horizons, trying to shift away.

And meanwhile down south, Dr. Thompson published the first (1984) edition of his “Freshwater Snails of Florida” [15].  Perhaps unsurprisingly he clung to the Clench & Turner seven-species model, citing the work of Chambers only once, superficially: “Recent studies on isoenzymes show that, in Elimia, shell characters are conservative indicators of genetic divergence (Chambers 1980, Dillon & Davis 1980).”  Regarding Dr. Thompson’s choice of genus, see note [16] below.
From Chambers [17]
After ten years in Washington, Steve was transferred to the regional office in Albuquerque, where he became involved with federal efforts to protect endangered elements of the charismatic megafauna such as the Red Wolf.  But he had one more snail paper in the pipeline, his 1990 masterpiece “The genus Elimia in Florida and adjoining drainage basins” [17].  Here Steve combined his modern understanding of allozyme, chromosomal, and morphological divergence in an enigmatic and misunderstood freshwater gastropod fauna together with a scholar’s appreciation for the old literature and the old museum collections into a symphony of Malacological virtuosity.  He convincingly demonstrated that there are four species of Goniobasis in Florida and South Georgiaboykiniana (including athearni, albanyensis, and REF), curvicostata, dickinsoni, and floridensis (including vanhyningiana and clenchi), and the matter would seem to be settled.

But way back on page 261 of Chambers’ 1990 work we read: 
“R. T. Dillon has sent me shells of Elimia catenaria Say from South Carolina and suggested that my E. boykiniana may be a synonym of E. catenaria.  Although there is considerable merit in this suggestion, I decline to combine these Georgia and South Carolina populations with E. boykiniana at this time because they occur in major drainages for which genetic data are not available.”
Yes, after single years in Washington and at Rutgers, that farm boy from Kansas [8] had arrived in Charleston, SC, and begun work to tie his 1982 dissertation research in VA/NC together with the remarkable body of knowledge Steve Chambers had developed in FL/Ga.  Could direct conflict with Dr. Fred Thompson be avoided?  Tune in next time.


[1] Fred Gilbert Thompson (November 13, 1934 – December 27, 2016).  The Shell-O-Gram 58(1): 4-5.

[2] I did too, once.  When I was young.

[3] Goodrich, C. (1942) The Pleuroceridae of the Atlantic Coastal Plain.  Occas. Papers Mus. Zool. Univ. Mich. 456:1 – 6.

[4] Clench, W. J. & R. D. Turner (1956) Freshwater mollusks of Alabama, Georgia and Florida from the Escambia to the Suwannee River.  Bull. Florida State Museum 1:1 – 239.

[5] Chambers, S. M. (1977) Genetic divergence during speciation in freshwater snails of the genus Goniobasis.  Ph.D. dissertation, University of Florida, Gainesville. 59 pp.

[6] Ayala, F. J., M. L. Tracey, D. Hedgecock & R. C. Richmond (1974) Genetic differentiation during the speciation process in Drosophila.  Evolution 28: 576-592.

[7] Chambers, S. M. (1980) Genetic divergence between populations of Goniobasis (Pleuroceridae) occupying different drainage systems.  Malacologia 20: 63 – 81.

[8] Virginia, actually.  And he wasn’t born on a farm, and he never grew up, if you ask his wife.  For more, see…
  • The Clean Water Act at 40 [7Jan13]
[9] Chambers, S. M. (1978) An electrophoretically detected sibling species of “Goniobasis floridensis” (Mesogastropoda; Pleuroceridae).  Malacologia 17: 157 – 162.

[10] Thompson, F. G. (1982) On sibling species and genetic diversity in Florida Goniobasis.  Malacologia 23: 81 – 82.  
       Chambers, S. M. (1982) Sibling species and genetic diversity in Florida Goniobasis: A reply.  Malacologia 23: 83 – 86.

[11] Mayr, E. (1963) Animal Species and Evolution.  Belknap Press, 797 pp.  The definition of sibling species is found on page 34. 

[12] Chambers, S. M. (1982) Chromosomal evidence for parallel evolution of shell sculpture pattern in Goniobasis.  Evolution 36: 113 – 120.  
        Chambers, S. M. (1987) Rates of evolutionary change in chromosome numbers in snails and vertebrates.  Evolution 41: 166 – 175.

[13] Chambers, S. M. (1986) Two new bulimulid land snail species from Isla Santa Cruz, Galapagos Islands.  Veliger 28: 287 – 293. 

[14]   C. Schonewald-Cox, S. Chambers, B. MacBryde, and L. Thomas (1983) Genetics and Conservation: A Reference for Managing Wild Animal and Plant Populations.  Benjamin Cummings, Menlo Park.  722 pp.

[15] Thompson, F.G., 1984. The freshwater snails of Florida: A manual for identification. University of Florida Press, Gainesville, FL. 1-94.

[16] I suppose I should also mention that, separately but essentially simultaneously, J. B. Burch was working on his North American Freshwater Snails.  The Burch & Tottenham “Species List, Ranges and Illustrations” was published in 1980, with the full EPA Manual following in 1982 and the stand-alone separate work republished in 1989.  This work proposed a hybrid between the Goodrich and Clench & Turner classifications of the Florida and South Georgia pleurocerid fauna, which did not evolve through its extended publication history: athearni, clenchi, curvicostata, dickinsoni, induta, and boykiniana with three subspecies.  The recognition of a separate floridensis, while keeping vanhyningiana as a subspecies of catenaria, yielded a list of ten species and subspecies for the region, which Burch renamed to “Elimia.”

[17] Chambers, S. M. (1990) The genus Elimia (= Goniobasis) in Florida and adjoining drainage basins (Prosobranchia: Pleuroceridae)  Walkerana 4: 237 – 270.