Freshwater Gastropods of Georgia (Atlantic)

I'm very pleased to report that the Freshwater Gastropods of Georgia (Atlantic drainages), by R. T. Dillon, W. K. Reeves, and T. W. Stewart is officially up and open for business!  Check it out:
http://www.cofc.edu/~fwgna/FWGGA/ [1]

The database analyzed includes 845 records from 264 sites throughout eastern and central Georgia - primarily from our own original collections or those of the Florida Museum of Natural History in Gainesville.  We document 37 species of freshwater gastropods inhabiting the region, 10 new and 27 shared with our sites for South Carolina and North Carolina already on line.

The FWGGA site is designed to integrate smoothly with our pre-existing FWGSC and FWGNC sites.  Habitat, distribution, ecology, life history, taxonomy and systematics given are for each species, as well as PDF downloads of range maps.  The Georgia site also features a photo gallery, a clickable dichotomous key, and a section on conservation recommendations.

Special notes of appreciation are due to our colleagues Doug Florian for help with the mapping and Steve Bleezarde for his web wizardry.  If anybody spots any errors, glitches, or broken links, we'd appreciate a heads-up.  Now our undivided attention turns toward Virginia!

Note

[1]  This link is obsolete, as of 2010.  New link:
http://www.fwgna.org/FWGGA/

Goodrichian Taxon Shift

Editor's Note.  This essay was subsequently published as: Dillon, R.T., Jr. (2019c) Goodrichian Taxon Shift. Pp. 7-10 in The Freshwater Gastropods of North America Volume 3, Essays on the Prosobranchs.  FWGNA Press, Charleston.

I'm pleased to report that an illustrated Key to the Pleuroceridae of Virginia, recently developed by Brian Watson, Tim Stewart and myself, is now available on the FWGNA site. The resource includes notes on habitat, distribution, ecology, life history, taxonomy and systematics for all 12 pleurocerid species inhabiting the Commonwealth. It was made possible by a contract from the Virginia Division of Game and Inland Fisheries, and is part of our larger "Freshwater Gastropods of Virginia" project currently ongoing.

The focus of the VDGIF contract was on the diverse Goniobasis populations that inhabit the Tennessee River tributaries of southwest Virginia, a taxonomically difficult fauna about which some conservation concern has been raised. Our survey of the genetic divergence among 12 populations at 11 allozyme-encoding loci confirmed three species in the region - Goniobasis clavaeformis, G. simplex, and G. arachnoidea (1). We combined our fresh data on the Tennessee drainage pleurocerid fauna with previously-accumulated observations from the New River and Atlantic drainages to construct the statewide key that is now open for business.

The most interesting result of the VDGIF genetic survey was the extreme shell-morphological variation we documented in G. clavaeformis. Typical G. clavaeformis, as described in February of 1841 by Isaac Lea, bear rather vanilla shells of moderate thickness and rounded whorls. Goodrich (2) realized, however, that upstream populations can have more slender shells with a pronounced mid-whorl carination, a form that was originally described as "G. acutocarinata" by Lea, two pages after he described clavaeformis. Interestingly, it was acutocarinata that Pilsbry & Rhoads chose as the type of the genus Elimia, since that nomen appeared first in the alphabetical list of odd-lot taxa gathered into the genus by H. & A. Adams (1854). That acutocarinata was subsequently recognized as a synonym of clavaeformis by Goodrich had no bearing on the taxonomic controversy that ensued (3).

Although I was well aware of the acutocarinata/clavaeformis connection, I must admit to being fooled by the dazzling variety of shell phenotypes that met my eye when I first peered into the clear, cold waters of Indian Creek, a tributary of the Powell River in Lee County, at the southwestern tip of Virginia. I honestly thought I could distinguish six pleurocerid species. In the headwaters I recognized G. carinifera in addition to G. simplex and G. arachnoidea. Further downstream I found Leptoxis praerosa and Pleurocera unciale. And Goniobasis clavaeformis appeared to range throughout Indian Creek, headwaters to mouth, in all manner of shapes and sizes.

The gels soon showed that there was no genetic difference between samples I'd originally identified as G. carinifera and G. clavaeformis. In retrospect, I'm not sure what led me to draw a distinction between carinifera and Goodrich's acutocarinata. But a much greater surprise greeted me when I examined the gels run on samples taken from the mouth of Indian Creek, where it joins the Powell River just over the border in Claiborne County, Tennessee. There is no genetic difference between Goniobasis clavaeformis and downstream samples of Pleurocera unciale (PDF available at Note 4).

The shells borne by snails described as Pleurocera unciale by Haldeman in October of 1841 are broader and heavier than those of G. clavaeformis, their whorls marked with a prominent anterior angulation. Nineteenth-century taxonomists considered such an anterior angulation (and the notch it creates in the shell aperture) so significant that the genus Pleurocera was erected to contain any species distinguished by it. That two samples appearing to represent distinct genera of snails might in fact comprise a single randomly-breeding population seems improbable in the extreme.

But it has been reported before. Last month we reviewed the life and work of Calvin Goodrich, whose 20-year study of pleurocerid diversity in the American South led him to a modern understanding of evolutionary biology much earlier than almost any of his contemporaries (5). And it will be recalled that between 1934 - 1941 Goodrich published a series of papers entitled, "Studies of the Gastropod Family Pleuroceridae," describing extensive intraspecific variation in shell morphology along stream gradients.

In "Studies" Number I (6), Goodrich documented a striking gradient in shell "obesity" (measured as the maximum shell diameter divided by the length of the last two whorls) in the Lithasia geniculata population of the Duck River in central Tennessee. Populations of the typical subspecies L. geniculata geniculata (#14 above) collected downstream showed high obesity, populations from middle reaches (subspecies fuliginosa, #10 above) demonstrated intermediate obesity, and populations from the headwaters (subspecies pinguis, #11 above) were quite slender. In fact, such pinguis populations differed in their shell morphology so greatly from typical Lithasia that they were originally placed by George Tryon in the genus Anculosa.

Here I offer a new term, "Goodrichian Taxon Shift," and define it as intraspecific variation in freshwater gastropod shell morphology along an environmental cline of such magnitude as to prompt the erroneous recognition of multiple nominal taxa. And as Exhibit A, I offer the G. clavaeformis population of Indian Creek (see image at Note 7, below). I predict that many additional examples of Goodrichian taxon shift, likely ecophenotypic in origin, will come to light as modern genetic techniques are applied to natural populations of freshwater gastropods in all groups. And I think that Calvin Goodrich would hardly have been surprised.

References

(1) Dillon, R. T. & J. D. Robinson (2007a) The Goniobasis ("Elimia") of southwest Virginia, I. Population genetic survey. Report to the Virginia Division of Game and Inland Fisheries, contract 2006-9308. 25 pp. [PDF]

(2) Goodrich, C. (1940) The Pleuroceridae of the Ohio River system. Occas. Pprs. Mus. Zool. Univ. Mich. 417; 1-21.

(3) See my post, Goniobasis and Elimia [28Sept04]

(4) Dillon, R. T. & J. D. Robinson (2007b) The Goniobasis ("Elimia") of southwest Virginia, II. Shell morphological variation in Goniobasis clavaeformis. Report to the Virginia Division of Game and Inland Fisheries, contract 2006-9308. 12 pp. [PDF]

(5) The Legacy of Calvin Goodrich [23Jan07]

(6) Goodrich, C. (1934) Studies of the gastropod family Pleuroceridae - I. Occas. Pprs. Mus. Zool. U. Mich. 286:1 - 17. Goodrich, C. (1940) The Pleuroceridae of the Ohio River drainage system. Occas. Pprs. Mus. Zool. U. Mich. 417: 1 - 21.

(7) The image below shows Goniobasis clavaeformis from the mouth of Indian Creek, Claiborne County, TN. U = unciale form, U/T = intermediate between unciale and typical, T = typical form, T/C = intermediate between typical and carinate, C = carinate form.

The Legacy of Calvin Goodrich

Editor's Note.  This essay was subsequently published as: Dillon, R.T., Jr. (2019c) The legacy of Calvin Goodrich.  Pp 1-5 in The Freshwater Gastropods of North America Volume 3, Essays on the Prosobranchs.  FWGNA Press, Charleston.

My essay of November 2006 focused on Frank Collins Baker, a modest man of modest means who rose to prominence in twentieth-century American malacology. This month we'll look at the life and contributions of Calvin Goodrich, a contemporary whose career offers a number of interesting comparisons.

Like Baker, Calvin Goodrich came from a middle class background and held no advanced degree. He was born in Chicago in 1874 and spent his youth in Kansas, graduating from the University of Kansas in 1895. Goodrich then embarked on a career in journalism, serving as a reporter and then editor for The Kansas City Star, The Cleveland Leader, The Toledo Blade, The Detroit Journal, and the Newark Star-Eagle.

It was during his tenure with The Toledo Blade (1908 - 1917) that Goodrich initiated correspondence with the two gentlemen who shaped his second career, A. E. Ortmann of the Carnegie Museum and Bryant Walker of Detroit. Van der Schalie (1) reports that during this period Goodrich began riding the street cars out of Toledo into the surrounding countryside to collect mollusks. And in 1913 he arranged to join Ortmann on a field trip to southwest Virginia, an event that seems to have profoundly affected his life, at age 39. Goodrich began publishing short papers on pleurocerid snails in The Nautilus, obtained appointment as an honorary curator at the University of Michigan Museum of Zoology in 1924, and became a full-time curator at UMMZ in 1926, when he formally retired from the newspaper business.

From 1924 until his (second) retirement from the UMMZ twenty years later, Calvin Goodrich traveled widely in the American south and published about 50 works of scholarship, almost entirely on our mutually favorite family of gastropods (2). Between 1934 and 1941, he published a series of eight remarkable papers which deserve to be better known, his "Studies of the Gastropod Family Pleuroceridae." In those works we see a taxonomist born into 19th-century typology struggling with, and ultimately accepting, a modern understanding of intrapopulation variation.

In "Studies" Number IV (1935), for example, Goodrich focused on the Coosa River of Georgia and Alabama, inhabited by six "forms recognizable as subspecies" of Goniobasis caelatura. He tabulated variation in shell sculpture (two categories of plication and three categories of striation) and in overall shape (ratio of shell diameters measured at two spots). He observed that "In a general sense, the variation from conic to cylindrical shape is in a downstream direction. The same thing is true of variation from smoothness to sculpture." He concluded that "These several forms, however unlike one another they sometimes appear, are nevertheless of the same genetic stock, and they constitute a single, fairly compact group of mollusks." For 1935, such an insight was genuinely prescient.

Today Goodrich's reputation rests primarily on the review of the Pleuroceridae of North America he published as a series of brief works - the first six between 1939 and 1942 "in preparation for a molluscan check list undertaken by the American Malacological Union," the ultimate fate of which I am not aware. Two additional works were added to the series in 1944. These papers are short and spare - they include no descriptions, figures, or indeed biological information of any sort, except ranges. What Goodrich did, however, was to boil something in excess of 500 specific nomena of pleurocerids down into a bit more than 100. Many names were synonymized, without comment, and many others were simply omitted. The 100 nomena recognized by Goodrich have survived in the malacological literature to the present day, while those that Goodrich synonymized or ignored have essentially disappeared, except as dusty labels in the forgotten drawers of historic collections (3).

One might argue that such an approach was arbitrary, and heavy-handed. But Goodrich's judgments were informed by the seven-year study of morphological variation in the Pleuroceridae that preceded them, which he published separately. He was one of the first American malacologists to understand intrapopulation variation, and it was on the basis of his 1934 - 41 "Studies" that his 1939-1944 checklists were compiled.

And Goodrich's review has proven to be of great use to malacologists working in American freshwaters today. My 25 years of research on the population genetics of pleurocerids in the South suggests to me that the total number of biological species in this country will prove to be far less than 500, and indeed less than 100. I haven't found a biological species that Calvin Goodrich missed.

Goodrich's career followed that of F. C. Baker by almost exactly a half generation - he was born seven years after Baker and trailed him in death by 12 years, in 1954. This was an important half-generation. Because from the late 1930's to the mid-1950's, the architects of the "modern synthesis" were fashioning the stones cut by Darwin and Mendel into the science of evolutionary biology as we know it today. Frank Collins Baker, for all his tremendous talent, training, and experience, always considered species to be the subjective constructs of taxonomists such as himself. Any new specimen not matching a previously-described type was, to Baker, a new species. But Goodrich was beginning to think of species as populations or groups of populations, not as individual types. And populations vary. And with that revelation came the dawn of modern evolutionary science.

Keep in touch,
Rob


Notes

(1) Van der Schalie, H. (1955). Calvin Goodrich 1874 - 1954. Nautilus 68: 135-142.

(2) Goodrich's complete bibliography published by Rosewater J. (1959) Calvin Goodrich; a bibliography and catalogue of his species. Occas. Pprs. Mollusks, Mus. Comp. Zool., Harvard 2(24): 189-208. A partial bibliography is available from Kevin Cuming's website at INHS

(3) For a complete catalogue of pleurocerid names, see 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.

The Classification of The Lymnaeidae

Editor's Note. This essay was subsequently published as Dillon, R.T., Jr. (2019b)  The classification of the Lymnaeidae.  pp 7 - 11 in Freshwater Gastropods of North America Volume 2, Essays on the Pulmonates.  FWGNA Press, Charleston.

Last month I reviewed the legacy of Frank Collins Baker (1867-1942), the "freshwater Pilsbry" whose deep personal connection to the biology of the organisms he studied remains vivid through his writings now three generations after he passed away. But this month I suggest that, as much as we may admire it, Baker's (1911) monograph of the North American Lymnaeidae is simply obsolete, and cannot continue to be the basis of the classification system we use today.

Baker recognized about 113 species and subspecies of lymnaeids in North America. These he divided into seven genera: Acella, Bulimnea, Lymnaea, Pleurolimnaea (a fossil taxon), Pseudosuccinea, Radix and Galba. The first six genera were monotypic in North America, or nearly so, while "Galba" was immensely complicated, and further subdivided into five subgenera and three "groups" (1). Baker synthesized a tremendous amount of data in support of this classification, and his science was as good as any malacologist working in 1911. But he was innocent of the Modern Synthesis, and his species concept was typological. And his understanding of the Lymnaeidae has today been superseded by something much, much finer.

I don't know much about Bengt Hubendick – he worked at the Riksmuseum in Stockholm from 1950 to 1959, then moved to the Natural History Museum in Goteborg, where he remained active until about 20 years ago. He wrote a number of very important works, including his (1955) Phylogeny of the Planorbidae, his (1964, 1967, 1970) Studies on the Ancylidae, and the (1978) chapter he wrote on the systematics of the Basommatophora for the set of volumes on Pulmonates edited by Fretter and Peake. But for sheer beauty, no monograph ever written on any group of mollusks before or since has ever topped Hubendick's 1951 masterpiece, "Recent Lymnaeidae, Their Variation, Morphology, Taxonomy, Nomenclature and Distribution (2)."

This (223 page, 369 figure) work of genius is absolutely worldwide in scope, which is amazing for any era. Hubendick used thin-sectioning techniques as well as gross dissection to construct detailed diagrams showing longitudinal sections of the male copulatory anatomy. His plates I, II, and III are amazing – entirely comprised of photos of Lymnaea peregra, showing dozens of diverse forms from scores of localities. He pioneered the use of the "mean photograph," superimposing the images of as many as 20 specimens on top of each other for each figure. Intraspecific variance was not a nuisance to Bengt Hubendick - it was the stuff of evolution, and it was to be cherished.

Hubendick concluded that, while the Lymnaeidae as a family demonstrate "great morphological uniformity, there is a wide range of variation within the various species." He recognized about 40 valid species worldwide, which he saw no reason to subdivide, placing all in the typical genus Lymnaea (3). Here in North America Hubendick admitted humilis, cubensis, bulimoides, catascopium (?), emarginata, columella, megasoma, utahensis (?), haldemani, and arctica, plus the holarctic species stagnalis, palustris, and (perhaps) truncatula (4). With Baker's 113 species reduced to about a dozen, the continued recognition of seven genera would seem difficult to justify.

Hubendick was undeniably correct about that "great morphological uniformity" thing, especially when the lymnaeids are compared to the physids or planorbids. Inside the shell, lymnaeids are all the same snail, varying only in their age at maturity and ecological adaptation.   Outside the shell, their intraspecific variation can be so great as to swamp the interspecific.

Why his clean, modern, rigorous, elegant classification system did not immediately sweep the world is something of a mystery to me.  Burch continued to advocate a modification of Baker's seven-genus system in his (1980, 1982) "North American Freshwater Snails." He wrote,

"The genus Lymnaea has been used variously to include nearly all members of the Lymnaeidae (eg Hubendick 1951) or only Lymnaea stagnalis and several very closely related species (eg F.C. Baker 1928). In this latter system, the family contains a number of species groups (genera) equal in rank to Lymnaea s.s. A third system, more or less a compromise between the previous two, uses Lymnaea as a large inclusive genus, but recognizes various subgeneric groups within it. These subgenera correspond to the genera of the F.C. Baker scheme. As a convenience for species-group separation, the less conservative scheme is used here."

Burch went on to recognize 55 species and subspecies of lymnaeids in North America (3), which he divided into the same genera recognized by Baker (1911, 1928): Acella, Bulimnea, Fossaria, Lymnaea, Pseudosuccinea, Radix and Stagnicola.

Hubendick's system was received more warmly in Europe, adopted in Belgium by Adam (1960) and in England by Macan (1977), although not in Germany by Gloer (1994) nor in the Czech Republic by Beran (2002). The greatest authority on the Lymnaeidae active today must be Poland's Maria Jackiewicz (5), who prefers the "compromise" system mentioned by Burch - Lymnaea as a large inclusive genus with subgenera Stagnicola, Radix, Galba, and so forth.

Let's go with the compromise, shall we? Hubendick's marvelous monograph has convinced me both that the number of valid biological species of lymnaeid snails is small, and that as a family they collectively demonstrate broad morphological uniformity. But I hate to lose the indexing functions of the old genus names. Higher taxonomic categories can play an important roll in information retrieval, and such terms as "Stagnicola" or the "fossarine" lymnaeids have been around for decades.  So for the FWGNA project, I have referred all the lymnaeid species to the genus Lymnaea, with subgenera according to Baker and Burch. I'd invite you all to join me.

And have a Happy New Year!
Rob


Notes

(1) Baker modified this system slightly for his (1928) "Freshwater Mollusca of Wisconsin." The smaller-bodied half of Galba he elevated to the genus Fossaria, and the larger-bodied half became Stagnicola.

(2) Kungl. Svenska Vetenskapskademiens Handlingar. Fjarde Serien Band 3. No. 1. Stockholm: Almquist & Wiksells Boktryckeri AB. I don't understand any of this - I'm just copying it off the cover page.

(3) Excluding the weirdo limpet-shaped Lanx and its relatives.

(4) Hubendick seems to have missed Lymnaea caperata, which I do think is likely valid. He wasn't a god.

(5) Jackiewicz, M. (1998) European species of the family Lymnaeidae. Genus 9: 1 - 93.

Freshwater Gastropods of North Carolina

I'm very pleased to announce that a new website entitled "The Freshwater Gastropods of North Carolina" (R.T. Dillon, B. T. Watson, & T. W. Stewart) is now up and on line!

The new "FWGNC" web resource documents 35 species of freshwater snails inhabiting the Atlantic drainages of North Carolina, using a format intended to be seamlessly compatible with the South Carolina site we've had on line since 2003.  You'll find a photo gallery, a dichotomous key, PDF and jpeg maps showing species distributions, a tabulation ranking all species by their statewide abundance (5,645 records!), and conservation recommendations.

The addition of a second state has prompted reorganization of the entire Freshwater Gastropods of North America website.  The FWGNA site has now moved, and features expanded information resources and many other improvements.  Here's that link:
http://www.cofc.edu/~fwgna/fwgnahome.htm [1]

Update your bookmarks!  And drop by for a visit today - we'd be interested to hear your comments, suggestions, and reports of broken links.  We expect the Atlantic drainages of Georgia to be up in six months, with Virginia close behind.

Special appreciation is due to Dr. Art Bogan and Ms. Jamie Smith for graciously hosting us at the NC State Museum in Raleigh.  Doug Florian was the GIS consultant and Steve Bleezarde the web wizard.  The site was made possible by a grant from the Sierra Club Board of Directors.  Thanks to all of you!

Note

[1] This link obsolete, as of 2010.  New link:
http://www.fwgna.org/FWGNC/

The Legacy of Frank Collins Baker

Editor's Note. This essay was subsequently published as Dillon, R. T., Jr. (2019b)  The legacy of Frank Collins Baker.  pp 1-5 in The Freshwater Gastropods of North America Volume 2, Essays on the Pulmonates.  FWGNA Press, Charleston.

Frank Collins Baker (1) is a hero of mine. Born in Warren, Rhode Island in 1867, he grew up playing with seashells brought to him by his seafaring grandfather. He attended a small business college and spent a year at Brown University before getting his big break, a Jessup Scholarship to the Academy of Natural Sciences in Philadelphia in 1889. At the ANSP he studied under Henry Pilsbry and took part in an expedition to Mexico. Then after several years working for "Ward's Natural Science Establishment" in Rochester (NY), Baker was offered a curatorship at the Chicago Academy of Science (1894 - 1915), where he produced his two-volume "Mollusca of the Chicago Area" (1898, 1902) and his monograph on the Lymnaeidae (1911). A change in research climate at the Chicago Academy sent Baker to the newly-established New York College of Forestry on the campus of Syracuse University for three years, during which time he completed his monumental study of Oneida Lake. In 1918 he accepted a curatorship at the University of Illinois Museum of Natural History (Urbana), where he crowned his productive career with his "Life of the Pleistocene or Glacial Period" (1920), the two-volume "Mollusca of Wisconsin" (1928), his "Fieldbook of Illinois Land Snails" (1939), and his monograph on the Planorbidae, published posthumously (1945).

Baker came from a middle class background, and had just the B.S. degree he earned at the Chicago School of Science in 1896. He was described by H. J. van Cleave (2) as "slight in stature, unpretentious in attitude, mild in disposition, kindly and charitable." Yet his lifetime bibliography extends to 360 titles, including several works to which students of American malacology often refer today.

I can still remember the marvel I felt when, as a graduate student at the ANSP 25 years ago, I first pulled Baker's Oneida Lake monograph off the dusty shelves of the malacology library. The volume was actually three publications of the New York State College of Forestry bound together: Technical Publication #4 (1916), Technical Publication #6 (1918), and Circular #21 (1918). The first work ("The Relation of Mollusks to Fish," 366 pages) was a meticulous description of the diet and habitats of every element of the Oneida Lake molluscan community and a catalogue of all their "enemies," piscine and otherwise. The second work ("Productivity of Invertebrate Fish Food ... with Special Reference to Mollusks," 264 pages) reported Baker's quantitative survey of Lower South Bay, concluding that his study area contained "4,704 million mollusks, and 3,062 million associated animals." The third work ("The Relation of Shellfish to Fish," 34 pages) was an abstract of the two larger works, intended for wider circulation.

Baker's Oneida Lake research was at the vanguard of the new science of Ecology. He took quantitative samples using an Ekman grab, a device so new that he felt obliged to figure it and describe it in detail. His publications featured gigantic fold-out maps of his study areas and equally gigantic fold-out data tables recording the raw counts of every snail, bug, and glob of algae he collected in all CDXII samples he took from Lower South Bay. There are scores of charming photographs where he spilled out the entire catch from selected samples for the camera ... hundreds of tiny little chironomid larvae pushed to one corner and wads of macrophyte knotted up below. The work was a labor of love, and a lighthouse for future studies of benthic ecology (3).

But for lasting influence, few works in American malacology can rival Baker's monographs on the Lymnaeidae (1911) and Planorbidae (1945). Both of these works featured meticulous scholarship and detailed anatomical drawings executed with great skill. His "Lymnaeidae of North and Middle America, Recent and Fossil" (Chicago Academy of Sciences Sp. Publ. #3) ran to 539 pages plus 60 plates, providing descriptions of shell, radula, and genitalia, as well as ranges and life history notes for the 95 species and subspecies he considered valid, organized into seven genera.

His "Molluscan Family Planorbidae" (University of Illinois Press) was intended to be worldwide in scope, with Part I ("Classification and General Morphology," 212 pages) providing complete descriptions of the 36 genera he recognized and Part II ("Planorbidae inhabiting North & South America," 21 pages) describing 26 new species and "varieties."  Baker's plan to provide detailed accounts of all the planorbid species then recognized in the Americas was cut short by his death. But his editor (van Cleave) was able to assemble 60 plates which would have accompanied the body of Baker's Part II text, together with their explanations, and add them to the 81 plates the author had intended for Part I.

F. C. Baker was innocent of the modern synthesis. It was in 1942, the year Baker died, that Ernst Mayr first formally proposed (and forcefully advocated) the biological species concept (4). Even in his last work, Baker was still attaching Latin nomena to "varieties" of gastropods, as for example, "Helisoma subcrenatum perdisjunctum is similar to disjunctum but is much smaller, about the size of oregonense, but lacks the characteristic shape of the aperture of the last named form."

Baker did not enjoy the understanding of intraspecific variation that informs the research of most evolutionary biologists today. But while he kept one foot firmly planted in 19th century typology, Baker strode forward to the 20th, bearing a profound appreciation for the biology of the animals he was classifying - anatomy, physiology, ecology, and more. I think of him as the "freshwater Pilsbry."  His contributions rank second only to those of Thomas Say in their impact on our understanding of the pulmonate gastropods inhabiting lakes and rivers in America today.

Keep in touch,

Rob

Notes

(1) A photo of the older Baker, together with a brief bio and partial bibliography, is available from Kevin Cummings' site at INHS.

(2) The biography above is based largely on "A Memorial to Frank Collins Baker" by H. J. van Cleave, published as pages xvii - xxxvi in Baker (1945). Baker's complete bibliography is available in that work as well. Van Cleave also published briefer obituaries in Science 95: 568 (1942) and The Nautilus 56: 97-99 (1943).

(3) To learn more about the Oneida Lake molluscan fauna, and its sad fate, see Harman & Forney (1970 - Limnol & Oceanog 15:454), Dillon (1981 - Am. Nat. 118:83) or my book (Dillon 2000) Chapter 9.

(4) For a nice historical review of the biological species concept, see Coyne (1994 - Evolution 48:19).

FWS Finding on the Idaho Springsnail

To the FWGNA group:

Most of you will probably recall my series of messages last year regarding the Idaho springsnail controversy - a tangle of competing proposals to delist the federally endangered "Pyrgulopsis idahoensis," or to keep P. idahoensis on the list and add several other very similar Pyrgulopsis populations in Oregon, Washington, and Wyoming. See April 2005, December 2005, and January 2006.

Now it would appear that a decision is finally at hand. This past Thursday, Sept 28, the US Fish & Wildlife Service published in the Federal Register a "Notice of two 12-month petition findings and a proposed rule to delist the Idaho springsnail." A press release was simultaneously issued from the Snake River Office bearing this headline: "Protection Not Warranted for Four Springsnail Species." See below.

I'd guess this pretty much settles the matter. Ahead is yet one more comment period, and a final decision by Interior Secretary Dirk Kempthorne, who (ironically) used to be Governor of Idaho. But in all honesty, the Conservation Community has already spent way too much time on this, "the largest single population of freshwater snails ever documented." North America is home to scores of other freshwater gastropod species much more deserving of protection than the Idaho springsnail. Let's move on.

Speaking of which, the website of the Snake River FWS Office features two other news releases of interest - one having to do with the (much more endangered!) Pyrgulopsis bruneauensis and the other regarding the (undescribed) "Banbury Springs Lanx." If anybody on this list has additional information regarding either of these two species, contact the FWS.

And keep in touch!
Rob


NEWS RELEASE
Snake River Fish and Wildlife Office
1387 S. Vinnell Way, Room 368 • Boise, Idaho 83709
http://idahoes.fws.gov/
September 28, 2006
Contact: Meggan Laxalt Mackey

PROTECTION NOT WARRANTED FOR FOUR SPRINGSNAIL SPECIES
Public Comments Accepted through November 27, 2006 on Service’s Proposal to Delist Idaho Springsnail

The U.S. Fish and Wildlife Service today announced the Idaho springsnail, currently listed as endangered under the Endangered Species Act (ESA), has been found to be the same species as three other groups of freshwater springsnails, none of which warrants protection under the ESA. The Service therefore today published a proposal in the Federal Register to remove the Idaho springsnail from the Federal list of threatened and endangered species.

This decision is based on the new taxonomic information and other available scientific information that resulted in four groups of springsnails being classified as one species, Pyrgulopsis robusta. The four groups of springsnails are the Idaho springsnail that inhabits Idaho’s Snake River, the Harney Lake springsnail in southeastern Oregon, the Jackson Lake springsnail in western Wyoming, and the Columbia springsnail from the lower Columbia River between Oregon and Washington.

“The 1992 listing of the Idaho springsnail as endangered was based on the best information available at that time on the species and the threats it faced,” said Ren Lohoefener, director of the Fish and Wildlife Service’s Pacific region. “New scientific information became available that prompted the Service to closely examine the status and classification of the species and other closely related springsnails. The examination of all the information currently available led us quite clearly to this decision.”

Comments from all interested parties regarding the proposal to delist the Idaho springsnail will be accepted by the Service until close of business November 27, 2006. Requests for public hearings must be received on or before November 13, 2006. Comments may be submitted by e-mail to: fw1srbocomment@fws.gov, by fax to 208-378-5262, or by mail or hand-delivery to the Service’s Snake River Fish and Wildlife Office at 1387 S. Vinnell Way, Boise, Idaho 83709. Please include the title “ISS RIN1018-AU66” in the subject line.

The Service conducted a comprehensive 12-month review of the four groups of springsnails. The Service’s review was prompted by two separate petitions concerning the four springsnails: one petition by the Idaho Governor’s Office of Species Conservation and the Idaho Power Company seeking the delisting of the Idaho springsnail, and another from a group of academics and environmental organizations requesting the listing of all four springsnail populations.

The Service also included in this process a five-year review of the Idaho springsnail, a process required by the ESA for all listed species to determine whether the species is properly classified. The five-year review of the Idaho springsnail is available from the FWS Pacific Region website. Pyrgulopsis robusta is a small (4-6mm) freshwater snail species. It may be found in various habitats from small springs and spring-fed creeks to reservoirs and large river systems. The snails feed primarily on algae, bacteria, fungi, diatoms (small plants), and protozoa (small animals) on the surface of rocks or gravel in the water.