George Morgan Davis (1938 – 2024): The Director’s Cut

Dillon, R.T., Jr., G. Rosenberg & P.M. Mikkelsen (2025) George Morgan Davis (1938 - 2024): Life, work, and legacy.  Malacologia 68: 3 - 26. [pdf]

The email was delivered to my inbox in the mid-afternoon Thursday, June 20, 2024, and I don’t remember being terribly surprised by the news.  Dr. George M. Davis, my major advisor at Penn 1977 – 1982 was dead.  His wife, my old friend Elaine Hoagland, was asking me to pass the word along to the wider malacological community.

King George was the last monarch of American Malacology, guiding our discipline from the classical typology of the 19th century to the neoclassical typology of the present day.  I posted single-paragraph tributes on Facebook and the Mollusca list server early that next morning, and was gratified to see how rapidly the word spread.

George M. Davis (1938 - 2024)

Gary Rosenberg, George’s successor at the Academy of Natural Sciences in Philadelphia, stepped forward to offer a memorial at the annual meeting of the American Malacological Society, upcoming just six weeks away in Pasadena, and President Pat Krug found room on the program.  Gary also began to compile a bibliography of George’s published works at that time.

Elaine also mentioned to me, in the days immediately following her husband’s death, that she had separately contacted our colleague Chuck Lydeard, George’s immediate successor as Editor-In-Chief of Malacologia, about a special volume of to be published in his memory as well.  I suppose I had assumed that Gary would take the lead on George’s professional obituary for that volume, just as he had taken the lead for the oral tribute at the AMS. 

So, the email from Chuck that arrived in my inbox 15Aug25 surprised me quite a lot.  Chuck confirmed that an entire issue of Malacologia, dedicated to the memory of my influential mentor, was indeed nearing completion.  Paula Mikkelsen and Alan Kabat had joined Gary Rosenberg to compile an extensive GMD bibliography, now grown to 200 titles, together with lists of all the species and higher taxa he had described, all the species named for him, and all the articles written about him, for publication as the lead article in that memorial issue.  But none of our distinguished colleagues apparently felt called to write a biography of the man to explain all the fuss. Nor could anybody else alive on God’s green earth be found to volunteer.

 

And so at this late date, 14 months into a 15-month project, with every other option exhausted and time running out, pushed beyond the point of despair, Chuck Lydeard was contacting me to ask if I would take the lead authorship of the lead article in a George M. Davis memorial issue of Malacologia.  My coauthors Rosenberg and Mikkleson would contribute the bibliography half, and I the biography half.  My deadline would be the end of September, six weeks away.

 

And I felt duty-bound to accept.  I didn’t like George Davis.  I don’t know anybody who did.  And although I did work closely with him during that brief window 1977 – 1982, and got to know him better than I cared to during those five years, after I left Philadelphia we really did not keep in touch.  Our areas of research interest just barely intersected, and our philosophies of science not at all.

 

But George Davis helped me.  He administered the best facilities for malacological research in the world at that time, and he made the superb ANSP collections, the wonderful library, the modern laboratory, and rooms full of equipment and cabinets bulging with supplies all available to me, free and without obligation, and otherwise left me alone, and that is exactly what I needed.

 

So, I went to work on a tribute for my major advisor.   And I confess I did not quite make my deadline, but by the first week of October had a first draft ready to share with my coauthors.  Divided into six sections, that first draft reviewed George’s 60-year career, contextualized his manifold scientific contributions, and transmitted some feeling for the outsized influence he wielded in American Malacology during the latter half of the 20th Century, an influence that continues to the present day.  And in an effort to humanize the man, each of those six sections was introduced with a story, told in the first person, in which I reminisced about my own personal experiences with my major advisor in the late 1970s.

 

And I added a top layer as well.  Looking back on 20th Century Malacology from a 25-year perspective, I found myself able to trace the evolution of our entire discipline in the career of Dr. George M. Davis.  Working on his dissertation at the University of Michigan Museum of Zoology 1960 - 1965, Davis was born into classical, 19th-century typology.  But the Modern Synthesis of Darwin + Mendel was ascending, hypothesis-driven, grounded in the theory of Fisher, Haldane, and Wright, rigorously scientific.  And just dawning at that very moment was a new age of molecules and computer analysis, ultimately to devolve into a 21st-century neoclassicism just as typological as the 19th.

 

Davis did, in fact, include a set of rigorously-designed breeding experiments in his dissertation research, published in 1967, as well as classical morphological studies and remarkably forward-looking protein electrophoretic analysis.  Yet even though his breeding studies answered the question his dissertation addressed, and neither the morphology nor the molecules contributed, Davis never tested another hypothesis throughout the remainder of his 60-year career.  Quoting myself directly: 

“George Davis had been baptized Presbyterian-form in modernity with his early laboratory experiments on hybridization, sprinkled not dunked.  Then finding objective, hypothesis-driven science too constraining, he jumped over the deeper waters of Fisher, Mayr, Dobzhansky and Simpson to the Neoclassical now, drawing our discipline along with him.”

 Well, such vivid language is never going to find publication in a scientific journal, and well I knew it, even as the Christian denominations were flying off my fingertips.  That first draft of early October was edited significantly by my coauthors, as is their right.  Paula, in particular, objected to my use of the first person in the reminiscences, which I certainly understand.  But conversion to the third person and the passive voice, much color is sacrificed.  And awkwardness left in the wake.

 

And Paula, Gary, and I kept Chuck Lydeard on the cc line as we passed the drafts around.  Rather early in the process Chuck alluded to sending our manuscript out for peer review, which seemed a wry comment on the vitality of our discipline.  “I’m not saying our science is dead, but in Malacology, obituaries are peer-reviewed.”  I needn’t have worried about that step, however, as Alan Kabat ultimately contributed a couple excellent insights which I was pleased to incorporate.

 

So, the George M. Davis Memorial issue of Malacologia was published in December, and the lead article by Dillon, Rosenberg and Mikkelsen is downloadable from this [pdf] link.  Although the paper as ultimately published does convey the overall flavor of the Scotch pie I baked in September, much of the spice has been lost.

 

Then for those of my readership who prefer their monthly dose of Rob Dillon redolent of creekbank, ancient tome, and spoiled ethanol, this month I have made FWGNA Circular #9 available for download from the website, here:

 

George Morgan Davis (1938-2024): The Director’s Cut

 

This version is very nearly that first early-October 2025 draft of my manuscript, unexpurgated, with a sprinkling of subsequent improvements.  It is missing the section on George Davis’ personal life, which was subsequently added by Gary Rosenberg, as well as the complete GMD bibliography and supporting sections as contributed by Gary and Paula working together. It does conclude with a literature-cited section, to the references cited in text only.  Also included are several nice figures not appearing in the published journal article, and a couple really good stories.  Bon appetite.

My buddy, Bob

Editor’s Note – This essay was subsequently published as: Dillon, R.T., Jr. (2023b)  My Buddy, Bob.  Pp 203 – 210 in The Freshwater Gastropods of North America Volume 6, Yankees at The Gap, and Other Essays.  FWGNA Project, Charleston, SC.

Although we studied at different institutions, Bob Hershler was my best friend in graduate school.  He was enrolled at Johns Hopkins way down in Baltimore, and I at the University of Pennsylvania in Philadelphia.  But we shared George Davis as our major advisor, so for several years we both spent most of our hours many of our days in the Malacology Department at the Academy of Natural Sciences.

In many ways, Bob was closer to George than I was, ultimately completing his dissertation on a morphological study of an endemic radiation of hydrobioids in northern Mexico [1], really rather similar to George’s masterpiece in southeast Asia [2].  Meanwhile, I was in the lab running allozyme gels on the opposite of an endemic radiation in the American pleurocerids [3].

Rob & Bob in Philadelphia, 1977

Bob often slept on the carpet under the big table in the malacology library.  And there was a night watchman, stationed at the desk at a side door downstairs, who patrolled regular rounds inside the building.   The watchman knew Bob was sleeping under there, which he most certainly was not supposed to be doing, but nevertheless turned a blind eye.

One evening Bob was running a set of exploratory electrophoresis gels, and after he got his samples on, decided to go out for supper.  And he walked by the watchman at the side door and out onto 19th Street and was gone for perhaps an hour or so.  And when he returned to the 19th Street door, the watchman was on his rounds.  And the door was locked.

And Bob panicked.  Fearing that his gels would cook and might ultimately (I suppose?) start a fire, he pulled the fire alarm on the side of the building.  I will leave to your imagination what response might follow a fire alarm at a 150-year-old building crammed with rotten alcohol and animal skins in downtown Philadelphia in the middle of the night.

Bob was awarded his Ph.D. in 1983, the same year that I was awarded mine, and by the blessing of divine providence, was able to win a curatorial position at the U.S. National Museum a couple years later.  Such plums are few and far between for scientists with our very specialized (and quite possibly obsolete) backgrounds.  My self-effacing buddy Bob confided to me shortly thereafter, “It was a weak field” [4].

But at the USNM Smithsonian, Bob’s career flourished.  His research focused almost entirely on the hydrobioids, of which he became the unchallenged North American authority, publishing over 100 peer-reviewed papers, many monographic in their size and scope.  His malacology was neoclassical, featuring lovely, detailed drawings of the dissected animals themselves, in the style of a Henry Pilsbry or our shared mentor George.  But he often whistled modern notes.  At his best, Bob was able to recognize intrapopulation and interpopulation variation along with the most evolutionary of his contemporaries.  His unification of Pyrgulopsis robusta, for example, will have a lasting impact [6].

The 1990 monograph of the phreatic genus Fontigens he published with John Holsinger and Leslie Hubricht [7] was a thing of beauty.  It opens with a marvelous and detailed examination of morphological variation in the most common species, Fontigens nickliniana, depicting shells of all shapes and penial complexes of in all their elaborate forms and twisted wonder.  His section subtitled “material examined” lists hundreds of distinct springs and spring runs in ten different states where populations of F. nickliniana can be found, with locality data of sufficient quality so that any subsequent worker can go and look at the things himself, if he has any questions.  That’s the best part of the monograph.

F. nickliniana penis variation [8]

Then moving forward from a rock-solid understanding of morphological variation in Fontigens nickliniana, Bob proceeded by rigorous methodology to distinguish nine additional species in the genus.  Four of these are more or less sympatric with nickliniana in the Valley-and-Ridge Province of Virginia, most strikingly F. orolibas, which even co-occurs mixed with F. nickliniana, lending credence to the hypothesis of reproductive isolation.  Again, Bob offers lovely figures of shell, radula, and reproductive anatomy for each, with excellent locality data.  And he iced the cake with a dichotomous key to cleanly distinguish among the ten total.  This work is easily on par with that of a Hubendick or a Meier-Brook [9].  Bob’s 1990 Fontigens monograph is as good as classical malacology ever got or can get.

Almost as good, equally important, and even larger was his 1994 monograph on the North American Pyrgulopsis [10].  I keep a copy by my desk and refer to it often.  In this 115-page tour-de-force Bob recognized 11 Pyrgulopsis species from Eastern North America as well as the 54 that (by that early date) had been described from the American West.  Of the 54, Hershler had himself described 21.  We will have much more to say about both the Eastern subset (now referred to Marstonia) and the western subset (still Pyrgulopsis) in coming months.

I think it is fair to say, by a margin of approximately 54 to 11, that Bob’s greatest love was his first love – the hydrobioid fauna of the American West.  Running my finger through the “Annotated Checklist of Freshwater Truncatelloidean Gastropods of the Western United States” he published with Hsiu-Ping Liu in 2017 [11], I count 43 papers with Hershler as the senior author, including such major contributions as the (63 pg) review of the Arizona hydrobiids he published in 1988, his (140 pg) Cochliopine monograph of 1992, his second (132 pg) Pyrgulopsis monograph of 1998, his (41 pg) Fluminicola monograph of 1996, and his (53 pg) Tryonia monograph of 2001 [12].

From his hypotheses regarding evolutionary relationships among western hydrobioid populations he developed a secondary interest in biogeography, publishing reconstructions of ancient drainage systems [13].  He had little interest in population biology, ecology, or the broader malacofauna beyond the hydrobioids, however.  When I approached him about collaborating on the FWGNA project back in 1998, he replied, “I don’t do checklists” [14].

It should not surprise you, after having read the anecdote with which this essay opened, that Bob did not run any sort of genetic laboratory.  But in 1999 he developed a tremendously productive relationship with someone who did, Dr. Hsiu-Ping Liu, and for 20 years they made beautiful malacological music together.  Of the 43 papers on the western hydrobioids I counted above, 30 were published after 1998, and of those 30, Hsiu-Ping is also listed as an author on 26.  In addition, the Literature Cited section of the Hershler & Liu catalog also includes 10 papers with Hsiu-Ping as lead author, Bob’s name following.

Pyrgulopsis penis from Hershler [15]

Alas, the rising tide of mtDNA sequence data that began to slosh across Bob’s desk in the 2000s pushed his species concept in a more typological direction, as became common in the generation that followed us.  How many species did he ultimately describe?  I do not know.  Bob and Hsiu-Ping’s 2017 catalog listed 126 species of Pyrgulopsis inhabiting the waters of the Western United States, 107 of which (I count) Bob Hershler was the author.  To put that figure in perspective, the total number of valid, biological species of freshwater gastropods inhabiting all U.S. Atlantic drainages, plus all the drainages of The Ohio, including the Tennessee/Cumberland, summing all families, including all pulmonate species and all pleurocerid species, as well as all the hydrobioids, across all or part of 17 eastern U.S. states, amounts to exactly 107.

For many years, I took it for granted that Bob would be behind his desk at the USNM, replying to my occasional email requests for consultation, and to the emails (many greater in their number) I forwarded to him from others.  And then, one day, he was gone.

Bob retired in 2018, the same year I myself would have retired from the College of Charleston, had all gone according to best-laid plans [16].  I tried emailing, and I tried calling, and nothing.  Finally, I called the front office at the USNM Department of Invertebrate Zoology where I learned the news.  He hadn’t left any forwarding info.

So now, perhaps you, my readership, can dimly see why I led this blog post with such a potentially-embarrassing anecdote about such a shy colleague, who is famous in the small world we share for many things, but not for his sense of humor.  Where are you, Bob?  Do you want to set me straight on any of the slander I have broadcast above?  I have a surprise for you, Old Buddy.

Notes

[1] Hershler, R. (1985) Systematic revision of the Hydrobiidae (Gastropoda: Rissoacea) of the Cuatro Cienegas Basin, Coahuila, Mexico.  Malacologia 26: 31 – 123.

[2] Davis, G. M. (1979)  The origin and evolution of the Gastropod family Pomatiopsidae, with emphasis on the Mekong River Triculinae.  Monograph of the Academy of Natural Sciences of Philadelphia 20: 1 – 120.

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

[4] I didn’t apply for the 1986 Smithsonian opening.  I had accepted an assistant professorship at The College of Charleston in 1983, which was a mighty quick turnaround, but that wouldn’t have stopped me.  Curatorships at the USNM are very, very sweet plums [5].  What did stop me from applying for the position at the Smithsonian that Bob Hershler ultimately won in 1986 might be grist for a future post on the FWGNA blog.

[5] I happened to notice on a website called federalpay.org that at his retirement, Bob was earning a base salary of $161,900.  That is way more than twice what I ever made, in 33 years of labor at a mid-sized college of regional reputation, grading thousands of lab reports written by entitled 19-year-old sorority girls.

[6] Hershler, R. & H-P. Liu (2004) Taxonomic reappraisal of species assigned to the North American freshwater gastropod subgenus Natricola (Rissooidea: Hydrobidae). The Veliger 47: 66-81.  Hershler, R. & H-P. Liu (2000) A molecular phylogeny of aquatic gastropods provides a new perspective on biogeographic history of the Snake River region. Molec. Phyl. Evol. 32: 927-937.  For a review of the tempest these two papers stirred up in the teacup of freshwater gastropod conservation, see:

• Idaho springsnail showdown [28Apr05]
• Idaho springsnail panel report [23Dec05]
• When pigs fly in Idaho [30Jan06]

[7] Hershler, R., J.R. Holsinger & L. Hubricht (1990) A revision of the North American freshwater snail genus Fontigens (Prosobranchia: Hydrobiidae). Smithsonian Contributions to Zoology 509: 1-49.  For an appreciation, see:

• Springsnails of The Blue Ridge [26July06]

[8] This is a scan of Figure 7 from Hershler, Holsinger & Hubricht [7].  It shows camera lucida outline drawings of penes from ten different populations of F. nickliniana, collected mostly from Virginia.  Pl = proximal penial lobe, Dl = distal penial lobe, Pf = Penial filament.

[9] For appreciations of the work of those two icons of neoclassical malacology, see:

• The classification of the Lymnaeidae [28Dec06]
• Character phase disequilibrium in the Gyraulus of Europe [4Feb22]

[10]  Hershler, R. (1994)  A review of the North American freshwater snail genus Pyrgulopsis (Hydrobiidae).  Smithsonian Contributions to Zoology 554: 1-115.  I mentioned idly  “leafing through” this important work back in 2016, for example:

• Marstonia letsoni, quite literally obscure [5Feb16]

[11] Hershler & Liu (2017) Annotated Checklist of Freshwater Truncatelloidean Gastropods of the Western United States, with an Illustrated Key to the Genera.  US Bureau of Land Management Technical Note 449: 1 – 142.

[12] Hershler, R. 1998. A systematic review of the hydrobiid snails (Gastropoda: Rissooidea) of the Great Basin, western United States. Part I. Genus Pyrgulopsis. Veliger 41:1-132.  Hershler, R. 2001. Systematics of the North and Central American aquatic snail genus Tryonia (Rissooidea: Hydrobiidae). Smithsonian Contributions to Zoology 612:1-53.  Hershler, R., Frest, T.J. 1996. A review of the North American freshwater snail genus Fluminicola (Hydrobiidae). Smithsonian Contributions to Zoology 583:1-41.  Hershler, R., Landye, J.J. 1988. Arizona Hydrobiidae. Smithsonian Contributions to Zoology 459:1-63.  Hershler, R., Thompson, F.G. 1992. A review of the aquatic gastropod subfamily Cochliopinae (Prosobranchia: Hydrobiidae). Malacological Review Supplement 5:1-140.

[13] Hershler, R., D.B. Madsen, and D.R. Currey (eds) Great Basin Aquatic Systems History. Smithsonian Contributions to the Earth Sciences 33: 1 – 405 (2002).

[14] Ironic in light of the title of his 2017 publication, cited at note [11] above.

[15] This is a detail from Figure 2 of Hershler [10].  It depicts three views of a whole mount penis dissected from Pyrgulopsis californiensis.  Tg = terminal gland, Pg = penial gland, Vd = ventral gland, Dg = dorsal glands. 

[16]  I was banned from campus and forced into retirement for a Woodrow Wilson quote four weeks into the spring semester of 2016.  A press release issued by the Provost’s Office at the College of Charleston on the afternoon of February 21 stated, in part, “We have endured that sanctimonious asshole for 33 years, 5 months, 21 days, 13 hours and 15 minutes, and cannot stand him for one second more.”  For a review, see:

• Inside Higher Education [8Aug16]

I used my settlement from the lawsuit to set up the FWGNA as a sole proprietorship.

Bill and Ruth and Jack and Virginia, and Campeloma

Editor’s Note – This essay was subsequently published as: Dillon, R.T., Jr. (2023c)  Bill and Ruth and Jack and Virginia, and Campeloma.  Pp 85 – 95 in The Freshwater Gastropods of North America Volume 7, Collected in Turn One, and Other Essays.  FWGNA Project, Charleston, SC.

Bill Clench was already well into the mascot phase of his career when I first met him at the 1976 AMU meeting in Columbus, Ohio.  Colleagues, students, and friends ushered him front-row-center for the annual society photo, Joe Morrison [1] and Leslie Hubricht [2] trailing in his wake.  I found Dr. Clench to be a warm and outgoing gentleman, still alert at age 78.  Please Lord, take me home before anybody calls me “alert.”

William J. Clench was born in New York in 1897 and grew up in the Boston area, collecting bugs, snails and shells around the Fenway, the Blue Hills and the local beaches [3].  Charles W. Johnson, the noted marine malacologist at the Boston Society of Natural History, was an early influence.  Clench graduated from Michigan State University in 1921, earned his MS at Harvard in 1923, then moved on to the University of Michigan to work on his doctorate [4], where Bryant Walker, quoting Tucker Abbott’s remembrance [5], “lit the malacological fires within Bill and was largely responsible for his first love, the freshwater mollusks.”

American Malacological Union 1976 [6]

From Michigan Clench accepted the mollusk curatorship at Harvard’s Museum of Comparative Zoology, where he served for 40 years, 1926 – 1966, mentoring many students who would become quite influential themselves.  Clench’s most famous student was R. Tucker Abbott, who succeeded Henry Pilsbry as curator at the ANSP and editor of The Nautilus, but we should not fail to mention the unionid guys Dick Johnson and Sam Fuller, or Arthur Clarke, whose landmark work on the Canadian freshwater molluscan fauna [7] sits handy by my desk, here 40 years after its publication.

Clench’s bibliography lists 420 scientific papers, covering the breadth of malacology: marine, terrestrial, freshwater and fossil, focused on North America but ultimately worldwide [8].  Most of his better papers were coauthored by Ruth Turner, another former student, with whom Clench’s life was “entwined,” to borrow Dick Johnson’s carefully-chosen verb.  We have previously featured on this blog the 1956 Clench and Turner monograph on the freshwater mollusks of Florida/Georgia Gulf drainages [9], which was an important contribution.

Clench’s malacology was early-modern, rooted in the old typology but with a growing appreciation of genetic variation within and among populations.  Looking down from my 2021 freshwater-gastropod-centric perspective, his greatest contribution was his two-part series on the North American Viviparidae, published in 1962 [10] and (with Sam Fuller) in 1965 [11].

The taxonomic history of the North American Viviparidae is identical to the taxonomic history of the North American Pleuroceridae, minus one order of magnitude coming into the 20^th century, and two going out [12].  Digging through the musty tomes on the shelves and the dusty shells in the cabinets of the MCZ in the early 1960s, Clench was able to uncover 49 Latin nomina assigned to the genus Campeloma, Isaac Lea [14] tying C.S. Rafinesque for the lead with six each.  Of those 49 nomina, 35 he discarded for cause or synonymized, little rationale given or expected, at the close of the era when such good works were still possible.  Clench did not preface his work with an exhaustive study of shell morphological variation, as did my hero Calvin Goodrich for the North American pleurocerids in the 1940s [15].  But I don’t think he missed any viviparid nomina either, as Goodrich simply skipped hundreds of pleurocerids.  I think Clench got them all.  Thank you, Bill.

Alas, Clench did not explain why he spared the 14 specific Campeloma nomina that survived his 1962 monograph, any more than he explained why he cut the other 35.  That burden was shouldered 20 years later by Dr. John B. Burch [16], with an obscure contribution from Dr. Virginia A. Vail.

From the Burch/Vail key [16]

The “Family Viviparidae” header in Burch’s dichotomous key, way back on page 227, carries an asterisk.  And at the bottom of page 227 is printed, “*From Burch & Vail (1982).”  But no work by Burch & Vail is listed among the references, nor was one ever published subsequently, to my knowledge [17].

Burch’s bibliography does, however, list six papers published by Virginia Vail at that point in her career, all solo, and they are good ones.  She was an excellent scientist, about whom I have been able to discover little.  She was born in Schenectady, NY, in 1945, earned her B.A. at Hartwick College (NY) and her M.S. and Ph.D. at Florida State University, graduating in 1975 [18].  From thence Vail went directly to the Tall Timbers Research Station north of Tallahassee, where she spent the rest of her career.

In 1977 and 1978 Virginia Vail published a two-part series comparing the reproductive anatomy and life history of Campeloma, Lioplax, and Viviparus in Florida.  Her first paper [19] was anatomical, featuring very nice drawings of male and female reproductive systems for all three taxa, and her second paper [20] ecological, detailing seasonal reproductive cycles.  The viviparids are quite conservative anatomically; Virginia was able to document only negligible difference in the plumbing of the three genera [21].  But here 40 years later, we still await a finer contribution to the comparative biology of the North American Viviparidae.

Virginia Vail identified the Campeloma population she selected for her study as C. geniculum (Conrad).  Interestingly, that particular population, inhabiting the Chipola River about 60 miles NW of Tallahassee, seems to have been entirely sexual, males and females (apparently) in roughly equal proportion.  She made only passing reference to asexual reproduction in her 1977-78 papers, noting that Mattox [23] had documented parthenogenesis in Campeloma rufum [24] as early as 1937.

Vail [19] figs. 5 & 10 [25]

The next year, Vail described Campeloma parthenum from Lake Talquin, an impoundment of the Ochlockonee River west of Tallahassee.  She distinguished that population both by its apparent absence of males and by the contour of the outer lip of the shell [26].  But she seems to have been struggling with species concepts, even as she was describing new ones.  Here is the title and abstract of the talk she gave at the August 1979 meeting of the American Malacological Union in Corpus Christie, TX:

“CHAOS IN THE GENUS CAMPELOMA (GASTROPODA: VIVIPARIDAE)

A poor understanding of environmentally induced shell variation, anatomical characteristics and the animal’s biology makes species identification difficult.  The occurrence of both dioecious and parthenogenetic populations (races? species?) and their peculiar geographic distributions further complicate the problem.  Observations on southeastern populations are offered to illustrate the problem and suggest solutions.”

I could not have said that better myself.  Fascinatingly, this was neither the title nor the abstract ultimately published in the Bulletin of the American Malacological Union for 1979, page 67.  The version that saw print was much more tamely entitled, “The Species Problem in Campeloma,” and featured a relatively measured critique of reliance on shell character, noting “the fact that reproduction can occur either parthenogenetically or sexually.”  As of the publication of her 1979 abstract, Virginia Vail was only counting two Campeloma species in Florida and Georgia combined, C. geniculum and “C. limum (includes C. floridense).”

I seem to remember [27], here 40 years later, that the solution Virginia Vail suggested on that August morning at La Quinta Royale Hotel in Corpus Christie, TX, recognized just those two species, a heavily-shelled C. geniculum (sexual) and more lightly-shelled C. limum (parthenogenetic).  That was certainly the direction Fred Thompson was tending by the 1990s with his “Identification Manual for The Freshwater Snails of Florida [30].”  Thompson listed four Campeloma species for The Sunshine State (geniculum, limum, floridense and parthenum), but observed, “in view of the inconsistency of shell characters, these last three forms may represent only a single species, Campeloma limum.”

American Malacological Union 1979

But returning to the thread of our story.  It was sometime during the late 1970s that Jack Burch signed a contract with the EPA to deliver his illustrated key to the North American Freshwater Snails [16].  And somehow [31] he linked up with Virginia Vail, during the full flower of her career.

The Burch/Vail key to the North American Viviparidae that ultimately saw publication in 1982 proceeds unremarkably through its first ten couplets, guiding us to the genus Campeloma on page 228, where we are referred to supplemental note (4).  That endnote – on page 268 now – begins with a brief review of Clench’s signal (1962) contributions to our understanding of the genus Campeloma [10].  Then four more nomina are subtracted from Clench’s list of 14 species on the authority of Arthur Clarke [32]: leptum Mattox 1940, tannum Mattox 1940, integra (Say 1821) and milesi (Lea 1863).  That brought our continental fauna down to 10.

Returning to the main key, on page 229, we find an earnest effort to distinguish, by shell morphology alone, eight species of Campeloma.  Three of the ten species surviving Burch’s endnote (4) did not survive the perilous transfer forward from page 268 to page 229.  The specific nomina brevispirum (Baker 1928), exilis (Anthony 1860), and gibba (Currier 1867) seem to have vanished [33].  But one brand new species of Campeloma was added, Vail’s [26] parthenum, bringing our total continental Campeloma fauna to N = 8 canonical species, as of 1982.  In the order of their description:

• Limnaea decisa Say 1817.  Clench speculated “Delaware River?”
• Campeloma crassula Rafinesque 1819.  The Ohio.
• Paludina genicula Conrad 1834.  Flint River, GA.
• Paludina regularis Lea 1841. Coosa R, AL.
• Paludina lima Anthony 1860. South Carolina.
• Melantho decampi Binney 1865. Decatur, AL. [34]
• Campeloma floridense Call 1886.  Wekiva River, FL.
• Campeloma parthenum Vail 1979.  Lake Talquin, FL.

The Burch/Vail key to the Campeloma begins with aperture color (white vs brown), then moves on to shell shoulders (angled vs rounded) then moves on to shell profile (broadly ovate vs narrowly ovate), and so forth.  It is a valiant effort, and I do not mean to diminish the contribution of its authors.  Just the opposite.

Science is the construction of testable hypotheses about the natural world.  It is not about being right, it is about being testable.  The Burch/Vail dichotomous key to distinguish the eight canonical species of North American Campeloma is science.

Next month, we test it.

Notes

[1] For my remembrance of J.P.E. Morrison, see:

• Joe Morrison and the Great Pleurocera Controversy [10Nov10]

[2] For a bit more about Leslie Hubricht, see:

• The Most Cryptic Freshwater Gastropod in the World [6Aug17]

[3] Most of the biographical details relayed above were gleaned from: Turner, R. D. (1985)  William J. Clench October 24, 1897 – February 22, 1984.  Malacological Review 18: 123-124.

[4] Surprisingly, Clench did not finish.  He was ultimately awarded honorary doctorates from both Michigan and MSU in 1953.

[5] Abbott RT (1984). "A Farewell to Bill Clench". The Nautilus 98 (2): 55–58.

[6] This is a detail from a scan of the original 8x10 glossy in my files.  The back is stamped, “Dept. of Photography & Cinema, The Ohio State University,  No. 191231-1, Please Give Credit”  Done.

[7] Clarke, A.H. (1981) The Freshwater Mollusks of Canada. Ottawa: The National Museums of Canada.

[8] Johnson, R.I. (2003)  Molluscan taxa and bibliographies of William James Clench and Ruth Dixon Turner.  Bulletin of the Museum of Comparative Zoology at Harvard College 158: 1- 46.

[9] Clench, W.J. & R.D. Turner (1956)  Freshwater mollusks of Alabama, Georgia, and Florida from the Escambia to the Suwannee River. Bull. Fla. State Mus. (Biol. Sci.), 1: 97-239.   For more, see:

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

[10] Clench, W.J. (1962) A catalogue of the Viviparidae of North America with notes on the distribution of Viviparus georgianus Lea. Occasional Papers on Mollusks 2(27): 261-287.

[11] Clench, W.J. & S.L.H. Fuller (1965) The genus Viviparus (Viviparidae) in North America. Occasional Papers on Mollusks 2(32): 385-412.

[12] Graf [13] has catalogued “nearly 1,000” specific nomina historically applied to North American freshwater gastropods of the family Pleuroceridae.  Between 1934 – 1944 my hero Cavin Goodrich was able to pare these down to approximately 150.  For more, see:

• The Legacy of Calvin Goodrich [23Jan07]

[13] Graf, D. L. (2001) The cleansing of the Augean Stables, or a lexicon of the nominal species of the Pleuroceridae (Gastropoda: Prosobranchia) of recent North America, north of Mexico. Walkerana 12 (27) 1 - 124.

[14] For more about the “Nestor of American Naturalists,” see:

• Isaac Lea Drives Me Nuts [5Nov19]

[15] For the further exploits of my hero, see:

• Goodrichian Taxon Shift [20Feb07]
• Mobile Basin II: Leptoxis Lessons [15Sept09]
• CPP Diary: The Spurious Lithasia of Caney Fork [4Sept19]

[16] This is a difficult work to cite.  J. B. Burch's North American Freshwater Snails was published in three different ways.  It was initially commissioned as an identification manual by the US EPA and published by the agency in 1982.  It was also serially published in the journal Walkerana (1980, 1982, 1988) and finally as stand-alone volume in 1989 (Malacological Publications, Hamburg, MI).

[17] The parallel between the careers of Virginia Vail and George Te is inescapable here.  George Te was a Burch student in the late 1970s and seems to have ghost-written Burch’s entire treatment of the Physidae, as Virginia Vail ghost-wrote the Viviparidae.  For more on George Te, see:

• To Identify a Physa, 1975 [6May14]
• To Identify a Physa, 1978 [12June14]

[18] Abbott, R.T. (1975)  American Malacologists, Supplement.  American Malacologists, Greenville, Delaware. 

[19] Vail, V.A. (1977)  Comparative reproductive anatomy of 3 viviparid gastropods.  Malacologia 5: 519 – 540.

[20] Vail, V.A. (1978)  Seasonal reproductive patterns in 3 viviparid gastropods.  Malacologia 6: 73 – 97.

[21]  The viviparids have evolved [22] quite a few unique adaptations that separate them from all other living gastropods, including a weird operculum and even weirder radula.  The right tentacle of the male has been modified into a simple, external penis and the pallial gonoduct of the female modified into a marsupium, capable of nursing fertilized eggs until their hatch into impressively large crawl-away juveniles.  But within the family, their anatomy is as boringly uniform as the pleurocerids.  You crack a Viviparus shell, or a Lioplax shell, or a Campeloma shell, and look inside, and it’s basically viviparid guts.  Every time.

[22] “Retained” might be a better verb here.  The worldwide family Viviparidae seems to be ancient.  They share their peculiar concentric operculum with the Ampullaridae, which suggests that the two families are sisters.  But the viviparids have absolutely no living marine antecedents.  I take this as evidence of an hypothesis I advanced back in 2009, that evolution is slower in fresh waters than in the marine environments from which all life originated.  Like the pleurocerids, the viviparids are “living fossils.”  For more, see:

• The snails the dinosaurs saw [16Mar09]

[23] Mattox, N.T. (1937) Oogenesis of Campeloma rufum, a partheogenetic snail.  Zeitschrift fur Zellforschung und Mikroskopische Anatomie 27: 455 – 464.

Mattox, N.T. (1938)  Morphology of Campeloma rufum, a parthenogenetic snail.  Journal of Morphology 62: 243-261.

[24] Mattox sampled his study population from a tributary of the Wabash River in eastern Illinois.  Clench [10] subsequently synonymized Campeloma rufum under C. crassulum (Raf.) 

[25] Abbreviations from Vail [19] figures 5 and 10: AG = albumin gland, CM = columellar muscle, DG = digestive gland, M = mantle, O = ovary, OD = oviduct, PMC = posterior end mantle cavity, PO = pallial oviduct, PR = prostate gland, RT = right tentacle, SR = seminal receptacle, SV = seminal vesicle, T = testis, V = vagina, VD = vas deferens,  VD’ = pallial vas deferens.

[26] Vail, V.A. (1979) Campeloma parthenum (Gastropoda: Viviparidae), a new species from north Florida.  Malac. Rev. 12:85-86. 

[27]  Isn’t it interesting the way we can remember small vignettes from 40 years ago, but cannot remember what we had for supper last night [28]?  Virginia Vail gave her talk in the freshwater session of the Corpus Christie AMU meeting at 11:00 Thursday morning, August 9, 1979.  Young Rob Dillon, then listed as a graduate student at the University of Pennsylvania, gave his talk at 11:15, “The Goniobasis of southern Virginia and northwestern North Carolina: Electrophoretic and shell morphological relationships [29].”  At approximately 11:31, Old Joe Morrison jumped up and lectured me with great passion about obscure details of pleurocerid taxonomy and systematics.  At about 11:35, I said, “Easy, big fella.”  For more, see:

• Joe Morrison and The Great Pleurocera controversy [10Nov10]

[28] It was a chicken casserole, with cashews sprinkled on top.  I just looked in the refrigerator.

[29] That was just the second presentation I had ever made at a national meeting.  The research was ultimately published as: Dillon, R.T., Jr and G.M. Davis (1980) The Goniobasis of southern Virginia and northwestern North Carolina: Genetic and shell morphometric relationships. Malacologia 20: 83-98. [PDF]

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

[31] No, it was not at an AMU meeting.  Jack Burch was never a member of the AMU/AMS during his entire professional career, as far as I know, until being elected an honorary life member in 2009.  His election was not unanimous.

[32] Clarke, A.H. (1973) The freshwater mollusks of the Canadian Interior Basin.  Malacologia 13: 1 – 509.

[33] The nomina brevispirum (Baker 1928), exilis (Anthony 1860), and gibba (Currier 1867) were not actually forgotten.  If you look forward into Burch’s “Species List, Ranges, and Illustrations” on page 92, you will find them synonymized under Campeloma decisum.

[34] “Huntsville or Stevenson, Alabama.”  This was corrected to Decatur, AL by: Clench, W. J. and R.D. Turner (1955) The North American genus Lioplax in the Family Viviparidae.  Occasional Papers on Mollusks, Museum of Comparative Zoology, Harvard. 2(19): 1 -  20. 

Dr. Henry A. Pilsbry was a Jackass

Editor's Notes - The only reason I am posting the brief essay that follows is that I need to blow off some steam.  This is entirely personal.  I apologize in advance.

This essay was subsequently published as: Dillon, R.T., Jr. (2023c)  Dr. Henry A. Pilsbry Was A Jackass.  Pp 283 – 286 in The Freshwater Gastropods of North America Volume 7, Collected in Turn One, and Other Essays.  FWGNA Project, Charleston, SC.

In the winter of 1888-89, while he was yet a student at Brown University [1], my hero Frank Collins Baker [2], just 21 at the time, undertook an expedition to the little fishing community of Micco, Florida, on the Indian River Lagoon.  There he “had the opportunity to compare the species which are common to both the Northern and Southern shores.”  He seems to have collected at least 19 lots of shells during his explorations around Micco, including land snails as well as marine gastropods and bivalves, to judge by collections subsequently catalogued into the ANSP.

From Clench & Turner [3]

The following summer Baker accepted a Jessup Scholarship to work with Dr. Henry A. Pilsbry and moved to Philadelphia, carrying the collections he had made around Micco with him.  And in September of 1889, his mentor Pilsbry published Baker’s first scientific paper [4] in the very first volume of The Nautilus [5], entitled “Notes on Floridian Shells” [6].  Here is the third paragraph of that four-paragraph note, quoted in its entirety:

“I was very much surprised to find in one of my hauls with the dredge, a number of very perfect specimens of Turbonilla interrupta Totten, associated with Odostomia interrupta Say, and also Nucula proxima Say.”

Those “Turbonilla” shells, subsequently reidentified as Truncatella pulchella (Pfeiffer) [7], can still be seen in the ANSP collection today, catalogue numbers 60124 for adults and 60344 for juveniles.  Their locality data remain exactly as Baker wrote on his labels in 1888, and as he published in 1889, “off Micco, Fla., Indian River.”

The image below was posted on Facebook back in April of 2019 by our good friend Paul Callomon, collection manager at the Academy of Natural Sciences of Philadelphia.  It shows an outrageous slander on the name of my hero, written in the unmistakable chicken-scratch of Dr. Henry A. Pilsbry.

This is inexcusable.  I cannot find any reason to doubt that F. C. Baker collected those two lots of shells from the Indian River at Micco.  But if Pilsbry did have doubts, he could have broached them with his young protégé right there man-to-man, as Baker was standing in front of his editorial desk, manuscript in one hand and shells in the other.  There can be only one reason that Pilsbry accepted Baker’s paper, published it, and then impugned the young man’s character because of it.  Dr. Henry A. Pilsbry was a jackass.

Thanks, I feel better.

Notes

[1]  H. J. Van Cleave (1945) “A memorial to Frank Collins Baker (1867 – 1942).  Pp xvii – xxxvi in, Baker, F.C., The Molluscan Family Planorbidae.

[2] For a brief biography of my hero, see:

• The Legacy of Frank Collins Baker [20Nov06]

[3] Clench, W.J. & R. Turner (1948) The genus Truncatella in the Western Atlantic.  Johnsonia 2: 149 – 164.

[4] He had one prior publication – a description of the Conchology Department in “The Old Curiosity Shop” of California.

[5] Volumes I and II were entitled “The Conchologist’s Exchange.”  Pilsbry picked up the subscription list of The Conchologist’s Exchange, and hence began publication of The Nautilus in May of 1889 with Volume III.

[6] Baker, F. C. (1889) Notes on Floridian shells.  Nautilus 3: 53 – 54.

[7] The sample that Baker identified as the pyramidellid “Turbonilla interrupta” turns out to be Truncatella pulchella, an amphibious gastropod of Florida and Caribbean coastal environments [8], apparently washed from its near-shore habitat into the deeper waters of the Indian River.

[8]  It is certainly possible that Pilsbry subsequently realized that Baker’s little sample of snails were truncatellids, not pyramidellids.  Then why not assume that Baker was mistaken in his identification, as the curatorial staff at the ANSP does today?  Why imagine that Baker lied about his collection locality?  Arrogant jackass!  Takes one to know one.

The Emperor Speaks

Editor’s Note – This essay was subsequently published as: Dillon, R.T., Jr. (2023c)  The Emperor Speaks.  Pp 253 – 260 in The Freshwater Gastropods of North America Volume 7, Collected in Turn One, and Other Essays.  FWGNA Project, Charleston, SC.

I will begin my essay this month confessing an error that I committed in the late summer of 2018, as relayed to this group two months ago.  Here is a direct quote from my 5Oct20 essay on the “Flat-topped Helisoma of the Everglades”

“So, reading Wetherby’s description in the calm of my office here one sunny morning in Charleston a couple years ago, I was stricken with the impression that the type locality of Helisoma duryi might could use a bit of narrowing-down.  And I swiveled my chair and pulled my well-thumbed copy of F. C. Baker’s (1945) “The Molluscan Family Planorbidae” off the bookshelf.”

That was lazy of me.  I should have consulted Baker's mentor, the Elderly Emperor [1] Dr. Henry A. Pilsbry (1862 - 1957).

His regal ghost still flickered, dimly, through the hollow corridors of the mollusk collection at the Academy of Natural Sciences during my years as a graduate student in Philadelphia.  When I arrived at that venerable institution in the summer of 1977, the first stop on my first tour was the “Pilsbry Chaos,” a pile of boxes, papers, and shells through which curatorial assistants were still laboring, 20 years after the great man’s death. 

From H. B. Baker [2]

He was born on a small farm near Iowa City and seems to have developed his interest in land and freshwater shells at the nearby University, from whence he was awarded his B.Sc. in 1882.  Pilsbry then became a newspaper man, briefly, as was his contemporary Calvin Goodrich [3], moving to New York City as a proofreader in 1887.

He rocketed to malacological stardom almost immediately thereafter, at the age of 25.  On Thanksgiving Day of 1887, Pilsbry was invited to Philadelphia by George W. Tryon [4], who offered him a job as his assistant.  When Tryon died suddenly in February of 1888, Pilsbry inherited Tryon’s position as Conservator the Conchological Section, and Editor of the Manual of Conchology.  Pilsbry sat behind that high desk at the ANSP for 70 years, until they carried him out on a plank [5].

In 1889 he founded “The Nautilus,” the Volume 70 galleys of which were on the cluttered desk around which his plank was wedged.  Frank Collins Baker also came to Philadelphia to work with Pilsbry in 1889, and left the next year profoundly affected [6].  In 1890 Pilsbry organized the American Association of Conchologists, the first of several precursors to the American Malacological Union, of which he was elected first president.  In 1899 he was awarded a doctorate of science by his alma mater, the University of Iowa, the first of three doctorates he was ultimately to receive [7].

Pilsbry’s primary interest was in the North American land snails.  H.B. Baker characterized his (1895) “Guide to the Study of Helices” as “the most brilliantly original, iconoclastic book that ever has been written about the subject.”  Of land snails.  His four-volume “Land Mollusca of North America” (1939 – 1948) is the alpha of the American terrestrial gastropod fauna even unto the present day and may ultimately (I fear) prove to be the omega as well.

But the Elderly Emperor was widely published in freshwater, marine, and fossil malacology as well, from all over the world.  No one can count the sum of his works.  His biographers wrote, “an estimate between 3,000 and 4,000 possibly might cover the number of published articles that flowed from his facile pen.” Paging through the Burch canon [8], I count 57 species or subspecies of North American freshwater gastropods described by Pilsbry surviving even unto 1980, plus eight Pilsbry genera and three Pilsbry subgenera, in eight families.  Accepting Burch’s estimate of approximately 500 species, Pilsbry may be credited with describing over 10% of our freshwater gastropod fauna.  Not bad for a secondary interest.

So early in his career Pilsbry began taking regular vacations to Florida [10].  And in 1934 he published a large and wide-ranging paper in the Proceedings of the ANSP entitled “Review of the Planorbidae of Florida, with notes on other members of the family [11].”  The first 17 pages of that work were subtitled “I. The Large Planorbes of Florida,” which since not followed by a second section subtitled “The Small Planorbes of Florida,” turns out to have been what he meant by “The Planorbidae of Florida” in his main title, screw all those little ones [12].  The second 20 pages of Pilsbry’s 1934 paper were subtitled “Notes on Other Planorbidae,” which turned out to be an ambitious review of planorbid systematics across The Americas, with descriptions of a bunch of new species from three continents.  He described his new genus Australorbis about halfway through that second section, assigning Say’s (1818) glabratus to it, not helping [13].

Paratype lot of H. duryi in the UMMZ [9]

But it is the first half of Pilsbry’s 1934 paper that has brought him to our attention this month.  He began with Helisoma trivolvis, which (of course) is widespread throughout North America, which he allocated to the Dall subgenus Pierosoma.  He then undertook to describe a new subgenus, Seminolina, with Helisoma scalare (Jay 1839) as the type [14].  He also assigned to his new Seminolina two fossil species of Dall (conanti and disstoni) and “the Helisoma duryi complex.”  In the duryi complex he recognized, in addition to Wetherby’s typical subspecies of 1879, intercalare (Pilsbry 1887), preglabratum (Marshall 1926), and three new subspecies: seminole, normale, and eudiscus.  We touched briefly upon all this taxonomic churn back in October.  Sorry to bring it up again.

And regarding the type locality of Helisoma duryi, Pilsbry wrote: “Wetherby’s locality “Everglades of Florida” was vague and doubtless inexact.  I am informed by Mr. Ralph Dury [15] that in the trip of 1874 his father visited places along the coast of Volusia County – Tomoka River, Port Orange, Daytona, Halifax River […] It seems likely therefore that H. duryi was found somewhere along the eastern border of Volusia County.”

D’oh!  Back in 2018, with F. C. Baker’s 1945 monograph open in my lap [16], I had convinced myself that a good typical (if not necessarily type) locality to sample H. duryi might be located on the Tamiami Trail at the 40-mile bend.  That is what sent me dodging airboats way down in The Everglades in October of 2018 [17], and that is why I had such high hopes for Cindy Norton’s 2019 breeding experiments [18].  In retrospect, I should have consulted The Elderly Emperor first.

In my own defense, here is the verbatim quote from Wetherby: “This shell was given me several years ago, by Mr. Charles Dury, who brought it from the Everglades of Florida.  It was also among the shells received from the Miami country.”  Volusia County is not in The Everglades, even under the most expansive modern definition of that term.  And Volusia is 250 miles north of Miami, and always has been.

Excuse logged.  Now go back to Florida, Dillon, and do your job right.

Digging into the Pilsbry paper further, it materializes that The Elderly Emperor examined Wetherby’s actual type lot, which Bryant Walker got hold of somehow, which sits in the UMMZ collection to this day.  That set of shells comprises a holotype (UMMZ 83501) and nine paratypes (UMMZ 83502) as figured above.  Pilsbry measured and figured four of the ten, including the holotype, which is where I got “19.5 mm” for footnote [3] of my October post.  And if you can believe it, Pilsbry split one of the shells out of Wetherby’s type lot of Helisoma duryi duryi into his own newly-described Helisoma duryi seminole.  See figure #4 in the Pilsbry montage below.

Wetherby’s type lot. #2 = holotype, #4 = H. d. seminole

Now would be an opportune time, I suppose, to make explicit what has, to this point in my essay, been implicit.  Henry Pilsbry was innocent of the Modern Synthesis.  The only species concept of which he was aware was the nineteenth-century “organism or group of organisms recognized as distinct by a competent taxonomist.”  Which Pilsbry, without question, was.  So, if His Imperial Majesty recognized a species, then it was a species, by definition.

And exactly the same for subspecies.  Under today’s modern synthesis of evolutionary thought, we define subspecies as “populations of the same species in different geographic locations, with one or more distinguishing traits [19].”  Pilsbry never considered that “different geographic locations” thing.  Subspecies were what he recognized as subspecies, just the same as species were what he recognized as species, only with less of whatever that species juice might be.

So although Pilsbry examined the type lot in Bryant Walker’s collection, it materializes that he never had any fresh Helisoma duryi duryi from anywhere in Volusia County in front of him.  Nor did his protégé Baker.  The type locality remained only slightly less mysterious to The Elderly Emperor than to me, reading his words in the calm of my office a couple months ago.

Volusia County is today home to approximately a half-million residents, 122 motels, 5 Walmart Supercenters, and the World Center of Racing.  Next month, we race off to Daytona!

Notes:

[1] R. Tucker Abbott (1958) coined that sobriquet on page 103 of his contribution to the Pilsbry festschrift: "From the Pilsbry Chair of Malacology."  Nautilus 71: 100 – 103.

[2] I have gleaned most of the biographical details relayed here from Baker, H.B. (1958) Henry Augustus Pilsbry 1862 – 1957.  Nautilus 71: 73 – 83.

[3] Calvin Goodrich (1874 - 1954) was an early-modern malacologist, Pilsbry the paragon of the late pre-modern.  For more, see:

• The Legacy of Calvin Goodrich [23Jan07]

[4]  We explored the relationship between George Tryon (1838 - 1888) and his immediate predecessor at the ANSP in:

• Isaac Lea Drives Me Nuts [5Nov19]

[5] Not really, but darn close.  He suffered a heart attack at his desk in September of 1957 and died in his sleep in October.

[6] For a bit of background on my malacological hero, see:

• The Legacy of Frank Collins Baker [20Nov06]

We will hear much more about the relationship between Baker and Pilsbry in coming months.

[7]  Pilsbry was ultimately awarded doctorates of science by the University of Iowa (1899), the University of Pennsylvania (1940), and Temple University (1941).

[8] This is a difficult work to cite.  J. B. Burch's North American Freshwater Snails was published in three different ways.  It was initially commissioned as an identification manual by the US EPA and published by the agency in 1982.  It was also serially published in the journal Walkerana (1980, 1982, 1988) and finally as stand-alone volume in 1989 (Malacological Publications, Hamburg, MI).

[9] We thank Taehwan Lee of the University of Michigan Museum of Zoology for braving the perils of the worldwide Coronavirus panic to assemble and photograph for us the lovely montage of H. duryi paratype lot 83502 reproduced above.

[10] Here’s a quote from T. L. McGinty (Nautilus 71: 97 – 100):  “Early in 1937, Dr. Pilsbry secured a cottage in Lantana, Florida, and each succeeding winter visit to his Florida home brought the Doctor new friends.”

[11] Pilsbry, H. A. (1934)  Review of the Planorbidae of Florida, with notes on other members of the family.  Proceedings of the Academy of Natural Sciences of Philadelphia 86: 29 – 66.

[12] Helisoma duryi, you may recall from my essay of October, was originally described by Wetherby (1879) as neither large nor small, but rather “medium-sized.”  Pilsbry (1934) folded the medium-sized planorbes in with the large.  I suppose we, the students who follow in the great man’s footsteps, should be grateful.

[13] Here’s a direct quote from H. B. Baker’s Pilsbry obituary [2]:

“Very rarely, when in a Puckish mood, did he (Pilsbry) wield his prestige to establish dubious cognomens; thus he argued against the use of Mesomphix instead of Haplotrema, but contrarily replace Planorbina guadaloupensis by (Biomphalaria) Australorbis glabrata (1934).”

I don’t know what that means, but it sounds important, so feel obligated to pass it along.

[14] John Clarkson Jay (1839) spelled the species name “scalaris” to agree in gender with the feminine Paludina.  Pilsbry re-spelled it scalare when he moved the specific nomen under the neuter noun-construct Helisoma in 1934.  Thanks to Harry Lee for the insight.

[15] We tipped our hat to Mr. Charles Dury in October footnote [4].  His son Ralph E. Dury (1899 – 1984) was Director of the Cincinnati Museum of Natural History for almost 60 years.

[16] Baker, F.C. (1945) The Molluscan Family Planorbidae. Urbana: University of Illinois Press. 530 pp.

[17] If you haven’t read it already, you might be entertained by:

• The Flat-topped Helisoma of The Everglades [5Oct20]

[18] And as long as you’re reviewing my previous posts, you might as well bring yourself up to date:

• Foolish Things With Helisoma duryi [9Nov20]

[19] To refresh your memory on the definition of the word “subspecies” as adopted by the FWGNA Project, see:

• What Is A Subspecies? [4Feb14]
• What Subspecies Are Not [5Mar14]

The Return of Captain Lyon

Editor’s Note – This essay was subsequently published as: Dillon, R.T., Jr. (2023b)  The Return of Captain Lyon.  Pp 81 – 88 in The Freshwater Gastropods of North America Volume 6, Yankees at The Gap, and Other Essays.  FWGNA Project, Charleston, SC. 

“One man only remained behind.  It was Captain Sidney S. Lyon, of Jeffersonville, one of Indiana’s most gifted engineers.  As the Federal army moved away, he sat down upon a rock and waited.  Beside him lay a black, snake-like rope, the end running into the steep side of the mountain.  It was a fuse.  He had mined the mountain and filled the hollow with all the powder the fleeing army could spare…The tramp of the marching died away; the commissary stores were burning, and still he sat, as the night fell over the heights and the darkness filled the ravines.  Were the Confederates coming?  He heard the faint hoof-beats, the rumble of a great force of men coming from the Tennessee side.  There was the sparkle of a match, the splutter of powder, and a man fleeing down the mountain toward Kentucky for his life, and then…”

I honestly don’t know to what extent The Blowing of Cumberland Gap is true [1], but it does not matter, because it’s a great story, and I would recommend that you look up the Lyon family history [2] from google books and read pp 185 – 192, if you want to know how it turned out.  Or, if you would rather read my tenth (and final!) essay on the shell morphological variation and taxonomic confusion in the pleurocerid snails of the Tennessee/Cumberland in twelve months, soldier on below.

Sidney Smith Lyon (1808 - 1872)

In the fall and winter of 2016, a duller 154 years later, I spent a couple weeks in Frankfort, Kentucky, working with our good friend and colleague Ryan Evans on the freshwater gastropods of the Bluegrass State.  I had been surveying the region in preparation for the roll out of our new FWGO web resource for several years and, at that juncture, fancied myself familiar with most elements of the freshwater gastropod fauna of the Ohio River basin.  But what I found in KYDOW macrobenthic samples from little tributaries of the Green River, and even in some little creeks draining directly into The Ohio in western Kentucky, surprised me.  To the point that I can still remember it four years later.  Which at my age, is saying a lot.

I was already aware that P. simplex populations, of diverse shell morphology, extended across the entire drainage of the Tennessee/Cumberland from SW Virginia to Chattanooga, west through Tennessee and north into the Ohio River tributaries of Kentucky.  But my goodness!  Crawling around together with the P. simplex in the little creeks of western Kentucky, way out beyond Louisville, the KYDOW teams were also collecting pleurocerids indistinguishable to my eyes from Pleurocera troostiana, or arachnoidea, or spinella, or strigosa, or striatula, or whatever else people have called that pleurocerid bearing light, slender, striate shell with small body whorls in upper Tennessee River tributaries for almost 200 years.  The entire malacofauna of little creeks in Western Kentucky was indistinguishable from that of East Tennessee.

The shells of the particular population I was examining that afternoon were not costate, although several nearby populations were.  That is what especially struck me about those KYDOW samples, as I sat at the lab bench in Frankfort back in 2016, in addition to all the above.  I was as stricken by what I was not seeing, as what I was.  And this thought suddenly dawned on me.  If a pleurocerid species bearing a fat-smooth shell, like P. simplex, ranges through little creeks across four states, why can’t a pleurocerid species bearing a skinny-striate shell, like P. troostiana?

Western Kentucky or East Tennessee?

Let me back up for a bit of context.  Throughout Kentucky, and indeed through Middle Tennessee as well, streams of all sizes and descriptions are often choked with large and morphologically diverse populations of Pleurocera laqueata (Say 1829).  They bear shells with large body whorls that are sometimes striate but always, always costate.  I’d been looking a bottles-full of Pleurocera laqueata from central Kentucky all week.  Is it possible that some of those samples might be P. troostiana, slightly costate, not P. laqueata, slightly striate?

Have P. laqueata and P. troostiana been confounded in this part of the world for 200 years?  Have they been confounding each other [3]?  Might the nineteenth-century literature contain dozens of names for both, subsequently scrambled by synonymy?  If so, what are workers calling the tall-skinny-striate pleurocerids in Kentucky today [4]?

Following Branson’s [6] “Keys to the aquatic Gastropoda known from Kentucky” one finds four Goniobasis species “with longitudinal plicae” (i.e. striations) at any point in their shell growth: laqueata (Say 1829), curreyana (Lea 1841), costifera (Hald. 1841), and plicata-striata (Wetherby 1876).  The type shells borne by all four of these species are primarily costate, and I think the last three nomina will prove junior synonyms of Say’s laqueata.  Bookmark that question.  One day we’ll come back to it.

Reference to the primary literature, however, returns an excellent paper by Bickel [7] on the biology of pleurocerid populations he called “Goniobasis curreyana lyoni (Lea 1863)”.  I think that’s it.

Goniobasis lyonii then [10] and now.

Isaac Lea published a brief, Latinate description of Goniobasis lyonii – note two eyes in the original spelling [8] – in his ANSP Proceedings paper [9] “ordered to be published” on May 27, 1862 (although it says 1863 on the title page), followed by a more complete English description and figure in his ANSP Journal paper [10] “ordered to be published” on May 26, 1863.  We have touched on this pair of publications several times in recent months, featuring them in May.

Lea’s acknowledgement in the ANSP Journal paper [10] was: “I dedicate this with great pleasure to Mr. Lyon, Civil Engineer and State Geologist.”  War had not yet broken out between the states at the date of Lea’s writing, and Mr. S. S. Lyon had served as the topographer for the first (1854 – 57) Kentucky Geological Survey.

But war did come, carrying all American topographers along with it.  And my loyal readership will remember that it was Capt. Sidney Smith Lyon who stooped to capture two populations of rebel pleurocerids at Cumberland Gap in the summer of 1862, which he posted back behind the lines to Isaac Lea in Philadelphia.  Lea described that sample under four names in the smaller PANSP paper of 1863 [11] that kicked off this entire series of blog posts, way back in [4Aug19].

I counted 20 hits to S. S. Lyon in Isaac Lea’s bibliography [12], first as Mr. Lyon, then as Capt. Lyon, and ultimately as Maj. Lyon, after August of 1863.  According to the family history published in 1907 [2], Lyon was an artist, a crinoid paleontologist, a naturalist, and singlehandedly saved the army of General George Morgan when it was surrounded at Cumberland Gap in September of 1862.  The account extracted at the top of the present essay was written by one Col. James Keigwin, and lovingly passed down to us by the Lyon family.

But back to the snails.  Lea’s description of Goniobasis lyonii noted that the shell was “very much drawn out” and “striate above.”  He did not mention costae but his figure (#156, reproduced above) shows light costation as well as light striation on the slender shell.  Lea gave the habitat as “Grayson County, Kentucky, S. S. Lyon.”  Its subsequent taxonomic history is Byzantine [14], but it was the substance of Bickel’s 1968 paper that lyonii should be resurrected as a subspecies of curreyana, which I don’t think it is [15], but I do give thanks for the resurrection.

Bickel restricted the lyonii type locality to Spring Fork Creek, a tributary of the Rough River in Grayson County, Kentucky.  He also reported populations in seven other small streams in four other counties: Breckinridge, Meade, Hardin, and Larue.  I sampled Spring Fork at three locations on the morning of 15May19 and was most disappointed by the quality of the environment, which in the last 50 years has declined to the status of muddy ditch, and by the population of pleurocerids dwelling therein, which has dwindled to zero.  Providentially, my database contained three other Grayson County sites at which lyonii populations had been collected by the KYDOW, and I found a decent population in a tributary of Big Run Branch, 5 km W of Leitchfield (37.5029, -86.3411).  See the example figured above.

Attributed to S. S. Lyon [13]

Pleurocerid populations bearing slender shells of the lyonii phenotype are not uncommon in small tributaries of the Tennessee, Cumberland, Green and Ohio Rivers in Kentucky west of Louisville and in Tennessee west of Nashville.  I cannot find any argument counter to the suggestion that lyonii (Lea 1862) is a junior synonym of troostiana (Lea 1838).  But let’s continue to recognize this set of populations with a subspecific nomen, Pleurocera troostiana lyonii, shall we?  And again, please review the definition of the word “subspecies” as adopted by the FWGNA from footnote [17] below.

Some populations of P. troostiana lyonii bear shells with light costation around their apex, rendering them effectively indistinguishable from P. troostiana perstriata, and some do not, rendering them indistinguishable from P. troostiana troostiana.  I struggled with the suggestion offered above.  Ultimately, I decided to preserve the S. S. Lyon patronymic in the pleurocerid canon more to protect the small (but not insignificant) literature associated with the taxon than to further any grand evolutionary hypothesis.  Besides which, I have grown rather fond of the gentleman.  He was not some pompous jackass sitting high-and-mighty behind a walnut desk in Philadelphia.  Capt. Sidney Smith Lyon did his duty.

Now at last, the time to summarize has arrived, woohoo.  Available from the link below is a pdf download entitled:

The four subspecies of Pleurocera troostiana (Lea 1838),with synonymy.

Here I have listed and figured all 13 of the specific and subspecific nomina I have synonymized under Isaac Lea’s Pleurocera troostiana since last August, together with type (or typical) localities and references.  I have not listed all the dozens of older synonyms under these nomina, as catalogued by Tryon and Goodrich. Quoting the latter, “I have not had the heart.”  This is FWGNA Circular Number 2 (July 6, 2020).

Notes:

[1] Or maybe they’re lyon?  Sorry, I couldn’t resist.

[2] Lyon, S.E., et al. (1907) Lyon Memorial: Families of Connecticut and New Jersey, including records of the descendants of the immigrants Richard Lyon of Fairfield and Henry Lyon of Fairfield.  Detroit: William Granham Printing.  Pp 185 – 192.

[3] I really think Pleurocera troostiana and P. laqueata hybridize broadly, anywhere and everywhere their ranges overlap.  I mentioned this back in essay of [15Apr20] and asked you to bookmark it.  And I'm mentioning it again here, and asking you to bookmark it, again.

[4] Back in November we reported that Isaac Lea described 505 species of pleurocerids.  Well, Dan Graf [5] cataloged approximately 500 more, contributed by other authors.  Those 1,005 names are a slough of despond into which I do not intend to fall.  So my approach for 40 years has been first to work out the biology, and then to work out the taxonomy.  First, my evolutionary intuition suggests that there are approximately 40 biological species of pleurocerids in North American waters, with another 20 subspecific nomina that may prove of some utility.  So second, I am trying to figure out what 60 names to call those populations or groups of populations.  Those 60 names may not be the oldest, nor the most familiar; they may be a compromise between age and modern use.  But if I live to be 107, I will never get to task #3, what the heck most of those other 945 names mean.

[5] Graf, D. L. (2001)  The cleansing of the Augean stables.  Walkerana 12(27): 1 - 124.

[6] Branson, B.A. (1987)  Keys to the aquatic Gastropoda known from Kentucky.  Trans. Kentucky Acad. Sci. 48: 11 - 19.

[7] Bickel, D.  (1968)  Goniobasis curreyana lyoni, a pleurocerid snail of west-central Kentucky. The Nautilus 82: 13 - 18.

[8] To quote The Eagles, “You can’t hide your lyon eyes,” pleural.  OK, I’ve got all the corny jokes out of my system, for now.

[9] Lea, Isaac. (1862) Description of a new genus (Goniobasis) of the Family Melanidae and eighty-two new species. Proc. Acad. Nat. Sci., Phila., xiv, pp. 262-272.

[10] Lea, Isaac (1863) New Melanidae of the United States.  Journal of the Academy of Natural Sciences of Philadelphia 5: 217 – 356.

[11] Lea, Isaac (1863) Descriptions of fourteen new species of Melanidae and one Paludina.  Proceedings of the Academy of Natural Sciences of Philadelphia 15: 154 – 156.

[12] Scudder, N. P. (1885)  Bibliographies of American naturalists – II. The published writings of Isaac Lea, LL.D.  Bull. US National Museum 23: 1 – 278.

[13] For more about the “White Horse” painting attributed to S. S. Lyon, see:

Clark County Museum looking for help to restore historic painting.  News and Tribune (Clark County, Indiana).  February 12, 2018. [html]

[14] I quote Bickel [7] verbatim: “This animal is Goniobasis lyoni Lea, 1863, a species that Tryon (1865) placed in the synonymy of Goniobasis glauca (Anthony).  It was subsequently transferred along with G. glauca to the synonymy of Goniobasis athleta (Anthony) by Tryon (1873), and Goodrich (1940) shifted it to the synonymy of Goniobasis laqueata (Say).  Goniobasis lyoni is a form of Goniobasis curreyana (Lea) and is distinct enough to merit recognition.”

[15] It is hard for me to understand why Dave Bickel thought lyonii was a subspecies of curreyana.  Lea’s (1843) figure clearly depicts Melania curryana as a laqueata-type, bearing an aperture “about one-third the length of the shell,” not a troostiana-type, with aperture smaller.  And Lea specifically stipulated [16] that the shell of curreyana “is without striae,” focusing instead on the “large and strong folds.”  Lea’s (1841) Melania curreyana is clearly a junior synonym of laqueata (Say 1829) and has nothing to do with his Goniobasis lyonii of 1862.

[16] Lea, Isaac (1843) Description of New Fresh Water and Land Shells.  Transactions of the American Philosophical Society 8: 163 – 250.

[17] Subspecies are populations of the same species in different geographic locations, with one or more distinguishing traits.  For an elaboration, see:

• What is a subspecies? [4Feb14]
• What subspecies are not [5Mar14]