Dr. Rob Dillon, Coordinator





Showing posts with label Endangered Species. Show all posts
Showing posts with label Endangered Species. Show all posts

Wednesday, December 10, 2025

Art, science, and public policy: A dialogue in three languages

I have a lot of friends and family who enjoy travel, and spend a substantial amount of time and money doing it, and imagine that their hobby returns some sort of profit to the intellect or character.  Isn’t travel broadening?  Don’t we learn from exposure to new lands, new cultures, new value systems, new ways of looking at the world?  Yes, of course.  But I found countless worldviews, cultures, and value systems trumpeted in the morning headlines at the end of my driveway this morning, ripe for engagement in the aisles of the grocery store across the street this afternoon.  No further travel is required.

As my loyal readership will attest, I have long been fascinated by the diversity of worldviews I encounter in my workday life, and the relationships between them.  The index item at the right of your screen labeled “Worldview Collision” will link to eight essays on the subject when I push the “submit” button on the present post.

The majority of my seven previous essays have explored the relationship between the worldview of science and the worldview of law or public policy, among the most common of the culture clashes in my experience. To understand their proper relationship I have adapted an analogy I first developed about twenty years ago during the height of the most recent creation/evolution controversies between science and religious faith.  Scientists play baseball, lawmakers and regulatory agencies play banjo.  Those worldviews are not incompatible, in the sense that my father was both a banjo picker and a catcher.  But not compatible, either.  Nobody ever tries to integrate one with the other. 

I favor the baseball/banjo analogy because most of my colleagues in the world of science seem to have a better grasp on the proper relationship between sports and art than between science and anything else, possibly because we are more objective.  We don’t write grant proposals to shortstops, nor debate fiddlers over the 10th grade biology curriculum.

 

Indeed, I rarely interact with artists in my professional life [1].  Thus, an email I received this past August (18Aug25) from one Ms. Julia Galloway [2] fell upon my eye as dew drops from heaven.  She introduced herself as: 

“… a professor and ceramic artist based in Montana. I’m currently working on a project focused on raising awareness about endangered species. For this project, I am creating ceramic urns to represent each threatened, endangered, recovered, and extinct species in the U.S. The project will culminate in the creation of approximately 1,200 urns, with completion expected by 2027.”

Ms. Galloway went on to explain that she had created “400+” urns as of that mid-August date and had (apparently) worked down from the California Condors to the freshwater gastropods, finding her supple hands now poised over a throw of clay to be entitled, “Anthony’s Riversnail.”  And googling about the internet for inspiration, she had found my “fabulous photo” posted on the FWGNA site.  And requested permission to use.

 

How fascinating!  The concept of “endangerment” is purely a matter of law, of course, and “raising awareness” a political objective.  Here an artist addresses a scientist over a matter of public policy. A conjunction of three worlds.

 

On 21Aug25 I replied to Ms. Galloway certifying, as I always do, that all of the images on the FWGNA site are freely available for anybody to use for any purpose whatsoever, and offering to help her in any other way I could.  And I concluded, doing my best to address her in her native tongue: 

“Notice that there's a photo of Leptoxis crassa (“Anthony’s Riversnail”) in situ at the bottom of (the species) page [here].  Which brings me to my final point.  Bless your heart!  These things are brown bumps on a rock.  God made them by throwing little balls of clay at dirt.  If you can make art out of Leptoxis crassa, my cap's off to you.”

 And to the bottom of my email of 21Aug25 I added, “PS – 1,200 urns? Are you nuts?”


Brown bumps in Limestone Creek, AL.

Here the challenge of communication across cultures was on full display.  At no point in her initial email to me, nor indeed in any of her subsequent correspondence, did Ms. Galloway mention the rich symbolism just below the surface of her artwork.  Her urns are modeled after the funerary urns of ancient Egypt.  But, quoting from her website, “displayed empty as a sign of hope.”  Anthony’s Riversnail may be on its sickbed, by this metaphor, but it ain’t dead yet.

 

Ms. Galloway replied immediately thanking me for my “delightful and thoughtful reply,” confessed that she does feel a little bit nuts at times, and asked me if I would like to see images of the urn upon completion. And I responded immediately in the affirmative, and suggesting that she just “throw a couple balls of clay at a flowerpot” to expedite the process.

 

I was tremendously impressed to receive the set of three jpegs below on the afternoon of 22Aug25, less than 24 hours after I had granted Ms. Galloway permission to use FWGNA imagery.  Maybe I shouldn’t have been so surprised – with 400 urns down and 800 to go, she must be nothing if not efficient.  Three individual snails were depicted various faces of the Anthony’s Riversnail urn, one snail crawling to the left (a) and two crawling to the right (b, c), as from the vantage point of a wary stonefly.  None of these images was modeled, as far as I could tell, from anything on the FWGNA site.  At least overtly.

 

Although starkly beautiful in their execution, I immediately noticed a couple significant technical problems with this urn.  Or at least, they seemed significant to me.  So on 23Aug25 I addressed my third email to Ms. Galloway.

And this time, I felt compelled to speak a little bit of science.  I began with some background on gastropod coiling, so as to introduce the adjectives “sinistral and dextral,” and noted that no case of sinistrality has ever been documented in the Pleuroceridae, as far as I know.  I then wrote:

“You have depicted three individual snails on your urn, am I correct?  The images of which you labeled on the jpegs you sent me yesterday afternoon AO22-a, AO22-b, and AO22-c, yes?  Snail (b) is bearing a dextral shell, correct as you have sculpted.  But snail (a) and snail (c) are bearing sinistral shells.  Not only is that uncomfortable to my eyes, it is unscientific.”

That seemed harsh.  So I added, to soften the blow:

“Perhaps you were working from (somebody else's) closeup photograph of a pleurocerid animal, reversed in some popular publication or on the internet?   Amateurs often publish reversed images, careless of the difference.  In fact, it seems highly unlikely to me that anybody other than a professional malacologist would ever notice the chirality of the snails you have sculpted on your urn.  So if you want to let it go (as Queen Elsa would suggest) I would certainly understand.”

Well, a couple weeks passed.  And I honestly did not think I would ever hear from Ms. Galloway again.  At the rate of one urn per 24 hours, I imagined that she must be well into the unionid mussels by that point, gastropods let go, gone, and forgotten.  But on 2Sept25, I was most pleased to receive yet another lovely communication from my artist-pen-pal:

“Thank you so much for bringing chirality to my attention! I loved reading your email and learning more about gastropods. It's always a treat when I get to hear someone talk passionately about their field of study (especially when they're brown bumps on a rock).

 

Sometimes species have very few images, which was the case with this species. Not knowing about chirality, I flipped some of the images to give myself more viewing angles and compositional options. Now that I know about how snail shells grow, I would like to remake the urn. Would you happen to have any additional images of Anthony's River Snail that you could share?

 

Thank you again for taking the time to share this information with me and for your support! Take care and I hope to hear from you again soon!”

Well no, I myself did not have any additional photos of L. crassa showing anything that the couple images already posted on the FWGNA species page did not.  But on a whim, I executed simple a google-image search on “Athearnia anthonyi” and landed, of course, on the Wikipedia page [3].  And good grief.  The image at the top of the Wikipedia page is sinistral!

Wikipedia. From Dick Biggins, USFWS.

So on the evening of 4Sept25 I addressed my fourth email message to Ms. Galloway, apologizing on behalf of the entire profession of freshwater malacology, re-assuring her that Dick Biggins (the donor of the photo) is a careful worker and confessing that I could not imagine that he himself would upload a mirror-reversed image.  But regardless of how the error occurred, I was sorry that some member of our extended community had not fixed it by now.  Not it.

 

I then suggested three remedies: (1) Sculpt the Wikipedia image from a mirror, (2) Adjust the Wikipedia image using the Photoshop mirror-image-reverse button, or (3) model from some other pleurocerid. Honestly, at the resolution of a funerary urn, all pleurocerid bodies are indistinguishable.  I sent her a good image of some other pleurocerid individual crawling to the viewer’s right, so that she would have both left-travelling dextral and right-travelling dextral models to work from.

 

And on 7Sept25 Ms. Galloway thanked me once again, and asked me if I would like to see images of “the new and improved urn when it is completed?”  To which I replied in the affirmative.  But I never heard from her again.  And that’s OK.

 

The philosopher Ian Barbour (1923 – 2013) has suggested [4] that there are four ways in which worldviews might relate: (1) independence, (2) dialogue, (3) conflict and (4) compatibility.  Independence is the unexamined status quo, compatibility a pipe dream, and conflict is right out.  I love, love, love dialogue.

 

The relationship I have demonstrated above is (2) a dialogue between the worldview of science and the worldview of art.  One might subclassify it as (2a) science helps art.  Ms. Galloway asked me to help her.  I did everything I could to do so.

 

Science and art are not incompatible, here obviously.  But mark me well.  Science and art are not compatible either.  They are very simply, and very profoundly, different.  Ms. Galloway is an artist, and she took the lead in this interaction, and I (a scientist) did what I could to help her create a work of art, and at no point did anything that happened between us during our entire two-week interaction have anything to do with science whatsoever.  The pleurocerid images she carved into that pot could bear sinistral shells, or dextral shells, or polka-dot shells, it does not matter.  And in fact, she never asked me for any of the free advice I offered her at any point, and in retrospect, I may have been interfering with the creative process, and if I ever hear from her again, I will apologize for butting in.

 

But I can’t help it, I love that sort of thing. The proper relationship between our worldviews is one of dialogue.  A dialogue with a fine artist is like an expedition into the bush with a Hottentot, from the seat of my own kitchen table.

 

And here is the most interesting thing about my two-week dialogue with Ms. Galloway.  This particular artist’s motivation was not artistic, but political. 

 

The Oxford Dictionary defines art as “an expression of human skill producing works to be appreciated primarily for their beauty or emotional power.”  But to quote from Ms. Galloway’s website, she considers her work “a catalyst for social change.” 

 

She selected Anthony’s Riversnail as one of her 1,200 subjects because that particular gastropod was entered onto the Federal Endangered Species list on April 15, 1994, see 59 FR 17994-17998 [5]. And “By creating an urn for each (such) species, (she) is making (rarely-seen) species visible, and through this awareness, compassionate action is possible.”  In other words, Ms. Galloway apparently thinks that the worldview of art and the worldview of politics are compatible, such that the former can influence the latter.

 

Bless her heart.  You will have by now noticed that no representative from the world of politics or public policy walked into the bar with the artist and the scientist to this point in my essay, nor will one subsequently appear.  I myself was awarded a AAAS Congressional fellowship many years ago, and learned just enough of the language spoken on Capitol Hill 1981 – 82 to appreciate my limitations.  And I do know quite a few hardworking biologists employed by natural resource agencies, both state and federal. And speaking now for all the legislative bodies and all the regulatory agencies and all the departments of natural resource management involved in all the endangered species conservation nationwide, as well as the entire [6] gastropod fauna of Limestone Creek, Madison County, Alabama, thanks for the pot.  Not a great likeness, but it’s lovely, dear, it really is.


Dave Michaelson & Randy Sarver

Art and Public Policy have different languages, different cultures, different values, and different ways of looking at the world. That they are not incompatible is witnessed by Ms. Galloway herself, who is both an artist and a social activist.  But the worldviews are not compatible, either.  Neither art nor public policy can affect the other, any more than the marching band affects the halftime score, or the halftime score the marching band.

 

But lest we condescend.  Of all the holders of all the worldviews of all the world – Art, Science, Business, Finance, Law, Medicine, Engineering, Religious Faith, Harry Potter, or Star Trek, we scientists are the most arrogant.  The notion of science-based public policy is just as absurd as pottery-based public policy, and none of us seems to realize it.

 

On the morning of 4Sept25, the same day I was to send my fourth email to Ms. Galloway, I met my good friend Randy Sarver in the parking lot of the Missouri Department of Natural Resources, Jefferson City. Randy is an excellent biologist, and we have developed a warm relationship over many years, and he helped me unload four flats of empty black-capped vials that used to hold macrobenthic samples collected by the MoDNR 2015 - 2017, and swap them for a fresh batch of MoDNR macrobenthic samples 2017 - 2018. 

 

I am sure that Randy and Dave Michaelson and all our friends at the MoDNR do a great job monitoring the water quality of the Show-Me State, and I would never dream of offering them any advice whatsoever, because I can’t, any more than they would dream of offering me any advice about malacology, because they can’t.  Randy and I are in dialogue.  That’s the thing I love.

 

Notes:

 

[1] I actually did post one previous essay on the relationship between science and art, way back in 2011:

  • When Science and Art Collide [4Feb11]

 [2] Learn more about Ms. Galloway from her lavish web presence:

  • Julia Galloway [home]
  • Endangered Species Project [direct]
  • Wikipedia [page]

 [3] Wikipedia, accessed 10Dec25 [link]

 

[4] Here I am generalizing Barbour’s thought on the science – religion relationship to the relationship between worldviews of any sort.  His “fourfold typology” was most clearly stated in:

  • Barbour, I. G. (2000) When Science Meets Religion: Enemies, Strangers, or Partners?  Harper, 205 pp.

[5] For the Democrats among my readership, who imagine that findings such as those published in the Federal Register on 15Apr94 have anything to do with science whatsoever, please refresh your memory with the ten essays I have written on the Snake River Physa scandal to date.  Actually, you could skip the first six, if you want, and go straight to:

  • The SRALP and the SRNLP: A new hope [14May24]
  • The SRALP and the SRNLP: Physa acuta were found [11June24]
  • The Twelve Phascinating Physa of Bliss [2July24]
  • Cytoplasmic Male Sterility in the Snake River Physa [7Aug24]

[6] Fourteen species comprise the exuberant gastropod fauna of Limestone Creek: four pleurocerids, three hydrobioids [7], three viviparids, and four pulmonates.  I feel certain that all have benefited from the endangered status of their most-famous member.


[7] Counted among the Limestone Creek hydrobioids is a population of Marstonia olivacea, which is a senior synonym of Marstonia ogmorhaphe, which was the other gastropod entered onto the Federal Endangered Species list 15Apr94.  And hence that hydrobiid population should be every bit as federally-protected as the pleurocerid "Athearnia anthonyi," but it isn't, because public policy has absolutely nothing to do with science.  For more, see:

  • Is Marstonia olivacea extinct? [19Sept23]


Wednesday, August 7, 2024

Cytoplasmic Male Sterility in the Snake River Physa

Editor’s Note – This is the fourth installment of my three part series on the Snake River Physa controversy.   So if you’re interested in all the public policy craziness surrounding the nominally endangered Physa natricina, I would recommend that you back up to my posts of [14May24], [11June24], and [2July24] before proceeding.  If you’re just interested in the science, on the other hand, you might want to refresh your memory with  my [9June22] essay on cytoplasmic male sterility.

Last month we reviewed a mtDNA sequence dataset collected by Mike Young and colleagues [1] from “The Twelve Phascinating Physa of Bliss,” a small sample taken by IPC/FWS biologists from the tailwaters of the Bliss Dam in the Snake River of Idaho in 2019.  That sample included N = 4 snails bearing a CO1 sequence so strange and divergent that our buddy Mike hesitated even to assign it to the genus Physa.  He called the four snails bearing that sequence “Snake River Candidate Species 3,” or SRCS3 for short.

Blasting Mike’s SRCS3 sequences against the entire worldwide NCBI GenBank, we confirmed that OK510774 (taken as typical for the set of four) was more similar to a planorbid from Bangladesh than to any other physid sequence ever reported, “with five phascinating exceptions.”  SRCS3 was approximately 93% similar to two Physa sequenced in Singapore [2], and three Physa sequenced in South Africa [3].

With those little scraps of data in front of us last month, I speculated that SRCS3 might be a strain of Physa acuta demonstrating cytoplasmic male sterility (CMS).  And at that point I abruptly shifted focus to a remarkable paper published in 2022 by my good friend Patrice David and colleagues, reporting very similar levels of mtDNA sequence divergence in a French strain of Physa acuta unable to mate in the male role, but nevertheless fully fertile as female [4].  Could the Snake River – Singapore – South Africa sequence signal the existence of a second case of CMS, a mitotype strikingly different both from the normal, simultaneously hermaphroditic (N) strain and the divergent, male-sterile (D) discovered by Patrice and his team?

Yes.  Two days after I posted last month’s essay I sent an email to Patrice, subject line “Shout-out on the FWGNA blog,” the entire body of which simply read, “I thought you might be interested to see how your 2022 paper on cytoplasmic male sterility could have an impact on conservation biology here in the USA.”  And Patrice replied with a pdf reprint that knocked my socks off [5].

The paper was published in Evolution by Patrice’s student Fanny Laugier, working with a team of nine coauthors from Montpellier and Villeurbanne [6].  Although available in preprint since March, it just appeared in the physical journal last month.  But let’s back up two steps before we leap forward three.

Cytoplasmic male sterility is well studied in plants.  And given the accelerated rates of mutation demonstrated by mitochondria hosting CMS genes, I do not suppose it is surprising that natural populations of gynodioecious plants often host multiple, independently evolving CMS lines.

And as I mentioned in my essay of [9June22] natural populations of gynodioecious plants hosting CMS mitochondria have also often evolved nuclear genes that restore male fertility.  This results from the selective advantage that a fully functional hermaphrodite has over other members of its population reproducing strictly as a single sex [7].

So, reasoning from numerous well-documented cases in plants, Fanny and her team returned to the Physa populations around Lyon originally studied by Patrice and his colleagues to look for additional CMS strains.  And sure enough, prospecting around with a clever PCR test, they found seven individual snails in two populations bearing a “mitotype K,” startlingly different from both normal hermaphroditic N and the CMS mitotype D they had reported in 2022.

My readership might also remember from [9June22] that CMS mitotype D was, around the entire mitochondrial genome, approximately 44% different from the normal Physa acuta mitotype N.  The 20% CO1 sequence divergence between those two mitotypes was actually less than average.  So the newly discovered mitotype K ultimately proved to be 35% - 57% divergent from N and 35% - 57% divergent from D (ten genes), with CO1 sequence divergences 23% and 28%, respectively.  This is the phenomenon I have termed “mitochondrial superheterogeneity” in many previous posts on this blog [8].

And can you smell what our French chefs are cooking?  Focusing now on the CO1 sequence, the French CMS strain bearing the newly discovered mitotype K matched Ting Hui Ng’s [2] Singapore sequence and Molaba’s [3] South Africa sequence 100%, almost identically.  And the match with Mike Young’s [1] SRCS3 sequence from the Bliss Rapids was 93%, just as I reported last month.  Fanny Languier, Patrice David, and their colleagues had discovered a strain of Physa acuta bearing a wildly divergent mitochondrion worldwide in its distribution.

Laugier [6] Figs. 1B and 1C, modified.

But that is not the headline news.  Here is the headline.  Mitotype K Physa acuta were not male-sterile!  Our French colleagues cultured up a big batch of Mitotype K Physa and were easily able to cross them with their albino N laboratory line [9], both lines copulating readily in both the male and the female role.

So again, reasoning from many years of accumulated research on cytoplasmic male sterility in plants, Fanny and her team suspected that nuclear genes might have evolved in the mitotype K line to restore male function.  And they launched a 17 – generation introgression experiment to insert mitochondrial lineages derived from their wild-caught (pigmented) K line into the “naïve” nuclear background of their (albino) laboratory line N.

And sure enough!  After just 5 generations of introgression, 69% of the naïve snails bred to bear K-type mitochondria had lost their ability to mate as males, with no such loss whatsoever in control lines bearing the N-type mitochondria.  After 11 generations, the proportion of naïve male-sterile K snails stabilized at around 60%.  No loss in female function was ever detectable in any line.  In my 5July24 email back to Patrice, I wrote:

“Wow, nuclear restorers!  It's a shame we work with crap-brown little trash snails.  If snails were maize, wheat, or rice [10], we'd be rich.”

What wonderful science!  Deductive reasoning, tested by rigorous experimentation carefully designed to proceed from the known to the unknown.  Conducted for the joy of it, for the exploration of evolutionary mechanisms, for the pushing back of the darkness.  The construction of testable hypotheses about the natural world, period, full stop, nothing more and nothing less.  Pure and unsullied science, utterly useless and wonderful!

Laugier [6] Fig. S1, modified.

And yet, it turns out, completely by accident, useful.  Ting Hui’s study in Singapore was directed toward invasive species, the Molaba study in South Africa was parasitological, and Mike Smith’s study on the Snake River directed toward conservation.  Fanny’s results contribute toward an understanding of all those results, and more.

And here is Lesson Number One.  Gene sequences are not species.  A species is a population or group of populations reproductively isolated from all others [11].  Sequence divergence is typically correlated with reproductive isolation [12], of course, no different from morphological divergence.  But gene trees are not species trees, and genes are not species.

So, as I pointed out in last month’s essay [2July24], the Snake River line of Physa acuta that Mike Young and colleagues called SRCS3 is not the same as the French mitotype K.  The two lines are 7% different, as though they arose from a single mutation in some mitochondrial DNA repair gene sometime in the past and have subsequently diverged.  Could such a mutation have occurred more than once in the evolutionary history of the Physidae, here in North America, where Physa seem to have first evolved?  You betcha.

There is no reason that the CO1 gene sequence that Gates, Kerans and colleagues [13] recovered from the stunted Physa below the Minidoka Dam, roughly 15% different from the N line at Bliss (OK510580), couldn’t be evidence of yet another CMS strain in Physa acuta.  Gates and Kerans identified that sequence as “Physa natricina.”  But gene sequences are not species.  Species are defined by reproductive isolation.  And we have no data on reproductive relationships between the Minidoka population and the dirt-common Physa acuta downstream whatsoever.  And plenty of evidence otherwise [14].

Nor is there any reason in the world that the peculiar mitotype shared by Mike Young’s SRF14, the Owyhee Wet Rock Physa of eastern Oregon [15] and scattered in odd lot Physa populations from California to British Columbia couldn’t be evidence of a CMS strain in Physa gyrina.  MtDNA sequence data, and the gene trees we make with them, are (at best) weak null models of population relationships, correlated with speciation, nothing more [16].

Do not misunderstand me.  I am not demanding controlled breeding studies between every pair of the 2.2 x 10^6 species described from Planet Earth.  But for nominally endangered species, such as Physa natricina, before we enact pages of Federal regulations and spend millions of dollars on conservation, we could at least run a couple of $12 allozyme gels and test for evidence of assortative mating [17]  with a nuclear polymorphism or two, am I right?

Notes

[1] Young, M.K., R. Smith K.L. Pilgrim, and M.K. Schwartz (2021)  Molecular species delimitation refines the taxonomy of native and nonnative physinine snails in North America.  Scientific Reports 11: 21739. https://doi.org/10.1038/s41598-021-01197-3

[2] Ng, T.H., Tan, S.K. & Yeo, D.C. (2015) Clarifying the identity of the long-established, globally-invasive Physa acuta Draparnaud, 1805 (Gastropoda: Physidae) in Singapore. BioInvasions Rec. 4, 189–194.

[3] Molaba, G.G. et al. (2023) Molecular detection of Fasciola, Schistosoma and Paramphistomum species from freshwater snails occurring in Gauteng and Free State provinces, South Africa.  Veterinary Parasitology 320: 109978.  https://doi.org/10.1016/j.vetpar.2023.109978

[4] David, Patrice, Cyril Degletagne, Nathanaëlle Saclier, Aurel Jennan, Philippe Jarne, Sandrine Plénet, Lara Konecny, Clémentine François, Laurent Guéguen, Noéline Garcia, Tristan Lefébure, Emilien Luquet (2022) Extreme mitochondrial DNA divergence underlies genetic conflict over sex determination.  Current Biology 32: 2325 - 2333.  https://doi.org/10.1016/j.cub.2022.04.014.  For a review, see:

  • Cytoplasmic Male Sterility in Physa! [9June22]

[5] Patrice went on to apologize “I told my PhD student Fanny a dozen times to send this to you, that you would probably make good use of it because it was solving a controversy, apparently she didn’t, so I do it myself.” Yes, please send me your reprints!  I lost my library privileges when I was banned from campus in 2016.  I really haven’t had access to most of the scientific literature published since.

[6] Laugier, Fanny, Nathanaëlle Saclier, Kévin Béthune, Axelle Braun, Lara Konecny, Tristan Lefébure, Emilien Luquet, Sandrine Plénet, Jonathan Romiguier, and Patrice David (2024) Both nuclear and cytoplasmic polymorphisms are involved in genetic conflicts over male fertility in the gynodioecious snail, Physa acuta.  Evolution 78 (7): 1227–1236. https://doi.org/10.1093/evolut/qpae053

[7] Yes, mitochondria bearing CMS genes are “selfish” in the sense of the Richard Dawkins (1976) classic.   Don’t get me started on Richard Dawkins.

[8] Mitochondrial superheterogeneity (mtSH), where two or more of the members of a single population demonstrate greater than 10% divergence in any single-copy mtDNA gene, not sex linked, seems to be remarkably common in freshwater gastropods.  In pulmonate populations, I wouldn’t be surprised if most or all mtSH is ultimately traceable to CMS.  In prosobranch populations, however, I think mtSH is a signature of great age, plus low-frequency long distance dispersal, the “Jetlagged Wildebison Model.”  Here is a sample of my previous posts on mtSH:

  • The Snails the Dinosaurs Saw [16Mar09]
  • Mitochondrial superheterogeneity: What we know [15Mar16]
  • Mitochondrial superheterogeneity: What it means [6Apr16]
  • Mitochondrial superheterogeneity and speciation [3May16]
  • Mitochondrial heterogeneity in Marstonia lustrica [3Aug20]

[9] Dillon, R.T. and A.R. Wethington (1992) The inheritance of albinism in a freshwater snail, Physa heterostropha. Journal of Heredity 83:208-210. [pdf]  For nice review, see:

  • Albinism and sex allocation in Physa [5Nov18]

[10] The clever manipulation of cytoplasmic male sterility, together with nuclear restorers, has been one of the more important methods by which plant breeders have achieved the outcrossing of normally self-pollinating crop plants.  I am quite sure that a lot of money has been made with that genetic technology.  With Physa acuta, however, the prospects are not as lucrative.

[11] This is the biological species concept, most closely associated with the work of Ernst Mayr.  It should need no restatement, much less a defense.  But the best paper Jerry Coyne ever wrote was a defense of both Mayr and his species concept, here:

  • Coyne, J. A. (1994) Ernst Mayr and the origin of species.  Evolution 48: 19 – 30.

[12] Dillon, R. T., A. R. Wethington, and C. Lydeard (2011)  The evolution of reproductive isolation in a simultaneous hermaphrodite, the freshwater snail Physa.  BMC Evolutionary Biology 11:144. [html] [pdf].  For a review, see:

[13] Gates, K. K., B. L. Kerans, J. L. Keebaugh, S. K. Kalinowski & N. Vu (2013) Taxonomic identity of the endangered Snake River physa, Physa natricina (Pulmonata: Physidae) combining traditional and molecular techniques.  Conserv. Genet. 14: 159-169.  For a review, see:

  • The Mystery of the SRALP: No Physa acuta were found [2May13]

[14] Rogers, D.C. & A.R. Wethington (2007) Physa natricina Taylor 1988, junior synonym of Physa acuta Draparnaud, 1805 (Pulmonata: Physidae).  Zootaxa 1662: 45 - 51.  For a review, see:

  • Red flags, water resources, and Physa natricina [12Mar08]

[15] Moore, A.C., J.B. Burch, and T.F. Duda Jr. (2015) Recognition of a highly restricted freshwater snail lineage (Physidae: Physella) in southeastern Oregon: convergent evolution, historical context, and conservation considerations.  Conservation Genetics 16: 113 – 123.

[16] I have made this argument as many times as the number of essays listed under the label “Gene trees” above.  For an overview, see:

[17] For an explanation of gametic phase disequilibrium, its power to distinguish species, and its extension to character phase disequilibrium, see:

  • What is character phase disequilibrium? [4Jan22]
  • Character phase disequilibrium in the Gyraulus of Europe [4Feb22]
  • Just 125 Species of Pyrgulopsis in the American West [7Sept22]

Tuesday, July 2, 2024

The Twelve Phascinating Physa of Bliss

Editor’s Note – This is the third episode in a three part series on the Snake River Physa controversy, prompted by the recent paper of M. K. Young and colleagues [1].  If you’re new to this blog, and seriously interested in the science, I would recommend that you back up two months and read my essays of [14May24] and [11June24] before continuing onward.  I’ve also written six older posts on the subject 2008 – 2013, but no point in going back that far, screw it.

In August of 1979 the Idaho Power Company (IPC) received a permit to study the construction of a proposed A. J. Wiley Hydroelectric Project on the Snake River at RM 565, about 1.5 miles west of the little town of Bliss [2].  The proposed dam would have impounded a narrow reservoir as deep as 85 feet nearly eight miles upstream, to the foot of the Lower Salmon Falls Dam.  Inundated would have been the childhood home of a lichen biologist named Dr. Peter Bowler.

By 1992, Bowler had developed a second career as a malacologist, thick as thieves with Dwight Taylor, Terry Frest, and Ed Johannes.  A new species of Physa had been described, Physa natricina, which although found as a fossil throughout northern Utah and southern Idaho, was known alive only from RM525 to RM571 of the Snake River [3].  The “Snake River Physa” had been fast-tracked onto the Federal Endangered species list [4], and the A.J. Wiley Project had been permanently shelved.

The Snake River at RM 570

So last month we reviewed the exciting story of heroic efforts by IPC biologists working in the tailwaters of the Swan Falls Dam, RM 420 – 449, ultimately yielding hundreds of small physids matching the Physa natricina phenotype in at least some respects [11June24].  Because the IPC sample was taken 76 miles downstream from the type range, however, we referred to this population as “Snake River natricina-like Physa,” or SRNLP for short.  Ultimately, the SRNLP from RM 420 – 449 proved a genetic match with Physa acuta, a trash snail invasive on five continents [1].

This is not the situation at RM 675, however.  Previously we have reviewed very similar efforts by biologists engaged by the U.S. Bureau of Reclamation in the tailwaters of the Minidoka Dam 100 miles upstream of the type range of Physa natricina [5].  There, the SRNLP bear a unique mtDNA haplotype, not matching Physa acuta.

Between the Swan Falls Dam and the Minidoka Dam, in the actual type range of Physa natricina, previous research results have been equivocal.  Rogers and Wethington examined hundreds of physids sampled from this stretch of river by IPC biologists 1995 -2007, ultimately unable to distinguish the entire lot from Physa acuta [6].  John Keebaugh disagreed, selecting a small subsample as bona fide Physa natricina.  But the genetic results from the Keebaugh subsample were dubiously reported and never vouchered [7].

Are there no reliable genetic data for Snake River physids sampled between RM 525 and RM 571?  Well, yes, there are.  Twelve very interesting and extraordinarily diverse mtDNA sequences from the tailwaters of the Bliss Dam at RM 559, to be precise.

The Snake River in Idaho, modified from Gates et al [7]

It materializes that the Idaho Power Company has interests that extend beyond the conservation biology of freshwater gastropods.  Sport fishing is an important part of the local economy of the small communities along the Snake River, for example, and mercury levels in the game fish a matter of some concern to the IPC [8].  IPC biologists also have an ongoing project to survey mercury levels in the macrobenthos, as a component of the diet of the white sturgeon.

And so it came to pass that on 17Sept19 a team of IPC and USFWS biologists gathered to sample macrobenthos from the tailwaters of the Bliss dam at RM 559 with a suction dredge.  Their sample came from wadable depths – not the middle of the river – and Physa were not the target [9].  But the 12 individual Physa recovered as bycatch were sent to Mike Young and colleagues at the National Genomics Center for Wildlife and Fish Conservation in Missoula for analysis, along with that big sample of P. acuta from the Swan Falls tailwaters we obsessed over last month [1].  And the results were fascinating:

P. gyrina, Snake R. [10]

N = 1 Physa gyrina
.  This is the most widespread of the Snake River physids, inhabiting the main stem throughout the state of Idaho, upstream beyond the Minidoka Dam and into many of the tributaries.  Populations of P. gyrina are not as invasive as P. acuta, however, demonstrating slower growth rates, larger sizes at first reproduction, and lower fecundities per unit body mass.  The CO1 sequence similarity between the singleton P. gyrina recovered by IPC biologists at Snake River mile 559 (SRP199 = OK510769) and the type P. gyrina sequence from Council Bluffs, Iowa (AY651187) is 99.0%.  Good.  No surprises there.

N = 5 Physa acuta.  Arbitrarily selecting the first-listed of the five sequences in this most common category of physids recovered from the Bliss tailwaters, the divergence of SRP001 (= OK510580) from the P. acuta type sequence (River Garonne, AY282589) is 4.7%, and the divergence with our typical P. acuta sequence from the Swan Falls tailwaters (SRP051 = OK510624) is 3.4%.  Apparently, a population of very ordinary, typical [11] Physa acuta was found, again.  This is additional confirmation, if any was needed, that the range of a trash snail invasive on five continents does in fact extend 76 miles up the Snake River from Swan Falls to the type locality of Physa natricina.  Fine.  No surprises there, either.

N = 4 Snake River Candidate Species 3 (“SRCS3”).  Four individual physids sampled from the Snake River at RM 559 bore a haplotype so divergent from all other sequences in the worldwide database that Mike Young, bless his heart, considered the evidence “consistent with their assignment to a separate genus.”  Ultimately, however, Mike decided simply to nominate this subset of four as #3 in his slate of 18 new “candidate species” for the physinine malacofauna of North America.

P. acuta, Snake R. [13]
Picking the first SRCS3 sequence from my buddy Mike’s Supplementary Table 2 as typical (SRP206 = OK510774), we can immediately confirm nearly negligible CO1 sequence similarities of 76.1% with Bliss acuta (OK510580) and 75.8% with Bliss gyrina (OK510769).  Indeed, blasting OK510774 against the entire NCBI GenBank database, we discover that Mike’s SRCS3 is more similar to Indoplanorbis exustus from Bangladesh than to (essentially) any other physid sampled from anywhere else in the world, with five phascinating exceptions.

Mike Young’s SRCS3 sequences are 93.2% similar to a pair of CO1 sequences uploaded by Ting Hui Ng and colleagues from their famous survey of freshwater gastropods in the ornamental pet trade of Singapore (KP182981 and KP182982).  I reviewed Ting Hui’s paper shortly after its 2016 publication [14], adding a footnote to her earlier paper focusing on Singapore Physa in particular [15], and a footnote connecting the Singapore Physa to the situation in the Snake River of Idaho, which now looks genuinely prophetic [16].

Mike Young’s SRCS3 sequences are also 92.1 – 93.9% similar to a set of three sequences very recently uploaded to GenBank by Gantshe Molaba and colleagues [17] from a 2023 parasitological study in South Africa (ON953193, ON953197, ON953200).  The South African sequences and the Singapore sequences are virtually identical.

What is going on here?  In Singapore and in South Africa and now in the Snake River, physids bearing an extravagantly divergent haplotype have been recovered together with Physa acuta bearing a typical haplotype, and in none of these cases did the authors of the papers report any corresponding morphological distinction between divergent and typical snails.

This situation looks precisely like the situation in France, and indeed right here in Charleston, South Carolina, elegantly demonstrated by our good friend Patrice David and his colleagues [18] to be a signature of cytoplasmic male sterility.  Patrice has hypothesized that mutation in some component of the mitochondrial DNA repair mechanism has simultaneously increased mitochondrial mutation rates in such lines and robbed their bearers of fertility in the male function.  But because they retain female function and mate freely with typical P. acuta (serving as male), they have not speciated.  They are still Physa acuta.

From David et al. [18]
Male-sterile variants of P. acuta are also at least as invasive as typical P. acuta, possibly even more so.  It is impossible to resist pointing out that the middle Snake River Valley is home to scores of fish hatcheries and fish farms, jostling each other for every cubic foot of the spring water that gushes forth from its scenic canyon walls, yielding truckloads of rainbow trout, cutthroat trout, salmon and steelhead.  It was almost certainly these hatcheries, and the fisheries they support, that introduced Potamopyrgus into the American West back in 1987.  Surely, they could bring exotic Physa into this country as well, yes?

Sure, maybe.  But we should also immediately note that Physa acuta is a North American native, and it is just as likely that we exported a male sterile variant to Singapore as Singapore exported one to us.  And the Snake River strain is not the Singapore strain.  They are 6.8% different, as though they sprang from a single mitochondrial mutation, and have subsequently diverged.

And the hypothesis that SRCS3 really is a valid biological species cannot be ruled out.  I’ve been footnoting the possibility of a second acuta-like Physa in the Pacific Northwest for years now – even suggesting Haldeman’s (1843) concolor as a possible name for the critter [19].  Wouldn’t it be great to do some old-fashioned breeding studies here?  Informed by old-fashioned nuclear markers – allozymes, microsatellites, anything?  I have a dream.

N = 2 Snake River Form 14 (“SRF14”).  Mike Young, bless his heart, declined to nominate the two individual snails (SRP200 and SRP201) recovered from the Bliss tailwaters bearing CO1 sequence OK510770 as a “candidate species” unto themselves, instead referring to them as “Form 14” of the 34 “forms” of physinine snails he ultimately distinguished in his stupendous gene tree.  His OK510770 sequence is 92.6% similar to Bliss gyrina (OK510769) and just 83.6% similar to Bliss acuta (OK510580).

Blasting the CO1 sequence Mike Young obtained from his two SRF14 individuals against the entire NCBI GenBank database, we discover that OK510770 is 95 - 96% similar to a set of 13 sequences obtained by University of Michigan (UMMZ) researcher A. C. Moore, with Jack Burch and Tom Duda, from a “highly restricted freshwater snail lineage” they introduced to the world in 2014 as the “Owyhee Wet-Rock Physa,” or OWRP for short [20].

The Owyhee River enters the Snake River in eastern Oregon at RM 395, downstream 165 miles from the Bliss Dam.  (See map below.)  It would be quite the adventure, but if one could float those 165 miles down the Snake to the Oregon border then pioneer back up the Owyhee approximately 130 miles through deep, arid canyons one would arrive at a geothermal spring complex inhabited by a population of Physa that apparently reminds everybody, both in their morphology and in their life habit, of the federally endangered Physa zionis of Utah.

Fig 1 of Moore et al [20] modified
Both the Owhyee Wet-rock Physa and Physa zionis are tiny, stunted little things, with grotesquely expanded body whorls, even more stunted and even more grotesque than Physa natricina.  What is the chance that Mike Young’s SRF14 might be Dwight Taylor’s long-sought and much-endangered species?

The evidence does not point in that direction.  As most of my (highly specialized) readership will be aware, pulmonate snails of the family Physidae bear several distinct types of penial morphology.  The most common type was called “Type C” by George Te in 1975 [21] – bearing a one part, muscular penial sheath.  Type C physids include Physa acuta, Physa natricina [22], and Physa zionis.  Also quite common in North America are physids bearing “Type B” penial morphology – a two part penial sheath – such as demonstrated by Physa gyrina.  If that distinction is unfamiliar to you, and you have more than a casual interest in the Snake River Physa controversy, I would assign the 2007 paper of Wethington & Lydeard [23] for homework.

Wethington & Lydeard showed that penial morphology correlates with molecular phylogeny.  So, although neither the IPC/USFWS team that fished them from the Bliss rapids nor Mike Young, bless his heart, dissected any of the 12 fascinating physids under discussion here, nor (indeed!) did the UMMZ group dissect any of their Owyhee Wet Rock Physa, it turns out that both the SRF14 sequence and the 13 OWRP sequences all clustered together with Type B Physa gyrina, not with Type C Physa acuta/natricina/zionis.

So, the next four paragraphs will be a digression, but indulge me.  The UMMZ group sequenced 13 Owhyee Wet-rock Physa for three genes, ultimately obtaining five slightly different CO1 sequences (KF305393 – 405).  While a match to P. zionis was not confirmed, those sequences did demonstrate a 95% match to a set of 11 CO1 sequences (KF305406 – 416) obtained from a population identified as Physa gyrina inhabiting Aqua Fria Creek, about 20 miles west of Yosemite National Park in central California.

And in 2018, four years after the publication of the OWRP paper, our hardworking friends at the Centre for Biodiversity Genomics (CBG) in Guelph uploaded to GenBank a “Physidae sp.” sequence collected from a beaver pond by the Similkameen River in southern British Columbia (MG421809) that also demonstrated a 95% match to the OWRP.  The Similkameen River flows south into the Okanogan River, joining the Columbia River in central Washington.
Now for the punch line.  It materializes that Mike Young’s SRF14 sequence, collected by IPC/USFWS biologists from the Bliss Rapids of the Snake River, is a stunning 99.7% similar to that “Physidae” sequence collected by the CBG from the beaver pond by the Similkameen 500 miles to the northwest.

Do the four populations mapped above, inhabiting wet rocks on the eastern edge of Oregon, a creek in central California, a beaver pond in southern British Columbia, and now discovered in the Bliss rapids of the Snake River of Idaho, represent a single previously unrecognized species of Type B Physa, extraordinarily variable, widespread across western North America?  Or might they have evolved from a male-sterile variant of Physa gyrina, as SRCS3 seems to have evolved from Physa acuta?

Again, as was the case with SRCS3 and P. acuta, Mike Young’s SRF14 population is sympatric with a population of Physa gyrina of the typical phenotype, bearing mitochondria of the typical haplotype.  Dare I dream that some worthy and industrious graduate student might someday survey these populations for variation at some sort of nuclear marker or markers and look for reproductive isolation, or lack thereof?  Followed by some breeding studies, perhaps?

There was a time in my career when I could have answered every question posed in this entire essay with a day in the field, four days in the lab, and $100 for reagents and expendable supplies.  Get a nice, big sample of Physa from the Bliss Dam tailwaters, paying attention to microhabitat.  Examine their anatomy and shell morphology critically, sort them into Type B and Type C subsets.  Within those subsets, are nuclear markers segregating in Hardy-Weinberg equilibrium?  How many biological species of Physa are we dealing with here?  Give me another six months and a student or two, and the breeding experiments done would have been done.  How many million dollars and years of wandering in a wilderness of ignorance and superstition might timely reference to a scientist, who actually knows something about the biology of the living, breathing animals he is studying, have saved?

N = 0 Minidoka SRNLP.  Although otherwise bearing a stunningly diverse array of mitochondria, our little sample of 12 physids from the tailwaters of the Bliss Dam did not include any bearing a CO1 sequence matching that of the peculiar little misshapen Physa recovered from the Minidoka Dam tailwaters 116 miles upstream, ascribed by Gates, Kerans and Keebaugh to Physa natricina [7].  True, the IPC/USFWS biologists did not sample midstream, where Physa bearing such haplotypes were recovered at Minidoka.  This is a weak test, indeed.

But these are all the relevant data we have, for 18 years of effort by multiple teams of hardworking, well-meaning biologists and millions of dollars flushed down the toilet.  So, since Physa acuta do indeed appear to be common in the shallows of the Snake River in the type range of Physa natricina, and since populations identified as Physa natricina in the Swan Falls tailwaters, both in the shallows and at midstream, are synonymous with P. acuta, the strongest hypothesis at present remains that the population Dwight Taylor described as Physa natricina in its RM 525 – 571 type range is a junior synonym of Physa acuta, as suggested by Rogers and Wethington in 2007.  And the SRNLP population in the roiling tailwaters of the Minidoka Dam is Something-Else-God-Knows-What.

To summarize.  On [12Mar08] I posted the following overly optimistic observation:
“Science is a self-correcting process. It is gratifying to see two of our own, Rogers and Wethington, designing the research program and publishing the paper that has turned us back from our 20-year blunder. But at such a cost! Literally millions of dollars have been wasted monitoring, managing, and protecting a snail that anyone on six continents could find in the ditch behind his local McDonalds, licking special sauce off the hamburger wrappers.”
Now 16 more years have passed, the Rogers and Wethington paper ignored, and millions of additional dollars wasted.  In 2008 I was laboring under the assumption that the world of science and the world of public policy could find some common frame in which to communicate.  Although I went on to write, “Science and politics do not mix,” in retrospect, I should have written that science and public policy cannot mix.

I am playing bluegrass music.  The natural resource agencies are playing baseball [24].  And the Idaho Power Company, just trying to run a business heaven help them, is standing in the batter’s box with a fiddle.

Notes

[1] Young, M.K., R. Smith K.L. Pilgrim, and M.K. Schwartz (2021)  Molecular species delimitation refines the taxonomy of native and nonnative physinine snails in North America.  Scientific Reports 11: 21739. https://doi.org/10.1038/s41598-021-01197-3

[2] Malde, H. E. (1981)  Geologic factors pertinent to the proposed A. J. Wiley Hydroelectric Project, No. 2845, Bliss, Idaho.  U.S. Department of the Interior, Open File Report 81-569.  75 pp.

[3] Taylor, D. W. (1988) New species of Physa (Gastropoda: Hygrophila) from the western United States. Malacological Review 21: 43-79.

[4] US Fish & Wildlife Service (1992). Endangered and threatened wildlife and plants; Determination of endangered or threatened status for five aquatic snails in south central Idaho. 50 CFR Part 17. Federal Register 57(240)59244-57. (December 14, 1992)

[5] Gates, K. K., and B. L. Kerans (2010) Snake River Physa, Physa (Haitia) natricina, Survey and Study.  Report to the US Bureau of Reclamation under agreement 1425-06FC1S202.  87 pp.  For a review of this report, see:

  • The Mystery of the SRALP: A bidding… [5Feb13]
  • The Mystery of the SRALP: A twofold quest! [1Mar13]
  • The Mystery of the SRALP: Dixie Cup showdown! [2Apr13]

[6] Rogers, D. C. & A. R. Wethington (2007) Physa natricina Taylor 1988, junior synonym of Physa acuta Draparnaud, 1805 (Pulmonata: Physidae). Zootaxa 1662: 45-51.  For a review, see:

  • Red Flags, Water Resources, and Physa natricina [12Mar08]

[7] Gates, K. K., B. L. Kerans, J. L. Keebaugh, S. K. Kalinowski & N. Vu (2013) Taxonomic identity of the endangered Snake River physa, Physa natricina (Pulmonata: Physidae) combining traditional and molecular techniques.  Conserv. Genet. 14: 159-169. For reviews, see:

  • The Mystery of the SRALP: No Physa acuta were found [2May13]
  • The SRALP and the SRNLP: A New Hope [14May24]

[8] Willaker, J.J., C.A. Eagles-Smith, J.A. Chandler, J. Naymik, R. Myers and D.P. Krabbenhoft (2023) Reservoir stratification modulates the influence of impoundments on fish mercury concentrations along an arid land river system.  Environmental Science & Technology. DOI: 10.1021/acs.est.3c04646

[9] This is a personal communication from my friend Jim Chandler at IPC.  No reports or publications are as yet available for the sturgeon/food web/methylmercury research project.

[10] The specimen of P. gyrina figured above was collected by RTD from wetlands below the Minidoka Dam on 19Sept2010.

[11] Actually, I think the Snake River population of P. acuta may represent the “Clade B” form of Ebbs, Loker, and Brant [12].  The sequence similarity of OK510580 to Clade B sequence MF694449 from Montana was 98.2%.

[12] Ebbs, E.T., Loker, E.S. & Brant, S.V. (2018) Phylogeography and genetics of the globally invasive snail Physa acuta Draparnaud 1805, and its potential to serve as an intermediate host to larval digenetic trematodes. BMC Evol Biol 18, 103. https://doi.org/10.1186/s12862-018-1208-z

[13] The specimen of P. acuta figured above was collected by RTD from the Snake River at Bliss, 19Sept2010.

[14] Ng, Ting Hui, Tan SK, Wong WH, Meier R, Chan S-Y, Tan HH, Yeo DCJ (2016) Molluscs for Sale: Assessment of Freshwater Gastropods and Bivalves in the Ornamental Pet Trade. PLoS ONE 11(8): e0161130. https://doi.org/10.1371/journal.pone.0161130.  For a review, see:

[15] Ng, T.H., Tan, S.K. & Yeo, D.C. (2015) Clarifying the identity of the long-established, globally-invasive Physa acuta Draparnaud, 1805 (Gastropoda: Physidae) in Singapore. BioInvasions Rec. 4, 189–194.

[16] If you write five crazy things per month for 25 years, eventually at least one will turn out to be a prophecy.  See footnote [6] of my [9Oct17] post for an example.

[17] Molaba, G.G. et al. (2023) Molecular detection of Fasciola, Schistosoma and Paramphistomum species from freshwater snails occurring in Gauteng and Free State provinces, South Africa.  Veterinary Parasitology 320: 109978.  https://doi.org/10.1016/j.vetpar.2023.109978

[18] David, Patrice, Cyril Degletagne, Nathanaëlle Saclier, Aurel Jennan, Philippe Jarne, Sandrine Plénet, Lara Konecny, Clémentine François, Laurent Guéguen, Noéline Garcia, Tristan Lefébure, Emilien Luquet (2022) Extreme mitochondrial DNA divergence underlies genetic conflict over sex determination.  Current Biology 32: 2325-2333.  https://doi.org/10.1016/j.cub.2022.04.014.  For a review, see:

  • Cytoplasmic Male Sterility in Physa! [9June22]

[19] The hypothesis “that some physid bearing a type-C penial morphology, but not correctly identified as either P. natricina or as P. acuta, might inhabit rivers of the Pacific Northwest” was Hypothesis #2 (of 3) in my post of [14Sept10].

[20]  Moore, A.C., J.B. Burch, and T.F. Duda Jr. (2015) Recognition of a highly restricted freshwater snail lineage (Physidae: Physella) in southeastern Oregon: convergent evolution, historical context, and conservation considerations.  Conservation Genetics 16: 113 – 123.

[21] Te, G. A. (1975) Michigan Physidae, with systematic notes on Physella and Physodon (Basommatophora: Pulmonata).  Malacological Review 8: 7-30.  For a review, see:

  • To Identify a Physa, 1975 [6May14]

[22] Figure 6 in Dwight Taylor’s original description [3] clearly shows a type-C penial morphology.  And Jack Burch’s Fig 2.3 confirms that the SRNLP population in the Minidoka tailwaters are also type-C Physa [5].

[23] Wethington, A.R., & C. Lydeard (2007) A molecular phylogeny of Physidae (Gastropoda: Basommatophora) based on mitochondrial DNA sequences.  Journal of Molluscan Studies 73: 241 - 257. For a review, see:

  • The Classification of the Physidae [12Oct07]

[24] The relationship between the worldview of science and the worldview of public policy is analogous to the relationship between music and sports.  They have different languages, cultures, and values.  They are not “compatible” in any sense, neither are they “incompatible” in any sense, because they are absolutely, utterly different.  Nothing I have written in this essay, nor indeed  any scientific findings of any sort, can ever have any effect on public policy.  I play the banjo, and the USFWS plays baseball.

Most of advocates for science in public policy are harmless naïfs, enamored of big government, Democrats the lot of them.  I worked with a building full of that sort 1982-83, as a AAAS Congressional Fellow on Capitol Hill.  They write reports which nobody reads.  No harm in that.

But should any person step forward, claiming to be a scientist, promising to conduct research to inform some issue of public importance, in exchange for contract money or grant, beware!  This is a charlatan, and pseudoscience is his snake oil.